I.

McPartla
and I:
-3
 Hemp Diseases and Pests
Management and Biological Control
 Hemp Diseases and Pests
Management and Biological Control
a

An Advanced Treatise

J.M. McPartland
Assistant Professor, University of Vermont

Robert Connell Clarke

Projects Manager, International Hemp Organization
Director, Pharmtex Consultants,Inc., San Diego, Calqornia

and

David Paul Watson

Director, HortaPharm B.V.
Chairman, International Hemp Organization,Amsterdam

CAB1 Publishing
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A catalogue record for this book is available from the British Library, London, UK.

Library of Congress cataloging-in-publication data

McPartland, J. M.
Hemp diseases and pests : management and biological control / J.M. McPartland, R.C.
Clarke, D.P. Watson
p. cm.
Includes bibliographic references (p.)
ISBN 0-85 199-454-7 (alk. paper)
1. Hemp--diseases and pests. I. Clarke, Robert Connell, 1953- 11. Watson, David Paul.
111. Title

ISBN 0 85 199 454 7

Disclaimer:
This book contains information about pesticides and biocontrols obtained from research publications, reference texts
and highly regarded experts. This book does not make recommendations for pesticide use, nor does i t imply that the
pesticides discussed here have been registered. Pesticides must be registered by appropriate government agencies
before they can be recommended for use on a plant. Recommendations regarding Cannabis are double-disclaimed in
the USA, since the cultivation of this plant is illegal. The authors and the publisher cannot assume responsibility for the
validity of all information, or for the consequences of information application.
Trademark notice: product or corporate names may be trademarks or registered trademarks, and are used only for
identitication and explanation, without intent to infringe.
The publishers have made every effort to trace the copyright holders for borrowed material. If they have inadvertently
overlooked any, they will be pleased to make the necessary arrangements at the first opportunity.

Printed and bound in the UK by Cromwell Press, Trowbridge, UK, from copy supplied by the authors.
Book design by AMRITA Press and DeGraffics.
 Contents
Preface ................................................................................................................................................................ vii
...
Acknowledgements ........................................................................................................................................\rlll
Forewords ........................................................................................................................................................... ix
H o ~ 7to use this book ........................................................................................................................................ xi

Section One: Prerequisite Information about Crop Biology

1. PRINCIPLES OF PLANT PROTECTION ...........................................................................................1
Introduction to IPM concepts and keys for identifying problems
2. REQUIREMENTS FOR GROWTH ...................................................................................................... 9
Something about soil, moisture, temperature, and light
3. TAXONOMY AND ECOLOGY .......................................................................................................... 13
Classification of organisms associated with Cannabis, their ecological relations,
and symptoms of parasitic interactions

Section Two: Diseases and Pests of Cn~zlzabis

4. INSECTS AND MITES ......................................................................................................................... 25
Cnlztz~zbisis not insect-pollinated, so these organisms primarily act as parasites,
often causing major destruction
5. FUNGAL DISEASES ............................................................................................................................93
Some fungi become Cannabis symbionts, but most cause disease, sometimes death
6. OTHER CANNABIS PESTS AND PATHOGENS .......................................................................... 137
A menagerie of viruses, bacteria, nematodes, other plants, protozoa, slugs,
non-insect arthropods, and vertebrate pests
7. ABIOTIC DISEASES ...........................................................................................................................155
Descriptions of mineral deficiencies and toxicities, sunscorch, other unsuitable
environmental conditions, and herbicide damage
8. POST-HARVEST PROBLEMS ........................................................................................................... 167
Featuring hemp retters and rotters, seed eaters, and marijuana contaminants

Section Three: Control of Diseases and Pests

9.CULTURAL & MECHANICAL METHODS OF CONTROLLING DISEASES AND PESTS... 175
Old and new techniques for reducing damage from pests and disease
10.BIOLOGICAL CONTROL .................................................................................................................181
Useful predators, parasites, microbial pesticides, and companion plants
11.BIORATIONAL CHEMICAL CONTROL ....................................................................................... 189
T11e use and abuse of organic pesticides

Appendix 1: Synthetic chemicals .................................................................................................................. 205

Appendix 2: A dichotomous key of diseases and pests ............................................................................. 211
Appendix 3: Conversion factors .................................................................................................................... 219
References ......................................................................................................................................................... 221
Index ...............................................................................................................................................................245
 Preface
As the 21st century begins, we are favoured with a worldwide resurgence in hemp cultivation. As
global forests dwindle to rampikes and timber limits, we see a glimmer of hope for our future, thanks to a
renewed interest in this ancient and humble source of food and fibre.
This book is our contribution to hemp's revival. Much of the literature regarding hemp diseases and
pests dates back 50 years or more. Further, these publications are frequently buried in obscure agronomy
journals. Cultivators of illicit Cannabis have published high-calibre research in the last 25 years, but they
published in semi-clandestine "grey journals" such as Sinsenzilla Tips. Our primary effort was to collect
this scattered bibliography and assimilate it into a comprehensive and readable format.
Our second effortwas to manoeuvre the control of diseases and pests into the 21st century. Most
hemp research dates to the days when DDT was considered a glamorous panacea. We must find new
control methods for sustainable hemp cultivation. Many "new" pesticides are old, such as pyrethrum, a
popular insecticide before the days of DDT. Biological control also is old; the use of biocontrol against
hemp pests began around 1886, in France. ~ t r une
e t@tea Papineau, we see biocontrol and hemp
resurging together.
This book was written by amateurs, in the sense that none of the authors earns a living primarily as
an entomologist or plant pathologist. Several Canadian and European professionals reviewed our work
and honoured us with unequalled advice. For this book to have a second edition, we ask you, the reader,
to share the results of your experience. Please send us feedback, via our research centre-the International
Hemp Association, Postbus 75007, 1070 AA Amsterdam, The Netherlands.
The format and organization of this text benefited from George Lucas's Diseases of Tobacco, Cynthia
Wescott's Plant Disease Handbook and Gardeneu's Bug Book, the APS Conzpetzdiuin series, and the many-
authored Diseases and Pests of Vegetable Crops it1 Canada, edited by Ronald Howard, J. Allan Garland, and
W. Lloyd Seaman. We hope our efforts to catalogue the C~znnabispest-control literature will reward the
reader, and senreas a platform for launching further investigations. We mind J.D. Bernal's remark, "It is
easier to make a scientific discovery than to learn whether it has already been made."

-J.M. McPartland, R.C. Clarke, D.P. Watson

 Acknowledgements
McPartland: to Tom Alexander, editor of the first American journal dedicated to Cannabis agronomy. Thank you to
Patty for her patience and support, to my brother for Roget's Tl~esalirtis,and to my folks for growing me in the
Garden State. Best regards to Ralph VonQualen and Don Schoeneweiss for plant pathology.
Clarke: to my parents for their unwaivering support and Mother Cannabis for her continuous supply of academic
challenges.
Watson: to my mother, Evelyn Watson, for interesting me in gardening, Allen Chadwick for teaching me
biodynamic farming, and the SSE for its conservation work.

Many individuals graciously reviewed parts of our manuscript (or the entire text - noted by *): J. Bissett, M. Cubeta,
J. Dvorak*, W. Elmer, J. Goodwin, K. Hillig, S.J.Hughes, D. Jackson", D. Lawn, J. Leland, P.J. Maddox,
P. Mahlberg, S. Miller, S.J.Miller*, J. Moes*, G. O'Toole*, D. Pimentel, P.L. Pruitt*, W. Ravensberg, M. Robiner*,
D.P. Rogers, G. Scheifele*,R.E. Schultes, K. Seifert, E.G. Simmons, J.B. Sinclair*, E. Small, B. Sutton, F. Uecker,
D. West*, D. Wirtshafter*. Our 1991 edition was greatly improved by R. Rodale; we miss his equivocal wit.

We are pleased to acknowledge the financial sponsorship provided by three companies in producing the colour
illustrations for this book. All three companies have a practicai interest in the production of organically grown
Cannabis species.
Koppert Biological Systems have produced clear nuclear populations of beneficial organisms in convenient
application systems, which can be placed on affected plants. Beneficial insects can be positioned close to the
focus of pest infestation, and the beneficials conveniently disappear when their food supply is exhausted,
leaving no residues to compromise quality. Contact details: tel. +31 (0)10 5140444; +31 (0)lO 5115203; e-mail
info@koppert.nl;http://www.koppert.com/
GW Pharmaceuticals Ltd grows Cannabis under license from the Home Office in commercial quantities for
medicinal purposes. Organic methods of growing are used and biological pest control is a feature of the
production of medicinal Cannabis. The methods of organic growing ensure high yields and elimination of
pesticide residues. Contact details: tel. +44 (0)1980 619000; fax: +44 (0)1980 619111; e-mail
info@medicinal-cannabis.org; http: / /www.medicinal-cannabis.org/
Hortapharm BV breeds unique Cannabis varieties for medicinal or industrial applications. Hortapharm has
cultivated Cannabis under IPM conditions for a decade, and is responsible for breeding the medical varieties
utilized by GW Pharmaceuticals. Contact details: tel. +31 (0)20 6185591; fax +31 (0)20 6185726; e-mail
Horta@euronet.nl

We donate 25% of our royalties to the International Hemp Association, Postbus 75007,1070 AA Amsterdam, the
Netherlands. Tel./fax: +31 (0)20 6188758; e-mail iha@euronet.nl

B I O L O G I C A L SYSTEMS

'WPharmaceuticals Ltd
 Forewords
Hemp is grown for a variety of uses and there are records of its production and conversion into useful
products for at least 5000 years. Important raw materials are still available from different types of plant in the
genus Cnnilnbis. It is a renewable starting material for many industries, including textiles, paper, rope, as a
foodstuff and fuel, used from ancient times to the present. More recently it has been used in cosmetics and is
being investigated as the starting point for a new generation of biodegradable plastics. Depletion of feed-
stocks from fossil fuels and the need to access alternative sources of cellulose and fixed oils has refocused
attention on renewable sources of industrial starting materials obtainable from Cannabis.
The use of Cnnnnbis in medicine has had a chequered career throughout history. Until the early part of the
twentieth century it was regarded as a valuable medicine in general practice. At present, Cannabis is classified
as a Schedule 1 ("Class A" in the UK) drug and its use is proscribed under legislation following the UN Single
Convention 1961. Recently, approval has been given by the UK Home Office for Cannabis to be grown for
medicinal purposes so that the safety and efficacy of Cannabis as a medicine can be critically re-evaluated,
with a view to its reinstatement as a prescription medicine.
Cnrzlznbis has been grown from antiquity and methods of producing it have been developed in many
geographically different communities. Control of diseases and pests is a necessary part of good husbandry,
and it is useful to have this information critically evaluated and brought together in one volume.
The resurgence of interest in medical and non-medical uses of Cannabis further underlines the need for an
authoritative and comprehensive book on diseases of this valuable crop. The emphasis on biological methods
of control is a recurrent theme throughout the book and reflects the move towards organic methods of horti-
culture for crops grown in the field and under glass. The authors have met this need by providing a compre-
hensive guide based on practical experience and academic expertise.
- Brian A. Whittle

Scientific Director, GW Pharmaceuticals Ltd

This comprehensive volume is an important resource for researchers, crop management consultants, and
growers who wish to increase their knowledge of the pest management of hemp. Many of the pests and
diseases profiled in this book are widely known to crops throughout the world. This book focuses on their
damage to hemp and the specific solutions for this crop. Particularly useful is the clear, systematic approach to
pest and disease descriptions. This is accompanied by an extensive overview of biological, natural, and
chemical solutions to the pests and diseases of hemp, with details on products, application methods, and
dosages. The emphasis on realistic biocontrol options is an outstanding feature . It is rare that such extensive
irxformation on disease and arthropod control has been combined in one volume. Moreover, it will be very
useful for anyone working with pest management in other high-value crops.
Biological control of pests and diseases has evolved rapidly in the past 25 years, and many of the tech-
niques described here have been adapted from tried-and-true strategies that have proved successful for
growers of other crops. 111 fact, Koppert Biological Systems has been collaborating with one of the authors and
\ye have successfully controlled many different pests on hemp with natural enemies routinely used in green-
house vegetables.
Just as the science surrounding hemp continues to evolve, so does the science of biological control. The
authors have succeeded in creating a complete, up-to-date collection of pest management information for
hemp. Ultimately, the techniques described in this volume will be adapted and improved by growers them-
selves and I am confident that many growers of hemp will benefit greatly from the knowledge contained in
this volume.
- Willem J. Ravensberg
Head, R & D Microbials, Koppert Biological Systems
 How to Use This Book
You need not begin this book at page one. If you tentatively identified a pest or pathogen infesting
your plants, look its name up in the index. See if the problem in hand matches its description in the book.
Proceed to read all about the problem and its control measures. Control measures for common pests and
pathogens are highlighted with charts, illustrations, and explicit instructions.
If you face an unknown assailant, turn to the identification keys in Chapter 1 and Appendix 2. If you
dislike keys, most common pests can be diagnosed by the "picture-book methodu-flip through the
illustrations until you recognize your problem.
This book has 110 separate glossary; technical terms used in this text are defined within it. If a techni-
cal term is unfamiliar to you, use the index to locate the term's definition. Terms in bold print indicate
they are being defined in that paragraph. We define technical terms in Chapters 1,2, and 3, to build the
precise language needed for pest identification using pest morphology (form and structure). We are
unapologetic morpl~ologists,and agree with Wheeler (1997), "no credible theoretical reason exists why
molecular data should be more informative or less homoplastic than morphological data."
To control diseases and pests, we consider chemicals a last resort. We prefer cultural techniques,
mechanical methods, and biological controls. Cultural techniques alter the farmscape, making it less
favourable for pests and disease organisms (see Chapter 9). Mechanical methods utilize traps, barriers,
and other ingenious techniques (see Chapter 9). Biological controls employ beneficial organisms to
subdue pests and pathogens (see Chapter 10). Although we discourage the use of chemicals, technical
information regarding their use is presented in Chapter 11 and Appendix 1. This information is presented
in the spirit of harm-reduction, not as a green light for chemical abuse. Use as little as possible.
For current availability of specific biocontrol organisms and biorational chemical controls, please
obtain the Annllnl Directon/ published by BIRC (Bio-Intergral Resource Centre), P.O. Box 7414, Berkeley,
California 94707, telephone (510) 524-2567.
 " I n order to control nzosqulfoes, one m u s f learn to think liken mosqtrito."
-Samuel Taylor Darling

Chapter 1: Principles of Plant Protection

INTRODUCTION condition favouring crop damage. The longer each side, the
Diseases and pests cost farmers many millions of dollars larger the the greater the crop Damage Is
in losses every year, ~~~i~~(1997)estimated that 13% fibre limited by the triangle's shortest side-either an absence of
crops lost to insects, llyo M,ere lost to diseases, and 7% parasites, a healthy environment, or a resistant host. All our
lvere lost to and other organisms, addition to these control methods manipulate one or more sides of the triangle.
losses in the field, Pimentel et al. (1991) added another 9% in Host is manipulated by breeding crop plants for
postharvest storage losses. These percentages would soar if lesistance diseases and Pests. Breeding is simply the
crops were not managed for diseases and pests. mixing of genes and the selection of hybrids with resistant
Our challenge is to sustain crop yields as we shift from traits. The larger a crop's gene pool, the better chance of
chelnical control to biological control. We want products finding resistant genes. Callflabis still has a large gene pool.
made fron1 healthy Canlanbis-no weakened or discoloured It has been bred for aeons all over Asia, and each geographic
fibre, rancid seed oil, or aflatoxin-laden medical marijuana, pocket of plants developed resistance to its local diseases
But we don't nrant pesticide-tainted products, either. and Pests.
Everyone benefits from nontoxic pest control-consumers,
cult~vators,Cnil~~abi?, and our environment. Nontoxic pest
control 1s the keystone for sustainable agriculture.
Sustainability requires a shift in societal consciousness, as
tvell as an agronomic transformation, and we feel Ca;znnbis
cultil-ation can inspire the shift.
Some experts clalm that hemp suffers no pests or
d~seases(Herer 1985, Conrad 1994). Their view may be
ske~ved.LYe have gropvn hemp (McPartland under DEA
permit, Clarke and Watson under Dutch and British licenses),
and we can say with confidence that hemp suffers diseases
and pests. As long as we insist on cultivating large acreages
of intensive, high-output, monocropped plants, our crops
~villattract problems.

DISEASES A N D PESTS
Do insects cause disease? No. According to Whetzel,
q ~ ~ o t eind \Vestcott (1990), disease is damage caused by the
coiitiriiieil or ~ ~ ~ r s i s tirritation
~ilt of a causal factor (e.g., a
nutnent deficiency or a pathogen). Pathogens are organisms
that cause c o i ~ t l i z ~ ~irritation,
cd such as viruses, bacteria, fungi,
parasitic plants, and nematodes.
Injury, in contrast, is caused by transient irritations.
Insects are transient; they cause feeding injury, not disease.
Hail stones cause injury not disease. Injury, however, may Figure 1.2: The expanding gene pool: sources for genetic
lead to srcoil~lar!,iilscose. Hail stones injure plants, and injured resistance to parasites and pests (adapted from Cook
tissues may become infected by disease organisms, such as Qualset'996).
the grey niould f ~ n g u (s B ~ t y tcitlerea).
i~ Aphids injure plants,
and they often transmit viruses as they hop from diseased
When we relocate plants, they are attacked by new pests.
plants to healthy plants.
We must breed new resistance. To breed ~ i-~ y.n l f i c nresistance
nt
requires casting into a larger gene pool-perhaps crossing
THE CROP DAMAGE TRIANGLE
two different cultivars, such as the French culti~.ar'Fibrimon
Any discussion of plant protection should begin with
21' and the Hungarian culti~.ar'Kompolti' (Fig 1.2). Larger
the "crop damage triangle" (Fig 1.1).Crop damage requires
yet, we can breed different landraces, such as dwarf Northern
the presence of: Side 1) a susceptible host, Side 2) a pest or
Russian hemp and giant Southern Chinese hemp. LYe can
pathogen, and Side 3) an environment conducive to crop
even breed different species, such as Cnizilnbis satma and
darnage. The length of each side of the triangle represents a
Caizilabis ajglznnica. Beyond species lies the breeding frontier:
intergeneric crosses between Carlnnbls sotlua and Hl~iiuiliis
l l ~ p l ~ l z(hops).
~s Coniiabis and H l ~ i i ~ i ~are
l i ~ easily
s grafted
together (Crombie & Crombie 1975), but cross-pollination
does not produce viable seeds. Cereal scientists have created
intergeneric crosses since the 1950s (Sears 1956), combining
X-ray irradiation with embryo rescue on artificial growth
Figure 1.1 : The crop
media. But X-rays are now passe. Genetic engineering is the
damage triangle.
1
2 Hemp Diseases and Pests

latest way to mo\.e genes from one plant to another. Engi- crop damage triangle, and utilizes cultural, biological, and
neers have used a bacterium, Agrobacteril~rrltllnzefaciens, to chemical contrd methods (Stern et al. 1959). Researchers in
splice DNA between plants (for a description, see A. tllniefa- Canada and Europe have used terms such as "comple-
cirlls in Chapter 6 ) . More recently, A. ti~?nelfacie>zs has been mentaryUand "coordinated," but the Australian concept of
replaced by microprojectile bombardment devices known as "integrated management" has won the acronym battle
"bioblasters" or "gene guns" (Cl~ristou1996). IPM began as a solution to eco1ogical and econoillic prob-
Bioengineers can introduce genes from closely related lems associated with pesticides. It replaces the concept of
genera (i.e., H u r n u l i ~ s ) or
, distantly related plants (e.g., eradication with the concept of coexistei~ce(McEno 1990). IPM
~ e ~ ~ , o i a d e n dgigantellin).
roi~ Genetic engineering can even practitioners (IPMers) coexist with pests as long as pests re-
produce interplzyletic crosses, splicing genes from bacteria, main below economically-damaging levels. What constitutes
fungi, or animals. In the future, engineers n~illsynthesize "economically-damaging" differs from pest to pest and plant
"designer genesupthe ultimate, infinite gene pool, with to plant. A gardener growing for a flower contest may con-
unkno~\.nbenefits and dangers lurking in its depths. sider damage by a single budworm intolerable. Fibre crops,
Despite this "lurking unknown," we see bioengineered in contrast, endure nlanv budworms before economic thresh-
crops being planted across the USA and Canada, so North olds are reached. ~ i o i o n t r o lalone can keep most pest
America has become a large-scale transgenic field study. populations under economic thresholds.
- -

Unanticipated catastrophes have already arisen, both eco- IPM integrates ideas from conventional agriculture with
logical (Losey et al. 1999) and medical (Nordlee et a1. 1996, ideas from organic farming. Organic farming is defined by
Ebven & Pusztai 1999, Fenton et 01. 1999). the National Organic Standards Board as "an ecological farm
Resistance against insects and diseases does not last for- management system that promotes a n d enhances
e\,er. Pests and pathogens are moving targets; they mutate biodiversity, biological cycles, and soil biological actilrity. It
and undo the work of plant breeders. Pests can adapt within is based on minimal use of off-farm inputs and on
three >ears of a ne1.v cultivar's introduction (Gould 1991). management practices that restore, maintain and enhance
Paradoxically,crops with partial resistance may last longer than ecological harmony." From an economic and polit~cal
strongly-resistant varieties. For more information on breeding standpoint, the minimal use of products from comlnodity
resistance, see Chapter 7 (Genetics) and Chapter 9 (Method 5 ) . corporations removes organic farmers from the corporate
Environment, the base of the crop damage triangle, is food chain. This places organic farmers at odds bvith multi-
manipulated by cultural and mechanical methods. Glass- national giants; this also places organic farmers at odds with
house environments are easily manipulated. To control grey the USDA, which depends upon research money from the
mould, lower the humidity and increase the temperature. Monsantos of the world.
To control spider mites, raise the humidity and lower the The organic process was postulated by Goethe, and
temperature. Outdoor environments may also be manipu- elaborated by Rudolf Steiner, the founder of bio-dynamic
lated. Careful site selection influences the immediate farming. Bio-dynamic farmers control diseases and pests by
microen\.ironment around plants. Choosing a shady site stimulating natural processes and enhancing healthy
protects against sun-loving flea beetles but exacerbates grey ecological relationships (Steiner 1924, AGOL 1998). The same
mould. Prevent root rot by avoiding low-lying, heavy soil year Steiner founded bio-dynamic farming, Sir Albert
(or lightening tlie soil and planting in raised beds). Howard began developing his "Indore Process" for
Fleld locations should be selected with an eye toward maintaining soil fertility (Howard 1943). In the CSA, J.I.
neighbouring crops-if European corn borers are a problem, Rodale began using methods that paralleled the research of
do not plant near maize. The previous season's crop also Howard and Steiner.
dffects a site-expect white root grubs if rotating after sod. Organic farmers focus on soil fertility lnuch more than
Lastly, host density affects the microenvironment and plays 1PMers. Natural soil fertility has many benefits. Crops
a significant role in crop protection (De Meijer et 01. 1994). growing in organically-managed soils suffer less pests than
Pests and pathogens, the third side of the triangle, are crops growing in conventionally-managed soils (Phelan et
controlled by cultural, mechanical, biological, and chemical al. 1996). Organic crops cause less pharyngitis and laryngitis
methods. Traditionally, agronomists often cite "the two E's" in marijuana smokers than crops cultivated with chemical
of pest and pathogen control: excll~sionand emdicnfion. fertilizers (Clarke, unpublished research 1996). Marshmann
Exclusion is why the USA government established the et 01. (1976) compared organically-managed crops to crops
Plant Quarantine Act in 1912-to exclude foreign parasites. grown with chemical fertilizers, and found the former con-
Farmers exclude parasites in three ways: 1) cleaning all tained more A'-tetrahydrocannabinol (THC).Similarly stud-
equipment before entering a field, glasshouse, or grobvroom, ies with Papaver showed that poppies sprayed with pesti-
2) sterilizing or pasteurizing soil before bringing it into a cides produced less opiates than controls (Wu et nl. 1978).
glasshouse or growroom, and 3) using certified seed which Unfortunately, the term "organic farming" has degraded
1s disease- and pest-free. into a marketing tool. The new USDA certification standards
Eradication requires the elimination of parasites once are weaker than standards previously established by indi-
they have arril~ed.Some problems can be eradicated by vidual states in the USA. The term organic farming is also
cultural and mechanical methods (e.g., pruning diseased confused with another USDA buzzword, sustainable agri-
branches, starving pests via crop rotation:heat treatment of culture. Sustainable agriculture shares goals with organic
seeds). Tot01 ei~dicntio?lusually requires the use of heavy farming, but is difficult to define in absolute terms.
pesticides. But spraying one pest may increase the population Sustainable agriculture does not prescribe a concrete set of
of another pest, resulting in more crop damage. "Eradicntion" regenerative &chnologies, practices, or policies. It is more a
llrcolrlcs ail utirenlistic niz~fsetf-defeatirlggoizl. Eradication is not process of education (Roling & Wagemakers 1998).
a concept embraced by practitioners of IPM. Organic farmers and IPMers differ in their approaches
to pesticides. Organic farmers eschew synthetic pesticides,
INTEGRATED PEST MANAGEMENT (IPM) as well as many natural pesticides (see "The National Llst"
IPM is a holistic approach to controlling pests (and in Chapter 11). IPMers, in contrast, use any pesticide that
disease pathogens and weeds) that integrates all sides of the works. But IPMers reject the cot~velitioilnlapproach of spray-
 Chapter 1: Principles of Plant Protection 3

ing pesticides, which is rigidly based on the calendar date. with their hosts, eventually becoming dependent on a single
Instead, IPMers spray on a schedule determined by three host species ("Fahrenholz's Rule"). This is why many pests
aspects: 1) pest monitoring, 2) climate monitoring, and 3) attackone species and disregard others. This happekewith
the presence of beneficial organisms. W i t h IPM, carefill obser- Cannabis pests. Some pests attack hemp plants but cannot
i l ~ t l ( replaces
~i~ tile b r ~ ~ t e f o r cofe conventional cheinicnl zuarfare. feed on marijuana plants (McPartland 1992,1997~1). Are these
Farnzers llsilzg IPM vzzlst closely inonitor crop conditions, hosts different species? Rothschild & Fairbairn (1980) found
hlocontrol organisnls, the iueather, and all pests it? tile area, not the insect pest Pieris brassicae could distinguish between
just single target species. IPM is pest inanagevlent for the Turkish and Mexican strains of marijuana. Are these hosts
infirnlatioiz age. different species?
IPMers must understand a complex web of ecological re- Human taxonomists differentiate between plants by
lationships, such as parasitism, mutualism, and competition. genetic, chemical, and morphological characteristics. ~ h ;
The centre of this web is a crop. In our case, Cannabis. Animals genetic characteristics of Canizabis are currently under close
and other non-photosynthetic organisms have a parasitic re- scrutiny (see "Genetics" section in Chapter 7). The chemical
lationship with Cannabis. There are exceptions, such as Hoino taxonokv of Cannabis is complex; ~a;lnabisis a veritable
sapieiis, who maintains a mutualistic relationship with chemical factory. Cannabis uniquely produces the
Cannabis. Mutualism occurs when both species benefit from cannabinoids, a family of C2,terpenophenolic compounds,
their interaction. Hoino sapiens nurtures the plant's growth including THC, cannabidiol (CBD), cannabinol (CBN),
(cultivation) and disperses seeds (zoochory); the plant cannabichromene (CBC), cannabigerol (CBG), and at least
provides us with fibre, food, oil, and medicaments. Caizizabis 60 other cannabinoids (Turner et al. 1980). The uniaue smell
has a competitive relationship with most other plants, com- of Cannabis, however, is not from cannabinoids, but from
petition for the raw materials of photosynthesis-sunlight, terpenoids. Terpenoids are polymers of a C3 isoprene pre-
\trater, and soil nutrients. cursor, such as the monoterpenoids (with C,,, skeletons),
Relationships change xvith space and time. When two sesquiterpenoids (C,?),diterpenoids (C2,),and Giterpenoids
~nsectsmeet on a plant, the competition becomes intense in (C,,). Cannabis produces over 150 terpenoids, including
a space-limited env~ronmentsuch as the hollow stalk of a caryophyllene, myrcere, humulene, limonene, and several
hemp plant. But insects don't have to meet tocompete. Plants pinenes (Hood et al. 1973, Hendriks et al. 1975,Ross & ElSohly
damaged by leaf-chewing insects are avoided by leafminers 1996, Mediavilla & Steinemann 1997). Collectively,
for many months (Faetl~ 1986).Chewed plants produce more terpenoids are called the essential oil or volatile oil of the
defence chemicals, such as THC, making the remaining plant. One terpenoid, caryophyllene oxide, is the primary
lealres less desirable to leafminers. Also, leafminers feeding volatile sniffed by narcotic dogs (Brenneisen & ElSohly 1988).
on previously-damaged leaves suffer greater parasitism than Interestingly, hemp varieties produce more caryophyllene
leafminers feeding on unchewed leaves. This is because some oxide than drug varieties (Mediavilla & Steinemann 1997).
parasitoids (parasites of pests) use chewed leaves as clues Cannabinoids and terpenoids have pesticidal and repellent
to locate their hosts. Amazingly, leaves damaged by wind or properties (McPartland 199%).
other mechanisms do not attract parasitoids unless oral se- This book will follow a polytypic approach to Cannabis
cretions from pests are added (Turlings et al. 1990). Further- taxonomy. The key below describes four prominent Canna-
more, parasitoids can distinguish between leaf damage bis segregates that we can tell apart on a morphological ba-
caused by their hosts and leaf damage caused by other her- sis. The morphological key is adapted from work by Schultes
bivores (DeMoraes et al. 1998). Plants emit different volatile et al. (1974),Emboden (1973),Small & Cronquist (1976),and
chemicals in response to different pests, and parasitoids clue Clarke (1987):
into the differences. The communication between plants and
parasitoids is more sophisticated than previously realized 1. Cannabis sativa (=C. sativa var. satizia):
(DeMoraes et al. 1998). Plants tall (up to 6 m), stems smooth and hollow, laxly
The final outcome of any interaction is often arbitrated branched with long internodes; petioles short, usually
by the en\~ironment,or the microclimate. Apest may flourish 5-9 leaflets per leaf, leaflets lanceolate, largest leaflets
on a plant's lower leaves, shaded and protected, but not averaging 136 mm long (length/width ratio = 7.5);
silrvive in the harsh environment near a plant's apex. This is racemes have long internodes, and achenes are
especially true in Canriabis, where flowering tops accumulate partially exposed; achenes (seeds) usually >3.7 mm
THC-a chemical with pesticidal activity. long, somewhat lens-shaped with a blunt base, surface
These few paragraphs indicate how complex IPM can dull light-to-dark green and usually unmarbled, seeds
become. In practice, IPM methods are arranged in a hierarchy tlsllally adherent to plants at maturity. Cultivated for
depending on pest populations, crop density, a n d fibre, oil, and sometimes for drugs.
environmental concerns. The primary IPM strategy is
sclectlgity. A control method should selectivel!j kill pests and 2. Catznabis irzdica (=C. sativa var. indica):
not beneficial organisms. Selective timing and selective Plants shorter (under 3 m), stems smooth and nearly
treatment applied to selective infested plants (not the entire solid, densely branched with shorter internodes;
field) minimize collateral damage. Selectivity requires careJ111 petioles shorter, usually 7-11 leaflets per leaf; leaflets
idelltification of pests and patlzogens. Know thy enemy narrow lanceolate, largest leaflets averaging 92 mm
long (l/w ratio = 10); achenes averaging 3.7 mm long,
CANNABIS less lens-shaped, with a more rounded base, surface
Also know thy host. Unfortunately, the taxonomy of Call- green-brown and marbled or unmarbled, with or
nabis remains in flux. The genus may be monotypic without an abscission layer. Cultivated primarily for
(consisting of one species according to Small & Cronquist drugs but also used for fibre and oil.
1976), or polytypic ( ~ , i t ltwo
i or more species according to
Schultes et al. 1973 and Ernboden 1974). 3. Cannabis ruderalis (=C. satlva var. spontanea):
Knowing your host becomes particularly important when Plants small (usually under 0.5 m), stems smooth and
dealing with pathogens and pests. Many parasites coevolve hollow, occasionally unbranched; petioles short,
4 Hemp Diseases and Pests

usually 5-7 leaflets per leaf, leaflets elliptic, largest items of diagnostic equipment come into play, and are de-
leaflets averaging 60 mm long (l/w ratio = 6); achenes scribed below. These items can be purchased from sources
small lvith a pronounced abscission structure at the listed in the annual directory published by BIRC (Bio-Integral
base, surface dull green and marbled, abscission layer Resource Centre), USA telephone 1-510-524-2567. For diffi-
fleshy with oil-producing cells, seeds readily shed from cult-to-find items like aspirators and beat sheets, contact
plant. Not cultivated. Gempler's for their agricultural catalogue (USA 1-800-382-
8473, web site http:/ /www.gemplers.com).
4. Cannabis afghanica (=C. sativn var. nfglznnica):
Plants short (under 1.5 m), stems ribbed and nearly Insects and other artlzropods
solid, densely branched with short internodes; petioles Most insects can be collected with tweezers or a hand
long, usually 7-11 leaflets per leaf, leaflets dark green trowel, plus a flashlight (many are nocturnal). An aspirator
and broadly oblanceolate, largest leaflets averaging 130 (venturi suction trap) is useful for collecting small, mobile
mm (1/w ratio 5); racemes have short internodes, and insects. A penknife may be needed to extract recalcitrant
achenes are not exposed; nested, compound bracts individuals from protected places. Since smashed insects are
sometimes produced; achenes usually <3.0 mm long, difficult to identify, a collection jar keeps them incarcerated
nearly round with a blunt base, surface shiny grey and for closer scrutiny. Knock insects out of foliage with standard
marbled. Culti~ratedexclusively for drugs, primarily muslin sweep nets or beating sheets. Insect traps baited with
hashish. foods or pheromones allow you to monitor pests 24 hours a
day. Insect traps suitable for IPM monitoring are discussed
In our opinion, researchers frequently misname these in Chapter 9 ("method 12").
Cni~ilal~is segregates. C. iizdicli is frequently misnamed C.
sntica, and C. n~ylzniiicnis frequently misnamed C. iildicn.
Clarke (1987)attempted to correct the confusion by elevating
C. ~7&11nizzcnVavilov from its original subspecies level (=C.
snt~i,nf. nj;Ciznilicn Vavilov 1926). Clarke noted that Schultes
ct nl. (1974)lumped C. nf-ghanicawith C. indica. Unfortunately,
Caflruab~sfrom Afghanistan has come to typify C. indicn,
especially in the eyes of marijuana breeders. This is incorrect;
Lamarck (the botanist who named C. iridicn) was entirely 0Yo 10O/o 20O/o 50
unfamiliar \\,ith Afghan Cnni~ab~s. His taxon refers to the Figure 1.3: Visual scale for damage assessment by fungi or
blotype from India (indica). Marijuana breeders' use of the leaf-eating insects on a Cannabis leaf (McPartland).
name "indicn" for the l?fgl~ariicabiotype has become
entrenched, causing extensive confusion. Some breeders (e.g., A magnifying lens (lox to 16x)may be needed to identify
Schoenmakers 1986)double the confusion by callingnfghnnicn small insects (aphids, thrips, mites). Some species require
plants "ul~dernllsspecies". microscopic examination of their genitalia, usually after
The cannab~noidcontent within each Cnilnabls segregate chemical clearance with potassium hydroxide. Immature
\.aries greatly. Reducing THC has occupied hemp breeders larvae (caterpillars, grubs, maggots, etc.) may prove
for years (Bredemann et 01. 1956). Cnnnabis segregates can impossible to identify. With care, captured larvae can be
interbreed and hybridize, exemplified by XC. intersitu Sojak nurtured into adulthood for proper identification.
and by hybrids illustrated by Schoenmakers (1986) and Kees Many Cannabis pests can be identified by the damage
(1958). C. s~ltioaand C. iizdica escape cultivation and grow they cause. A poplllation of insects can also be monitored,
wild, like C. r~ideralis(Small & Cronquist 1976). either directly (by counting them) or indirectly (b?.assessing
their damage). Degree of damage can be estimated with
IPM STEPS visual scales (Fig 1.3). The American plant pathologist
TPhf consists of five steps: 1) identifying and monitoring Nathan Cobb first devised visual scales. Cobb worked vvith
diseases and pests, 2) monitoring the environment, 3) disease damage, but his scales can be used for estimating
deciding the proper IPM intervention, 4) implementing the insect damage. Tehon & Stout (1930) illustrated a variety of
intervention, 5) post-inten7entionreassessment. Cobb scales.
Monitoring methods vary from casual hearsay between
neighbours to daily quantitati1.e trap sampling. Your moni- Fungi and bacteria
toring effort should match the severity of your problem. Few fungi and bacteria are readily identified in the field.
Somev\rhere along the line, keeping a logbook becomes Many must be identified with a microscope. "Immature"
essential. (nonsporulating) fungi, like immature insects, may defy iden-
Monitoring requires the regular inspection of plants, tification. Specimens should be kept moist for a few days to
~nsecttraps, or soil samples. The larger the crop, the greater promote spore development, or they may need to be isolated
number of samples. Be sure to rnonitor hard-to-see spots, and raised on artificial media in petri plates. Hundreds of
like centres of crop fields or back corners of glasshouses. Also different agar-based artificial media are commercially
monitor glasshouse "hotspots" located near doors and available. Many selective media only allow growth of
window \.ents. In your logbook, record the date, time, and specific pathogens.
location of any crop damage observed. If pests are present, Advances in biotechnology may move petri plates to
estimate their n u m b e r s (qualitatively-"many," or the basement (Miller 1995).Two biotechnical approaches are
quantitati\.ely-"average of 5 aphids per leaflet"). Recording available. Inznzl~~iodiagiiosiic
iiietizodolog~esinclude immuno-
the temperature, humidity, and time of day is helpful. Mark fluorescence, dot immunobinding, and enzyme-linked
infested plants with a bright-coloured pole or flag so they immunosorbent assays (ELISA). Niiclric acid-based
can be relocated. ineti~odologies
include nucleic acid probes, restrictive fragment
A seasoned entomologist or plant pathologist can length polymorphism analysis, and polymerase chain
identify a problem at first glance. For the rest of us, several reaction techniques. Immunodiagnostic ELISA tests are the
 Chapter 1: Principles of Plant Protection 5

Keys are indispensable tools for diag-

nosing crop problems . The best keys are Most common svmDtoms with their most common causes:
crop.specific . Some diseases and pests pre- no seedlings-old seeds. cold soil. damping off fungi. eaten by pests
vail in fibre crops. other problems predomi- wilting-too little or too much H 2 0 . leaf-sucking insects. root insects.
nate in drug crops (McPartland 1996a.b) . wilt fungi. nematodes
Keys should be site-specific. because dis- mould on buds or leave-grey mould. brown blight. downy mildew. pink
eases and pests change between indoor rot. powdery mildew (don't confuse mould with webbing)
and outdoor crops . Problems also change webbing-spider mites. budworms. hemp borers. leaf-eating
as plants grow from seedlings to flowering caterpillars
adults . Diseases and pests vary geographi- spots on leaves-leaf-sucking insects. leaf fungi. leafminers. too much
cally-many virus diseases. for instance. fertilizer
are limited by the range of their insect vec- brown and curling leaf margins-too much fertilizer. dry air. too little K.
tors . Different keys can be constructed for brown blight
all these different scenarios . holes in leaves-caterpillars. beetles. grasshoppers. bacteria
There are basically 2 types of identifica- pale green or yellow leaves-not enough N. poor pH. nematodes.
tion keys: synoptic and dichotomous . soil fungi. leaf-sucking insects
Synoptic keys rely on pattern recognition. lumpy stems-European corn borers. hemp borers. beetle grubs.
while dichotomous keys are structured de- canker fungi. stem nematodes
cision trees . Watson prefers a synoptic spindly stem-not enough light. too much yellow light. temperature too
"Simpleton's Key." Clarke prefers the six hot. soil too wet. not enough N or K or Ca
synoptic "Top 10" lists presented below. disfigured roots-soil fungi. nematodes. broomrape. grubs. maggots.
McPartland prefers a dichotomous key. rodents
relegated to Appendix 2 . tips of limbs or tops missing-rodents. rabbits. deer. cattle. humans

Six "Top 10" lists of common disease & pest problems. indexed with page numbers:
Seed & seedlina problems Root problems
Damping-off fungi ...................... . . .......................97 Root knot nematodes ....................... ................ ..... 137
Overwatering ........................ . ............................164 Rhizoctonia root rot ............................................. 102
Cutworms .............................................................. 54 Beetle grubs .................... . . . ...............................65
Crickets ................... .. ........................................ 87 Fusarium root rot .................... . . . ......................108
Rodents .................................................................153 Broomrape .................... . . . ............................. 150
Slugs and snails ................... . . . . ................. 151 Cyst nematodes .................................................. 138
Flea beetles .......................................................... 65 Texas root rot ...................................................... 125
Birds ......................................................................
152 Root maggots ............................. .......... ...................85
Old seed ................................................................ 169 Rodents ............................................................... 153
Insufficient light, water, or temperature .................164 White root grubs .................... . . . .........................68

Flower & leaf p r 0 b l e m ~ outdoors

. Pacle Whole-plant problems Pacle
Grey mould ..............................................................93 Fusarium wilt ....................................................... 109
Aphids .....................................................................31 Charcoal rot .......................................................... 112
Yellow and brown leaf spots ...........................101,104 Verticillium wilt .................... . .............................. 122
Nutritional diseases ............................................. 155 Nutritional problems ..............................................155
Flea beetles ......................... . . ..............................65 Overwatering ...................... . . ....................... 164
. .
Budworms & leaf-eating caterpillars .................51, 57 Bacter~alw ~ l ........................................................
t 146
Downy mildrew .................................................... 106 Virus diseases ...................
....... .................... 142
Plant bugs ............................................................... 73 Dodder ..................................................................148
Brown blight .................... ................... ....................114 Armyworms ...................... . . . . ............................54
Bacterial leaf diseases ............................... 144 Thieves with guns .......................
................ ............23
Deer ....................... .
. ...................................... 153

Stem & branch problems Pacle Indoor problems (alasshouses & arow rooms) &gg
European corn borers ...................
.................... ...... 44 Nutritional diseases and overwatering .......... 155. 164
Grey mould .............................................................. 93 Spider mites ...................... ......... .....................25
Hemp borers .................... . ................................... 48 Aphids ......................... . . ................................ 31
Hemp canker ........................................................... 96 Whiteflies ............................ ...... ..........................39
Beetle & weevil grubs ..................................... 70, 72 Grey mould .............................................................. 93
Fusarium canker ..................... . .. ... ............ 107 Thrips .......................... . . ..................................... 60
Rhizoctonia sore shin ...................
.................... ..... 102 Powdery mildrew ................... . . . . .......................111
Anthracnose ....................... . . ...........................121 Fungus gnats .......................... ... ...................89
Striatura ulcerosa (bacteria) ................................ 145 Leafhoppers ...................
.................... .................. 79
Stem nematodes ................................................. 139 Virus diseases .......................................................142
6 Hemp Diseases and Pests

most popular. They are based on monoclonal antibodies and development of a pest or pathogen varies very little-it
provide portable, on-site testing with rapid results. always equals the sum of temperature (between thresholds)
Commercially available ELISA kits can instantly identify and time. Thus, we can predict the development of problems
fungi that cause disease in Cannabis (e.g., Botrytis cinerea, by summing the growing season's accumulated heat. This is
Rllizoctonin solani, Sclerotinia sclerotiorilnz, Filsariilnz measured in degree days ("Cd or "Fd, for centigrade or
g r ~ ~ n l i r z e a r ~ These
~ ~ i z ) . kits detect microscopic fungi before fahrenheit). Methods of measuring "Cd vary in complexity.
symptoms appear. But they are expensive-single tests cost A daily maximum-minimum thermometer can estimate the
between US$d to $50 (Sutula 1996). approximate "Cd at your location. More accurate
determinations of heat require a calculus of minute-by-
A'Einatodes minute measures of air and soil temperatures.
Some nematodes, such as female root knot nematodes, Starting in early spring, record the average temperature
can be identified by simply inspecting roots. Other for the day (maximum plus minimum divided by 2). From
nematodes may require special techniques, such as boiling the average temperature you then subtract the low threshold
infested roots in lactophenol mixed with cotton blue stain, ("base") temperature. For hemp, Van der Werf (1994) used a
then decolourizing roots in plain lactophenol and viewing base temperature of O°C, and began measuring on the day
roots under a microscope. Agrios (1997) described three he sowed seed. For example, the day he sowed seed, the
methods for separating nematodes from soil or plant tissues: maximum temperature reached 15"C, the minimum was 5"C,
the Baermann funnel method, sieving method, a n d equalling an average temperature of 10°C. Subtract 0°C as
centrifugal or sugar flotation method. The Baermann is the base temperature. By this calculation, the seeds
simple and produces a high yield of nematodes, but takes accumulated ten degree days (10 "Cd) the first 24 hours after
two to five days. Centrifugal flotation can be more expensive planting. Daily degree days are continuously added to
and may yield fewer nematodes than the Baermann method, calculate the accumulated "Cd for the season. Van der Werf
but takes much less time (only ten minutes). (1994) determined that hemp seedlings required 88.3 "Cd
before they emerged from the soil. Slembrouck (1994) also
Viruses used a base of O°C, and began measuring on the day seeds
Identifying viruses is not easy. Their symptoms can be were sown. She calculated that 'Fedora' plants began
confusing. To see them requires an electron microscope. flowering at 1350 "Cd, whereas 'Futura' plants required 1400
Cross-inoculation studies with other plants are time- "Cd before flowering.
intensive. Serological tests and fluorescent antibody For pests and pathogens of temperate crops, many
techniques are expensive and not available for HSV (hemp experts use 10°C as the base temperature. In Vermont (USA),
streak \,irus) or HhlV (hemp mosaic virus)-the most using 10°C as the base temperature a n d beginning
common viruses on Cai~nnbis.Viral inclusion bodies can be measurements on March lst, adult flea beetle emerge from
detected with a light microscope by using special dyes soil to chew on seedlings at 90-110 'Cd (data is less accurate
(Christie rt al. 1995), techniques described in Chapter 6. for soil insects, because we measure air temperature, not soil
temperature). The first generation of European corn borer
Nutritional diseases moths lays eggs around 250-275 "Cd. This year Vermont
Mineral deficiencies are usually diagnosed by observing suffered record-breaking temperatures all spring, so pests
characteristic symptoms in foliage. Soil and plants can be developed early Some farmers were caught by surprise when
chemically tested for mineral deficiencies. Agricultural egg-laying bollworm moths appeared a month before usual,
agents and extension services often provide inexpensive soil but the moths were right on time by our "Cd estimations, so
and water tests. Cheap kits for testing soil are available, but we were ready with our Tricltogra~itnzawasps and Bt sprays.
may provide inaccurate readings.
IMPLEMENTING IPM STRATEGIES
The ultimate aid for identifying pests and diseases is an The person in charge of monitoring pests should also
expert. The USA government has employed public be the decision-maker who implements control strategies. If
consultants since 1854, when Asa Fitch was hired as an not, then the monitor and decision-maker must keep in close
entomologist. The State of New York hired the first communication. Similarly, if the decision-maker and the
professional plant pathologist, Joseph C. Arthur, in 1882. implementor are separate, communication is key. Good IPM
Today, the USDA maintains a network of county extension decisions frequently require outside support, as this book
agents across the country; find your local office in the yellow hopes to provide.
pages. Of course, until Cannabis cultivation is legalized in
the USA, asking extension agents for assistance may be LAST STEP: POST-INTERVENTION MONITORING
hazardous.
During- the 1940s and 1950s, farmers sprayed
. , DDT and
were done, knowing their pests were dead. Not any more.
ENVIRONMENTAL MONITORING Today, monitoring pests and pathogens must cont~nueafter
While monitoring diseases and pests, also keep an eye you have intenrened with control methods, ~ ~ ~ t - i ~ t ~
on the weather. Weather affects the severity of diseases and monitoring provides feedback f o r evaluating the
pests, and the efficacy of control methods. For instance, if effectiveness of the IPM programme. Biocontrol methods
cutworms are a problem, cool, wet weather causes them to require very careful feedback,
proliferate. Wet weather hampers biocontrol of cutworms
1 ~ 1 t hTricl~ogrnn~nza
wasps, but enhances biocontrol with SCRIPT:SHOULDERS OF GIANTS
Stciizerizcnin nematodes. Our current work on Cannabis builds on earlier efforts
Heat controls the development of pests and pathogens. by hundreds of men and women. Allow us to briefly
Development cannot begin until the environment reaches a
highlight some previous researchers and their work. For a
certain temperature ( t h e low threshold), a n d pest
more complete history see the two-part series by McEno
development stops if the temperature climbs too high (the
(1987, 1988).
high threshold). The a m o u n t of heat required for
 Chapter 1: Principles of Plant Protection 7

Surprisingly,the first published de-

scription of a Cannabis disease did not
appear until 1832, and it was described
by an American, Lewis David von
Schweinitz (Fig 1.4). Schweinitz dis-
covered a fungus infesting hemp stalks
near Salem, North Carolina. He named
it Sphaeria cannabis. Schweinitz was an
interesting character-a cigar-smoking
minister, and the first American to earn
a Ph.D. (Rogers 1977).He described the
first fields of wild hemp growing in
North America (Schweinitz 1836).
In Europe, the first Cannabis
problems to receive attention were leaf
diseases: Lasch described Ascochyta
cannabis in 1846;Westendorp described
powdery mildew in 1854; Kirchner de-
scribed Depazea cannabis in 1856; and
two researchers independently
described Septoria cannabis in 1857.Our
modern-day plague, grey mould
(caused by Botrytis cinerea), was
originally described on hemp by
Hazslinszky in 1877. In the
entomology world, the first Cannabis-
specific insects were described in
1860-Aphis cannabis by Passerini in
Italy, and Psyche cannabinella by
Doumere in France.
One of the great workers emerging
from this period was Oskar Kirchner
(Fig 1.4). A "Renaissance man" from
Germany, Kirchner wrote about all
kinds of Cannabis problems-fungi
and insects, as well as nematodes,
bacteria, and parasitic plants (Kirchner
1906). His artwork was admired by
many, and frequently imitated (see
comments in the section on yellow leaf
spot, Chapter 5 ) . Figure 1.4: Outstanding Cannabis researchers: top row, left to right: Vavilov;
America's "Renaissance man" was Schweinitz; Charles; middle left: Kirchner; lower left: Roder; lower right: Dewey
Lyster HoxieDewey (Fig 1.4).Dewey's holding a male (staminate) plant growing next to a female (pistillate) plant.
career at the USDA began with critical
work on the Gramineae. He wrote about medicinal herbs, time, Dewey imported seed directly from China, and devel-
deploring the overharvesting of wild plants such as oped 'Chington.' He subsequently bred the first inter-vari-
goldenseal and ginseng. His ecological views were unique eta1 cultivar, 'Arlington,' by crossing 'Kymington' with
in the USDA. Dewey explained how destroying our native 'Chington.' Dewey also crossed 'Kymington' with the Ital-
prairie enabled tumbleweed (Russian thistle) tospread across ian variety 'Ferrara,' to create the celebrated hybrid
Midwestern rangelands. Dewey's contemporaries, in con- 'Ferramington' (Dewey 1928).Dr. Fleischmann, in Hungary,
trast, believed tumbleweed was a Russian plot to destroy bred a similar cultivar by crossing Dewey's 'Kymington,'
American agriculture [see ScientificAmerican 2645341. with 'F-hemp,' a variety of north Italian ancestry.
From 1899 to 1935 Dewey led fibre-plant investigations Fleischmann's stock sired many of the Hungarian cultivars
at the USDA. He became a champion of hemp, dedicating available today (De Meijer 1995).
his energies and talents to the advancement of Cannabis. Dewey was an ecologist as well as a plant breeder. He
Dewey imported seeds from all over the world, from fibre worked on ways to make paper out of hemp hurds. Long
and drug plants, and evaluated them on American soil. Bbcsa ago he lamented, "There seems to be little doubt that the
(1999) called Dewey "the first hemp breeder," inaugurating present wood supply cannot withstand indefinitely the
a distinguished lineage that includes Fleischmann, Grishko, demands placed upon it ... Our forests are being cut three
Bredemann, von Sengbusch, Hoffmann, Allavena, Virovets, times as fast as they grow" (Dewey & Merrill 1916).
Mathieu, and Bocsa himself. Unfortunately, Dewey lived to see his hemp efforts undone
Dewey (1914) called Coates Bull and Fritz Knorr the first by Harry Anslinger's anti-marijuana propaganda. Dewey
hemp breeders. This pair from St. Paul bred 'Minnesota NO. died several years after passage of the Marihuana Tax Act of
8' from the best Chinese landraces they could buy in 1937.
Kentucky. Dewey subjected 'Minnesota No. 8' to a decade Dewey was not expert in pests and pathogens. For these
of inbreeding and half-sib family selection to create his first problems he collaborated with Vera Charles (Fig 1.4). Vera
successful variety, 'Kymington' (Dewey 1928). At the same Charles was another USDA researcher based near Washing-
d Hemp Diseases and Pests

ton, D.C., but she collected Cannabis specimens from across thanks to the tireless efforts of Fcrri and Goidanich at Bolo-
the country. She also lived to see hemp cultivation outlawed, gna, and Noviello at Naples. Eastern Europe got busy, with
and saw her old hemp research on Potomac Flats re- publications from Poland, Romania, Bulgaria, the former
placed by the Pentagon. Yugoslavia, the former USSR, and especially Hungary. Hun-
Dewey's counterpart in Russia was Nikolai Vavilov (Fig garian research has been headed by Bbsca (an agronomist)
1.4).Among the great scientists of our century, Vavilov has and Nagy (an entomologist).
been lionized as an international statesman of agriculture The study of Cannabis parasites became divided during
and plant genetics (Menvedev 1969). Vavilov collected the 1970s with the rise of anti-marijuana biocontrol research.
Cniznnbis from around the globe, often journeying to central While Europeans tried to control pests and pathogens of
Asia, which he considered the centre of Cnli~iabisdiversity. hemp, USA researchers experimented with the same pests
His Caniiabis germplasm collection is preserved at the newly- and pathogens to control marijuana! Marijuana growers
renamed Vavilov Research Institute (VIR) in St. Petersburg. acquired their own disease and pest experts-Frank,
Preservation of Cannabis germplasm by the VIR has been Rosenthal, Rob Clarke, Sam Selgnij, the Bush Doctor, and
supported by the lnternati&al Hemp ~ssociation(Lemeshev Chief Seven Turtles, who published in journals such as High
e f 171. 1994). Tinzes and Sinsenzilla Tips. Arthur McCain, a biocontrol
Vavilov's research with drug plants elicited criticism from researcher at UC-Berkeley, says the anti-marijuana biocontrol
other Soviet agronomists. His-research was eventually ter- era ended when research was cancelled by the Carter
minated by political action, as was Dewey's research. But administration (Zubrin 1981). Unfortunately, the USA
Vavilov also lost his life. Shortly after his publication of The government may have caught the biocontrol bug again
Oulgzn ~ 7 the
f Cz~ltivatiurzof o l ~ rPrin~aryCrops, in Partict~larof (McPartland &West 1999).
Cl~ltlvate~f Henip, Vavilov locked horns with T.D. Lysenko. The 1980s and 1990s saw a renewal of hemp cultivation
Lysenko was famous for fabricating genetic theories based in Western Europe. This encouraged new publications
on Marxist doctrine. He became a powerful toady of Stalin. concerning diseases and pests (Spaar et a1. 1990, Gutherlet &
Lysenko had Vavilo\~arrested. Shortly before Vavilov died Karus 1995). Much recent phytopathological research has
in one of Stalin's gulags, he wrote for the ages, "We shall go come from Holland (DeMeijer 1993,1995;Hennink et al. 1993,
to the pyre, Itre shall burn, but we shall not renounce our Kok et al. 1994, Van der Werf 1994). McPartland (1981 st 0 1 . )
convictions." revived phytopathological research at the Uni~rersityof
Besides Dewey and Vavilov, many other Calzlzabis re- Illinois, which has a long history of Caililnbis research (e.g.,
searchers died during World War 11, including Kirchner and Tehon & Boewe 1939, Adams 1942, Hackleman & Doiningo
Klebahn in Germany, Curzi in Italy, Guilliermond in France, 1943, Tehon 1951, Boewe 1963, Haney & Bazzaz 1970, Haney
Lange in Denmark, and Komarov and Tranzschel in the & Kutscheid 1973, Haney & Kutscheid 1975). Biocontrol
USSR. One researcher that survived the carnage was Kurt research continues in India, where Cannabis is utilized to
Roder (Fig 1.4). Rbder published a half-dozen papers on control pests of other crops (Pandey 1982, Mojumder et al.
Cnizt~nbisviruses, fungi, and insect vectors, before his 1989, Kaushal &Paul 1989, Upadhyaya & Gupta 1990, Bajpai
laboratory in Berlin was destroyed. & Sharma 1992, Kashyap et al. 1992, Jalees et al. 1993, Vijai et
In the 1950s, Italy became a centre for hemp research a1. 1993, Sharma et al. 1997).The future looks promising.
 "You nezlev knozo zolzat is enotiglz illzless yoti klzoz~izuhnt is move than enotiglz."
-William Blake

Chapter 2: Requirements for Growth

Plants have 17 requirements for growth-moisture, warmth, You can estimate brightness with the light meter in your
light, alr, and 13 nutrients found in the soil: camera. Set the film-speed dial to ASA 200. Aim the camera
meter at a sheet of matt white paper placed near plants, and
MOISTURE orient the paper to receive maximum light. Position the cam-
Thanks to ~ t extenslrre
s root system, Cannabis tolerates era so the meter sees only the paper, and the camera does
dry conditions (although it does not thrive in dry conditions). not shadow the paper. Set the shutter speed at 1/500 second.
Lisson & Mendham (1998)detected water extraction by roots The f-stop setting for a correct exposure at 1/500 can be
130 cm deep in soil. On the other hand, Cantlabis grows converted to lumens using Table 2.1.
poorly in wetlands or saturated soil. Hemp growth peaks
when soil moisture is at 80% of soil field capacity (Slonov & Table 2.1: Converting f-stops to Lumens
I'etinov 1980).Duke (1982)summarized data from 50 reports
and found Cannabis does best in areas receiving 970 mm
rainfall per year (range, 310-4030). During the grozoing sea- f-stop: Lumens:
5011, Lisson & Mendham (1998) measured maximum fibre
1,ields in hemp receiving 535 mm water (rain + irrigation). f2.8 125 L
They calculated a hemp "water use efficiency" equalling 3 g f4 250 L
stem (dry weight) per kg water. f5.6 500 L
Plant hydration is expressed as water potential (w), f8 1000 L
gauged in MegaPascals (MPa) or bars. Older literature fll 2000 L
measures hydration as a percentage of total leaf shturation f16 4000 L
(TLS). Hemp growth peaks at a TLS of 85-93% (Slonov & f22 8000 L
Petinov 1980).This TLS approximates a y value of -0.3 MPa
( = -3 bars). During dry summer months, routinely drops
to -1.2MPa (= -12 bars). When y~ drops below -1.5 MPa, pho- The other way to measure light is energy, its ability to do
tosynthesis shuts down in 75% of maize plants. Note that work (e.g., drive photosynthesis). Engineers measure light
pl~otosynthesisstops before wilt symptoms are seen. Canna- in kilocalories per hour imparted per square metre, or Watts
bis probably shuts down below -1.5 MPa, but the exact per square metre (W m-2).Sunlight entering the Earth's up-
number awaits measurement. Inexpensive instruments for per atmosphere imparts 1350 W m-'. By the time sunllght
measuring w are becoming available. reaches the Earth's surface, its energy has dropped to 1000
Besides soil water and plant hydration, careful cultivators W m-2, at noon on a clear summer day near the equator. On
must account for atnlosplleric water. Cannabis grows best at a a cloudy day sunlight dissipates to 100 W m-2. Moonlight
relati\.e humidity (RH) between 40-80% (Frank 1988), but exerts only 0.01W m-2.Cannabis researchers have grown seed-
RH over 60"" promotes gray mould inafglzanica biotypes and lings under as little as 96 W m-2 of mixed incandescent and
their hybrids. So a RH between 40-6096 is optimal during fluorescent lighting (McPartland 1984). Frank & Rosenthal
floivering, to a v o ~ dgray mould. (1978) suggested flowering Cannabis under a minimum of
215 W m-2 (=20 watts ft-2). For C02-enriched growth
TEMPERATURE chambers, Rosenthal (1990) recommended up to 320 W m-2
In a meta-analysis of 50 studies, Duke (1982)determined to saturate Cannabis photosynthesis.
Cnnnabls growth peaks at a temperature of 14.3"C (range 5.6- Some Cannabis scientists have measure light energy in
27.5'C). For C02-enriched plants in a glasshouse or terms of emitted photons, as pE/m?/second, where 1000 Lux
growroom, the ideal temperature is higher-21-27°C during = 19.5 pE m.'s-' (Balduzzi & Gigliano 1985). Bush Doctor
the day and 13-21°C at night (Frank 1988). (1993b)described the energy emitted by different lights (fluo-
rescent, metal halide, high pressure sodium bulbs), and how
LIGHT they translate to watts m-2 and watts per dollar.
Light may be measured two ways-by quantity and Light energy depends on colour. Colour is a function of
energy. Light quantity is measured by the briglltness cast by wavelength, measured in nanometers (nm).Energy increases
a candle onto a square foot of surface one foot away (1 in proportion to wavelength. Short-wave light has less energy
footcandle or 1 Lumen). Light brightness is what the than long-wave light. For instance, purple light (short
"exposure meter" in your camera measures. In the metric wavelength 420 nm) requires 130 milliWatts to generate 1
world, light imparted by a candle upon a square metre one Lumen of brightness, whereas yellow (long wavelength 570
metre away equals 1 Lux (1 Lux = 0.093 Lumen). n m ) needs only 1.4 milliwatts to generate the same
Ordinary indoor light averages 150 L u , too dim for Can- brightness. Converting from light brightness (Lux) to light
iinbis, a plant that requires a lot of light. Researchers have energy (W m-2) is not simple, since most light represents a
grown Cnnriabzs in growrooms under as little as 600 Lux mixture of wavelengths. Rosenthal (1998) provides
(Sbringer & Nagy 1971). Paris et al. (1975) used fluorescent brightness-to-energy conversion factors for many different
and incandescent lamps emitting 14,000-18,000 Lux. The types of bulbs.
brightest sunlight yet measured is 100,000 Lux atop Mauna Plants prefer certain wavelengths. Plants reject (reflect)
Loa in Ha~vai'i. green-yellow light (500-600 nm). This is why they look green
9
10 Hemp Diseases and Pests

to us. Photosynthesis works best with Table 2.2: Soil nutrient extraction of different crops during one growing season.'
red and blue wavelengths. Bush Doc-
tor (1993b) described the wave- N P2O5 K20 CaO MgO S
lengths ernitted by commercial light CROP (kg ha-1)(kg ha-') (kg ha-') (kg ha-1) (kg ha-') (kg ha I)
bulbs.
Ultraviolet (UV) radiation Maize (Zm ~~zays) 302 130 302 93 123 37
consists of wavelengths shorter than 12,200 kg grain ha-1
deep purple light. UV-A contains Wheat (Tr~ttcunisp ) 152 61 184 34 35 23
wa\relengths from 420 to 315 nm, 5200 kg grain ha-1
UV-B ranges from 315 to 280 nm, and Oats (Aveiza snttva) 131 43 165 21 37 22
UV-C ranges from 280 to 100 nm. UV 3600 kg grain ha-1
radiation damages nucleic acids and Hemp-whole plant 177 53 184 199 35 18
proteins in plants a n d people, =200,000 dry kg ha-1
especially UV-C. Ozone in the Earth's Hemp-stems only 52 12 99 68 12 8
upper atmosphere absorbs all LV-C, 6000 kg ha 1
about 959; of UV-B, and about 50°/; Hemp-seeds only 33 18 8 3 6 9
of UV-A. It has been suggested that 700 kg ha-1
Caizizabis biosynthesizes THC as a UV Hemp-flowers only 56 30 15 6 10 9
protectant (Pate 1983, 1994). Indeed, 1200 kg ha-1
under conditions of high UV-B expo-
sure, Cni~iznbisproduces more THC 1. Data for rows 1-3 converted from Wolf (1999), rows 4-6
(Lydon et al. 1987). from Berger (1969), row 7 from McEno (1991).

Most experts describe Caizizabis

as a short-day pinnt. It flowers in the autumn, when the researchers argue that nickel (Ni), cobalt (Co), sodium (Na),
photoperiod drops below 12-13 hours per day, depending vanadium (V), titanium (Ti), and silicon (Si) are also plant
on the variety and its geographical origin. Actually, micronutrients (Jones 1998).
Cniztiabis is best described as a long-i~iglztplai~t-interrup- Cannabis places greater nutrient demands upon the soil
tion of dark periods by a short light period will completely than other crops. See Table 2.2. Fibre crops require high soil N,
prevent flowering, while an interruption of the light period high K, then in descending order: Ca, P, Mg, and
by even a long dark period will not prevent flowering. micronutrients. Seed crops, compared to fibre crops, extract
less K and more P from the soil. The nutrient extraction of drug
ATMOSPHERE crops has not been measured, but we present estimates in Table
Plants, like all living things, require oxygen to survive. 2.2. Drug crops have a high P requirement (Frank & Rosenthal
But unlike all other creatures, plants provide their own 0 2 1978, Frank 1988), and Mg, Fe, and Mn may play a role in the
as a by-product of photosynthesis. Atmospheric carbon enzyme regulation of THC synthesis (Kaneshima et al. 1973,
diox~de(COz)is often the limiting factor for photosynthe- Latta & Eaton 1975).
sis. Frank (1988) reported peak growth at COz levels of Storm (1987) described the function of Cniltzabis plant nu-
1500 to 2000 ppm (=1.5-2.O%),five or six times greater than trients in detail. Asummary is found in Table 2.3. Plants lacking
current atmospheric concentrations. nutrients produce telltale synptoms. For deficiency symptorn~
and their correction see Chapter 7.
SOIL Soil acidity, measured as pH, directly affects the
Soil science is an interdisciplinary field, the most availability of nutrients in the soil. See Fig 2.1 for an illustration
complex feature a farmer must manipulate. In the USA of this relationship in organic soils. In soils with insufficient
about 20,000 types of soil are recognized (Brady & Weil organic materials, pH has a greater influence on nutrient
1999). Soil series are named by their sites of discovery- availability (Wolf 1999).Duke (1982) summarized pH data from
my garden is dense Vergennes clay; up the hill, the soil 44 reports and suggested a soil pH of 6.5 is best. Test the pH of
lightens to a Covington silty clay loam. Cnilrzabis grows a tablespoon of wet soil by adding a pinch of baking soda. If it
best in a nutrient-rich, well-drained, well-structured, high fizzes, then pH < 5.0 (too acid). Then test a tablespoon of dry
organic matter, silty loam soil. To create this hypothetical soil by adding a few drops of vinegar-if it fizzes, then pH >
substrate, you have to evaluate the soil's nutrient content, 7.5 (too alkaline). Meters to measure p H are relatively
pH, type, and texture. inexpensive and accurate. Frank & Rosenthal(1978) provided
Macronutrients are elements required by plants in charts and tables for adjusting different soils to a proper pH.
relatively large amounts. Organic materials in soil provide Understanding soil, however, is more than measuring pH
three of the six macronutrients-nitrogen (N),phosphorus and nutrients. Digging up soil for chemical tests is like grinding
(P),and sulphur (S).Minerals in soil provide the other three up your finger and conducting the same tests-you learn a lot
macronutrients-potassium (K), magnesium (Mg), and about pH and chemistry, but nothing about structure and func-
calcium (Ca). tion of the soil.
Micronutrients (formally called trace elements) are Soil structure and function is determined by mineral par-
also essential for plant growth, but in relatively small ticles, organic material, and microbiology. The particle size of
amounts. Most micronutrients become toxic to plants if minerals determines the three major soil types-sand, silt, and
they exceed trace amounts. Minerals in soil provide all clay. Sand consists of relatively large particles, froin 2.0 to 0.05
seven micronutrients-iron (Fe), zinc (Zn), boron (B), mm in diameter (these sizes are USDA standards-the British
copper (Cu), manganese (Mn), chlorine (Cl), a n d standard is 2.0-0.06mm). Sandy soil feels gritty when rubbed
molybdenum (Mo). Some nutrients are needed in between the fingers. Silt consists of particles from 0.05 to 0.002
extremely tiny amounts. For instance, a plant needs a mm in diameter, with a floury feel. Clay particles are smaller
million N atoms for every Mo atom (Jones 1998). Some than 0.002 mm, invisible under light microscopes. Wet clay soil
 Chapter 2: Requirements for Growth 11

"slicks out," developing a continuous ribbon when pressed in clay compacts easily. Compacted clay drains poorly, be-
between the thumb and finger. Mixtures of the three soil types comes puddled, and dries into large hard clods. To improve
are called loam soils. clay soil, add organic material or sand. "No-textured soil"
Organic material, "humus," consists of decomposing is really hydroponics-soil science without the soil
plant and animal matter, mixed with microorganisms. solids-100% space. See Storm (1987) for an introduction
Microscopic bacteria and fungi convert the nutrients in humus to Carlilabis hydroponics. See Chapter 7 in this book for
into forms absorbable bv. plants.
- Bacteria and fungi also form more information on soils.
the base of an underground food chain, including protozoans,
arthropods, nematodes, and earthworms. Thus, soil organisms ASSORTED ECOLOGICAL CHARACTERISTICS
do Inore than nznke soil-they are the soil. The microorganisms Bush Doctor (1987a) found stressed plants growing
and bugs hold nutrients in t h e ~ bodies,
r keeping the nutrients at 3050 m above sea level. Sharma (1983) reported plants
from leaching away. As soil organisms die, they release surviving up to 3700 m in altitude. Ca~zrzabisgrows from
nutrients for plants. Soil with more than 10% organic material 0" (the equator) to approximately 63" latitude (nearl). the
is termed muck; if over 25% it becomes a bog or peat field. arctic circle, in Norway). Many hemp cultivars ha\-e been
Optimally, soil consists of 5-10% organic material and 40- developed for 40-55" latitude; they perform poorly at
4jo/b mineral material. The other 50% is suace. The lattice of semitropical and equatorial latitudes, d u e to early
space should contain equal parts air and water. The lattice flowering. Drug varieties were traditionally developed at
allows nutrients to flow, roots to grow, bugs to move around, semitropical and equatorial locations, and perform poorly
and everything to breath. at high latitudes (above 45") due to late flowering (frost
Coarse-textured soil is mostly sand. Sand maintains its kills them before flowering). Other ecological parameters
lattice of space; it does not compact. It drains well, sometimes described in the "triangular ordination" of Grime et al.
too well. TO improxre the water-holding capacity of sand, add (1988)-such as aspect, slope, bare soil index, and
organic material (e.g.,peat moss) or synthetic soil conditioners associated floristic diversity-have not yet been collected
(e.g.,vermiculite). Fine-textured soil is mostly clay. The space for Cannabis.

pH
t Strongly Acid

4.5 5.0 5.5 6.0 6.5

Neutral
I
7.0 7.5 8.0
t
8.5
Strongly Alkaline

9.0 9.5 10.0

1

Figure 2.1:
Effect of soil pH on availability of plant nutrients. Maximum availability is indi-
cated by the widest part of each bar. (McPartland redrawn from Thompson &
Troeh 1973)
12 Hemp Diseases and Pests

Table 2.3: Soil nutrients required for plant growth, and their function.

FORMS
TAKEN
NUTRIENT UP BY PLANT INFORMATION
NUTRIENT

Nitrogen NH,+ part of amino acids (proteins), nucleic acids (DNA and RNA),
N NO; enzymes, coenzymes, cell membranes, and chlorophyll

Phosphorus H,PO; a component of sugar phosphates (ATP), nucleic acids, lipids, and
P HPO,'- coenzymes, promotes root formation and flowering

Potassium K+ The primary intracellular cation and a major enzyme catalyst, fuels
K the "hydrogen pump" and drives stomata1 movement

Calcium Ca2+ cements the middle lamella in cell walls and regulates N
Ca metabolism, promotes healthy root and stem development

Sulphur required constituent of amino acids, enzymes, and coenzymes, involved

S in the formation of vitamins

Magnesium Mg2+ the core of chlorophyll, regulates P metabolism, and may activate
Mg enzymes that synthesize THC

Iron Fez+ occurs in respiratory enzymes and a catalyst of chlorophyll formation

Fe Fe3+ and perhaps THC synthesis

Boron B(OH),- appears in enzymes and regulates K and Ca metabolism, promotes root
B ~0,~- development and prevents tissue necrosis due to excess oxygen

Manganese Mn2+ a component of photosynthetic enzymes, perhaps THC-synthesis

Mn M n3+ enzymes, involved with N and Fe metabolism

Copper Cu2+ part of respiratory and photosynthetic enzymes, involved in cell

Cu wall formation and lignification

Molybdenum serves as a metal component of enzymes required for utilization of N

M0

Chlorine CI- major intracellular anion, activates photosynthesis, involved with K

CI in the regulation of osmotic pressure

Zinc Zn2+ required for DNA and protein synthesis and formation of auxin and
Zn Zn(OH), other growth hormones

Cobalt Co2+ enhances the growth of organisms involved in symbiotic N fixture,

Co constituent of vitamin B,,

Vanadium V+ promotes chlorophyll synthesis, functions in oxidation-reduction reactions

v
Silicon Si(OH), forms enzyme complexes that act as photosynthesis regulators, plays
Si a role in the structural rigidity of cell walls

Sodium Na+ involved in regulation of osmotic pressure

Na
L
 " O n l y when there is clnssificntioti cnlz there be ntznlysis."
-William Jaines

Chapter 3: Taxonomy and Ecology

bIvlost people, especially ecologists, find taxonomy tedious. Nematodes, fungi and parasitic plants occasionally
Ecologists and taxono~nistsexamine the organisms in a hemp vector viruses as they move (grow) from plant to plant.
field from different perspectives. Ecologists examine all
- -
Viruses commonly spread through vegetative propagation
species to study the way organisms interact with each other, (cloning) of infected "mother plants." Viruses may be
at one site. Taxonomists examine all ecosystems to study the transmitted through seeds. Viruses also spread plant-to-plant
way a species is related to other organisms, around the world via root grafts, r' by leaves rubbing in the wind, or by
(Wheeler 1997).Obviously, taxonomists and ecologists need workers moving among diseased and healthy plants. Tobacco
each other. mosaic virus (the virus originally discovered by Iwanowsky)
When it comes to pest identification, taxonomists have can even be transmitted by smoking infected cigarettes near
the upper hand. Once you know the name of a pest or uninfected plants.
pathogen (especially the scientific or Latin name), you can Viruses cause symptoms of stunting, chlorosis, and
research wavs to control it. Exact identification becomes overgrowth. stunting, also known as dwarfing, indicates a
crucial if you use biocontrol, which only works against slowing or cessation of plant growth. When a shoot becomes
specific pests. Applying biocontrol against a misidentified stunted, the internodes are shorter, which results in the
pest can be a waste of time and money. Sloppy identification crowding of foliage, known as a rosette. Viral destruction of
1s permissible if you use nonspecific pesticides. chlorophyll results in chlorosis-the yellowing of normally
Most of us learned a two-kingdom taxonomy in school. green plant tissue. Chlorosis over an entire leaf is called virus
Everything was jarnmed into the Plant Kingdom or the yellows. Chlorosis may form in circular patterns as ring spot,
Animal Kingdom. In 1969 R. H. Whittaker described a five- in stripes as hemp streak, or appear randomly over the leaf
kingdom taxonomy-Monera, Protista (now called as virus mosaic.
Protoctista), Fungi, Plantae, and Animalia. We describe six Symptoms of overgrowth caused by viruses include
kingdoms: Vira, Gttle bits of bad news wrapped in a protein hypertrophy and hyperplasia. These appear as distorted en-
coat; Monera, the prokaryotes, including bacteria, largements and thickenings of leaves and flowers, sometimes
phytoplasmas, and actinomycetes; Protoctista, unicellular called witch's brooms. The lainina and leaf margins of virus-
eukar! otes, lnclud~ngprotozoans, algae, s l ~ m moulds,
e and infected plants inay become distorted and these symptoms
oomvcetes, Fungi, - moulds, mildews, smuts, etc , Plantae, are called wrinkle leaf.
Cniiiinbis, and Animalia, ~ncludingnematodes, molluscs, Luckily, few viruses attack hemp. Why? Cailtznbis
insects, and vertebrates. See Fig 3.1 for an illustration of extracts and purified cannabinoids inhibit the replication of
Cnil?znl~isparasites representing most of these kingdoms. viruses (Blevins & Dumic 1980, Braut-Boucher et al. 1985,
Each klngdom 1s subdivided into Phyla, then Classes, Orders, Lancz et 02.1990,Lancz et al.1991).We know no plant viruses
Families, Genera, and Species ("King Phillip Came Over For causing disease in people; likewise, plants cannot catch the flu.
Gold Sovereigns") Many viruses infect insects. Insect viruses may contain
DNA or RNA. Garrett (1994) suggested insect-borne RIGA
VIRA viruses were originally plant viruses that, millions of years
Conceptually, the viral kingdom resides within the ago, infected insects as insects fed on plant nectar. One such
other kingdoms. Viruses only replicate as true obligate insect RNA virus is sold as a biocontkl agent-the Agrutis
parasites, in connection with a living host. Viruses cannot segetl~liz cytoplasmic polyhedrosis virus ( a b b r e ~ ~ i a t e d
"grow," they do not eat, they d o not have sex. They are AsCPV). DNA viruses are more common biocontrol agents,
complex molecules, entities between chemicals and life. such as the Kuclear Polyhedrosis Virus (NPV) and its many
Viruses contain DNA or RNA, which encode information strains (MbKPV, AcNPV, HzNPV, SeNPV, HcNPV, TnNPV,
for the re-production of identical chemicals. Viruses cause etc.), the Asrutis segetl~lrlgranulosis virus (AsGV), and the
disease by reprogramming their host's metabolic machinery, M e l a n o p l ~ ssatigi~i?lipesentomopoxvirus (MsEPV)
causing host cells to produce foreign (viral) proteins. All the aforementioned biocontrol viruses produce oc-
A Russian botanist, Dmitri Iwanowsky, discovered clusion bodies (OBs) in their hosts. OBs are viruses embed-
\.iruses in 1892 while studying diseased tobacco plants. He ded within a proteinaceous capsule. Viruses in OBs persist
found that the juice from diseased tobacco plants could pass longer in the environment than non-occluded viruses, which
through a bacterial filter and still be infective. Viruses were makes OB viruses more useful as biocontrol agents. OBs
not actually seen until the ad~rentof electron microscopy. formed by NPV viruses contain hundreds or thousands of
Viruses are transmitted (vectored) by aphids as the Lrirusparticles and grow to 20 pm in diameter (Hunter-Fujita
aphids move from diseased plants to healthy plants. To a et al. 1998).OBs of GVs are much smaller and only contain
les>erdegree, viruses are vectored by leafhoppers, whiteflies, one virus particle. When insects ingest OBs, the proteinaceous
~ n ~ t emealybugs,
s, thrips, and other insects with sucking capsule is dissolved by enzymes in the insect midgut. This
mouthparts. Viruses and their vectors work in a symbiotic releases the virus particles, allowing viruses to infect the host
relationship. Viruses induce a change in plant metabolism- and replicate. ~ v e n t u a l the l ~ insec: dies and its carcass dis-
a kind of premature senescence-which makes plant sap integrates, releasing more OBs onto leaf surfaces. Viroids
more nutritious for sap-sucking insects (Kennedy 1951, are similar to viruses, but have no protein coat. Plant viroids
Kennedy et nl. 1939).The insects, in turn, transmit the viruses were discovered in the early 1970s. About a dozen viroid
to new hosts. diseases have been described, none on Carli~abls.
13
14 Hemp Diseases and Pests

tobacco mosaic virus

Figure 3.1: Shapes and sizes of some organisms associated with Cannabis (McPartland redrawn from Agrios 1997).
 Chapter 3: TaIxonomy and Ecology 15

MONERA Cannabis extracts also kill human pathogens (Krejci 1950,

This kingdom contains two phyla, the Cyanophyta Ferenczy 1956, Ferenczy et al. 1958, Kabelik et nl. 1960,
(blue-green algae) a n d the Schizomycetes (bacteria, Radosevic et al. 1962, Gal et al. 1969, Veliky & Genest 1972,
actinomycetes, a n d phytoplasmas). Antoni v a n Veliky & Latta 1974, Klingeren & Ham 1976, Braut-Boucher
Leeuwenhoek discovered bacteria in 1683, shortly after he et al. 1985). Cannabinoids are bactericidal or bacteriostatic,
started making microscopes. Leeuwenhoek was visually- including THC and CBD (Schultz & Haffner 1959, Klingeren
oriented (JanVermeer was his best friend), and he was quite & Ham 1976), cannabidiolic acid (Kabelik et al. 1960, Gal et
a curious fellow. Leeuwenhoek temporarily blinded himself 01.1969,Farkas & Andrassy 1976), cannabigerol (Mechoulam
by observing the ignition of gunpowder magnified xl.60. & Gaoni 1965, Elsohly et al. 1982), and cannabichromene
After that he turned to more pedestrian materials such as (Turner & Elsohly 1981). The non-cannabinoid essential oil
the scum on his teeth, where he discovered bacteria. of Cn~inabisis also bacteriostatic, and the essential oil derived
Thomas Burrill first proposed that bacteria cause from hashish is more bacteriostatic than the essential oil
disease. He studied "fire blight" of pear trees at the derived from fibre cultivars (Fournier et 01. 1978).
University of Illinois in 1878. Burrill predated Pasteur, but Some bacteria act symbiotically with plants, not para-
Pasteur studied bacteria in people, so he got all the glory. Thus sitically. Rhizobium species live within legume roots and trap
\re don't speak of the Burrillization of milk. atmospheric nitrogen for plants. Azobacter, Azospivtlll~n~, and
Bacteria are far smaller than plant cells or animal cells. Klebsiella species live on the surface of roots and also fix
About 1000 bacteria laid side-by-side measure a millimetre. nitrogen. ~ b m have
e been isolated from Cannabis (Kosslak &
Exceptionally small bacteria approximate the size of large Bohlool 1983). Researchers have sprayed these bacteria on
l~iruses,but bacteria are more complex than viruses. Bacteria roots as nitrogen %iofertilizers" (Fokkema & Van Heuvel1986).
are enclosed by a membrane composed of proteins, Many bacteria colonize the surface of plants, forming
carbohydrates, and lipids, similar to plant cells and our cells. resident populations. Most of these epiphytic bacteria are
Unlike us (but like plant cells), bacteria have a cell wall. gram-negative, such as Erwinia, Psel~don~onas, Xatztl~omoi~as,
Bacteria can grow, eat, and have sex. Unlike organisms in all and Flavobacteril~inspecies. Gram-positive species occur less
other kingdoms, they only have one chromosome, and it is frequently ( B a c i l l ~ i s ,Lactobacilllls, and Coryiiebacteriilin
circular, without a nuclear membrane. Bacteria communicate species). Epiphytes live on leaf surfaces (the phylloplane)
~ r i t heach other by passing genes between themselves. more frequently than root surfaces (the rhizoplane). Most
Bacteria are extremely prolific. Some double their num- epiphytes colonize plants in a commensal relationship, living
bers every 20 minutes. If unchecked, one such bacterium off cellular leakage. Others are mutualistic-in exchange for
could multiply into a colony covering the Pentagon in 16.50 plant nutrients, the epiphytes protect plants from pathogenic
hours. Bacteria are very hardy. One plant-pathogenic species, organisms. Redmond et 01. (1987)applied an Er.iuinia epiphyte
Rac~lluscerei~s,can revert to a spore stage and survive in as a biocontrol against Botrytis, the fungal pathogen that
boiling water, frozen water, or no water at all. causes grey mould disease.
Here are some generalities about Cannabis pathogenic s o m i bacteria aid plants by killing insects. One such
bacteria: they are usually rod-shaped bacilli, gram-negative, species, Bacilllls tlzl~rirl~iensis (Bt)is a well-known insect killer.
aerobic, motile (moving with propeller-like flagellae), enter Spray Bt on plants and insects will die after they eat sprayed
plants through small wounds or other openings, generally leaves. Another insect-killing bacterium, X e i ~ o r l ~ n b d l ~ s
have a narrow host range, and rarely harm humans. nematophillls, lives within soil nematodes. You can purchase
About ten species of bacteria attack Cannabis. Their nematodes that contain X. ileitzatoplzii~~s and mix them into
sy~l~ptoins often begin as cl~lorosis-the same as symptoms insect-infested soil. The nematodes find insects, penetrate
from viruses. But the chlorotic tissue subsequently dies them, then release the bacteria. The bacteria kill the insects
(becomes necrotic), dries out, and turns brown. Leaf spots and the nematodes feed off the cadavers. Quite a delivery
are localized lesions of necrotic leaf tissue, more or less system.
circular. If lesions enlarge and become irregular in shape, Phytoplasmas are essentially small bacteria without
they become blotches. Blotches have indistinct chlorotic cell walls. ~ h e were
v disco~ieredina ahoto darkroom shared
margins, and may be accompanied by wilting. Wilting is a by plant and animal scientists. A plant scientist studying dis-
drooping of leaves or shoots, a loss of plant turgor, indicating eased plants was surprised by a lack of viruses appearing in
bacteria have obstructed plant xylem. Blights are symptoms his electronmicrographs. His veterinary colleague, who
of wilting and necrosis which involve whole shoots or studied m y ~ o ~ l a s m a l ~ ~ n e u m of o nswine,
ia said, " ~ u look
t
branches. If wilting becomes permanent, whole branches at all those mycoplasmas." The new microorganisms were
suffer dieback-the wilted leaves turn brown and dry out. subsequently called mycoplasma-like organisms (MLOs).
Dieback begins at the tip of a branch or shoot and advances Genetic studies have determined these organisms are related
backwards to~vardsthe base. Cankers are localized, sunken, to mycoplasmas, so now they are called Phytoplasmas
necrotic lesions on stalks or branches; cankers may cause (Agrios 1997).They spread by sap-sucking insects, which also
wilting and dieback. Rot is a brown liquefying necrosis, in- become infected by them. A Cannabis phytoplasma has been
dicating complete tissue destruction. A root rot indicates reported in India, causing rosette symptoms with
necrosis and collapse of part or all of the root system. Crown hypertrophy and leaf distortion (Phatak et al. 1975).
rot involves the crown (the transition zone between the root
and stalk). One species of bacteria causes hyperplasia and PROTOCTISTA
hypertrophy of plant cells, called a crown gall (a cancer-like This kingdom contains green, plant-like organisms, as
growth). well as organisms that move about like animals. Thus, some
Coiiliabls protects itself by producing many researchers split Protoctista into two kingdoms-Chromista
antibacterial compourtds. Extracts of Cannabis inhibit or kill and Protozoa (Hawksworth et al. 1995). Protoctistans are
plant-pathogenic bacteria (Bel'tyukova 1962, Vijai e f a!. 1993). unicellular or multicellular. They differ from bacteria by
An aqueous extract of h e m p or wild hemp, called having several chromosomes (surrounded by a nuclear
"cansantine" or "konsatin," was sprayed on potatoes and membrane), plus membrane-bound cell organelles (such as
tomatoes to kill bacteria (Zelepukha 1960, Zelepukha 1963). mitochondria, chloroplasts, etc.)
16 Hemp Diseases and Pests

The green algae (Chromista phylum Chlorophyta) de- Fungi cause more Cannabis disease than the rest of
serve mention, since they produce most of the Earth's earth's organisms combined. Some symptoms are similar to
atmospheric oxygen. Chrysophytids should always be those caused by viruses or bacteria, such as chlorosis, necro-
mentioned. They lived and died 20 million years ago. For sis, rosettes, wilting, leaf spots, blotches, blights, diebacks,
aeons their calcium microskeletons piled up on ocean cankers, root rots, and crown rots. Powdery mildew arises
bottoms. We now call these deposits diatomaceous earth, as a thin covering of white fungus upon the upper surfaces
and dust them on plants as an organic control against of leaves. Black mildew and sooty mould appear as black
aphids. growths on leaves; the latter is associated with aphid drop-
Protoctistans of the phylum Protozoa are animal- pings. Rust is distinguished by rust-coloured pustules of
like-such as the classic ~ n i o e b nand the bacteria-eating fungal spores.
Paramecinnz. One g e n u s of flagellated protozoans, The vast majority of fungi are saprophytes, living off
Plzytoinorlas, causes plant disease. It has not been found in already-dead material. They benefit us by decomposing
Caizrlnbis. Extracts of Caiznabis and purified cannabinoids organic matter and releasing nutrients back to the soil. Fungi
kill protozoans (McClean & Zimmerman 1976, Pringle et 01. also ruin our food and overrun leather, cotton, and paper in
1979, Nok et nl. 1994). damp places. A small group of fungi "go both ways" as
Members of the phylum Oomycota (also called saprophytes of dead plants and parasites of living plants.
Oomycetes, algal fungi, or phycomycetes) are difficult to Termed Facultative Parasites (FPs), they normally live as
differentiate from true fungi. They have cellulose in their saprophytes but can attack living hosts. Many of therot fungi
cell walls (like plants), instead of the chitin found in fungi. and damping-off fungi fall into this category. Some FPs also
T~vogenera, Pse~~doperonospora and Pytlziilnz, commonly attack us. Look in an old pint of cottage cheese. The fuzzy
infect Cn~ziznbis.They give rise to unique symptoms. fungus on the lid causes lung disease (geotrichosis),the black
Pseildoperonosporn species cause downy mildew, a blue- mould causes mucormycosis, and the orange slime on the
white felt that forins on the undersides of leaves. Pythiurn bottom causes meningitis and endocarditis in people with
species cause damping off, a rapid collapse of small AIDS. Facultative Saprophytes normally attack living hosts,
seedlings. but can feed on recently-dead ones when times are tough.
Harvey (1925) invented a technique for isolating Obligate Saprophytes only eat the dead. Obligate Parasites
Oornycetes, using steam-sterilized ~ n t i n n b i sseeds. He (OPs) only feed on the living, which poses a problem for
floated the seeds in pond water as Oiimycete "bait." Thanks scientists who study them, since OPs cannot grow on agar
to Harvey's technique, the scientific literature is filled with in petri plates.
reports of aquatic Ocimycetes infesting Cannabis seeds. None Some FPs live as symbionts within the nooks and
of these "baited" Oiimycetes cause problems unless you store crannies of leaves, feeding on cellular leakage, aphid
seeds in pond water. honeydew, pollen grains, and other airborne debris. These
Phylum Myxomycophyta includes the fungus-like phylloplane fungi live above the leaf epidermis (epiphytes)
slllne moulds-a trulv curious bunch. Some "individuals" or in spaces below the epidermis (endophytes).Phylloplane
can exist as eltiler one multicellular organism or a collection fungi protect their plant hosts by repelling pathogenic fungi
of single-celled organisms. If there is enough food around, and herbivorous animals (Fokkema &Van den Heuvel1986).
the single cells go about their business, growing and Mycorrhizae are symbionic fungi that live within plant
dividing like amoebae. But if starved, they aggregate into roots. They extend hyphae into the deep soil, drawing water
clumps and crawl off like a slug. Finding better conditions, and minerals (mostly phosphorus) back to their host's roots.
the slug erects a tall stalk topped by spores. The spores blow In return, the host supplies the mycorrhizae with
off, revert to amoebae, and go their separate ways. Some photosynthetic products. Plants enjoying this fungus
slime moulds are brightly coloured and visibly pulsate. They partnership grow faster t h a n their nonmycorrhizal
often make neighbourhood news when found crawling up neighbours (see "Mycorrhizae" in Chapter 5).
someone's house. Gzebenyuk (1984) found one species, Other "friendly" fungi act as hyperparasites and feed
D~dynlil~riiC ~ Q Z Jclimbing
~ L S , hemp stems. THC and CBN are on other fun@,strangle soil nematodes in nooses of hyphae, and
toxic to other slime moulds (Bram & Brachet 1976). infest insects. Over 700 species of fungi cause diseases in
insects (Roberts & Hajek 1992). One such fungus, Cordyceps
FUNGI sinensis, produces long, thin, black sclerotia, known as "dead
Classification and taxonomy get complicated here. man's fingers" (highly esteemed in Chinese medicine). Many
Even Linnaeus, the genius taxonomist who first coined Cordyceps anamorphs are sold for biocontrol of insects, including
"Cannabis satian," found the fungi frustrating. He lumped Hirstltella, Nonu~raea,Pnecilornyces, and Verticilliz~nzspecies.
rnany fungi in his genus Clzaos. Cannabis produces antifungal chemicals. In this
Fungi can be unicellular or multicellular. They contain capacity, hemp has been planted with potatoes to deter the
one, two, or many nuclei per cell. Fungi have cell walls potato blight fungus, Ph?ytoplzthorn iizfestnns (Israel 1981).
composed of chitin. Some are mobile. Most are sexual. They Concentrated Cannabis extracts are lethal to fungi (Vysots'kyi
are everywhere we look. In this aspect they are more 1962, Misra & Dixit 1979, Pandey 1982, Gupta & Singh 1983,
successful than insects-fungi have conquered the seas, but Singh & Pathak 1984, Grewal 1989, Kaushal & Paul 1989).
we find no insects in marine environments. Experts estimate Pure THC and CBD inhibit fungal growth (Dahiya & Jain
there are at least 1.5 million species of fungi, but only 10% 1977, Elsohly et al. 1982, McPartland 1984), as does
ha1.e been identified to date (Hawksworth et al. 1995). cannabichromene (Turner & Elsohly 1581),and cannabigerol
Fungi have no digestive system, so they absorb (not (Elsohly et al. 1982).Terpenoids and phenols are antifungal,
ingest) nutrients. Fungi grow into food, exude enzymes such as linalool, citronellol, geraniol, eugenol (Kurita et nl.
which cause digestion to occur around them, then they 1981), limonene, cineole, P-myrcene, a- and P-pinene
absorb the nutrients. Multicellular fungi grow in threadlike (DeGroot 1972, Wilson 1957), and these are components of
tubular filaments termed hyphae. Hyphae may or may not Cannabis essential oil ( T ~ ~ r nete ral. 1980).
contain septa, which are incomplete cross-walls. The body But fungi fight back. They can replicate. Fungi
of a fungus, its collection of hyphae, is called a mycelium. reproduce by budding (like bacteria) or by spores. here
 Chapter 3: Taxonomy and Ecology 17

are several kinds of spores. Conidia are spores produced by Most plant-pathogenic fungi produce tivo types of
mitosis. They are genetically haploid, In (like our sperm and spores. The nzeioticnlly-derived sexual spores are called the
ova). But unlike our sperm and ova, conidia can germinate teleomorph stage; these include zygospores, ascospores, and
by themselves, directly into whole In organisms. Hyphae of basidiospores. Mitotically-derived asexual spores are called
two 1n organisms can intertwine and fuse, forming diploid the anamorph stage, including conidia, chlamydospores, and
(211) organisms. This is fungal sex. In some cases when sclerotia. The.ability to produce two spore stages confuses
hyphae fuse, their nuclei remain separate-forming taxonomists: each spore stage may have its own name! The
dikaryotic (1+111) organisms. The hyphae of all these grey mould fungus, for instance, usually produces conidia
organisms-haploid, diploid and dikaryotic-are identical and is called Botytis cinerea. But the fungus sometimes pro-
In external appearance. Only their nuclei know for sure. duces ascospores, and this stage is named Botryotinin
Diploids (2n) can produce spores by meiosis. These fuckeliaizn. TWOnames, one species. How could this happen?
spores may form at the site of haploid fusion (at the zygote, The two spore stages were discovered by different scientists
hence "zygospores"). Spores also arise distally in a sac (the who did not recognize the stages were related to each other.
sac is called an ascus and spores arising in the ascus are called Fungi are classified by their teleoi?zorplz (sexual) stage,
ascospores) or spores arise distally on a club (the club is and placed the Chytridiomycota, Zygomycota, Ascomycota
called a basidium and spores arising on the basidium are and Basidiomycota. The teleomorph stage of some fungi,
called basidiospores). Most Cannabis-attacking fungi however, has not yet been discovered; these fungi are only
produce millions of lnicroscopic spores. Spores come in known by their an an lor pi^ (asexual) stage. Two Frenchmen
assorted sizes and shapes (Fig 3.2). called these nonsexual organisms "imperfect fungi," and the
Some spores, asci, a n d basidia form within term stuck. Imperfects are placed in their own phylum, the
reproductive structures called fruiting bodies. Magic Deuteromycota. Hawksworth et al. (1995) abandoned the
mushrooms, for instance, are fruiting bodies of the Deuteromycota as "an artificial assemblage of fungi." He
basidiomycete Psilocybe cubensis. The rest of P, cubensis is an does not classify members of this large group of fungi, but
underground network of hyphae. Fruiting bodies of Cannabis merely calls them "mitosporic fungi." We disagree with
pathogens are less spectacular. Most look like the period at Hawksworth and retain the phylum for its three useful
the end of this sentence. Figure 3.3 illustrates some fruiting classes (see Table 3.1).
bodies found on hemp-sporangia, apothecia, pycnidia, Few areas in biology evoke more controversy than the
acervuli, synnema (coremia), cleistothecia, perithecia, classification of fungi. Within the phylum Ascomycota, for
spermigonia (=pycnia), aecia, uredia, and telia. Fruiting instance, different researchers recognize three to six classes,
bodies and spores serve as the most useful means of whereas Hawksworth et nl. (1995)recognize none. We fol-
identifying fungi, by their size, shape, colour, and arrangement. low Ainsworth et al. (1973),who recognize six classes. Nam-
Spores may bud directly off specialized hyphae, ing fungi (nomenclature) is governed by the International
without the protection of a fruiting body. Many fungi also Code of Botanical Nomenclature (Greuter et 01. 1994). The
produce chlamydospores, which are hyphae with hard, code is periodically updated and has become rather compli-
thickened walls that survive inhospitable conditions. After cated. As von Arx once said, "nonspecialists have difficul-
bad conditions pass, these survival units germinate and ties in understanding the code and adhering to its provi-
regenerate hyphae. A mass of these survival units, frequently sions."
rounded into a ball rvith a rind-like covering, is called a In this text, we are primarily concerned with plant-
sclerotium. pathogenic fungi. Thus we will bypass many othernrise-im-
portant fungi, such as truffles, the
Eurotiales (human pathogens), a5
well as one whole phylum-the
Chytridiomycota (chyt-rids).Table
Aiternaria 3.1 contains a hierarchical outline
aiternata of plant-pathogenic fungi, ar-
ranged by phylum, class, and or-
der.
Some fungi attack only Can-
ilnbis, while other species plague
a wide variety of plants. Some
fungi switch hosts during their
life cycles, producing different
fruiting bodies on different hosts.
For instance, rust fungi form up
to five different fruiting bodies on
different hosts. None resembles
another. At one time they were
considered five different organ-
isms, rather than fi\,e different
forms of one organism. Cl~aos.

PL ANTAE
We split the plant kingdom
Lasiodiplodia theobromae
into two phyla. The Bryophyta
consists of primitive nonvascular
plants with flagellated sperm
Figure 3.2: Spores of several Cannabis-pathogenic fungi, drawn to scale with a cells, such as liverworts and
glandular leaf hair (McPartland).
18 Hemp Diseases and Pests

Rust
Fungi:

Figure 3.3: Assorted reproductive fruiting structures of Cannabis-pathogenic fungi (McPartland).

mosses. Tracheophyta plants are vascular (containing xy- The most pernicious antagonists of Caniznbis are para-
lem and phloem) with roots, stems, and leaves. Their sperm sitic plants. These plants contain little or no chlorophyll. They
cells (pollen) have no flagella. The Tracheophyta includes leech off other plants by sending modified roots (haustoria)
five classes: psilopsids, club mosses, horsetails, ferns, and into roots or stems of their hosts. More than 2500 species of
seed plants: seed plants are split into two orders, the parasitic plants are known around the world. Luckily, less
G y m n o s p e r m a e (cyads, ginkos, conifers) a n d the than a dozen species leech off Catznabis, collectively they are
Angiospermae (monocots and dicots). Among the latter known as dodder and broomrape. To add injury to insult,
we find Cniznnbis, as well as some of its antagonists. parasitic plants may infect their hosts with viruses.
Antagonist plants that compete with crops for light,
water, and nutrients are called weeds. For a competitive ANIMALIA
edge, evolution has armed some plants with arsenals of The animal kingdom is divided into 33 phyla. Four
poisons and repellants (termed allelochemicals). Many herbivorous (plant-eating) phyla are discussed below: the
weeds inhibit Cniziznbis seed germination (Muminovic Neinatoda (nematodes), Moll~rsca(snails a n d slugs),
1990). According to Good (1953), Cannabis grows poorly Artliropoda (insects and their ilk), and Chor~fatn(e.g., birds
near spinach (Spiizncin olerncen), rye (Secnle cereale), and a n d mammals). Herbivorous animals are nitrogen-
garden cress (Lepidil~nzsativllnz). In retaliation, Caiznabis challenged. Plants consist mainly of carbohydrates, whereas
suppresses neigl-tbouring plants, whether they are weeds animals consist of protein. The difference is nitrogen.
such as purple nutsedge (Srivastava & Das 1974), Animals are 7-14% nitrogen by dry weight (dw).Plant rarely
quackgrass (Muminovic 1991),and chickweed (Stupnicka- contain more than 6% nitrogen (dry weight), except for
Rodzynkie~vicz1970), or crop plants such as maize actively growing tissues and reproductive parts (see Table
(Pandey &k Ivlishra 1982), rice (Vismal & Shukla 1970), 3.2).Thus young shoots and seeds become very attractive to
lupine, beets, brassicas (Good 1953), wheat, rye, and oats nitrogen-starved herbivores (Mattson 1980).
(Schwar 1972).
Catlizabis has weed mimics. Weed mimics imitate Phylum Nematoda (Aschelminthes)
crop plants, so farmers overlook them while weeding. Nematodes (roundworms, eelworms) that feed on
Seedlings of hemp nettle (Galeopsis species), for instance, plants are nearly microscopic. Nematodes are not related to
are very difficult to distinguish from Cannabis seedlings. earthworms. Built on a much simpler scale, nematodes have
Male plants of the dye plant Dntiscn caizrlabina are no respiratory or circulatory systems. They have a few
remarkable mimics of Cnizizabls, fooling even professional muscle cells, which enable nematodes to wiggle out of a
plant taxonomists (Small 1975). predator's grip, but the muscles cannot coord~natemove-
 Chapter 3: Taxonomy and Ecology 19

Table 3.1 : Taxonomy of fungi associated with Cannabis

PHYLUM ZYGOMYCOTA (hyphae rarely have septa; Leptosphaeria, Ophiobolus, Pleospora species
teleomorph produces zygospores) Order Dothideales: pseudothecia are uni- or
Class Zvgomvcetes: produce a profuse mycelium, multiloculate, asci ovate in shape and usually
much of which is immersed in the host; they repro- scattered within locules; Cannabis pathogens:
duce asexually by sporangia Mycosphaerella, Leptosphaerulina, Schiffnerula
Order Mucorales: saprophytic (as storage moulds) species
or weakly pathogenic on Cannabis: Mucor and PHYLUM BASIDIOMYCOTA (hyphae septate with
Rhizopus species clamp connections; teleomorph produces basidiospores)
Order Endogonales: soil fungi forming mycor- Class Teliomvcetes: simple septa present;
rhizal associations with plants, Cannabis basidiospores borne on promycelia and teliospores
symbionts: Glomus species Order Uredinales: obligate parasites with
Order Entomophthorales: parasitic on insects, complicated life cycles spanning several spore
biocontrol agents against pests: Erynia, types; known as the Rust fungi; Cannabis
Conidiobolus, Entomophthora species pathogens: Aecidium, Uromyces, Uredo species
PHYLUM ASCOMYCOTA (hyphae septate; teleomorph Class Hvmenomvcetes: dolipore septa present;
produces ascospores) basidia and basidiospores borne on a hymenium
Class Plectomvcetes: asci unitunicate (walls have (fertile layer lining a fruiting body)
one layer), scattered within a closed cleistothecium or Order Agaricales: fruiting bodies are monomitic
gymnothecium, with single-celled ascospores (one type of thin-walled hypha), hymenium
Order Erysiphales: produce a profuse mycelium, often hidden by a veil-the Mushrooms.
mostly on leaf surfaces; anamorph spores formed Order Aphyllophorales: fruiting bodies are
in chains-the powdery mildews; Cannabis monomitic to trimitic, hymenium exposed;
pathogens: Leveillula, Sphaerotheca species Cannabis pathogens: Athelia, Thanatephorus
Class Pvrenomvcetes: asci unitunicate, asci borne FORM-PHYLUM DEUTEROMYCOTA (hyphae aseptate
in a single layer within an ostiolated perithecium, with or septate; anamorph stage)
single or multicelled ascospores Form-Class Hvphomvcetes: mycelium bears
--
Order Sphaeriales: perithecia dark, carbon- conidia directly on special hyphae (conidiophores),
aceous, with or without a stroma. Anamorph conidiophores free or bound in tufts (coremia) or
spores bud directly off hyphae or form within cushion-like masses (sporodochia); Cannabis
pycnidia or sporodochia; Cannabis pathogens: pathogens: Alternaria, Aspergillus, Botrytis,
Chaetomium, Gibberella, Diaporthe, Hypomyces, Cercospora, Cephalosporium, Cladosporium,
Melanospora, Nectria, Phyllachora species Curvularia, Cylindrosporium, Epicoccum, Fusarium,
Class Discomvcetes: asci unitunicate, borne in a Myrothecium, Penicillium, Periconia,
single layer within a cup-like apothecium, with Phymatotrichopsis, Pithomyces, Pseudocercospora,
paraphyses and single- or multi-celled ascospores Ramularia, Sarcinella, Stemphylium, Thyrospora,
Order Helotiales: asci open with a pore or tear Torula, Trichothecium, Ulocladium, Verficillium
(inoperculate) to release ascospores; Cannabis species
pathogens: Sclerotinia, Orbilla, Hymenoscyphus Form-Class Coelomvcetes: conidia borne on
--
species conidiophores enclosed in pycnidia or acervuli;
Class Loculoascomvcetes: asci bitunicate (have 2 Cannabis pathogens: Ascochyta, Botryodiplodia,
layers), developing in unwalled locules within a Colletotrichum, Coniothyrium, Diplodina,
stroma; ascospores usually multicellular; anamorph Macrophomina, Microdiplodia, Phoma, Phomopsis,
stage produces conidia free or within pycnidia Phyllosticta, Rhabdospora, Septoria, Sphaeropsis
Order Pleosporales: pseudothecia usually species
uniloculate, asci clavate (long cylindrical) in shape --
Form-Class Aaonomvcetes: "Mycelia sterilia,"
and usually arranged in a single layer; mycelium with no reproductive structures; Cannabis
pseudoparaphyses present; Cannabis pathogens: pathogens: Rhizoctonia species
Botryosphaeria, Didymella, Leptospora,

rnent in a specific direction. Their nenrous system is so simple white fibrous material from stunted wheat in England. To
it can be described at the level of individual cells: Caeilorlzabditis his amazement, the fibrous material began to wiggle when
rlegnizs, for instance, has exactly 302 neurons. Acomplete wiring soaked in water. The fibres were matted larvae of A~zgllina
diagram of its nervous system has been compiled. The entire tritici, the wheat gall nematode.
DNA sequence needed to build C. elegans has been described- Most nematodes are dioecious, with males and fe-
19,000 genes, a 97-megabase genomic sequence. The physical males required for reproduction. Males are usually smaller
characteristics of typical plant-pathogenic nematodes are than females. Males of one species, Trichosoinoides
~llustratedin Fig 3.4. crassicallda, are so small they live in the female's uterus!
Nematodes are extremely abundant-a fistful of soil may Some nematodes are hermaphroditic (females with
contain tl~ousandsof them. They occupy every earthly niche additional male g o n a d s ) . Others eliminate males
from mountain top to sea bottom. Nematode crop losses tend altogether, and reproduce parthenogenetically.
to be underestimated because of the nematodes' small size and At least seven nematode species attack Cannabis. All
their unseen (mostly underground) damage. Indeed, the first are polyphagous, which means they attack many different
nematode to be discovered ~ 7 a san odd species that attacks crops. They feed with a hypodermic-like stylet, which
plants above-groliizd. In 1743 Turberville Needham extracted a resembles a hollow spear. Nematodes thrust stylets into
20 Hemp Dis:eases and Pests

Table 3.2:Parts of plants and the nutrients they contain1 Agriolinzas and Arion species) can be nasty pests. They kill
seedlings and can whack older plants. Cannabinoid recep-
Plant part Nitrogen2 Carbohydrate2 Water3 tors and anandamide have been found in several molluscs,
including M y t i l u s edtllis, an edible marine bivalve (Stefano
leaves 1-5 5-1 7 (40)4-90 et al. 1998).
stalks <1 505 30-50
seeds 5-1 0 30 30-50 Phylum Annelida
pollen 5-50 very low 30-70
This group deserves a mention for our nightcrawler
phloem sap 0.004-0.6 5-20 80-95 friend, L ~ ~ n t b r i c it e~rsr e s t r i s . We owe earthworms an
xylem sap 0.0002-0.1 <0.06 >98
incalculable debt for serving as "intestines of the earth," to
quote Aristotle. Earthworms digest organic material, their
l adapted from Mattson (1980), and Young (1997);2~itrogen and
Carbohydrate concentrations measured as % dry wei ht, except tunnels aerate soil, and their manure castings constitute one
for sap which is measured as % weight/volume; Water ex- 3 of the finest fertilizers available. Earthworms can be quite
pressed as % wet weight; 4droughted leaves may have very low prolific-in prairie soil they weigh up to 6738 kg ha-1 (6000
water levels; 5hemp fibres contain high levels of cellulose, an Ibs/acre), but in adjacent corn fields, their numbers drop to
indigestible carbohydrate. an average of 88 kg ha-1 (78 lbs/acre)-earthworms do not
tolerate ploughing and other soil disturbances (Zaborski
1998). Pesticides kill earthworms, particularly carbamates
(benomyl, carbaryl, zineb); over 200 pesticides have been
mouth tested for acute toxicity (Edwards & Bohlen 1992).
stylet muscle knobs
stylet Earthworms are usually 180 m m long and 4 mm wide,
,median bulbs of oesophagus Q
but can grow up to 300 mm long. Earthworms should not be
confused with nematodes, although 19th century experts er-
roneously urged farmers to destroy both. Parkinson (1640)
described how an aqueous extract of macerated hemp leaves,
"powred into the holes of earthwormes, will draw them
forth, and fishermen and anglers have used this feate to get
wormes to baite their hookes." This feat is still practised in
eastern Europe (Kabelik et al. 1960).Cannabinoid receptors
and anandamide have been found in a related annelid
species, Hirudo nledicinalis, the leech (Stefano et al. 1998).

Phylum Arthropoda
Over 800,000 arthropods have been described, wlth
dozens of new species named every week. Arthropods
Figure 3.4:Male and female nematodes showing account for 80% of the animal species on earth, and some
important parts (McPartland). wreak havoc on agriculture. Arthropods can be distinguished
by their jointed chitinous exoskeletons and seginented
bodies. Six classes are hemp herbivores: the Crustacea
(including pillbugs, wit11 five to seven pairs of legs),
plant cells, then suck out cell cytoplasm. Stylets vary in size Symphyla ("garden centipedes," with 12 pairs of legs),
and shape between species and are useful for identifying Chilopoda (true centipedes, with one pair of legs per
nematodes. Nematodes can feed from outside roots as segment), Diplopoda (millipedes, thousand-leggers, with
ectoparasites (e.g., P a ~ a 1 0 1 1 ~ i d onzaxitnl~s)
r~~s or enter plants two pairs of legs per segment and many, many segments),
to feed as endoparasites: Endoparasites are sedentary, Arachnida (spiders, ticks, and mites, with four pairs of legs),
remaining embedded in roots (e.g.,Meloidogyne, Heterodera and the Insecta, with three pairs of legs.
species) or migratory, moving root-to-root (e.g., Pratylenclzi~s Insects are the largest class. Here's some entomology
perzctrans) or migrating to above-ground plant parts (e.g., terminology: the body of an insect is segmented into the
D i t ~ ~ l c z ~ c lLz ~I Ir s~ S L I C ~ ) . head, thorax, and abdomen. Exter~zally,thehead may contain
A ~ o z ~ c - ~ symptoms Y o ~ L ~ z ~from nematodes mimic symp-
. . one pair of compound eyes, one or more pairs of simple eyes
toms of root injury-stunting, chlorosis, and insipient (ocelli), one pair of segmented antennae, breathing tubes
wilting (drooping of leaves during midday with recovery at (tracheae), and mouthparts. The thorax may sport-one or
night). B e l o i u - ~ r o l ~ i l symptoms
d are more distinctive, two pairs of wings, and three pairs of segmented legs. Legs
inEluding root knots or rhotgalls. are jointed, with a hip (trochanter), upper leg (femur),lower
Nematode populations are naturally biocontrolled by leg (tibia), and a foot (tarsus).Insect abdomens may exhibit
viruses, bacteria, protozoans, other nematodes, and killer vestigial legs (prolegs, the fleshy unjointed stubs you see on
fungi (which strangle nematodes in constricting hyphal caterpillars), tympana (thinned sections of abdomen which
loops).Some plants, including Cannabis, ooze metabolites that senreas "ears"), genitalia (modified as stiilgers in bees), and
repel or kill many nematodes (Kir'yanova & Krall 1971, cerci (caudal appendages serving olfactory or tactile
Haseeb et 01. 1978, Vijayalakshmi et al. 1979, Goswami & functions). Internalltl, the head houses brains, a blood vessel,
V~jayalakshrni1986, Mojumder et a1. 1989, Kok et al. 1994, and the oesophagus. The thorax contains nerve ganglia
.Matee\la 1995).Awormrelated to nematodes, Dugesia tzgrlna, ("sub-brains"), the lower oesophagus (including the crop),
15 kllled by THC and CBD (Lenicque et al. 1972). two pairs of spiracles (breathing apparati), the aorta, and
muscles for locomotion. Insect abdomens contain insect
Phylum Mollusca hearts, digestive organs, more nerve ganglia, eight more
Here we have the snails and slugs. They are familiar pair of spiracles, excretory organs, and reproductive systems
creatures, seemingly benign. But garden slugs (Linzax, (Fig 3.5).
 Chapter 3: Taxonomy and Ecology 21

Moths, butterflies, flies, beetles, wasps, bees, and ants

&AD THORAX ABDOMEN
have four-stage life cycles-eggs, immature larvae (caterpil-
lars, maggots, grubs), pupae (cocoons, chrysalids), and adults.
This is called complete metamorphosis because larvae undergo
a dramatic change during the pupal state. Larvae of moths and
butterflies (caterpillars) may have up to 22 legs-six in front
are true legs, the rest are prolegs and disappear during meta-
rnorphosis. Beetle larvae (grubs) have six legs. Fly larvae (mag-
gots) have no legs. Larvae generally do not feed on the same
plants as adults.
Aphids, leafhoppers, thrips, plant bugs, grasshoppers,
and related insects only pass through three stages-eggs,
immature nymphs, and adults. This is called incomplete
metamorphosis, without any dramatic change in appearance.
Nymphs and adults often feed on the same host plants.
Figure 3.5:
To grow through their hard exoskeletons, immature in-
Typical insect showing important parts (McPartland)
sects undergo several moults, when they shed their old
exoskeletons. Stages between moults are termed instars (eggs
hatch into first instars, then moult into second instars, moult
into third instars, etc.). lnsects may be univoltine, producing Insects feed with sucking or chewing mouthparts.
one generation per year, or multivoltine, producing several Sucking insects insert syringe-like mouthparts into plants
generations per year. Insects are cold-blooded, so their growth to suck sap. They mostly feed on phloem sap, although
rate and reproduction is partially d e p e n d e n t o n the some also feed on xylem sap. Sap-suckers must absorb
temperature. Cold temperatures may induce winter dormancy huge amounts of sap to obtain their required nitrogen (see
or hibernation in insects. Generally only one stage hibernates Table 3.2); some suck fluids weighing 100-300 times their
(either eggs, larvae, pupae, or adults, depending on the own body weight per day. Excess water and carbohydrates
species).Some insects go dormant before temperatures become are excreted as sticky honeydew.
unfavourable. This is called diapause, and is frequently lnsects with chewing mouthparts may selectively
triggered by short photoperiods in autumn. feed on the delicate leaf tissue between veins, leawng

Table 3.3: A synopsis of the insect orders associated with Cannabis.

ORDER EXAMPLES CHARACTERISTICS

Collembola springtails simple metamorphosis, no wings, chewing mouthparts

Orthoptera crickets, grasshoppers incomplete metamorphosis, 4 wings, chewing parts

Dermaptera earwigs incomplete metamorphosis, 4 wings, chewing mouthparts

lsoptera termites incomplete metamorphosis, 4 wings, chewing mouthparts

1 Thysanoptera thrips incomplete metamorphosis, 4 wings, rasping-sucking mouthparts I

Hemiptera true bugs incomplete metamorphosis, "half-wings" (part chitinous, part membranous),
piercing-sucking mouthparts arising from front part of the head.

Homoptera aphids, scales, incomplete metamorphosis, wings either chitinous or membranous

whiteflies, leafhoppers (but uniform), piercing-sucking mouthparts arising from posterior part
of the head

Neuroptera lacewings incomplete metamorphosis, 4 wings, chewing mouthparts in larvae and

adults (larvae carnivorous, eating other insects)

Lepidoptera butterflies and moths complete metamorphosis, 4 wings, chewing mouthparts in larvae,
siphoning mouthparts in adults; larvae lack compound eyes 1
Coleoptera beetles and weevils complete metamorphosis, 4 wings (the front pair hardened into a sheath),
chewing mouthparts in larvae and adults; larvae lack compound eyes

Hymemoptera bees, wasps, ants, complete metamorphosis, 4 wings, chewing or reduced mouthparts in
sawflies larvae, and chewing-lapping in adults; larvae lack compound eyes

Diptera flies complete metamorphosis, 2 wings, chewing or reduced mouthparts in

larvae, and sucking-sponging in adults; larvae lack compound eyes
22 Hemp Diseases and Pests

behind a skeleton of leaf veins (skeletonizers). Other chewers

are less selective and gnaw large holes in leaves or notch leaf
edges. Chewing insects also bore into stems (stem borers),
roots (root borers), or even into the narrow spaces inside
leaves (leafminers).Seed eaters may puncture seeds and suck
out the contents, or break open seeds and shell out the
contents, or eat seeds whole.
Twenty-seven orders of insects are currently recognized
by entomologists. About'a third of them include plant pests.
Table 3.3 outlines the orders involved in Cannabis ecology.
Of course, many insects are our friends, like ladybeetles.
Some insects are even domesticated, such as silk moths
(Bombyx mori), which now exist only in captivity. More than
35 centuries of selective breeding have turned B. mori
caterpillars into fat, sedentary, silk-spewing machines, and
the adults have lost the ability to fly (Young 1997).
Cannabis is primarily wind-pollinated, so it need not
attract symbiotic, pollen-collecting insects. Vavilov (1926)
described a symbiotic exception-he found a red bug
(Pyr~hocovisaptevus, Fig 4.39) practising zoochory-the
animal dispersal of plant seeds. P. apterus is attracted to a fat
pad at the base of Cannabis vuderalis seeds, not the seeds Figure 3.6: Cystolith trichomes on the surface of a
themselves. In the process of fat pad feeding, the bug carries Cannabis leaf, seen with a scanning electron microscope
seeds "far distances" and facilitates the spread of Cannabis. (SEMx2000, McPartland).
Herbivorous insects destroy as much Cannabis as
bacteria, protozoans, and nematodes combined. Wounds
caused by insects serve as portals for fungal infections. Some Volatile chemicals may repel pests from neighbouring
insects actually carry plant-pathogenic viruses, bacteria, and plants. Riley (1885) noted that Cannabis sativa growing near
fungi from plant to plant. These pathogens often cause more cotton "exerted a protective influence" against cottonworms
damage than the insects themselves. (Alabama argillacea, he called them Aletia xylina). Similarly,
Plants repel insects by producing mechanical and hemp grown around vegetable fields safeguarded the fields
chemical deterrents. Cannabis leaves are covered by cystolith from attack by a cabbage caterpillar, Pieris brassicae (Beling
trichomes (Fig 3.6).These microscopic hairs resemble pointed 1932); potato fields were protected against the potato beetle,
needles of glass. They are in fact heavily silicified,embedded Leptinotarsa decemlineata (Stratii 1976); and wheat suffered
with calcium carbonate crystals. Cystoliths impede the less damage by the root maggot, Delia coarctata (Pakhomov
mobility of smaller herbivores, and damage the mouthparts & Potushanskii 1977).
of larger ones. Small insects actually impale themselves on Dried Cannabis leaves repel weevils from harvested
cystolith spikes-they may remain impaled and die, or suffer grain (Riley & Howard 1892, MacIndoo & Stevers 1924,
a morbid series of impale-and-escape episodes (Levin 1973). Khare et al. 1974).Scattering a 2 cm layer of leaves over piles
Cannabis is also covered by glandular trichomes (see of potatoes protected them from the tuber moth, Phthorimaea
Fig 3.2). Glandular trichomes secrete many chemicals, operculella, for up to 120 days (Kashyap et al. 1992).Prakash
including terpenoids, ketones, and cannabinoids. Glandular et al. (1987) mixed dried leaves into rice, 2% w/w, to control
trichomes may rupture and release their fluid contents when Sitophilus o y z a weevils in the laboratory; but this dose failed
damaged by insects. The fluid oxidizes into a dark, gummy to provide adequate protection in natural storage conditions
substance. This gummy substance accumulates on mouth- (Prakash et al. 1982). Dried leaves kill ticks (Reznik & Imbs
parts and limbs of insects and eventually immobilizes them. 1965), Varroa mites (Surina & Stolbov 1981), and drive off
The contents of glandular trichomes may also poison bedbugs when placed under mattresses (King 1854, Chopra
insects. Glandular terpenoids produced by other plants are et al. 1941). Sprays made from Cannabis leaves kill many
powerful insect poisons (e.g., ryania, azadirachtin, and insect pests (Bouquet 1950, Abrol & Chopra 1963, Reznik &
pyrethroids). Two terpenoids produced by Cannabis- Imbs 1965, Stratii 1976, Fenili & Pegazzano 1974, Bajpai &
limonene and p i n e n e a r e almost as potent. Limonene is sold Sharma 1992, Jalees et al. 1993, Sharma et al. 1997). Juice
as an insecticide (Demizeo).High-THC drug plants produce squeezed from leaves or seeds has removed vermin from
three to six times more limonene and pinene than most hemp the scalp and ears (Pliny 1950reprint, Culpepper 1814, Indian
varieties (Mediavilla & Steinemann 1997). The terpenoids Hemp Drugs Commission 1894)
humulene and caryophyllene also poison insects (Messer et If Cannabis is such a good insecticide, how do Cannabis-
al. 1990).Methyl ketones are synthesized by Cannabis (Turner eating insects survive? sap-sucking insects, such as aphids
et al. 1980)and repel leaf-eating insects (Kashyap et al. 1991). and mites, use their long stylets to bypass chemicals secreted
Cannabinoids, such as THC, possess insecticidal and on the surface of plants. Leaf-chewing insects, however,
repellent properties (Rothschild et al. 1977, Rothschild & cannot avoid cannabinoids and terpenoids. Perhaps they
Fairbairn 1980, McPartland 199%). intersperse marijuana meals with less-toxic lunches on other
Levin (1973) asked why these chemicals are secreted plants. Spilosoma obliqua caterpillars, for instance, often eat
atop glandular trichomes rather than sequestered in the Cannabis in India (Nair & Ponnappa 1974). But when
interior of the leaf. He hypothesized that trichomes serve Deshmukh et al. (1979) force-fed S . obliqua a pure Cannabis
as an "early warning system." Many insects have diet, the caterpillars died after 20 days.
chemoreceptors located on their feet (their feet smell).These Insects protect themselves from consumed toxins by
insects are repelled the moment their feet touch a chemical- rapid excretion, enzymatic detoxification,and sequestration.
laden trichome. The rest of the leaf avoids damage. Sequestration involves the bioaccumulation of toxins into
 Chapter 3: Taxoinomy and Ecology 23

impervious tissues or glands, such as fat deposits or the ex- The production of cannabinoids increases in plants un-
oskeleton. Monarch butterfly caterpillars are famous for der stress (Pate 1999).Other plants under stress increase their
feeding on milkweeds a n d sequestering poisonous production of terpenoids. For instance, tobacco plants dam-
pyrrolizidine alkaloids. The stored poisons are distasteful aged by budworms (Heliothis virescens) release much more
to birds and rodents and serve as predator deterrents. Pcaryophyllene and slightly more pfarnesene than undam-
Rothschild et ill. (1977) studied sequestration of aged plants. Plants damaged by bollworms (Helicoveya zea),
cannabinoids in caterpillars of the tiger moth, Arctia caja, and produce slightly more Pcaryophyllene and much more P-
nymphs of the stink grasshopper, Zoizocer~~s elegans. farnesenea-a reversed ratio. These different ratios can be
Caterpillars eating Caniiabis stored a significant amount of distinguished by the parasitic wasp Cardiochiles nigriceps,
THC and CBD in their exoskeleton. Rothschild estimated a which only attacks H. virescens (DeMoraes et al. 1998).
single caterpillar containing 0.07 mg THC could elicit Essentially, the tobacco plants utilize volatile terpenoids as
pharmacological activity in a predatory mouse. "smoke signals" to alert the enemy of their enemy.
Grasshoppers stored little THC and n o CBD in their Terpenoids also have their bad side, from our perspec-
exoskeletons, thev excreted most cannabinoids in their frass. tive. Terpenoids attract certain pests (Howse et al. 1998),and
Some insects metabolize poisons and convert them to they repel or harm some beneficial insects. See "Method 5"
other uses. The following examples are cited by Duffey in Chapter 9 for more information.
(1980): Beetles blend plant oils (e.g., myrcene) with
pheromones to aid the diffusion of their mating signals. Phylum Chordata
Spider mites (Tetranychus urticae) convert farnesol to Chordates contain internal skeletons and dorsal nerve
pheromones. Lirnonene, a-pinene, and eugenol accelerate chords. Most of the 15 chordate classes ignore Cannabis, in-
the reproductive maturation of locusts. Aphids use P- cluding Class Reptilia and Class Amphibia. Concerning
farnesene as an alarm pheromone (Howse et al. 1998). Can- Class Osteichthyes (fish), we have two reports: Fairbain
nabis produces all these volatile chemicals (Turner et al.1980). (1976) described anglers using Cntuzabis seeds for fish bait.
Perhaps cannabinoids signal insects as behavioural Anglers germinated the seeds and used them when root tips
pheromones. Some pheromones are heterocyclic structures emerged from seedcoats. At this stage the seeds resemble
(Howse et al. 1998). Rothschild et al. (1977) observed water snails. Clarke (pers. commun. 1997)reported European
paradoxical behaviour in Arctia caja exposed to THC. fishermen mixing Cannabis seeds in dough balls as carp bait.
Rothschild gave caterpillars a feeding choice of two Caiznabis Members of only two chordate classes regularly interact
plants-one plant rich in THC, the other plant with little with Cannabis. They act as herbivores, sometimes aiding
THC but wit11 high levels of cannabidiol (CBD). The plants by distributing seeds which pass per anurn:
caterpillars s h o ~ \ ~ ea ddefinite preference for high-THC Class Aves includes four orders of Caiznabis seed-eating
plants, e\.en though caterpillars feeding on high-THC plants birds: Galliformes (r.g., quail and pheasant), Columbiformes
grew stunted and died sooner than caterpillars feeding on (doves), Pickformes (woodpeckers), and Passeriformes (lin-
high-CBD plants. Rothschild noted, "...should these nets, starlings, grackles, sparrows, nuthatches, goldfinches,
co~npounds[cannabinoids] exert a fatal fascination for tiger etc.).As a historical postscript, early reports from Kentucky
caterpillars, it suggests another subtle system of insect control described the now-extinct passenger pigeon (Ectopistes
by plants." nzigratorilis) feeding on hemp seed (Allen 1908). Seed from
Subsequently, Rothscl~ild& Fairbairn (1980) studied feral hemp is an important food for several midwestern
egg-laying females of the large white butterfly (Pieris American game birds. This has pitted wildlife agencies
brass~cae). Females of this species normally oviposit on against police who eradicate feral hemp (Vance 1971).
cabbage leaves. Spraying cabbage leaves with extracts of Members of the class Mammalia are frequent Cannabis
THC-rich plants reduced egg laying by I! brassicae, compared pests, including the order Rodentia (mice, moles, field voles,
to cabbage leaves sprayed with extracts of CBD-plants. gophers, and groundhogs/woodchucks), order Lagomorpha
Furthermore, spraying leaves with a 1%THC solution (rabbits and hares), and order Artiodactyla (deer). Some
reduced egg laying by I? bmssicae, compared to leaves humans (Honzo sapiens) can be destructive. Cannabinoids are
sprayed with a 1% CBD solution. Both cannabinoids not very toxic to mammals. The oral LDSoof THC in mice is
signalled females as oviposit deterrents. >21,600 mg kg-I (Loewe 1946).Mixtures of cannabinoids are
Rothschild conducted her research before the disco~7ery even less toxic than pure THC (Thompson et al. 1973).THC
of cannabinoid receptors. Does THC exert behavioural may harm ruminants (cows, sheep, deer) because of its
changes in insects via cannabinoid receptors? Preliminary antibacterial activity. Deer rumen microorganisms are
results sav no, cannabinoid receptors (cB~) could not bk inhibited by two terpenes, cine01 and camphor (Nagy et al.
detected in brain tissue of a locust, ~chiskcercngregnria 1964). These terpenes are also biosynthesized by Caiznabis
(Egertova et al. 1998). (Turner et al. 1980).
 " A single man cannot help his tinze, he can only express its collapse."
-S~ren Kierkegaard

Chapter 4: Insects and Mites

Mostafa & Messenger at tips of leaves and flow-

(1972)listed 272 species of ering tops. Clusters of
insects and mites asso- diapausing mites may con-
ciated with Cannabis. Here tain many thousands of
we describe fewer organ- individuals and grow
isms, about 150. This quite large (Plate 2).
chapter represents a
critical review of t h e TAXONOMY
existing literature-we This pest may consist of
believe many publications one species (Ravensberg,
describing "Cannabis- pers. commun. 1998) or
insect associations" two species (Gill & Sand-
actually report insects and erson 1998). We see two
entomologists meeting species, separated by dif-
accidently in hemp fields. ferences in morphology
Insects, after all, wander. and ecology. The species
Mites and insects are are known by at least six
presented here in their common names-red spi-
approximate order of der mites, carmine spider
economic impact. Com- mites, two-spotted spider
mon names are standards Figure 4.1 : Life stages of female Tetranychus urticae mites, glasshouse spider
accepted by the Entomo- (McPartland modified from Malais & Ravensberg 1992). mites, simple spider mites,
logical Society of America a n d common spinning
(Bosik 1997). These standards may differ from common mite. To compound the confusion, about 60 Latin names ex-
names used in Europe (Wood 1989) or Australia (Naumann ist for these two species, described from different hosts
1993).Scientific (=Latin)names change occasionally; we have around the world. At least four of these taxonomic synonyms
included synonyms or earlier names for the sake of appear in the Cannabis literature.
continuity and reference to earlier literature.
1. TWO-SPOTTED SPIDER MITE
SPIDER MITES Tetranychus tlrticae ~ o c 1886,
h Acari; Tetranychidae.
=Tetranychus binzaculatus Harvey 1898, =Tetranychus telari~isof
These insidious arachnids are the most destructive pests various authors,=Epitrtranycus 1901
of glasshouse and growroom Cannabis. Indoor areas are Description: Eggs are spherical and 0.14 mm in diametre.
contaminated by bringing in done'from Initially translucent to white, eggs turn a straw colour just before
infested mother plants. Outdoor crops may also become hatching. Hatching larvae have six legs and two tiny red eye spots.
infested in warm climates; Cherian (1932) reported 50% Protonymphs become eight-legged and moult into deutonymphs.
losses in field crops near Madras, India. Spider mites bite Deutonymphs moult into yellow-green coloured adults. As adults
into leaves and suck up exuded sap. They usually congregate feed on chlorophyll-rich plants, two brown-black spots enlarge across
on the undersides of leaves, but in heavy infestations may their dorsum (Fig 4.1; Plate 3). Females mites average 0.4 to 0.5 mm
be found on both sides of leaves. in length. Males are slightly smaller, with a less-rounded posterior.
Their dorsal spots may not be as evident. The male's knobbed
aedeagi are at right angles to the neck and symmetrical. As winter
SIGNS & SYMPTOMS approaches, two-spotted spider mites turn bright orange-red (Plate
Damage is not initially evident. Each mite puncture Pro- 2), making them difficult to distinguish from carmine spider mites
duces a tiny, light-coloured leaf spot ("stipple"), which ap- or even the predatory mite Phytoseiultls persirnilis.
pears on both sides of the leaf. Stipples are grey-white to
yellow. They begin the size of pinpricks, then enlarge (see Life History & Host Range
Plate 1). Many stipples arise in lines parallel to leaf veins. T.tlrticae overwinters as an adult and emerges in the
Ultimately, whole leaves turn a parched yellow colour, droop spring. Females lay eggs on undersides of leaves or in small
as if wilting, then turn brown and die (Kirchner 1906). In- webs, one ata time. They lay as many as 200 eggs. Eggs hatch
specting the underside of leaf surfaces, particularly along into larvae, which moult three times before they are capable
main veins, reveals silvery webbing, eggs ("nits"), faecal of reproduction. Females arise in a 3:l ratio to males. Under
deposits ("frass"), and the mites themselves. Leaves near the optimum conditions for development (30°C with low hu-
bottom of the plant are usually infested first. midity), the life cycle of two-spotted spider mites repeats
Spider mites tend to infest crops in a patchy distribu- every eight days. Shortened photoperiods in autumn usu-
tion, so early infestations may be missed. Symptoms are the ally induce a reproductive diapause, where females stop feed-
worst during flowering, when whole plants dry up and be- ing, turn orange-red, migrate into clusters, then hibernate
come webbed together. Short days may induce diapause in under ground litter. The photoperiod that induces diapause
spider mites, causing them to migrate and cluster together will differ in mite populations from different latitudes. Warm
25
26 Hemp D i s?ases
~ and Pests

temperatures inhibit diapause (Fig 4.2), and food availabil- Table 4.1: Infestation Severity Index (ISI) for the
ity may play a factor. Cool temperatures also induce hiber- two-spotted spider mite
nation in outdoor populations.
T. i~rticaeis common in North American and European Light any mites seen
glasshouses. It attacks outdoor crops in temperate climates, often no symptoms
and infests fruit trees as far north as Canada. Frank &
Rosenthal (1978) incorrectly stated this species will not in- Moderate <5 miteslleaf (not leaflet)
fest female flowers. feeding patches present
2. CARMINE SPIDER MITE
Heavy >5 mitestleaf
Tetra~zychuscinnabarinus (Boisdural) 1867, Acari; Tetranychidae. feeding patches coalescing
=Tetmn!jchus telarius (Linnaeus) 1758
Description: Eggs and immature stages of T. cinnnbnrinus
closely resemble those of T urticne. Adults, however, become plum
Critical 225 miteslleaf
red to brick red, with dark internal markings. But in cooler climates, shrivelled leaves and webbing
adults turn green, and become difficult to distinguish from two-spot-
ted spider mites. Adult male's aedeagi are not always symmetrical,
having a rounded anterior side and a sharp posterior side.
season. Spider mites can be carried into growrooms on
Life History & Host Range plants, people, and pets; they can even float on air currents.
The life history of T. cinnabarinus is similar to that of T. Glasshouses should be surrounded by a weed-free zone at
iirticae, especially in its fecundity. Its geographic range dif- least 3 m wide. Chickweed (Stellaria species) is an important
fers, because T.cinnabarinus prefers higher temperatures- weed host (Howard et al. 1994). For growers working with
35°C and above-so it thrives in semitropical areas. In cool vegetative clones, infested mother plants are the most
climates the pest is limited to hot glasshouses. High humid- common source of new mite infestations. Once mites have
ity causes all stages (larvae, nymphs, adults) to stop feeding infested your growroom or glasshouse, you will never get
and enter a quiescent period. Hussey &Scopes (1985) called rid of them without removing everything from the space
T. cinnabnrilzlls "the hypertoxic mite" and considered it more and disinfesting the place with steam heat or pesticides.
dangerous than T. iirticae. Cherian (1932) claimed the car- Method 5 (genetic resistance) is a future goal-select plants
mine spider mite is attracted to female flowering tops. He that survive heavy mite infestations. Cherian (1932)
noted plants yielding the largest flowers were the most heav- compared six varieties of Indian ganja for resistance to T.
ily infested. cinnabarinus; the most resistant varieties produced the
smallest female flowers. Regev & Cone (1975)compared five
varieties of hops for resistance to T.iirticae; the most resistant
varieties produced the least amount of farnesol, a
sesquiterpene alcohol that is also produced by Cannabis
(Turner et a/. 1980).

BIOLOGICAL CONTROL (see Chapter 10)

Biocontrol sho~ildbe established beforespider mite popiilations
explode. If mite populations balloon, biocontrols never catch
up. A mixture of three biocontrols, Phytoseil~liispersimilis,
Neoseitilt~scaliforniciis, and Mesoseililtis longipes, provides
excellent biocontrol for most glasshouses. These predatory
mites are described below. For unique situations and out-
door crops, Galendromzis occidentalis, Galendromlis pyri, and
Neoseiiiliis fallacis are also described below. Other predatory
mites eat spider mites in the absence of their primary hosts
(see Neoseii~liiscl~c~imeris and lphiseii~sdegenerans, described
under thrips).
Spider mite destroyers (Stetkori~spicipes, described be-
Figure 4.2: Diapause response curves in Tetranychus urticae low) and midge maggots (Feltiella acarisuga, described be-
from Holland-the effect of temperature on photoperiod (Mc- low) are predatory insects that prefer eating spider mites.
Partland modified from Helle 1962). General predators include lacewings (Chrysoperla carnea,
described under aphids), mirid bugs (Macrolophiis mliginosus,
under whiteflies; Deraeocoris breuis, under thrips), pirate bugs
(Oriiis species, under thrips), lygaeid bugs (Geocoris piinctipes,
DIFFERENTIAL DIAGNOSIS under whiteflies), a n d predatory thrips (Aeolothrips
Damage by other mites can be confused with spider mite intermediiis, under thrips).
injury. See the next section. Early aphid damage, sudden fun- Two fungi have been used to control spider mites.
gal wilts, and nutrient deficiencies may be confused with Neozygitesfloridana is being developed for commercial use,
symptoms from spider mites. Late-season hemp borers and and Hirslitella thompsonii (Mycar'a)was previously registered
assorted budworms hide in webbing that is mite-like. Find in the USA. These biocontrols require high humidity. They
the mites for a positive diagnosis. work well against mites in humid vegetative propagation
chambers.
CULTURAL & MECHANICAL CONTROL Biocontrol of mites nzlist be aclrieved before flowering has
(method numbers refer to Chapter 9) begun. Gaining biological control after tlrephotoperiod drops below
Method 1 (sanitation) must be observed every growing 12 hours per day is nearly impossible (Watson, pers. commun.
 Chapter 4: Insects and Mites 27

1999). Diapausing spider mites cluster by the thousands; at spider mites. Of course, flowering plants cannot be misted,
that point, many biocontrols stop working (especially preda- because they may mould.
tory mites). NOTES: P, persirnilis is the most popular mail-order mite.
It does best on low-growing bushy plants, where plants touch
Phytoseitrltis persimilis "P-squared" each other in a closed canopy (Helle & Sabelis 1985, vol1B).
BIOLOGY: A predatory mite that feeds on Tetranychtls I? persirnilis remains active in high humidity, works in a wide
spider mites, and is native to subtropical regions. Its eggs range of temperatures, and reproduces fast. But the preda-
survive best in high humidity (70-95% RH). The adults work tors avoid areas of high temperature, such as flowering tops
best at moderate temperatures (20-30°C) and >70% RH. basking in bright light. The effectiveness of P, persirnilis de-
Reproduction stops <60% RH (eggs stop hatching), and creases when Cannabis begins flowering, because the preda-
feeding stops <30% RH. A related species from Florida, tors find it difficult to move across sticky flower resins. Fur-
Plzytoseiulus macropilis, is also becoming commercially thermore, their need for high humidity is not compatible with
available. mould-susceptible Cannabis flowers.
APPEARANCE: Adult mites are orange-red in colour, P. persinzilis can annihilate a pest population. But the vo-
pear-shaped or droplet-shaped, 0.5-0.7 mm long, with long racious predators subsequently die out themselves, because
legs and no spots (Plate 4). Compared to the spider mite, P. they have no alternative food source. Thus, P. persirnilis is
persitnilis is slightly larger, more elongate, and moves quicker. often combined with Neoseil~li~s californiclis and Mesosei~ilus
P. persirnilis eggs are oblong and twice the size of spider mite longipes, because these mites live longer without food. P.
eggs. persimilis is also compatible with Bt, Feltiella acarisliga, and
DEVELOPMENT There are five stages-eggs, six-legged most parasitic wasps. I? persimilis can coexist with Neoseililt~s
larvae, eight-legged protonymphs, deutonymphs, and ct~cllrnerisbut the two biocontrols prey upon each other in
adults. After the larval stage P. persirnilis feeds continuously. the absence of their primary hosts (Hussey & Scopes 1985).
The life cycle takes seven days in optimal conditions (Fig P. persirnilis has been mixed with lacewings (Chrysoperla
4.3).Adults live another 3 0 4 0 days in the lab, but less in the species), but lacewings may eat predatory mites. Avoid most
insecticides, miticides and even fungicides while utilizing I?
persimilis (Table 10.1). According to van Lenteren & Woets
(1988), some strains of P. persirnilis tolerate some miticides
T, lirticae (sulphur and fenbutatin oxide) and some insecticides
(diazinon and malathion). Benomyl sterilizes predatory mites
s o they stop reproducing. Allow previously applied
pesticides to break down for two or three weeks before
introducing predators.
P.persitnilis Neoseiulus (Amblyseius) califomicus
BIOLOGY:A predatory mite that eats spider mites, broad
mites, and pollen, is native to southern California and Florida,
and does best in moderate humidity (260% RH) and
moderate to high temperatures (18-35°C).
APPEARANCE & DEVELOPMENT: Adult mites are
Figure 4.3: A Monthlo'Mites. At 21°C, predatory mites droplet-shaped and translucent to beige in colour, depending
(Phytoseiulus persimilis) reproduce every 7 days, twice as on what they eat. The life cycle may take only six days in
fast as spider mites (Tetranychus urticae), leading to a
large buildup of predators (McPartland redrawn from
Olkowski et al. 1991). Table 4.2: I? persimilis release rate for control of
two-spotted spider mites.
field. Adults consume up to 24 immature spider mites or
30 eggs per day. Within a week of hatching, females start
laying four or five eggs per day, for a total of 60 eggs per I Is*
NUMBER OF PREDATORS RELEASED PER
M~ OF GLASSHOUSE CROP** I
lifetime-they convert 70% of ingested food into eggs, and
produce a daily egg biomass equal to their own body weight Preventative 5 m-2every 3 weeks
(Sabelis & Janssen 1994). P. persirnilis, unlike some
phytoseiid mites, does not enter diapause in the presence Light 10 m.2 initial release, then 5 m'2 every
of cool temperatures and short day lengths.
APPLICATION: Supplied as adults and nymphs mixed
with inert materials (vermiculite, bran, corn cob grit, etc.) I Moderate 25 m-2on trouble spots, 10 m-2else- (
in pour-top bottles or tubes. Some are shipped on bean where, then 5 m-2every 3 weeks
leaves in plastic tubs. As with all predatory mites, P.
persinzilis should be applied immediately; store only if I Heavy 200 m-2on trouble spots, 25 m.2 I
necessary. Store for a maximum of four days in a cool (8- elsewhere, then 10 m-2every
3 weeks
1OoC), dark place. P. persirnilis can be sprinkled into
distribution boxes hanging on plants (Plate 85), or sprin-
Critical mechanical & chemical control of
kled directly on plants, the closer to spider mites the better.
trouble spots, 25 m" elsewhere, then
Recommended release rates are presented in Table 4.2.
Increase rates when plants are taller than 1m, planted more
10 m-2 every 3 weeks
densely than six per m2, or when humidity is low (<45%
* IS1 = Infestation Severity Index of two-spotted spider mite, see
RH). Encourage P.persirnilis by misting plants with water Table 4.1.
during periods of low humidity Misting also discourages **effectivenessof P persirnilis decreases during flowering.
28 Hemp Diseases and Pests

optimal conditions. Females lay three eggs per day for up to NOTES: Strong & Croft (1996) used N.fallacis to control
14 days. Adults eat five to 15 adult spider mites per day, T.ilrticaeinfesting hops in Oregon, released at rates of 125,000
along with some eggs a n d larvae. New strains of N . per ha (50,000 per acre). N. fallacis migrates vigorously
c a l f o r n i c ~ ~d so not diapause in short days (Ravensberg, pers. across plants of all sizes, and uses aerial dispersal to blow
commun. 1999). from plant to plant. It is probably compatible with other
APPLICATION: Supplied as adults and nymphs mixed Neoseiulus species. N . fallacis tolerates more pesticides than
with vermiculite or sawdust in pour-top bottles. Store one any other predatory mite. Some strains are compatible with
to four days in a cool (8-10°C), dark place. N . calfornicus neem, sulphur, abamectin, and pyrethrin (and the synthetic
migrates fairly well a n d has been used outdoors. N . pesticides malathion, dicofol, and propargite).
cillifornict~stolerates lower humidity and lives longer without
prey than P. persinzilis, but N , calfornicus feeds slower than Galendromus (Metaseiulus) occidentalis
P. persinzilis and its populations build slower. These attributes BIOLOGY: A predatory mite that eats spider mites and
make N. calqornic~~s an excellent preventative, and fine for some russet mites, and is native to western North America.
treating light infestations (same release rates as P, persirnilis). It tolerates a range of humidities (40-80% RH) and
But for heavy or critical infestations, switch to P. persirnilis. temperatures (26-35°C). G. occidentalis has been used in
NOTES: N . californici~sis compatible with I? persinzilis and Washington State apple orchards since 1962.
all biocontrols compatible with P. persirnilis. N. calfornicus APPEARANCE & DEVELOPMENT: G . occidentalis
tolerates more insecticides than P. persinzilis, but is still resembles N . fallacis. The life cycle takes seven to 14 days
susceptible to most pesticides (even benomyl, a fungicide). depending on temperature. Adults eat one to three spider
mites per day or six mite eggs per day. Many strains of G.
Mesoseiulus (Phytoseiulus) longipes occidentalis go into diapause with short days and cold
BIOLOGY: ~ ~ r e d a t o mite
r y that feeds on many mites, temperatures (Helle & Sabelis 1985, vol lB), so their
is native to South Africa, and tolerates conditions that are effectiveness ends when Cannabis begins flowering.
warm (up to 3B°C) and dry (as low as 40% RH at 21°C). APPLICATION: Supplied as adults mixed in pour-top
APPEARANCE & DEVELOPMENT: Adults resemble P. bottles. In the Pacific Northwest, G. occidentalis can also be
persitnilis, as do their eggs. The life cycle can turn in seven obtained from nearby orchards by collecting apple leaves.
days at optimal temperatures. Female mites lay three or four Store a maximum of five days in a cool (8-10°C), dark place.
eggs per day (Sabelis & Janssen 1994),and live up to 34 days. On field crops, release 12,500 per ha (5000 per acre) at the
Short photoperiods may send M. longipes into diapause. first sign of pests, repeat every two weeks. Release 50 per
APPLICATION: Supplied as adults in pour-top bottles. m2 every two weeks during indoor infestations.
Adults should be used immediately upon delivery but can NOTES: G. occidentalis eats less than P. persinzilis and re-
be stored in a cool (6-10°C), dark place for a couple of days. produces slower, but lives longer without food, disperses
Release at the same rate as P, persInzilis. more rapidly (including aerial dispersal), and tolerates
NOTES: M. longipes thrives in temperatures that wilt P. semiarid conditions (it does not tolerate high humidity). G.
peusinzilis or even N . californicus, and it tolerates lower occidentalis does well on both short and tall plants, and has
humidities. It migrates better than many mites, making it been used outdoors. It is compatible with Bt and Galendmrnils
suitable for taller plants. M , longipes reproduces slower than pyri in apple orchards and hops yards. A new strain of G.
P. persimilis. M. longipes is compatible with the same occidentalis is photoperiod neutral (nondiapausing), so it can
biocontrols as P. persirnilis. Avoid insecticides, though some be used while plants are flowering. G. occidentalis tolerates
strains tolerate malathion. pesticides better than most predator mites; some strains
tolerate sulphur, pyrethroids, abamectin, and carbaryl.
Neoseiulus (Amblyseius)fallacis Release mites at least a week after spraying. G. occidentalis is
BIOLOGY: A predatory mite that feeds on spider mites, a small predator, so some T. z~rticaeadults may be too big for
red mites, and russet mites, and survives on pollen in the G. occidentalis to handle.
absence of prey. It is native to the Northwestern USA, and
does best in moderate to high humidity (60-90% RH) and Galendromus (Typhlodromtis) pyri
moderate temperatures (optimally 10-27"C, but up to 38°C BIOLOGY: A generalist predatory mite that feeds on
in high humidity). spider mites. It occurs naturally in temperate apple orchards,
APPEARANCE: Adults are vear-shaped and white until and does best in cooler, humid climates.
they feed, when they take on the colour of their prey, usually APPEARANCE & DEVELOPMENT: Adults are indistin-
pale red or brown. They are slightly - .
smaller than other guishable from the aforementioned species. G . pyri
predators, such as P. persimilis. reproduces slowly, taking nearly two weeks to mature, and
DEVELOPMENT: Adults overwinter in crevices of tree lays only one egg per day (Sabelis & Janssen 1994). Short
bark, but not as successfully as Galendrornt~spyri (described days send it into diapause.
below). The life cycle takes ten days in optimal conditions. APPLICATION: Supplied as adults in bottles or tubes.
Adults live another 20-60 days (four to six generations arise Store up to five days in a cool (8-10°C), dark place. Release
per year). Adult females eat about 15 mites per day, and lay the same rate as G. occidentalis.
a total of 40-60 eggs. Short days and cold temperatures NOTES: G. pjri overwinters better than G. occidentalis and
interact to send N. fallacis into diapause. Diapause is delayed other predatory mites (Helle & Sabelis 1985, vol 1B). Its
by warmer temperatures, and completely averted above evasive behaviours also enhance outdoor survival. G. pyri
27°C (Helle & Sabelis 1985, vol 1B). does best on medium-sized to tall plants. It is compatible
APPLICATION: Supplied as adults in bottles, or a mix of with Bt and Galendrornz~soccidentalis in apple orchards. A
adults and nymphs on bean leaves in plastic tubs. Store a New Zealand strain is resistant to pyrethroids.
maximum of two or three days in a cool (8-10°C), dark place.
Used preventively, release ten mites per m2, every two weeks, Stethorus species "spider mite destroyers"
two or three times. For light to moderate infestations, release BIOLOGY: Three species of tiny ladybeetles (S.
20-40 mites per m2 weekly until controlled. punctilh~rn,S. pllnctzlrn, S.picipes) prey on spider mites and
 Chapter 4: Insects and Mites 29

their eggs, as well as red mites, aphids, whiteflies, and other mites (Helle & Sabelis 1985, vol. lB), or they may not (Cherim
soft-bodied insects. All three species are temperate Ameri- 1998).Adults tolerate organophosphates and juvenile growth
cans and d o best in moderate humidity (60% RH) and tem- hormones (e.g., fenoxycarb, teflubenzuron), but larvae are
peratures (26-35OC). Other Stethorus species are found in seriously impacted by insecticides (Biddinger & Hull 1995).
tropical regions.
APPEARANCE: Adults are oval, black, shiny, pubescent, Feltiella acarisuga (Therodiplosis persicae)
1.5-3 mm long, with brown or yellow legs (Fig 4.4). Larvae BIOLOGY: Acecidomyiid gall midge similar to Aphidoletes
are cylindrical, dark grey to black (turning reddish before aphidimyza, the aphid predator. F, acarisuga feeds on all spider
pupation), covered with fine pale-yellow spines, 1-2 mm mites, including T. urticae and T,cinnabarinzls.Adult midges
long. Eggs are oval, <0.5 m m long, initially white but turn- d o best in >60% RH and 20-27"C, but the predatory larvae
ing grey before hatching, usually laid singly on undersides tolerate a wider range of conditions. The species is called T.
of leaves near veins. persicae by some suppliers, but the correct name is F.acarisuga
DEVELOPMENT: Adults overwinter in leaf litter on the (Gagne 1995).Arelated species, Feltiella occidentalis, is also
ground, and emerge when apple trees are blooming. They being developed.
lay eggs from May to mid-August. Females lay up to 750 APPEARANCE & DEVELOPMENT: Adults look like tiny
eggs in the presence of adequate prey. Larvae feed on eggs pink-brown mosquitos, with a wingspan of 1.6-3.2 mm.
and young spider mites before pupating in foliage. The life Adults only live three or four days, long enough for females
cycle takes 20-40 days. Adults live another 30-40 days. to lay 30 eggs on plants near spider mite colonies. Eggs are
Adults consume all stages of mites, up to 75-100 per day. tiny, oblong, and shiny-yellow. Hatching larvae are slug-like
One to three generations arise annually in temperate regions. and grow to 1.7-2.0 m m long. They are yellow, cream-
Short days may send S. picipes into diapause (Helle & Sabelis coloured, orange, or red; their colour depends on body
1985, vol. 1B). contents. Larvae sink their mandibles into all stages of spider
APPLICATION: Supplied as adults in pour-top bottles. mites, from eggs to adults. They feed on 30 nymphs and
Store up to a week in a cool (8-1O0C), dark place. Used adults per day or as many as 80 mite eggs per day (Gagne
preventively, release one or two beetles per m2 every month. 1995). In one week larvae stop eating, they spin white co-
For light to moderate infestations, release three or four beetles coons, and pupate on undersides of leaves. The entire life
per m2 every two weeks. For heavy infestations, double the cycle takes two to four weeks.
release rate. APPLICATION: Supplied as pupae on leaves or on pa-
NOTES: Stetlzort~sbeetles are good fliers and find infes- per. Store a maximum of one or two days in a cool (10-15"C),
tations easily. They feed greedily but migrate before their dark place. Sprinkle pupae on soil or in open containers
job is finished. To reduce migration, use strategies described attached to plants; protect against direct sunlight. Release
under Hippodamia convergens (in the aphid section). Dense rates for preventative control range from one pupa per m2 to
mite webbing impedes newly-hatched Stethort~slarvae (Helle 1000 pupae per ha. For moderate infestations apply five
& Sabelis 1985, vol. 1B).Stethorus species may eat predatory pupae per m2 per week or 7500 pupae per ha per week.

Figure 4.4: Assorted beneficial ladybug beetles (all x6). A,B,C. Adult, pupa, larva of convergent lady beetle (Hippodamia
convergens); D. Spider mite destroyer (Stethorus picipes); E. Seven-spotted ladybeetle (Coccinella septempunctata);
F & G. Adult & larva of vedalia (Rodolia cardinalis); H & J. Adult & larva of mealybug destroyer (Cryptolaemus
montrouzier~);K. Two-spotted lady beetle (Adalia bipunctata) (Courtesy USDA except H & J, McPartland).
30 Hemp Diseases and Pests

NOTES: Some strains of this "flying predator" d o not and feeds on all kinds of Cannabis, including European fibre
diapause, a welcome advantage in flowering crops. The cultivars, southeast Asian drug landraces, Afghan landraces,
larvae move well across heavily-trichomed leaf surfaces. F. and ruderals (Hillig, unpublished data 1994).The mite popu-
acarisz~galarvae eat more spider mites than P. persirnilis, but lation at I.U. was possibly imported on seeds from Nepal or
reproduce slower. The two biocontrols are compatible. F. northern India.
acarisilga tolerates sulphur pesticides. In Europe F. acarisi~ga
can be parasitized by Apkanognii~swasps, which hamper its SIGNS & SYMPTOMS
effectiveness (Gagne 1995). A . cannabicola feeds primarily on petioles and leaflets.
Leaflets curl at the edges, followed by chlorosis and necro-
BIORATIONAL CHEMICAL CONTROL (see Chapter 11) sis. Petioles become brittle and leaflets break off easily. In
Use chemicals for "spot treatment" of heavy mite infes- bad infestations the mites crowd plants by the thousands,
tations. Heavy infestations arise on stressed plants, on plants giving leaflets a beige appearance. The mites may also in-
located near glasshouse openings, and ~ n - ~ l a ngrowing
ts fest flowering tops; they selectively feed on pistils, render-
along the windward edge of fields. Direct all sprays at the ing female flowers sterile.
undersides of leaves. Since mite eggs arenot harmed by many
chemicals, repeat the treatment a week later. After every har- TAXONOMY & DESCRIPTION
vest in glasshouses or growrooms, spray all walls, floors, Aclilops (Vasates) cannabicola (Farkas) 1965, Acari; Eriophyidae.
ceilings, and equipment (pots, tools, etc.). Of course, strong Description:Eriophyid mites are soft-bodied, sausage-shaped,
pesticides must subsequently be washed off, to prevent and exceptionally tiny, with only two pairs of legs (Fig 4.5 & Plate
residues from harming biocontrols in the next crop. For grow- 5). Their pale beige bodies are composed of two sections: the
ers working" with clones, chemicals are recommended thrice: gnathosoma (mouthparts)and idiosoma (restof the body).The legs
on mother plants before clones are cut, onclones several days project from around the gnathosoma, near the front. The idiosoma
is covered with many minute transverse ridges and tiny spines. A.
after transplanting, and again on clones the day before flow- cannabicola females reach 200 pm (0.2mm) long and 45 pm wide-
ering is induced. less than half the size of Tetranychus urticae.
Horticulture oil, Safer's Soap, clay microparticles, and
even tap water are mildly effective, and described in Chapter Life History
11. Cherian (1932) killed spider mites with either a lime- We know little about A. cannabicola's life cycle. Outdoor
sulphur spray or fish oil soap. "Ganja" sprayed with lime- populations probably overwinter in contaminated seed. In-
sulphur "...was tested by veteran smokers who gave their door populations remain on plants year round. The mites
verdict against it," whereas ganja sprayed with soap passed move towards the top of dyingplants, where they spread to
the smoker's test. Pure sulphur (not lime-sulphur) may work, other plants by wind or splashing water. A turn of the life
but mite populations started developing resistance to sulphur cycle takes about 30 days under optimum conditions of 27°C
90 years ago (Parker 1913b).Rosenthal(1998) sprayed mites and 70% RH. Related eriophyid mites lay ten to 50 eggs dur-
with sodium hypochlorite (household bleach diluted to 5%). ing a life-span of 20-40 days.
Bush Doctor (198613) sprayed mites with a flour slurry, A related pest, Aciilops lycopersici (Tyron), the tomato
strained through cheesecloth. Parker (1913b) used a flour russet mite, vectors viruses. A. lycopersici goes into a feeding
slurry against T.i~rticaein hops, mixing 3.6 kg flour in 378 1 frenzy called "solanum stimulation," where it feeds until it
water (8 lbs/100 gallons). Flour solutions require frequent kills its own host (Lindquist et al. 1996).
agitation to keep sprayers from clogging.
Turning to botanicals, Parker (1913b) killed 99% of two- DIFFERENTIAL DIAGNOSIS
spotted spider mites with nicotine sulphate, mixing 190 ml The hemp russet mite's morphology, colouring, and lack
of 30% concentrate nicotine sulphate in 378 1water (6.5 oz/ of webbing make it easy to discern from spider mites.
100 gallons). T.cinnabarinus is highly susceptible to neem
seed extracts, u p to 58 times more susceptible than its CULTURAL & MECHANICAL CONTROL (see Chapter 9)
predator, P.persimilis (Mordue & Blackwell 1993). Use clean seed to keep mites out-see method 11. Once
Cinnamaldehyde, extracted from cinnamon mites infest an area, a grower's best efforts only decrease
(Cinnamonuni zeylanic~~nz), kills all stages of spider mites, in- populations, not eliminate them. Method 5 (genetic resist-
cluding eggs. B& it also kills beneficial mites and insects. ance) is a future goal.
Ironically, European researchers have controlled spider mites
with leaf extracts of Cannabis sativa (Fenili & Pegazzano 1974). BIOCONTROL (see Chapter 10)
Pyrethrum works (Frank 1988),but synthetic pyrethroids (such No effective biocontrol of A . cannabicola is known. Phyto-
as permethrin) rarely kill mites and actually induce egg seiultis persin~ilisdoes not feed on A. cannabicola (Hillig, un-
laying. Imidacloprid (a nicotine derivative) and abamectin published data 1994). Other russet mites are controlled by
(a fermentation product) kill spider mites. Zetzellia mali (a tiny predator mite not commercially avail-
Synthetic insect growth hormones, such as flucyclozuron able). Some Aciilops mites are susceptible to Verticillium lecanii
and methoprene, kill immature spider mites. Hexythiazox (Olkowski et al. 1991), a biocontrol fungus described under
is a growth hormone that selectively kills mites. A synthetic whiteflies. Two other possibilities are described below:
pheromone, trimethyl docecatriene (StirrupMm),attracts spi-
der mites. It markedly enhances the effectiveness of miticides. Homeopronematus anconai
Kac (1976) tested 25 synthetic and systemic pesticides for BIOLOGY: Royalty & Perring (1987) controlled Aculops
controlling mites in Slovenian hemp (see the appendix). lycopersici with this predatory tydeid mite. Royalty & Perring
found H. anconai resistant to abamectin, so the predator and
pesticide could be used together. H. anconai may prey on
HEMP RUSSET MITE Phytoseililt~spersimilis, so their compatibility is questionable.
Hemp russet mites have infested hemp in central Eu-
rope (Farkas 1965) and feral hemp in Kansas (Hartowicz et Hirsutella thompsonii
al. 1971).At Indiana University, the pest thrives in glasshouses BIOLOGY: H. thornpsonii infests several eriophyid and
 Chapter 4: Insects and Mites 31

tetranychid mites. The fungus was released in Cuba to con- faces. Upper leaf surfaces of infested plants turn yellow, then
trol an Actllops species, and sprayed in Florida citrus groves red-brown, then die. Feeding damage peaks in April-June
against other eriophyid mites (Lindquist et al. 1996). H. and September-November. The life cycle can be as short as
tlioinpsoriil does best in hot, humid conditions. two weeks. E. orientalis normally attacks Citrus species but
APPEARANCE & DEVELOPMENT Conidia quickly ger- infests a wide variety of hosts, including Cannabis in India
minate, directly infect mites, and kill them in one or two (Dhooria 1983, Gupta SK 1985).Oriental mites live in north-
weeks. Fungal hyphae emerge from dead mites. The hyphae ern Africa, the middle East, India, and southeast China.
bear solitary phialidic conidiophores and slimy, round-to-
lemon-shaped conidia. Under optimal conditions, mite ca- 2. PRIVET MITES
davers sprout long, white, hairlike synnemata, which bear a. Brevipalpils obovattts Donnadieu 1875, Acari; Tenuipalpidae.
conidiophores and conidia. b. Brevipalpus riiglllosils Chaudri, Akbar & Rasool 1974
APPLICATION: H. thoinpsonii is mass-produced in liq- Description: Eggs are bright red, 0.1 mm long. Larvae and
uid media fermenters, and supplied as conidia (109 conidia nymphs are bright red. Adults are oval in outline, average 0.3 mm
per g). Release rate for citrus and turf is 2.24.5 kg ha-1 (2-4 long and vary from light orange to dark red in colour (Fig 4.5).
lbs/acre) in optimal weather conditions (Lindquist et al.
1996). This fungus was previously sold in the USA (as Life History & Host Range
Mycar"), but Abbott Labs cancelled its registration in 1988. B. obovatus is a polyphagous pest found in temperate
zones around the world. B. r~lgulostlsis restricted to the In-
CHEMICAL CONTROL (see Chapter 11) dian subcontinent. Both species attacked Cannabis in India
Spraying oil or sulphur works against some russet mites, (Gupta SK 1985). Their toxic saliva causes severe leaf spot-
but not A. cannabicola (Hillig, unpublished data 1994). ting, on upper and lower leaf surfaces. Female privet mites
Cinnamaldehyde, a new product extracted from cinnamon lay eggs on undersides of leaves, one at a time. They
(Cirznanzorzuin zeylanicilnz), may be effective against russet overwinter as adults and their life cycle takes about 60 days.
mites. Hillig used abamectin (avermectin Bl). Lindquist et
al. (1996) used abamectin against Acillops lycopersici; 3. TA MA MITE
abamectin killed more russet mites than dicofol, yet spared Typhlodromus cannabis Ke & Xin 1983, Acari; Phytoseiidae.
the biocontrol mite Hoineopror~ematzlsanconai. Hexythiazox Ke & Xin (1983) found this new species on Chinese
is a growth hormone that selectively kills immature mites. hemp. The mite also infested horsetail rush (Pteridium
Hillig (unpublished data 1994) killed A. cantzabicola by en- aqtlilinunz), clematis (Clematis species) and Popt11i~sspecies.
closing potted plants in large plastic bags and filling the bags No plant symptoms or control measures were described.
with C 0 2for two hours. Spider mites and thrips can also be
killed with 100% C02,but require at least 12 hours of expo- DIFFERENTIAL DIAGNOSIS
sure. Take care while using C02 in growrooms and other These other mites can be confused with spider mites,
sealed spaces. described earlier.

BIOLOGICAL & CHEMICAL CONTROL

Damage is greatest in drought-stressed plants (Hill
1983).For oriental mites, Dhooria (1983) cited two biocontrol
organisms-a predator mite (Amblyseiusalstoniae Gupta) and
a thrips (Scolotkrips indicns Priesn.). Neoseizilzls californicils
feeds on E. orientalis but does better against mites that pro-
duce heavier webbing, such as spider mites (described there).
According to Ravensberg (pers. commun. 1998), privet
mites can be controlled by Neoseililtis californictls and
Neoseitllus fallacis (described under spider mites), and
Figure 4.5: Other mites. A. Aculops cannabicola (from Neoseillllis cilczlmeris and Neoseizllils barkeri (described under
Farkas 1965); B. Eutetranychus orientalis (from Hill 1994), thrips). Privet mites are susceptible to sulphur but resistant
C. Brevipalpus obovatus (drawings not to scale). to organophosphate and carbamate pesticides (Hill 1983).Try
soap or horticultural oil.

OTHER MITES APHIDS

Researchers cite other mites attacking Cannabis: the ori-
ental mite, two species of privet mites, and the ta ma mite. Aphids, or "plant lice," are tiny, soft-bodied, and pear-
Additionally, many species of mites parasitize hemp seeds shaped. They have relatively long legs and antennae. Some
in storage (see Chapter 8). species have antenna1 tubercles, important landmarks in pest
identification. Antenna1 tubercles are paired "bumps on the
1. ORIENTAL MITE head," located between antennae (see Fig 4.8). Protrusions
Eiitetranychils orientalis (McGregor) 1914, Acari; Tetranychidae. from the rear of aphids also serve in pest identification: A
=Ez~tetranychusorientalis (Klein)1936 pair of tubelike comicles (or siphons) project backwards,
Description: Eggs are subspherical,pale brown, 0.14 mm diametre. they resemble dual tailpipes. Between the cornicles, the rear
Larvae are light brown and SIX-legged.Nymphs emerge from moults end tapers to a pointed, tail-like caudum.
with eight legs, like adults. Female E. orientalis adults are round- Most adult aphids d o not have wings (these adults are
bodied, greenish-brown, up to 0.5 mm long, and covered with short called apterae), but some d o (alatae). The wings of alatae
spines (Fig4.5). Males are smaller and more red than females. are much longer than their bodies.
Aphids suck sap from a plant's vascular system, using
Life History & Host Range long narrow stylets. Most aphids are phloem-feeders, but
Females lay eggs along main veins on upper leaf sur- some also suck on xylem (Hill 1994). Besides sucking sap,
32 Hemp Disearses and Pests

aphids damage plants by vectoring fungi, bacteria, and es- ent. Fundatrices multiply parthenogenically-no sex needed.
pecially viruses. Viruses and aphids have a symbiotic rela- They are born fertile and within days begin giving birth.
tionship (Kennedy p t al. 1959). They give birth viviparously-eggs hatch within their re-
At least six aphid species attack Cannabis. Aphid dam- productive tract-so larvae, not eggs, are "born alive" (an-
age increases in warm, moist weather, with gentle rain and other discovery by Leeuwenhoek). Fundatrices bear 60-100
little wind. Damage decreases in hot, dry weather and in the fundatrigeniae. Soon the fundatrigeniae-which are apter-
presence of strong, dry winds (Parker 1913a). ous (wingless) females-begin giving birth to more live fe-
males (apterous viviparae or apterae). Thus stem mothers
SIGNS & SYMPTOMS may live to see their great-great-great-granddaughters. In
Aphids congregate on the undersides of leaflets and late spring the first alatae (winged aphids) develop. They
cause yellowing and wilting. Some aphid species prefer older, are called spring migrants and fly off to secondary hosts,
lower leaves (e.g.,Myzlis persicae), and some prefer younger, such ascannabis.
upper leaves (e.g.,Aphisfabae). Some species even infest flow- Aphids are weak fliers, with a flight speed of only 1.6-
ering tops (e.g., Pkorodon Izumuli, Plzorodon cannabis). Early 3.2 km h-1 in still air. To migrate, alatae fly straight up into a
damage is hard to detect-the undersides of leaflets develop moving air mass. The wind governs their direction of flight.
light-coloured spots, especially near leaflet veins (look closely In this way they can migrate 15-20 km. Aphids terminate
at Plate 8). Eventually, leaves and flowers become puckered migration by actively flying downward and settling on
and distorted (Kirchner 1906). Heavily infested plants may plants. Once settled on Cannabis the alatae give birth to
completely wilt and die. Surviving plants remain stunted. apterae; apterae undergo four moults in about ten days to
Honeydew exudes from the anus of feeding aphids. On reach sexual maturity. Each aptera gives birth to 30-70 young.
warm afternoons honeydew may be seen falling as a mist Crowded conditions or a lack of food induce new alatae,
from severely infested plants (Parker 1913a). Honeydew which fly off seeking unexploited Cannabis. They are called
causes secondary problems-ants eat it, and sooty mould summer migrants. At the end of the summer, special alatae
grows on it. For the love of honeydew, ants become "pugna- called sexuparae (or autumn migrants or return migrants)
cious bodyguards" of aphids, and attack aphid predators. fly back to the primary host and give birth to ten sexuales.
Ants must be eliminated for biocontrol to be effective. Sooty Sexuales are either females or males, alatous or apterous.
mould reduces plant photosynthesis and leaf transpiration, They mate and the females become oviparae and lay five to
and hinders the movement of aphid predators and parasites. ten eggs, which overwinter.
Hi11 (1994) claims that a single springtime fundatrix can
1.GREEN PEACH APHID give rise to up to 12 generations of aphids in one year-and
Myzlis persicae (Sulzer) 1776, Homoptera; Aphididae. theoretically 600,000 million offspring. In tropical regions and
=Phorodon persicae Sulzer in zuarm glasshol~ses,aphids do not migrate betzoeen hosts, tzor do
Description: Apterae are green (sometimes yellow-green or pink), they lay ouerzuintering eggs. They reprodme partheizogenically
oval in outline, averaging 2.0-3.4 mm in length (Plate 8). Antenna1 all year long, and remain on their secondaq host. Thus, some
tubercles are mammary-like bumps (less than half as long as the aphids that are normally heteroecious may become autoe-
first antenna1 segment), and point inward (converge towards each
cious in warmer climates. M. persicae, for instance,
other). Cornicles are the same colour as the abdomen, except for
darkened tips. Cornicles are long, thin, slightly swollen near the overwinters as adult females on secondary hosts in the south
midpoint, and grow twice as long as cauda. Cauda are lightly bris- or in warm glasshouses (Howard et al. 1994).
tled and constricted slightly at the midpoint. Alatae nymphs are of- M. persicae attacks dozens of plant species, and now
ten pink or red, and develop wing-pads. Winged adults have black- lives worldwide (Spaar et al. 1990).The pest is exceptionally
brown heads, a black spot in the middle of their abdomens (Fig 4.6 restless; alatae repeatedly land on plants, probe briefly, then
& Plate 6), and hold their wings in a vertical plane when at rest. take off for other plants. This behaviour makes M. persicae
"the most notorious vector of plant viruses" (Kennedy et al.
1959). It infests feral hemp in Illinois (Bush Doctor, unpub-
lished data 1981) and marijuana in India (Sekhon et al. 1979).
M ,persicae is the most common aphid in Dutch glasshouses
(Clarke &Watson, pers. commun. 1995).This species resists
a broader range of pesticides than any other insect pest.

2. BLACK BEAN APHID

Aphisfabae Scopoli 1763, Homoptera; Aphididae.
Description: Apterae are oval to pear-shaped, olive green to
dull black, averaging 1.5-3.1 mm in length (Fig 4.7 & Plate 7); legs
I I are light green to white; antennae tubercles not prominent; cornicles
relatively short (0.3-0.6 mm) and cylindrical; cauda are heavily bris-
Figure 4.6: Adult aphids, alatae (winged forms), mounted tled. Alatae nymphs have prominent white markings on their ab-
on a microscope slide and cleared with potassium hydrox- domen. Alatae are slightly smaller than apterae (1.3-2.6mm long),
ide. A. Myzus persicae; B. Phorodon cannabis, LM x 10. their bodies are dark green-black with variable white stripes, wings
are spotted with white wax. Alatae hold their wings vertically over
their abdomens when at rest. A. fabae colonies are regularly ant-
Life History & Host Range attended and they may congregate in great numbers.
The complicated life cycle of aphids requires special ter-
minology. Most outdoor aphids are heteroecious (or Life History & Host Range
holocyclic) -they migrate between t w o hosts. The A.fabae is heteroecious-it migrates between two hosts.
overwintering host is called the primary host. M. persicae The overwintering host is Euonymus a n d sometimes
overwinters on Prllnz~s,as shiny black eggs laid on tree limbs Virbllrnlinz species. In the spring, eggs hatch into fundatrices,
and under the axils of tree buds. Eggs hatch into rotund "stem then fundatrigeniae, then apterae (see the life history of M.
mothers" or fundatrices. Hatch time is temperature depend- persicae for these terms). Apterae give birth to spring mi-
 Chapter 4: Insects and Mites 33

at rest. Males are smaller and darker than females, averaging 1.6-
1.8 mm long. Overwintering eggs are ovate, shiny green-black, 0.7
mm long. Fundatrices are more oval than other apterae but slightly
shorter (1.8-2.4 mm), with long antennae.

Life History & Host Range

The life historv of P, cannabis is autoecious, sometimes
called monoecious-the pest never alternates hosts
(Balachowsky & Mesnil1935, Miiller & Karl 1976).Sexuparae
Figure 4.7: Aphis fabae. A. Adult aptera (wingless); of P, cannabis never fly away, so oviparous females lay eggs
B. Adult alata (winged); both x 10 (from lmms 1948, in the flowering tops of Catztzabis. Most eggs are destroyed
labelled "Aphis rumicis'~. when the hemp crop is harvested.
The bhang aphid is particularly damaging to female
grants, which migrate to Canrzabis and other secondary hosts. buds; "...it sits between female flowers and seeds, sucking
At the end of the summer, autumn migrants fly back to the plant sap" (Kirchner 1906).Bhang aphids vector hemp streak
overwintering host. virus (Goidanich 1955), hemp mosaic virus, hemp leaf chlo-
Blackman & Eastop (1984) listed Cartnabis as a host for rosis virus (Ceapoiu 1958), cucumber mosaic, hemp mottle
A. fabae. It is the second most common aphid in Dutch glass- virus, and alfalfa mosaic virus (Schmidt & Karl 1970). The
houses, and attacks outdoor crops in China ( ~ l a r k e& species is native to Eurasia (ranging from Britain to Japan),
Watson, unpublished data 1992) a n d South Africa and now lives in North America and northern Africa. Bantra
(Dippenaar et al. 1996). A. fabae may have been the "black (1976)suggested using the pest as a biocontrol agent against
aphid" infesting a seed-oil cultivar ('FIN-315') in Finland illicit marijuana crops. According to Miiller & Karl (1976),P.
(Callaway & Laakkonen 1996).The taxon A.fabae represents canrlabis infests C. sativa, C, itzdica and C. ruderalis. The pest
a complex of at least four subspecies; exactly which-subspe- also attacks hops (Blackman & Eastop 1984).
cies attacks Carznabis is unknown. Black bean aphids are According to Martelli (1940), bhang aphids are identi-
found in all temperate regions except Australia. hey infest cal to hops aphids (the next species described below). We
many crops and vector over 30 viruses. disagree-hops aphids have different-shaped heads, with
few bristles (Miiller & Karl 1976, Fig 4.8). The two species
also have different life histories-P. cannabis has reduced to
an autoecious life cycle (only one host), first noted by
T K T UT
Balachowsky & Mesnil (1935). Autoeciousness suggests P.
m_
. , / cannabis evolved from P. hlinzlili, not vice-versa.

4. HOPS APHID
Pllorodon h t ~ m l i l i(Schrank) 1801, Homoptera; Aphididae.
Description: Hops aphids closely resemble bhang aphids, ex-
T K T cept the head of I? humuli sports few bristles, if any (Miiller & Karl
1976, see Fig 4.8); the few bristles on antennal tubercles have blunt
or pointed apices, never knobbed; the midline knob is scarcely ap-
parent. Apterae are larger on Prunus in the spring (2.0-2.6 mm) than
apterae on Hunuulus in the summer (1.1-1.8 mm); alatae average
1.4-2.1 mm long.

Figure 4.8: Prominent head bristles on Phorodon can- Life History & Host Range
nabis (left) compared to Phorodon humuli (right), from P. htlnztlli is heteroecious; the overwintering host is
Miiller & Karl (1976)."T"arrows point to tubercles, " K points Prunlis. Eggs of P. cannabis hatch when the temperature
to midline knob. reaches 22-26°C. Eggs hatch into fundatrices, then
fundatrigeniae, then apterae, then spring migrants (see the
life history of M.persicae for these terms). In the California
3. BHANG APHID or HEMP LOUSE Bay area, spring migration of P. htlmuli peaks on June 1st
Phorodon cannabis (Passerini) 1860, Homoptera; Aphididae. (Parker 1913a); in southern England the flight peaks in late
=Aphis cannabis Passerini, = M y z u s cannabis (Passerini), June (Dixon 1985). P. h u m n l i migrants fly straight up into a
=Pavnphorodoncannabis (Passerini), =Diplzorodon cannabis (Passerini), moving air mass, and can travel 150 kmor more (Dixon 1985).
=Aphis satlzvze Williams 1911, =Plzorodon asacola Matsumura 1917,
=Capitophovus cannab~foliaeShillji 1924, ? =Serniaphoides cannabiarum They spend the summer on secondary hosts, then fly back to
Rusanova 1943 overwintering hosts. For I! liiinil~lithis autumn migration
Description: Apterne have flattened, elongate-oval bodies. They peaks in late September (Dixon 1985).
closely resemble M , pevsicne but are about 25% smaller, averaging Blunck (1920),Flachs (1936),and Eppler (1986) reported
1.9-2.7 mm in length. They are described as nearly colourless P. hunzlili infesting Cannabis. These pests normally attack
(Ceapoiu 1958) to bright green with darker green longitudinal stripes hops. P. h u m i l l i vectors many plant viruses a n d
(Kirclu~er1906).Their heads are covered with tiny bristles; the bris- Pset~doperonosporah u m i ~ l (a
i fungus causing downy mildew
tles have knobbed apices which differentiate bhang aphids from hops of hops and hemp, Sorauer 1958). P. htlmuli lives in Europe,
aphids (Miiller & Karl 1976, Fig 4.8). Antennae are 1.1-2.2 mm long;
central Asia, north Africa, and North America, but not Aus-
antenna1 tubercles are prominent (at least half as long as the first
antennal segment), converge slightly, and several bristles sprout tralia. In California P. h u m u l i is aided by a large black ant,
from each tubercle. Between the tubercles arises a smaller midline Formica stibsericea Say (Parker 1913a).
knob, also bristled. Cornicles are white, up to 0.8 mm long (nearly a
third of the body length), cylindrical, and taper towardstheir tips. 5. OTHER APHIDS
Cauda taper evenly to their tips. Alatae are slightly smaller than Cherian (1932) and Raychaudhuri (1985) cited the cot-
apterae, and develop black-brown patches on heads and abdomens ton aphid (also called the melon aphid, Aphisgossypii Glover
(Fig 4.6). They hold their wings vertically over their abdomens when 1877) on marijuana in India. A. gossypii adults are 1-2 mm
34 Hemp Diseases and Pests

Figure 4.1 0:
Life history of Myzus
persicae.

A. Fundatrix giving birth to

live fundatrigeniae;
B. Fundatrigenia shedding
skin of its last moult;
C. Aptera;
D. Alata (spring migrant);
E. Summer migrant;
F. Autumn migrant or
sexupara;
G. Sexuales (female on
left, male on right);
H. Egg-laying ovipara;
J. Egg;
[illustrations of
M. persicae (C-G)
by Hill 1994, other
illustrations by
Weber 19301.

long, have short, bristled cauda, and lack antennal tuber- confused with young whiteflies, young scales, even young
cles. Body colour ranges from light yellow to very dark tarnished plant bugs. A hand lens helps distinguish these
green. Cornicles are always black, regardless of body pests. The wings of aphids extend at least twice the length
colour (Fig 4.9). A. gossypii lives around the world and their bodies, which differentiates winged aphids from fun-
seriously damages cotton and cucurbit crops, but feeds gus gnats, whose wings are shorter (Gill & Sanderson 1998).
on almost anything. The species prefers high temperatures; Differentiating aphids from each other can be difficult.
at 27°C the aphids mature in seven days. A, gossypii vectors If apterae are light green, consider M ,persicae, P. hllrnuli, or
over 50 plant viruses and is often ant-attended. P. cannabis. M. persicae is about 25% larger than the other
Raychaudhuri (1985) cited Uroleucon jaceae two species, its antennal tubercles are short and rounded,
[Linnaeus) 1758 in India. U.jaceae is a large (2.5-3.5 mm its midline knob between the tubercles is small, its cornicles
long) reddish brown to brown-black aphid that infests develop a slight swelling, and its cauda is slightly constricted.
Compositae in Europe and Asia. P. cannabis tubercles are long (sometimes finger-like projec-
tions), heavily bristled, and P. cannabis sports the largest
DIFFERENTIAL DIAGNOSIS midline knob. P. hzlmz~lihas short tubercles, few bristles, and
Symptoms from aphids (wilting, yellowing) can be lacks a midline knob. Blackman & Eastop (1984) provided
confused with damage by spider mites and whiteflies. Turn an illustrated key for these species as they appear on hops.
over a leaf to see what lurks beneath. Young aphids canbe If apterae are dark green-black and lack attennae tuber-
cles, suspect A, fabae. But don't forget A. gossypii, which can
also be quite dark and lacks tubercles. The jet-black cornicles
of A.gossypii set it apart from A. fabae.

CULTURAL & MECHANICAL CONTROL

(method numbers refer to Chapter 9)
Method 1 (sanitation) is the workhorse. Mechanical re-
pelling (method 12a) works well against winged aphids.
Catching winged aphids with yellow sticky traps (method
12b) provides an early warning of aphids migrating into your
area. Screen all entrances to glasshouses (method 13).Method
Figure 4.9: Aphisgossypii. A. Adult aptera (wingless); B. Adult 5 (genetic resistance) glitters with future potential-but thus
alata (winged); C. Young nymph; all x 10 (courtesy USDA). far, only aphid-susceptible plants have been identified. BB
 Chapter 4: Insects and Mites 35

(1990) reported aphids "just love" afghanica cultivars

developed in Holland. Exclude or kill all ants in the area.

BIOCONTROL (see Chapter 10)

Aphids can be nearly eradicated with biocontrol organ-
isms. To control established aphid infestations, predators work
better than parasitoids. Parker (1913a)and Campbell &Cone
(1994)listed many natural aphid predators in California and
Washington hops yards. Most are now commercially avail-
able-green lacewings (Chrysoperla carnea), convergent
ladybeetles (Hippodamia convergens), two-spotted ladybeetles
(Adalia bipunctata), and the tiny slug-like predator Aplzidoletes
nphidimyza. These are described below. Predators that feed
on aphids in the absence of their primary hosts include
Cryptolaernlls montrouzieri, Harvlzotzia axyridis, and Rodolia
cnr~iinalis(discussed in the section on mealybugs), Stethorus
piclpes (described under spider mites), Orills species
(described under thrips), and Deraeocoris brevis (described Figure 4.11 : Green lacewing, Chrysoperla carnea, adult,
under thrips). "aphid lion" larva, and eggs (courtesy USDA).
The best preventatives are parasitoid wasps-Aphidius
n~atricariae,A . colenzatzi, and Aplzelinzis abdonziizalis (described and large jaws (Plate 8). They look like tiny alligators, 2-10
below). For some reason, parasitoids tend to prey on rose- mm long. Eggs are oval, white, and suspended from under-
coloured aphid colour morphs, and predators tend to attack sides of leaves by slender threads 1 cm long (Plate 9).
green morphs (Losey et al. 1997). DEVELOPMENT: Lacewings overwinter as pupae or
Fungi are the best microbial biocontrols of aphids. adults. Adults have strong migratory instincts and may fly
Viruses and bacteria must be eaten to be infective-and since for three or four hours before settling down on plants. They
aphids suck sap, there is little chance of their injesting viruses are attracted by the scents of tryptophan and caryophyllene
and bacteria sprayed on plant surfaces. Fungi need not be two components of aphid honeydew. Caryophyllene is also
injested. They work on contact, and directly penetrate aphid part of the aroma of Cannabis (Brenneisen & ElSohly 1988).
skin. Verticilliz~rnleratzii, Metarhiziurn anisopliae, Entornoplzthora Adults do not eat aphids, they feed on nectar or honeydew.
exitialis, and Erytzia neoaphidis are discussed below; Beauveria Females lay up to 600 eggs. Larvae consume 300400 aphids
bassinrza and Beazizleria globulifera are discussed under during their two-week larval life (Hill 1994).In optimal con-
whiteflies. ditions the life cycle takes 25 days. Adults live another 40
Yepsen (1976) suggested companion-planting with co- days; three to eight generations cycle per year. Short days
riander, anise, wormwood, or mint to drive away aphids. induce diapause in some populations of C . carnea (Tauber &
Aphids are also repelled by nasturtiums, marigolds, chives, Tauber 1993), rendering them ineffective in short-day flow-
onions, and garlic (Israel 1981). ering crops.
Table 4.3: Infestation Severity Index for aphids. APPLICATION: Eggs are the best, supplied in paper tubes
with intact egg-threads, or glued onto cards, or loosely mixed
Light any aphids seen with vermiculite in shaker bottles. Adults are supplied in
often no symptoms sealed tubes, larvae are supplied in divider boxes. Adults
should not be stored. Larvae can be stored up to two days,
Moderate < 10 aphidstleaf (not leaflet) in a cool (7-14"C), dark place; eggs can be stored up to four
feeding patches present days. Recommendations for release rates are presented in
Table 4.4. Place eggs or larvae at the base of plants, in the
Heavy 11-50 aphidslleaf
feeding patches coalescing Table 4.4: C. carnea release rate for control of aphids.

Critical > 50 aphidslleaf ISI* NUMBER OF PREDATORS RELEASED PER PLANT

shrivelled, discoloured leaves OR M-2 OF GLASSHOUSE CROP**
and honeydew
Preventative 5 larvae m-2 every 3 weeks
Chrysoperla (Chrysopa) carnea "Green lacewing" Light 5 larvaelplant OR 10 m-2 every 2 weeks
BIOLOGY: The lacewing larva (also known as the "Aphid
lion") is a nocturnal predator. It eats aphids, and to a lesser Moderate 25 larvae m-2 on trouble spots, 10 m-2
extent, feeds on whitefly nymphs, budworm eggs, thrips, elsewhere every week
and spider mites. Unfortunately it may also eat ladybeetle
eggs (Gautam 1994). The C . carnea species complex lives in Heavy 50 larvae m-2 on trouble spots, 25 m-2
temperate regions worldwide. C. carnea does best in moder- elsewhere every week
ate temperatures (24-27°C) and moderate humidity (55%
RH), but adapts to a wide range of humidity (35-75% RH). Critical 100 larvae m-2 on trouble spots, 50 m-2
APPEARANCE: Adults (called "lacewings") have slen- elsewhere every week, use ladybeetles
der pale green bodies 12-20 m m long, with delicate, lacy,
light green and bright eyes (Fig 4.11). * IS1 = InfestationSeverity Index of aphids, see Table 4.3.
like moths when disturbed. Larvae (called "aphid lions") **control of aphids with C. carnea is difficult during flowering (light
have yellowish-grey bodies with brown marks, tufts of hair, cycles below 12 hourslday)
36 Hemp Dise'ases and Pests

crotch of branches, or in distribution boxes hanging from mealybugs, scales, and other small insects. It is native to
plants. Spread them out to avoid cannibalism. Researchers North America, and does best in moderate humidity and
have coated eggs with acrylamide (sticky gel), so they can temperatures (>40% RH, 19-31°C).
be sprayed and adhere to plant surfaces. APPEARANCE: Larvae are slim, flat, alligator-like, dark
Do not irrigate plants for a couple days after release, to with orange spots, and up to 12 m m long (Fig 4.3 & Plate
avoid washing lacewings away (misting is okay and may 10). Adults are tortoise-shaped, 4-7 m m long, with a black
help young larvae survive). Lacewings d o not establish and white-lined thorax, and two white lines converging be-
themselves with ease. Entice adults to stay and lay eggs by hind the head (Fig 4.3 & Plate ll).Wing covers are orange,
providing artificial honeydew-mix honey and brewers yeast red, or yellow, typically with six black spots (the number
on sticks and distribute among plants (replace w h e n varies from none to 13).Eggs are orange, cigar-shaped, and
mouldy); commercial products include WheasP, Biodieta, laid in clusters of ten to 20 near aphid colonies.
or Formula 57E (see Chapter 9, method 14). DEVELOPMENT: Females lay up to 1000 eggs in spring
NOTES: Greeii lacezoiiig eggs are the most cost-effective and early summer. Larvae eat voraciously for a couple of
biocoiztrol of apliids cl~rreiztlyavailable. Lacewings can even weeks, pupate, then the adults also feed on aphids. Devel-
provide preventative control. For spot control of heavy opment from egg hatching to adulthood takes 28 days in
infestations, lacewings are not as effective as ladybeetles. C. optimal conditions. Adults live another 11 months. At the
carilea is compatible w i t h other Clzrysoperla species, end of the season adults migrate to hillsides and creek beds,
ladybeetles, Trichogranznza wasps, and predatory mites, al- where they aggregate and overwinter.
though some of these biocontrols may be eaten in the ab- APPLICATION: Supplied as adults in bottles, bags, or
sence of aphids. Normally, foliage sprayed with Bt does not burlap sacks. Store in a cool (4-8"C), dark place. Beetles pur-
affect C. carnea; but the genetically modified, truncated form chased before the month of May must be released by the
of Bt in transgenic plants may be harmful (Hilbeck et al. 1998). end of May; beetles purchased after early June can be stored
Some C. carnea strains tolerate malathion. for up to three months. Every couple of weeks remove them
from the cold, mist with water, allow to dry, and return to
Clzrysoperla (Chrysopa) rtifilabris storage (Cherim 1998). Used preventatively, release 4 1 (1
BIOLOGY: Another lacewing resembling the aforemen- gallon) of adults per 4000 m2 (=20beetles per m?), or refer to
tioned species. C. rllfilabrls is native to the Southeastern USA, Table 4.5. Watson (pers. commun. 1992) applied up to 5000
where it commonly inhabits trees. In optimal temperatures adults to an extremely infested, extremely valuable plant,
(24-27°C) the life cycle takes 23 days. C. rllfilabris reproduces and reapplied weekly. Extra adults flew off to the rest of the
better than C. carnae in humid environments (275% RH), it is crop. In a few weeks the aphids were 99% gone.
thought to be a more aggressive predator than C. carnae, pro- Commercial supplies of H. convergetzs have been har-
duces more eggs per female, may be found in higher num- vested from winter aggregation sites in California. They har-
bers towards the end of the season. C . rlifilabris lacks the bour a strong migrational instinct which requires them to
strong migratory wanderlust of C , carnea, so it stays in re- disperse before feeding and laying eggs. Thus, field releases
lease areas longer (Hoffmann & Frodsham 1993). Neverthe- of H.conzlergens are ineffective, because the adults quickly
less, establishing a season-long, self-replicating population disperse. They work in enclosed structures such as glass-
is difficult; multiple releases may be required. C. r~lfilabrisis houses and growrooms. Screen vents to keep them from es-
supplied like C. carnea, released at the same rate. caping. Delay migration releasing at night, and by "gluing"
their wings shut with a spray of sugared soda (Coke, Pepsi,
Other Chrysoperla and Chrysopa species etc.) mixed 1:l with water. After a week the sugar-water
BIOLOGY: Ckysoperla conlanche is new. It stays active later solution wears off.
in the season, after C. carnea enters diapause. C. conzanche is Entice ladybeetles to stay and lay eggs by serving them
more fecund than C. carnae (females lay up to 1100 eggs). "artificial honeydewu-see recipes in the Chysoperla carnae
Chrysopa o c l ~ l a t aa n d Chrysopa nigricornis are native section, above. Provide moisture by misting plants with
biocontrols in the USA, not yet commercially available. water before releasing (be careful misting flowering plants-
Adults of these species are predatory, an added advantage. they may develop grey mould). Some commercial suppliers
decrease the ladybeetles' migratory instinct by feeding them
Hippodamia convergens "Convergent ladybeetle" a special diet and allowing them to "fly off" in a controlled
BIOLOGY: A beetle that preys on aphids as well as environment.
NOTES: H,convergens is the best-known mail-order
predator of aphids. For heavy infestations, ladybeetles are tlie
Table 4.5: Hippodamia convergens release rate for control
best-they can beapplied in kiglz nillnbers as a "living itisecticide."
of aphids.
But ladybeetles are a poor choice for preventative control-
ISI* NUMBER
OF PREDATORS RELEASED PER ~2 without food they migrate away or die. Ladybeetles may
CROP**
OF GLASSHOUSE eat other predators in the absence of pests, but they are
compatible with parasitoids. Ladybeetles are not fully
Preventative 10 ladybeetles m-2 every 2 weeks compatible with some biocontrol fungi (e.g., Beanveria
bassiana and Paecilomycesfiimosorosel~s).Commercial supplies
50 ladybeetles m-2 weekly of H. convergens have been harvested from the wild, so they
have little tolerance for pesticides.

AdaIia bipunctata "Two-spotted ladybeetle"

BIOLOGY: A beetle that preys on aphids, is native to
Critical apply chemical controls, wait 2 days, North America, and does best in moderate humidity and
then release 200 ladybeetles m-2 twice temperatures. Adults have oval, convex bodies, with red to
weekly yellowish-orange wings marked by two black dots (Fig4.3).
They overwinter as adults; several generations arise per year.
* IS1 = Infestation Severity Index of aphids, see Table 4.3.
 Chapter 4: Insects and Mites 37

Coccinella undecimpunctata "Eleven-spotted ladybeetle" houses with concrete or plastic-covered floors, sprinkle peat
BIOLOGY:Another beetle that preys on aphids and other moss between rows of plants for pupation sites. A. aphidimyza
small insects, introduced to North America from Europe, and adults are not attracted to bright light (unlike lacewings and
does best in moderate humidity and temperatures. Adults ladybeetles), so they won't fly into bulbs and windows. But
have elongate-oval, convex bodies, with red or orange wings they may be attracted to yellow sticky traps. A, aphidimyza is
marked by 11 black dots, 5-7 mm long. A related European compatible with Aplzidius matricariae, but not with beneficial
species, Coccinella septempunctata, known as the seven- nematodes. The maggots tolerate some pesticides, but adults
spotted ladybeetle, has become established in eastern North are very sensitive. Adults can be released a week after spray-
America. It can accumulate in large numbers, displacing ing pyrethrum (Thomson 1992). Olkowski et al. (1991) sug-
native ladybeetles and becoming a nuisance to humans (even gest collecting wild maggots from Compositae weeds, but
biting people, according to Schaefer et al. 198%). field-collected maggots may be infested by A p h a n o g m ~ i s
fi~llneki and other hyperparasites, which occasionally infil-
Aplzidoletes apltidinzyza trate into commercial mass-rearing units (Gilkeson 1997).
BIOLOGY: A midge (fly) whose larvae prey on aphids, is
native to northern Europe and North America, and does best Aphidius inatricariae
in moderate humidity and temperatures (70% RH, 18-27OC). BIOLOGY: A braconid wasp, native to England; it does
A. apkidilnyza does poorly in hot, dry, windy locations. best in moderate humidity and temperatures (optimal 80%
APPEARANCE: Adults look like tiny mosquitos, 2.5 mm RH, 22-24°C). A. nzatricariae parasitizes over 40 species of
long. Eggs are oblong, shiny orange, 0.3 m& long. Larvae aphids. It provides good control of M. persicae a n d A. fabae,
are white upon hatching but turn orange as they mature, but poor control of A . gossypii (Steenis 1995) and Phorodon
reaching 3 mm in length (Plate 12). species (Neve 1991).
DEVELOPMENT: Larvae inject a paralysing poison into APPEARANCE & DEVELOPMENT: Adults are tiny black
aphids (Fig 4.12), then suck out all body contents. Westcott wasps, 2 mm long (Fig 4.13). Female wasps lay 50-150 eggs,
(1964) claimed the maggots kill an aphid a minute. Usually each egg is individually deposited within an aphid nymph.
the larvae kill ten to 100 aphids in a week or less, then pu- Eggs hatch into maggots which grow and pupate within
pate in soil. Adults are night fliers and feed on aphid honey- aphids. Aphids swell and stiffen into shiny, papery, light-
dew. Females lay 100-250 eggs near aphid colonies. The life brown mummies. Emerging wasps leave small,-round exit
cycle takes about four weeks, adults live another two weeks. holes. The life cycle takes two or three weeks in optimal tem-
A

Unfortunately, short photoperiods send maggots into peratures, and gdults live another two weeks.
diapause, ending their effectiveness when Cannabis flowers. APPLICATION: Supplied in vials or shaker bottles as
A. aplzidimyza overwinters in soil and may become estab- adults or pupae in mummies. Store for a maximum of one or
lished in glasshouses with soil floors. two days in a cool (8-1O0C), dark place. Release in shaded
areas, from open boxes hanging in plants or on rockwool
slabs. To prevent aphids, Reuveni (1995) recommended a
weekly release of one mummy per 10 m2 glasshouse. For
light infestations, release three to six mummies per m2 weekly
(Thomson 1992).
NOTES: A. matricariae works best as a preventative in
glasshouse situations. The species reproduces slowly and
cannot handle heavv infestations. Some researchers revort
A. matricariae wasps travel widely in search of prey, others
report they fly less than 3 m (Steenis 1995).Heavy honeydew
hinders them. They do n o t diapause during flowering.
Unfortunately, A. nzatricariae may be killed by its own
hyperparasites, especially in late summer. Hyperparasites
Figure 4.12: Aphidoletes, a midge that preys on aphids. infest A. nzatricariae within aphids, and leave aphids via exit
A. Adult female; B. Maggot biting an aphid on the knee. holes that may be confused with exit holes of A, n~atricariae.
(from Davis 1916) Exit holes bv A. nzatricariae are smooth and round. whereas
exit holes by hyperparasites have ragged, uneven edges
(Cherim 1998). Purchasing adults instead of pupae will
APPLICATION: Supplied as p u p a e mixed w i t h reduce or exclude hyperparasites.
vermiculite or peat in shaker bottles or tubs. Store up to four
days in a cool (8-15"C), dark place. Release pupae in humid,
shaded areas-not on surfaces receiving direct sunlight.
Hussey & Scopes (1985)released one predator per 25 aphids
at five-day intervals. Clarke & Watson (pers.commun. 1995)
released two to five pupae per m2 at the first sign of aphids,
or ten pupae per m2 in heavy infestations, repeating every
two weeks until control is achieved. Koppert (1998)
suggested releasing A. aplzidilrlyza pupae in piles of at least
10 cc (approximately 20 pupae).
NOTES: A. aphidinlyza may be used preventatively or re-
leased against established infestations. The maggots eat less
than lacewing larvae, but the maggots do not migrate away.
Adult females require an established aphid population be- Figure 4.13: Two wasps parasitizing aphids. A. Aphidius, a
fore they lay eggs. A. aphidimyza maggots need soil to pu- braconid (from van Leeuwenhoek 1700); B. Aphidencyrtus,
pate and become established. In soilless hydroponic glass- a chalcid (from Griswold 1926).
38 Hemp Diseases and Pests

A. matricariae is compatible with other parasitoid wasps, APPLICATION: Supplied as conidia formulated in pow-
Apliidoletes aphidinzyzn, ladybeetles, and lacewings. A . der, granules, or solid cultures in foil packets, plastic bags,
ntafricariae is not compatible with the biocontrol fungus or bottles. Store for weeks in a cool (2-3"C), dark place.
Beauueria bassiana, which kills larvae developing inside NOTES: This fungus is compatible with biocontrol
aphids. Adults wasps are attracted to the colour yellow, so parasitoids such as Aphidius matricariae and Aphelint~s
remove yellow sticky cards before releases are made. Avoid abdominalis (Roberts & Hajek 1992), but may infect preda-
insecticides, although some strains of A. matricariae tolerate tors, such as Orills insidiosis. Disadvantages include a slow
neem and malathion. onset of action, and M , anisopliae does not grow through the
soil (Leslie 1994). There are many strains of M. anisopliae,
Aphidius colemani and the successful control of specific insects may require spe-
BIOLOGY: This species has been confused with the afore- cific strains.
mentioned A. matricariae. A , colemani wasps lay twice as
many eggs as A. matricariae, and they reproduce much better Entomophthora exitialis
on A, gossypii (Steenis 1995).Several companies have dropped BIOLOGY: A cosmopolitan fungus that parasitizes
A, nzatricariae in favour of selling A. colemani. aphids. It does best in moderate to high humidity and mod-
APPLICATION: Supplied and applied like A. matricariae. erate temperatures.
NOTES: A related species, Aphidius ervi, works well APPEARANCE &DEVELOPMENT This fungus produces
against potato aphids such as Macrosiplztlm euphorbiae and several types of spores. Ballistospores are forcibly discharged
Alllacorthlun solani. from dead insects, surrounding the cadavers with a white
halo of sticky, mucus-covered spores. Resting spores arise
Aphelinus abdominalis within cadavers, and are mass-produced in liquid media
BIOLOGY: A chalcid wasp that parasitizes aphids, is na- fermenters. Spores that contact susceptible insects rapidly
tive to temperate Europe, and does best in moderate humid- invade their hosts (E. exitialis does not have to be eaten by
ity and temperatures. insects; it can penetrate insect skin). E. exitialis spores have
APPEARANCE & DEVELOPMENT: Adults are 2.5-3 mm been sprayed in California to control aphids (Yepsen 1976).
long, with a black thorax, yellow abdomen, and short Unfortunately, E. exitialis also infects aphid predators (but
antennae. Female wasps deposit eggs in adult aphids. Wasps not aphid parasitoids, according to Hajek 1993).
also feed on ovipositor wounds. In about two weeks, aphids
turn into black, leathery mummies. Larvae pupate within Erynia neoaphidis (= Entomophthora aphidis)
dead mummies and emerge as wasps, leaving behind a rag- BIOLOGY:This widespread fungus is related to the pre-
ged exit hole at the rear of the mummy. vious pathogen. It causes natural epidemics in P. htlmtlli and
APPLICATION:Supplied as adults in shaker bottles. Store other aphids (Byford &Ward 1968),and is being investigated
a maximum of one or two days at 8-10°C. The wasps migrate as a biocontrol agent. In France, E , neoaphidis predominates
poorly and should be released close to aphids. Release 50 in cool, humid conditions (Samson et 01. 1988).
wasps per infested plant (Thomson 1992).
NOTES: A. abdominalis works best as a preventative. It BIORATIONAL CHEMICAL CONTROL (see Chapter 11)
prefers potato a p h i d s (Macrosiyhtlnz eliphorbiae a n d Frank & Rosenthal (1978) repelled aphids with a
Alllacorthrtim solani). This wasp is compatible with other vegetable-based spray: grind up four hot with one
parasitoids. onion and several cloves of garlic. Let the mash sit in hvo
quarts of water for several days. Strain the liquid, and add a
~ e r t i c i l l i u k(Cephalosporium) lecanii half teaspoon of detergent as a spreader. ~ i r e c all t sprays
BIOLOGY: A fungus that parasitizes aphids (Vertalecm) and dusts at undersides of leaves. Kill young aphids and
and whiteflies (Mycotaln). V.lecanii works best against Myzus repel adults with insecticidal soap, diatomaceous earth, and
persicae and Aphis gossypii, and worst against Apkisfabae. The clay microparticles. Soaps have been sprayed up to a week
fungus provides excellent short-term control; it knocks down before harvest without any distasteful residues discerned
heavy aphid populations to a level managed by slower- on finished drv flowers (Bush Doctor 1985).
working biocontrols. For more information see the section Nicotine works par excellence against aphids, especially
on whiteflies. bhang aphids (Ceapoiu 1958) and hops aphids (Parker
1913a).Imidacloprid, a synthetic nicotine, works well against
Metarhizium anisopliae M. persicae, A. gossypii, and especially P, kumuli (LDgg = 0.32
BIOLOGY:Asoil fungus first tested in 1879 against beetles ppm), b u t not against A . fabae (Elbert et al. 1998).
in Russia (Samson et al. 1988). Different strains are used Cinnamaldehyde, extracted from cinnamon (Cinnamonl~n~
against aphids and whiteflies (BackOfP),termites (BioBlasP), zeylanicum), kills all aphids but also kills beneficial insects.
spittlebugs (Metaquinoa), cockroaches (BioPatha), thrips, Parker (1913a) and Yepsen (1976) controlled aphids with
beetles (chafers, weevils), ants, termites, and other insects. quassia, which kills P. hllmuli but not M . persicae.
M . atzisopliae does best in high humidity and moderate tem- Two synthetic insect growth hormones, buprofezin and
peratures (24-28°C). kinoprene, kill aphids. Deltamethrin, a synthetic pyrethroid,
APPEARANCE & DEVELOPMENT: M . anisopliae is a kills aphids, especially when sprayed insmall droplets rather
hyphomycete with branching, phialidic conidiophores and than large droplets (Thacker et al. 1995). Enhancing toxicity
simple oval spores (conidia) which aggregate into green by spraying small droplets may hold true for most insecti-
prismatic columns. Conidia in contact with insects quickly cides. Neem works poorly against aphids, and some aphids
germinate and grow into their hosts. M . anisopliae does n i t are resistant to pyrethrum. Rotenone kills insects with chew-
have to be eaten; it can penetrate insect skin, although it ing mouthparts, not aphids. Smother overwintering eggs of
usually enters through spiracles. Infected insects stop feed- M. persicae and P. lzllmuli by spraying neighbouring Prllnl~s
ing, then die in four to ten days, depending on the tempera- species with dormant oil.
ture. In humid conditions, M. atzisopliae reemerges from dead Winged aphids can be lured into traps baited with food
hosts to sprout more conidia and repeat the life cycle. attractants. P. hl~muli,for instance, is lured by a mix of volatile
 Chapter 4: Insects and Mites 39

oils distilled from Prllnus and Humulus plants (Losel et al. 1. GREENHOUSE WHITEFLY
1996).Adding food attractants to sticky traps works well. Trialetlrodes vaporariortlrn (Westwood) 1856, Homoptera;
Winged sexuales (sexual aphids, the autumn migrants) Aleyrodidae.
can be baited with sex pheromones, such as nepetalactol Description: Eggs are 0.2 mm long, pale yellow, oval or foot-
(Losel et al. 1996). Sexuales can be mass-trapped to break ball-shaped, with short stalks anchoring them to the leaf (Fig4.16).
the reproductivecycle. P. humtili males can sense pheromones Sometimes they are laid in circles or semicircles, covered with dust
6 m (20 ft) away from traps (Quarles 1999).Sex pheromones from the female's wings. Eggs turn purple-grey to brown-black be-
fore hatching into tiny 0.3 mm long "crawlers." These first-instar
do not attract nonsexual aphids, so pheromones have lim- larvae are almost transparent, oval in outline, nearly flat, and radi-
ited value against aphids infesting Cannabis during the spring ate a halo of short waxy threads from their bodies.Subsequent instars
and summer (Howse et al. 1998). Sexual pheromones may lose their legs and resdmble immature scale insects, reaching0.7 mm
attract beneficial insects that attack aphids, such as Aphidius in length. Late in the fourth stage, larvae change from transparent
matricariae, from surrounding areas (Quarles 1999). to an off-white colour by secreting an extra layer of wax to pupate
Neve (1991) mixed an aphid alarm pheromone, (E)-P within. Pupae project a halo of short wax threads from their pali-
farnesene, with pesticides or biocontrols. Alarm pheromones sade-like perimetre (Fig 4.15). Several pairs of longer wax threads
cause aphids to stop feeding and disperse across plants. Dis- may also arise from the top surface of pupae (Plate 15). Adult
whiteflies rarely measure over 1 mm in length. Their four wings are
persal increases aphid contact with pesticides and off-white,have rounded contours,and are held flat over their abdo-
biocontrols, resulting in better aphid control. The alarm mens almost parallel to the leaf surface (Fig 4.15 & Plate 13). The
pheromone is best applied with an electrostaticsprayer. (E)- wings may become covered with a white dust or waxy powder.
pfamesene has not been tested on Cannabis crops, but it may
not work-Cannabis produces pcaryophyllene, another
terpenoid which inhibits the pheromone activity of (E)-P
farnesene (Pickett et al. 1992).

WHITEFLIES
Whiteflies primarily cause problems in warm glass-
houses. They resemble tiny moths but are neither flies nor
moths. Whiteflies are related to aphids and leafhoppers.They
damage plants by sucking sap and vectoring plant viruses.
Three species of whiteflies infest Cannabis.

SIGNS & SYMPTOMS

Whiteflies produce few initial symptoms. This allows
their populations to build until you are suddenly engulfed
by a massive infestation.Whitefly symptoms resemble aphid
damageplants lose vigour, leaves droop, turn yellow, wilt,
and sometimes die. Leaves become glazed with sticky hon-
eydew, followed by a sooty-coloured fungus which grows
on the honeydew. The adults congregate on undersides of
leaves, out of sight to the casual observer. But if you look
closely, the adults look like tiny specks of ash (Plate 13).When
infested plants are shaken, a billowing cloud of whiteflies
fills the air for several seconds before resettling.
Figure 4.15: Pupae and adults of 2 whitefly species.
A. Greenhouse whitefly, Trialeurodes vaporariorurn;
B. Sweetpotato whitefly, Bernisia tabaci (McPartland
modified from Malais & Ravensberg 1992).

Life History & Host Range

Greenhouse whiteflies reproduce year-round. Females
lay 1100eggs, on undersides of leaves near the tops of plants,
often clustered in a circular pattern, anchored by short stalks
inserted into leaf stomates (Fig 4.16). Eggs hatch after seven
to ten days. First instar larvae crawl around plants search-
ing for suitable feeding sites. Once feeding begins, larvae
settle in one spot to suck sap, and eventually pupate. Larvae
take two to four weeks to reach adulthood, depending on
temperature. Adults live four to six weeks. In glasshouses
the generations often overlap, so all stages occur together.
The reproductive rate is dependent on temperature and host
plant. Optimal conditions for T. vaporariorlirn are 27°C and
75-80% RH. The pest infests growrooms around the world
(Frank & Rosenthal1978, Frank 1988).It attacks a wide range
Figure 4.14: Larva of sweetpotato whitefly, Bernisia of glasshouse crops. T. vaporariortlm vectors many plant vi-
tabaci, seen with an SEM (courtesy USDA). ruses, including the hemp streak virus (Ceapoiu 1958).
40 Hemp Diseases and Pests

Life History & Host Range

Silverleaf whiteflies have devastated growers in the
southern USA, outdoors and indoors. B. argentifolii appeared
in the USA around 1986. It was originally called the poinset-
tia strain of B. tabaci (so named because it first appeared in
Florida on poinsettia plants). Summer rains in the south-
eastern USA dampen the pest's damage there, but it recently
arrived in southern California. It can feed on anything the
Imperial Valley has to offer, and is resistant to almost all
pesticides. Its life history is nearly identical to that of B. tabaci
(except B. argentifolii females lay 10% more eggs), and it pre-
fers the same temperatures, conditions, and host plants.

DIFFERENTIAL DIAGNOSIS
Figure 4.16: Female Trialeurodes vaporariorum laying All Homopteran leaf damage looks similar, so symp-
eggs in a circle (from Weber 1930). toms caused by whiteflies can be confused with those of
aphids, scales, and hoppers. Immobile whitefly larvae may
be confused with immature scale insects.
2. SWEETPOTATO (TOBACCO) WHITEFLY Whiteflies are difficult to tell apart (Fig 4.15). Eggs of T.
Bernisla fabaci (Gennadius) 1889, Hornoptera; Aleyrodidae. vaporariorunz are white and turn purple or brown before
= Bentisin gossypiperdn Misra & Lamba 1929 hatching, while Bernisia eggs turn yellow. Pupae differ in
Description: Eggs are 0.2 rnm long, white, oval, with short stalks colour, hairiness, and the angle of their edges (edges of
anchoring them-to leaves. As eggs mature they turn yellow or amber. Bemisia pupae taper at a 45' angle to the leaf surface, whereas
Larvae are nearly colourless. Pupae have reddish-coloured eyespots, edges of T. vaporariorilnz pupae drop at 90" to the leaf sur-
their bodies are pale, marked by a caudal groove, and slightly pointed
face-like the edges of cookie dough cut with a cookie cut-
111 the rear (Figs 4.14 and 4.15). Margins of pupae taper to the leaf
surface, the margins lack a halo of short wax threads, and only a ter). T, vaporarior~irnadults are slightly larger and lighter col-
few longer hairs project from top surfaces (Plate 16). Pupae grow an oured than Bemisia adults, and hold their wings differently.
anterior wedge of wax at tracheal folds (Fig 4.15, solid arrow), and a B , tabaci and B. argentifolii are nearly identical, their mor-
posterior wax fringe which extends lateral to the caudal setae (Fig phological differences are described above, under B.
4.15, hollow arrow). Adults are light beige to yellow, with longitu- argentifolii. The two species are reproductively isolated (they
d ~ n a lstriations and angled (not rounded) wing tips. Adults hold cannot mate), genetically different (DNA sequences of the
Lilngs close to their bodies, in a tent-like position over their abdo- two species are as different from each other as they are from
mens (Fig 4.15).
Triale~irodesspecies), and biologically distinct (B. argeiltifolii
Life History & Host Range is more fecund and produces more honeydew).
Sweet potato whiteflies occur on Cannabis outdoors, in
southern Europe (Sorauer 1958), and Brazil (Flores 1958). CULTURAL & MECHANICAL CONTROL
Outdoor populations overwinter as eggs. The pest also in- (method numbers refer to Chapter 9)
fests glasshouses; indoor whiteflies reproduce year-round, Whiteflies are attracted to yellow objects, so use method
and have a life history similiar to that of T,vaporariorum, de- 12b to remove many adults. Method 1 (sanitation) is always
scribed above. Females lay 2100 eggs. Optimal temperatures important, especially indoors. Severely infested plants
for B. tabaci are 30-33°C. The pest attacks many crops around should be rouged (method 10).Anecdotal reports claim that
the world. air ionizers alleviate moderate whitefly infestations. Frank
(1988) sucked adults from undersides of leaves with a low-
3. SILVERLEAF WHITEFLY powered vacuum cleaner, early in the morning when
Beniisia argentifolii (Bellows & Perring) 1994, Hornoptera; whiteflies are cold and slow-moving. You can also shake
Aleyrodidae. plants and suck the whiteflies out of the air. Vacuuming is
= Bemisia tabaci poinsettia strain or strain B useful before rouging plants, otherwise you may leave be-
Description: B . tnbaci and B. argentqooiii show subtle morphological hind many flying adults. Outdoors, do not plant Cannabis
differences-B. avgentiJolii pupae have narrower wedges of wax at near eggplant, sweetpotato, tobacco or cotton crops. These
tracheal folds (dark arrow, Fig. 4.15),the posterior wax fringe does plants are whitefly magnets.
not extend lateral to caudal setae (light arrow, Fig. 4.15), and they
have one less pair of dorsal hairs. The adults are slightly larger than
those of B. tnbnci (Bellows et al. 1994).
BIOLOGICAL CONTROL (see Chapter 10)
Early whitefly infestations are hard to detect, so grow-
ers with a history of problems should release biocontrols
Table 4.6: Infestation Severity Index for whiteflies. before pests are seen. The best prophylactics are parasitic

I Light any adults seen when plant is shaken I wasps-Encarsiafornzosa, E. luteola, and Eretnzocer~tserenzicils,
described below. Predators include Delphast~ispltsillus,
Geocoris punctipes, and Macrolophtls caliginoslls (described
1 Moderate 5-10 adultsiplant, seen on more than I below), pirate bugs (Orius species, described under thrips),
one plant Deraeocoris brevis (described under thrips), and green
lacewings and ladybeetles (described under aphids). Re-
Heavy 11-20 adultsiplant, seen on many searchers are evaluating two predaceous phytoseiid mites,
plants Euseius hibisci (from California, described under thrips) and
Euseius scutalis (from Morocco, not currently available).
Critical > 20 adultsiplant OR sooty mould and Use combinations of parasitoids and predators. Heinz
leaf discoloration present & Nelson (1996) achieved better control of B, argentifolii by
OR a few adults on all plants releasing Encarsia fornzosa together with Delpkastus pztsillus
 Chapter 4: lnsects and Mites 41

Figure 4.17: Intertwining life cycles of the parasitic wasp Encarsia formosa and its host, Trialeurodes vaporariorum.
A. Normal life cycle of T: vaporariorum; B. Life cycle of E. formosa, beginning with egg insertion into 2nd instar nymph of
T vaporariorum (McPartland modified from Malais & Ravensberg 1992).

than by releasing either biocontrol alone. D. pilsillus feeds gots which slowly devour their host. Parasitized T.
on a few whiteflies containing young E. formosa larvae, but vaporariorttm pupae turn black, making them easy to spot
as E. forn~osa larvae mature within their hosts, D. pusillus (parasitized B. tabaci pupae turn amber brown). Elzcarsia
avoids them. maggots moult into adults after 15-20 days and emerge from
Fungi are the microbial biocontrols of choice, because carcasses, leaving behind an exit hole (Fig 4.17). They repro-
they infect whiteflies through their skin (other microbials duce without mating; adult females live another 30 days and
such as viruses and bacteria must be eaten, which is unlikely lay up to 120 eggs. Females also directly kill larvae by feed-
with sap-sucking insects). Choices include Verticilli~lmlecanii, ing on all instars (usually two or three per day), so this spe-
Bealiveria bassiana, Asckersonia nleyrodis, Pnecilomyces cies serves as both a parasitoid and a predator.
f~lnzosorose~~s(see below), and Metarhizizim anisopliae (dis- APPLICATION: Supplied as pupae in parasitized whitefly
cussed under aphids). Whiteflies are repelled by Nasttlrtium larvae, attached to cards, usually 80-100 E.formosa per card
species (Israel 1981) and shoo-fly, Nicandra pkysalodes (Frank (Plate 86). Store a maximum of four days in a cool (8-1O0C),
1988). Cucumbers (Clrci~missatiuus L.) serve as trap crops dark place. Hang cards horizontally, in shade near the bot-
for B. tabaci (Hokkanen 1991). toms of plants. The card surface with pupae should face
downward, to simulate the bottom of a leaf (although this is
Etzcarsia fornzosa not necessary). Recommendations for release rates are pre-
BIOLOGY: This wasp parasitoid does best in bright light, sented in Table 4.7.
50-80% RH, and temperatures between 18-28°C (optimally NOTES: E. formosa works best against T.vaporariorum, less
24°C).E.,fornzosa occurs throughout temperate regions of the so against B. tabaci/argentifolii. E. formosa works poorly in
northern hemisphere, but was almost wiped out by DDT. It heavy infestations, because the wasps spend more time clean-
has been raised for biocontrol since 1926 in England (van ing honeydew from themselves than hunting whiteflies. Thus
Lenteren & Woets 1988). E. forinosa works best as a preventative. The percentage of
APPEARANCE & DEVELOPMENT:Adults are tiny wasps, parasitized larvae (black larvae vs. uninfested white larvae)
0.5-0.7 mm long, with a black thorax and yellow abdomen can be monitored easily and should be 290% for effective
(Plate 14). Wasps lay eggs in whitefly larvae (second through control. Dense leaf hairs covering Cannabis may interfere with
fourth instars) and sometimes pupae. Eggs hatch into mag- the wasp's ability to locate prey. This problem also arises on
42 Hemp Dise'ases and Pests

Table 4.7: E. formosa release rate for control of glass- Store up to two or three days in a cool (8-1O0C), dark place.
house whiteflies. Scatter loose pupae on dry surfaces in direct sunlight.
Alternatively, place them in paper coffee cups with screened
bottoms and tape the cups to stalks or stakes. Disperse pu-
ISI* NUMBER
OF PREDATORS RELEASED PER PLANT
pae on cards the same way as E.formosa pupae. Do not al-
CROP**
OR ~2 OF GLASSHOUSE
low them to get wet. Used preventively, release three females
per plant per week in moderately humid areas like the North-
Preventative 1-5 pupae per plant or 10 m-2 every
east (Headrick ef al. 1995); double the rate in hot, dry cli-
2 weeks
mates. Koppert (1998) suggested releasing nine wasps per
m2 per week for moderate infestations,-or switching to
Light 20 m-2 every 2 weeks until 80% of
Delphastus pusilllls. Continue weekly releases until at least
whitefly larvae have turned black
75% of whitefly pupae are parasitized.
NOTES: E, eremicus works at higher temperatures than
Moderate 30 m-2 on trouble spots, 20 m-2
E. formosa, tolerates pesticides better, and lays more eggs.
elsewhere, then 10 m-2 every 2 weeks
But E. erenzicus harbours a greater attraction td yellow sticky
traps than E.formosa (Gill & Sanderson 1998). E. erenziclls is
Heavy 50 m-2 on trouble spots, 30 m-2
compatible with the same biocontrols as E. formosa.
elsewhere, then 20 m-2 every 2 weeks

Critical 100 m-2 on trouble spots, 50 m-2

Delphastus pusillus
BIOLOGY:Aladybeetle that preys on T. vaporariorunz and
elsewhere, then 20 m-2 every 2 weeks
both Benzisia species. In the absence of whiteflies, it eats spi-
* IS1 = Infestation Severity Index of greenhouse whitefly, see
der mites and baby aphids. D. pilsilltls is native to Florida,
Table 4.6. and does best in moderate humidity (70% RH) and moder-
ate to warm temperatures (19-32OC).
cucumbers. Cucumber breeders have reduced trichome den- APPEARANCE &DEVELOPMENTAdults are shiny black
sity in new varieties, improving E.formosa effectiveness (van and small, 1.3-2.0 mm long (Plate 15). Males have brown
Lenteren & Woets 1988).Bredemann ef al. (1956)tried breed- heads, females are all black. Larvae are elongated, pale yel-
ing mutant Cannabis plants without trichomes, but their work low, with a fuzzy fringe of hairs, 3 mm long. They pupate
was abandoned. on lower leaves or in leaf litter. Eggs are clear, elongate, 0.2
E. formosa adults have a bad habit of flying into HID mm long, and laid on leaves alongside whitefly eggs. The
lights and frying, which limits their use in growrooms. Yel- life cycle takes 21 days at an optimal 25-30°C. Adults live
low sticky traps may attract them. E. formosa is compatible another 30 to 60 days. Females iay three to four eggs a day,
with green lacewings (Ckrysoperla carnea), Delphastiispnsillt~s, for a total of about 75 eggs. The adults are voracious-they
and Eretmocerus erenziclls. It is also compatible with Bt (used devour as many as 10,000 whitefly eggs or 700 larvae. Lar-
against other pests). Most E. formosa strains tolerate some vae also consume whitefly eggs (up to 1000 during devel-
pesticides (Table 10.1), including short-acting insecticides opment) and larvae.
(soap, horticultural oil, neem, kinoprene, fenoxycarb), APPLICATION: Supplied as adults in bottles or tubes.
miticides (sulphur, abamectin), a n d fungicides Store up to four days in a cool (8-10°C), dark place. At the
(chlorothalonil, iprodione, and metalaxyl). Pyriproxyfen, an first sign of whiteflies, release one or two beetles per m2,
insect growth hormone useful against whiteflies, rarely repeating every two weeks. For moderate infestations,
harms adult wasps, but it kills immature E. formosa inside double the release rate and repeat weekly.
whitefly pupae. NOTES: D. ptlsilll~sworks best in moderate outbreaks.
At low whitefly densities the beetles stop reproducing and
Encarsia luteola (=Encarsindeserti) disperse (to thwart dispersal, see strategies outlined under
This related species is being investigated for control of Hippodamia convergens in the aphid section). In heavy
B. argentifolii. infestations D. pusillils bogs down in heavy honeydew. It
moves poorly across hairy tomato leaves-and may find
Eretmocerus eremicus hairy Cannabis leaves equally disagreeable. D. pilsilllls avoids
BIOLOGY: A wasp, sometimes called Eretmocerus nr eating whitefly larvae obviously parasitized by Encarsia or
culifornictls, that parasitizes T.vaporariori~m,B. tabaci, and B. Eretmocen~s,making these biocontrols compatible (Heinz &
arge)zfifolii. It is native to California and Arizona, and does Nelson 1996). D. pilsillus can also be released with green
best in moderate humidity (optimally 40-60% RH), and tem- lacewings (Chysoperla carlzea).
peratures (optimally 20-35"C, but tolerates up to 45°C).
APPEARANCE: Adults are tiny, 0.8 m m long, with yel- Geocoris punctipes "Big-eyed bug"
low-brown heads and bodies, green eyes, and short, clubbed BIOLOGY: A lygaeid bug that preys on whitefly larvae
antennae. Eggs are translucent and brown before hatching. and adults, aphids, spider mites and their eggs, and Heliofhis
DEVELOPMENT: Wasps lay eggs under sedentary eggs and young larvae (Hill 1994). G. pzltzcfipes is native to
whitefly larvae. Eggs hatch and larvae chew into their hosts. the southern USA and Mexico. It does best in moderate hu-
The hosts live long enough to pupate; infested pupae turn midity and temperatures. Nymphs and adults prey on pests.
an abnormal beige or yellow colour. The whitefly pupae die APPEARANCE & DEVELOPMENT Adults have large,
and E, ereinicils larvae pupate into adults within the cadav- bulging eyes and somewhat flattened brown bodies covered
ers. Adults emerge via small, round exit holes. The life cycle with black specks, 3-4 m m long (Fig 4.39 & Plate 16).
takes 17-20 days in optimal conditions. Adults mate and Nymphs look similar but without wings. Eggs are grey with
female wasps live an additional ten to 20 days, laying up to a tiny red spot appearing shortly after being laid. Under
eight eggs per day. Females also host feed. optimal conditions, the life cycle takes about 30 days.
APPLICATION: Supplied as pupae within whitefly pu- APPLICATION: In cases of mixed infestations, G.
pae on paper cards, or pupae mixed with sawdust in bottles. puncfipes will eat the largest pests first. So even if the
 Chapter 4: Insects and Mites 43

predators are surrounded by millions of tiny whitefly larvae, pers, beetles, termites, ants, European corn borers). B, bassiana
they would rather eat a plump, young budworm. G,punctipes lives worldwide. Several strains are available. The GHA
is compatible with Bt but not broad-spectrum insecticides. strain (BotaniGard ES@,Mycotrol0, ESC 1700) is registered
for use against whiteflies, thrips, aphids, and mealybugs. The
Macrolophus caliginosus ATCC 74040 strain, also known as the JW-1 strain (Naturalis@)
BIOLOGY:A mirid bug that preys on T. vaporariorunz, B. is registered against the aforementioned insects, beetles, and
tabaci, and B. argentifolii. In the absence of whiteflies it eats other soft-bodied insects. The Bb-147 strain (Ostrinila) is reg-
aphids, spider mites, moth eggs, and thrips. M, caliginosus istered against European corn borers. Uncharacterized prod-
works best on solanaceous crops; it survives on plant sap in ucts include Ago Bio BassianaE and Boverins. B. bassiana does
the absence of prey. best in high humidity (2 92% RH) and a range of tempera-
APPEARANCE: Adults are light green bugs, 6 m m long, tures, 8-35"C, optimally 20-30°C.
with long antennae and legs. APPEARANCE &DEVELOPMENT:Spores germinate and
DEVELOPMENT:Nymphs and adults attack all whitefly directly infect insects on contact, and kill them in two to ten
stages but prefer eggs and larvae, which they pierce to suck days. Under optimal conditions, insect cadavers sprout white
out body fluids. M. caliginosus may feed on 3 0 4 0 whitefly whorls of conidiophores, which bear globose conidia in
eggs a day. The entire life cycle takes about a month. Each zigzag formations (Plate 17). Both conidia and yeast-like
female lays about 250 eggs, inserted into plant tissue. blastospores are mass-produced in liquid media fermenters.
APPLICATION: Supplied as adults or adults and nymphs APPLICATION: B. bassiana is supplied as blastospores or
in bottles. Store one or two davs in a cool (8-10°C), dark conidia in emulsified vegetable oil or water-disbursable gran-
place. Gently sprlnkle onto infested leaves. For light infestations, ules. Store up to two years in a cool (8-10°C), dark place.
release five M. caligitlosus per m2; for heavy infestations, Spores are mixed with water and a wetting agent, then
release 10 per m2 and repeat the release two-weeks later. sprayed on all surfaces of pest-infested plants. Some strains
NOTES: M. caliginosus is compatible with E. fornzosa and have three to seven days of residual activity before reappli-
D. p~lsilb~s.Avoid insecticides while utilizing this control (espe- cation becomes necessary. Cherim (1998) described B. bassiana
cially pirimicarb).In the absence of prey, large populations of M. causing some phytotoxicity in vegetable crops, but Rosenthal
caliginosiis may suck enough plant sap to cause crop damage. (1999) reported excellent success with Cannabis crops.
NOTES: Beauveria spores are sticky and they infect pests
Verticillium (Cephalosporium) lecanii that brush against them; the spores do not have to be eaten
BIOLOGY:Acosmopolitan fungus that parasitizes aphids to be infective. Beal~veriaspecies are not as deadly as V. lecanii.
(Vertalecm) and whiteflies (Mycotalo). Other strains infect Nor are they as selective-Beauz~eria species kill ladybeetles,
scales, mealybugs, thrips, beetles, flies, and eriophyid mites. green lacewings, and other soft bodied predators. Effective-
! lecanii does best in moderate temperatures (18-28OC), and
l ness of B. bassiana depends on what pests are eating; accord-
prefers high humidity (180% RH) for at least ten to 12 hours ing to Leslie (1994), pests eating plants with antifungal com-
per day. pounds (such as Cannabis) become somewhat resistant to
APPEARANCE & DEVELOPMENT: Spores germinate and Beallvevia species. The fungus persists in soil, but organic
directly penetrate the insect cuticle. After infection the fungus material hinders survival, and nitrogen fertilizer kills it
takes four to 14 days to kill the pest. Under ideal conditions, (Leslie 1994). The fungus may persist as an endophyte in
dead insects sprout a white fluff of conidiophores bearing some plants (see the section on endophytes in Chapter 5).
slimy, single-celled conidia. Slime facilitates the adhesion of Some people develop allergic reactions to Beauveria after
conidia to passing insects. Blastospores are yeast-like spores repeated exposure (Hajek 1993).
produced within cadavers and mass-produced in liquid
media fermenters. Aschersonia aleyrodis
APPLICATION: Supplied as a wettable p o w d e r BIOLOGY:Asubtropical fungus that parasitizes whitefly
containing 1010 blastospores per g. Store in original, larvae (T. vaporariorl~nz)and some scales. A. aleyrodies rapidly
unopened package for up to six months at 2-6OC. Spores infects young larvae under conditions of high humidity. The
germinate best if soaked in water (15-20°C) for two to four fungus does best in high humidity and warm temperatures-
hours prior to spraying. Spray around sunset-dew and conditions found in vegetative propagation (cloning)
darkness facilitate spore germination. Be certain to spray chambers and warm glasshouses. In the USA, A. aleyrodis
undersides of leaves. Keep the spray tank agitated. The was commercially available in the early 1900s until it was
manufacturer recommends spraying a 500 g container of replaced by insecticides; now it is back (Gillespie &
spores over 2000 m2 of crop area (density and height of plants Moorhouse 1989).
not specified). APPEARANCE & DEVELOPMENT: A. aleyrodis covers T.
NOTES: V. lecanii works best as a short-lived agent to vaporarionln~cadavers with a slimy orange stroma, which
knock down heavy populations of pests, allowing E.formosa contains pycnidia with phialidic conidiophores and hyaline,
or other biocontrols to take over. Hussey & Scopes (1985) fusiform conidia. The teleomorph is an Hypocrella species.
claim a single spray of Mycotals will control greenhouse APPLICATION: Supplied as conidia on solid media.
whiteflies for at least two or three months. But optimal spore Hussey & Scopes (1985) report a dose of 2 x 108 A, aleyrodis
germination requires high humidity for 24 hours. Thus V. conidia per plant killed 75% of whiteflies, and did not infect
lecanii cannot be sprayed when plants are flowering and sus- the whitefly parasitoid Encarsiaformosa.
ceptible to grey mould. Encarsia fovnlosa can be used with V.
lecanii, but is slightly susceptible to the fungus at high humid- Paecilomycesfumosoroseus (=I? farinosus)
ity levels. Most other biocontrol organisms are unharmed. BIOLOGY:A fungus that parasitizes whiteflies, including
B. argentifolii. Some strains also kill spider mites, aphids,
Beauveria bassiana thrips, and mealybugs. P.fi~rnosoroseus is native to semitropi-
BIOLOGY:A fungus with a very wide host range, includ- cal areas and is used in the Philippines (PreFeRalO), Florida
ing sap-sucking insects (whiteflies, aphids, thrips, (PFR-97@), and in warm glasshouses in Europe (optimal con-
planthoppers, bugs, mites) and chewing insects (grasshop- ditions 2 68% RH, 22-33°C).
44 Hemp Diseases and Pests

APPEARANCE & DEVELOPMENT: Sprayed spores infect followed by horticultural oil (five days), soap and N.gossei
whiteflies and begin killing them in four or five days. In hu- (one day), and garlic spray (four hours).
mid microclimates, cadavers sprout hundreds of synnemata Neem mimics whitefly growth hormones and effectively
(bundles of conidiophores), which bear single-celled conidia. kills whitefly nymphs; it does not affect D.pllsillt~sbut may
NOTES: P.fi~nzosorose~~s
produces conidia in solid culture, harm Encarsia species (Mordue & Blackwell 1993).Two syn-
and yeast-like blastospores when cultivated in liquid fermen- thetic insect growth hormones, kinoprene and fenoxycarb,
tation tanks. Blastospores germinate faster than conidia. They also kill whitefly nymphs. The Rodale crowd recommends
have been added to mist irrigation systems in greenhouses. ryania (Yepsen 1976). For severe infestations, nicotine and
P.ftirnosorose~~s
is compatible with other fungi and with preda- pyrethrum kill more whiteflies, but also kill more biocontrols.
tory mites; the fungus sometimes kills ladybeetles. A new chlorinated derivative of nicotine, imidacloprid,
works very well against whiteflies. Biocontrol mortality can
CHEMICAL CONTROL (see Chapter 11) be reduced by shaking plants and then spraying whiteflies
Heavy whitefly infestations may require chemical con- while they hover in the air.
trol before biocontrol organisms can take over. Direct all sprays
at lilzdersides of leaves. Small plants in pots can be dipped in
spray solutions. Three or four applications (at weekly inter-
vals) kill whiteflies emerging from eggs that escaped initial EUROPEAN CORN BORERS
applications. These pests have "a-maize-ing" appetites. European
Bentz &Neal (1995)tested a sucrose ester extracted from corn borers (hereafter "ECBs") have been recorded on 230
Nicotiana gossei, a species of tobacco. A 0.1% spray of the es- different hosts, basically any herbaceous plant with a stalk
ter killed nymphs of T. vaporarior~~rn while sparing many E. large enough to bore into. ECBs are native to eastern Eu-
fornzosa wasps, so the spray and the biocontrol were compat- rope, where hemp and hops served as original host plants.
ible together. Liu & Stansly (1995) compared five sprays ECBs switched to corn after the introduction of maize into
against B. argentifolii in greenhouses: 96% of adults were killed Europe (Nagy 1976,1986).Nagy (1976)described ECB strains
by 2% Sunspray Ultra-Fine horticultural oil, 68% were killed in Hungary which prefer Cannabis over maize, with hemp
by bifenthrin (a pyrethroid), 26% by N. gossei sucrose ester, losses as high as 80-100% (Camprag et al. 1996).A Japanese
and 12% by M-PedeB insecticidal soap. After sprays dried strain also prefers Cannabis to corn (Koo 1940). ECBs were
on leaves, mortality from horticultural oil and bifenthrin accidentally introduced into Boston about 1908. They have
actually increased, whereas N. gossei ester stayed the same, migrated across much of the USA and Canada. Another
and soap lost all activity upon drying. Mortality against population was introduced into California via a cargo of Eu-
nymphs was not tested. In terms of repellency, bifenthrin ropean hemp (Mackie 1918).ECBs ruined wartime hemp in
was best (egg-laying adults were repelled for seven days), Illinois (Hackleman & Domingo 1943). ECBs attack both fe-

Figure 4.18: Approximate distribution of the European corn borer in the USA and Canada, circa 1997. Overlapping areas,
such as Kansas, containing overlapping populations; some Kansas ECBs may produce 2 generations per year, some may
produce 3 generations per year (McPartland update of Showers et a/. 1983).
 Chapter 4: Insects and Mites 45

ral hemp and cultivated marijuana all over the midwestern and local weather (Fig 4.18). In western Europe only one
corn belt (Bush Doctor 198%). Today, only southern Florida, generation arises north of about 45" latitude. Young (1997)
northern Canada, and sections of the western USA have yet predicts global warming will expand the range of ECBs, so
to be invaded (Fig 4.18). two generations may arise as far north as 52" latitude. Sum-
mers with high humidity and little wind favour egg-laying,
SIGNS & SYMPTOMS egg survival, and larval survival.
Young larvae (caterpillars) eat leaves until half-grown
(through the third instar). ECB larvae then bore into small DIFFERENTIAL DIAGNOSIS
branches. Their bore holes extrude a slimy mix of sawdust ECBs must be differentiated from hemp borers
and frass. Bore holes predispose plants to fungal infection (Grapholita delineana), see Fig 4.22. Nagy (1959)reported that
by M~zcroplzonlinaphaseolina (McPartland 1996) and Fusariunz 91% of ECB galls are located in the lower three-quarters of
species (Grigoryev 1998). Within one or two weeks of bor- Cannabis plants, while hemp borers tend to infest the upper
ing into small branches, ECBs tunnel into main branches and thirds of plants (Nagy 1967).ECBs make longer tunnels than
stalks (Nagy 1959). Their tunnels may cut xylem and cause hemp borers (Miller 1982). The grubs of weevils, curculios,
wilting. Stalks at tunnel sites may swell into galls, which are and other assorted beetles bore into stems and may be mis-
structurally weak, causing stalks to snap (Plate 18).Snapped taken for ECB larvae. Gall midges cause small galls in small
galls spoil the fibre and prevent seed formation. Ceapoiu branches and male flowers.
(1958) and Nagy (1959) photographed badly infested hemp
fields, showing half the plants toppled at odd angles. Top-
pled plants become tangled in harvest machinery, with yield
losses up to 50% (Grigoryev 1998). According to Emchuck
(1937),five to 12 ECBs can destroy a hemp plant.
ECB larvae born in late summer or autumn will change
tactics-instead of boring into stems, they infest flowering
tops, wherein they spin webs and scatter faeces. They selec-
tively feed on female flowers and immature seeds. Camprag
et al. (1996) reported seed losses of 40%.
Figure 4.19: Egg mass of 0. nubilasis (from Senchenko &
Timonina 1978).
TAXONOMY & DESCRIPTION
Ostriizia n~~bilalis
(Hiibner) 1796, Lepidoptera; Pyralidae.
= Pyraustrn nubilnlis Hubner, = Botys silacealis Hiibner 1796
Description: Eggs are less than 1 mm long. Just before hatching, CULTURAL & MECHANICAL CONTROL
the brown heads of lanrae become visible within the creamy white (method numbers refer to Chapter 9)
eggs. Caterpillars are light brown with dark brown heads (Plate Method 1 (sanitation) was written with ECBs in mind.
18).Along the length of their bodies are found several rows of brown Also apply methods 2a (deep ploughing), 3 (weeding, espe-
spot-like plates, each sprouting setae. Mature caterpillars may grow cially Artenzisia),4 (planting late in the spring), 6 (crop rota-
to 15-25 mm long. They spin flimsy cocoons and transform into
tion with red clover, Trifolillm pratense), 7a (proper moisture
reddish-brow11 torpedo-shaped pupae, 10-20 mm long. Female
moths are beige to dusky yellow, with irregular olive-brown bands levels), 9 (hand removal), and 12d (mechanical light traps).
running in wavy lines across their 25 mm wingspan (Plate 19).Males Virovets & Lepskaya (1983) cited Ukrainian cultivars with
are slightly smaller and darker, also with olive-brown markings on resistance to 0.nubilalis. Grigoryev (1998) found highest re-
their wings. Eggs are laid on undersides of leaves, stems, or crop sistance to 0. nubilalis in some Ukraine cultivars ('USO-27,'
stubble. They are laid in groups of 15-50, with the masses measur- 'USO-25'), and landraces from Italy ('Carmagnola'), France
ing 4-5 mm in diametre. Young eggs are translucent and overlap ('Chenevis'), and Yugoslavia ('Domaca Province'). Cultivars
like fish scales (Plate 20); with age they fill out (Fig 4.19). with less resistance included 'USO-14,' 'USO-12,' 'Yellow
Prior to 1850, scientific authors preferred the taxon silacealis to
Stem,' and 'Uniko B.'
nubilnlis, different names for the same species. Another ECB spe-
cies, Ostrinin scnpulalis Walker 1859, reportedly infests Ukrainian Grigoryev (1998)reported less infestation in dense hemp
hemp and does not attack maize (Forolov 1981). 0.scnpulalis larvae fields (4-5 million plants per ha) than in sparse plantings
and female moths look identical to 0. nubilalis. Male 0. scapulnlis (500,000 plants per ha). Crops cultivated in organically-
moths exhibit large, wide tibiae; this morphological feature sepa- managed soils suffer less ECB problems than crops cultivated
rates the species from 0.nubilalis (Forolov 1981). with conventional fertilizers (Phelan et al. 1996). Avoid
planting Cannabis near maize fields. Cover glasshouse vents
Life History & Host Range with screens and extinguish lights at night to exclude light-
Mature larvae overwinter in crop stubble near the soil attracted moths.
line. Springtime feeding begins when temperatures exceed
10°C. Larvae pupate for two weeks and then emerge as BIOCONTROL (see Chapter 10)
moths in late May (or June, or even August in Canada and The most successful parasitoids are Trichogramlria spe-
northern Europe). Females are strong night flyers, seeking cies, described below. Research on ECB parasitoids began
out host plants to lay eggs. They lay up to 500 eggs in 25 around 1886 in French hemp fields (Lesne 1920).The USDA
days (Deay 1950). Eggs are deposited on lower leaves of the has introduced over 20 species of ECB parasitoids into the
most mature (i.e., earliest planted) hosts. Artenzisia vulgaris USA. Only a few became established, such as the tachnid fly
is a common weed host (Grigoryev 1998). Eggs of first-gen- Lydella thompsonii (described below), and the braconid wasps
eration ECBs hatch in a week or less. Meteorus nigricollis and Macrocentrus gmndii.
Larvae feed for about three weeks, then spin cocoons Several predators feed on ECB eggs, such as green
and pupate. Moths emerge, mate, and repeat the life cycle. lacewings (described under aphids) and pirate bugs (Orills
A hard freeze late in the year kills all but the most mature species, described under thrips). Yepsen (1976) claimed that
larvae (those in their fifth instar). In Russia, ECBs live as far ladybeetles eat up to 60 ECB eggs a day. Podisus macl~liventris
north as 56' latitude (Grigoryev 1998). One to four genera- preys on young ECB larvae (described in the section on leaf-
tions of 0, nlibilalis arise each year, depending on latitude eating caterpillars).
46 Hemp Disea!ses and Pests

Bacillus tlzuringiensis (Bt) is the best microbial pesticide, Trichogramma species are supplied as pupae within
described below. The fungus Bealiveria bassiana (described parasitized eggs, attached to cards made of cardboard, pa-
under whiteflies) is lethal to first-instar ECB larvae (Feng ef per, bamboo, or within gelatine capsules. Store for up to a
al. 1985).B. bassiana is widely employed in China for control week in a cool (6-12"C), dark place. Cold-hardy species may
of ECBs (Samson et 01. 1988). B. bassiana strain 147 (Ostrinilm) tolerate longer storage. Avoid releasing in cold, rainy, or
is registered for control of ECBs in France. The fungus also windy conditions. If inclement weather cannot be avoided,
survives as an endophyte in some plants; maize plants in- the rate and frequency of releases must be adjusted upward
oculated with B. bassiana were protected against ECBs tun- (Smith 1996). The wasps cannot fly against winds stronger
nelling within stalks. Kolotlina (1987) killed ECB larvae in than 7 km h-1, which blow them out of release fields (Bigler
hemp fields with a mix of B, bassiana and B, globulifera. ef al. 1997).Trichogramma pupae can be manually distributed
Nosema pyraz~sta,a naturally-occurring control agent, is by hanging cards from plants in warm, humid places out of
described below. Don Jackson of Nature's Control (pers. direct sunlight. This approach takes about 30 minutes per
commun. 1997) suggested killing ECBs with beneficial ha (Smith 1996).Watch out for ants, which eat the pupae. In
nematodes such as Steinernemu carpocapsae (described under the presence of ants, place pupae in small cartons fitted with
cutworms). Inject the nematodes directly into stalks. protective screening, then attach cartons to plants. For a
large-scale approach, attach Trichogranzma pupae to carriers
such as bran, and broadcast them from tractors or airplanes.
Pupae and carriers can be coated with acrylamide sticky gel
so they adhere to plant surfaces.
According to Orr & Suh (2000), Trichogramma product
support in Europe is superior to that in the USA. European
suppliers maintain rigid quality control procedures, and ship
their products by overnight delivery in refrigerated trucks
or in containers with ice packs. Furthermore, European sup-
pliers provide their customers with extensive technical in-
formation, and even monitor temperature data and ECB
populations in areas where Trichogramma releases take place.
Customers are informed in advance of product delivery and
release dates. In contrast, nearly 50% of USA companies
shipped dead biocontrols or Trichogramma species other than
that which was claimed, shipped products in padded enve-
lopes by standard "snail mail," and provided little or no
product information or instructions (Orr & Suh 2000).
NOTES: Trichogramma species are compatible with Bt and
NPV. Avoid insecticides while utilizing the wasps.
Trichogramnza wasps search for eggs by walking across leaf
surfaces, so leaf trichomes slow them down. On tomato
Figure 4.20: Life cycle of Trichogramma minutum. leaves, the wasps get entangled in trichome exudates and
A. Female wasp laying egg within caterpillar egg, die, especially if exudates contain methyl ketones (Kashyap
B. View of egg within egg, C. Larva feeding within egg, et al. 1991). It is worth mentioning that Cannabis trichomes
D. Pupal stage, E. emerging adult, F. Adult male with open exude methyl ketones (Turner ef al. 1980), and probably
wings, G. Adult female with closed wings (McPartland hinder Trichogramnza wasps. Bredemann et al. (1956) at-
redrawn from Davidson & Peairs 1966). tempted to breed Cannabis plants without trichomes, but
their work was discontinued. Trichogrammawasps live longer
and lay more eggs if provided with food (e.g., wild flowers
Trichogramma species with nectaries, see Chapter 9, method 14).
BIOLOGY: At least 20 Trichogrnnznza species have been Different Trichogramma species vary in their effective-
mass-reared for field use. They efficiently kill ECB eggs, before ness against different pests. Some species work best against
the pests can damage crops. The eight most popular species ECBs, other work best against budworms. The relative
are described below. Most Triclzogramma wasps work best at effectiveness of different Trichogramma species against ECBs
20-29°C (range 9-36°C) and 40-60% RH (range 25-70% RH). is described below.
Species of a related genus, Trichogramatoidea, also provide T, evanescens This is a cold-hardy European species,
biocontrol against caterpillars. with popular strains from Moldavia and Germany. Adults
APPEARANCE & DEVELOPMENT: Adults are very tiny, are weak fliers and usually move less than 3 m from release
0.3-1.0 mm long, with a black thorax, yellow abdomen, red sites. Li (1994)tested 20 Trichogranzma species, and judged T
eyes, and shortmtennae (Plate 20). ema ales lay eggs in up evanescens the second-most effective control of ECBs (behind
to 200 pest eggs, which turn black when parasitized. Larvae only T dendrolini). At the first sight of moths in the glass-
pupate within pest eggs and emerge as adults in eight days house, release ten pupae per m2. T. evanescens can be re-
(Fig 4.20), adults live another ten days. leased at wider intervals than other Trichogramma species-
APPLICATION:There are two application approaches, in- every one to three weeks, using 200,000-300,000 wasps per
oculation or inundation (described in Chapter 10). The ha (Smith 1996).
inoculation approach often fails, because the wasps d o not T. pretiosum & T. minutum These polyphagous species
become established. The inundation approach requires the parasitize the eggs of many Lepidoptera. Mixtures of both
release of wasps when ECB moths lay eggs. ~ e c a i s wasps
e species are available. Clarke (unpublished research, 1996)
can only parasitize young eggs (one to three days old), and used a combination of T. pretios~im,T. minufunz, and T.
ECB moths lay eggs for several weeks, repeated inundations evanescens for best control in Dutch glasshouses. T. prefiosnm
become necessary. works best on plants under 1.5m tail, while T. min~itnmpro-
 Chapter 4: 11?sectsand Mites 47

tects taller plants. Neither species is very cold-hardy. AC- Thomson (1992), it is not compatible with T.minzltum, an
cording to Losey & Calvin (1995), T. pretiosum parasitized East Coast species. Losey & Calvin (1995) reported poor re-
ECB eggs better than 'Iminutum, but both were inferior to sults against ECB in maize.
T. nubilalis. Nevertheless, Marin (1979) used T.minutum to
protect 12,500 ha of Soviet hemp. Marin released T. minutum Bacillus thuringiensis "Bt"
from an airplane; release rates were not given. Camprag et BIOLOGY: The Bt bacterium was described by Louis Pas-
al. (1996)released 75,000-100,000 wasps per ha, and repeated teur in the 1860s (Flexner & Belnavis 2000), but Bt was not
the release a week later. Wasps were released from 50-60 field-tested against pests until the 1920s (the first trials were
locations per ha. Tkalich (1967) required 120,000-150,000 against ECBs). Bt is actually a family of bacteria-at least 35
wasps per ha, released when ovipositionbegan, and released varieties of Bt produce at least 140 types of toxins.
again when oviposition peaked. The active agent in Bt is a spore toxin, which exists in
T. brassicae (=maidis) This cold-hardy species is sus- the bacterium as a nontoxic crystalline protein (&endotoxin).
ceptible to mechanical injury, so wasp shipped on The endotoxin does not become toxic until the protein is dis-
paper cards often die before release. French researchers im- solved by proteinases in the gut of certain insects. The acti-
proved survival rates by gluing pupae to the inner wall of vated toxin binds to cell membranes in the gut. Part of the
capsules (Trichocapss). This formulation makes handling toxin forms an ion channel in the cell membrane, causing
easier, with less wasp mortality. It has made T,brassicae the selective cell leakage of Na+ and K+, resulting in cell lysis.
favorite biocontrol of ECBs in Europe. In maize, release Bacterial plasmids that encode toxin production (cry
200,000 wasps (400 capsules) per ha per week, whenever sig- genes) have been classified by amino acid homology. Gener-
nificant numbers of ECB moths are caught in traps or seen ally, Cryl, Cry2, and Cry9 toxins are active against lepidop-
laying eggs. Depending on the number bf ECB generations teran caterpillars (formerly grouped as CryI), Cry3, Cry7,
per season, the number of required releases ranges from one and Cry8 toxins kill beetle grubs (formerly grouped as
(Orr & Suh 2000) to four to nine (Smith 1996). Bigler et al. CryIII), and Cry4, CrylO, and Cry11 toxins kill fly maggots
(1997) found that T. brassicae heavily parasitized ECB eggs (formerly grouped as CryIV).
within 8 m of each release site. APPEARANCE &DEVELOPMENTAfter eating Bt, insects
T. ostriniae & T. dendrolimi These species have been stop feeding within the hour. They become flaccid, and a
introduced from China. T. dendrolini is cold-hardy, T. ostriniae foul-smelling fluid (liquefied digestive organs) trickles from
is not (T. ostriniae cannot survive winters in Vermont). Re- their mouths and anuses. They shrivel, blacken, and die in
lease rates and intervals are similar to those of T.brassicae. In several days (Plate 21). Dead insects can be found hanging
maize, T. ostriniae prefers to parasitize eggs found in the from leaves. Unfortunately, Bt bacteria rarely multiply in pest
lower and middle parts of plants-not flowering tops (Wang populations; they must regularly be reapplied.
et al. 1997). Li (1994)judged T.dendrolimi the most effective APPLICATION: Bt is formulated as a dust, granule, sand
Trichogramma control of ECBs, out of 20 tested species. Ad- granule, wettable powder, emulsifiable concentrate, flowable
ditionally, T.dendrolimi is polyphagous, so in the absence of concentrate, and liquid concentrate. Commercial products
ECBs it will parasitize other caterpillars. do not contain viable bacteria. Most Bt formulations contain
T. nubilale This species is native to North America and the toxin and spore, but some contain only the toxin. Store
readily parasitizes 0. ntlbilalis. Losey & Calvin (1995) re- powders in a cool, dry place. Store liquid formulations in a
ported best results with T. nubilale against ECB in maize. refrigerator, for a maximum of one year. Immediately before
According to others, however, T. nubilale may not reduce ECB spraying, mix with slightly acidic water (pH 4-7). For better
populations consistently (Andow et al. 1995); 'Inubilale was results add a spreader-sticker and UV inhibitor. For best re-
15% less effective than 'Iostrinia. Simultaneous release of sults add a feeding stimulant (Entice@,Konsumes, PheasP).
the two species produced worse control-so T. nubihle and Spray all foliage uniformly and completely.
T,ostrinia should not be used together. Bt works best against young, small larvae-so spray at
T. platneri is native to the USA west of the Rockies. It the first sign of caterpillar hatchout or crop damage. Bt on
has been used in orchards and vineyards. According to foliage is degraded by UV light within one to three days, so
spray outdoors in late afternoon or on cloudy (not rainy)
days. Repeat application at least weekly while pests are out
and about. Spraying Bt every ten to 14 days prevents ECB
infestations, althotlgh suvface sprays will not kill ECBs already
inside stalks. Frank (1988) used a large-bore syringe to inject
Bt into stalk galls.
Bt variety kurstaki is the most popular Bt, on the mar-
ket since 1961. Over 100 products contain Bt-k, largely de-
rived from the HD-1 strain. Popular trade names include
Agrobac@,Biobitm, BMP 123@,Condor@,Cutlass@,Dipel@,
Full-Bac", and Javelins.Bt-k kills caterpillars, including ECBs,
hemp borers, and budworrns. Genetic engineers have trans-
ferred the Bt-k toxin gene CrylA(b)to Pset~domonasflorscens.
This bacterium has a thick wall, so the P.florscensproduct is
marketed as "microencapsulated" (M-Trakm, MVP 118, M-
Peril@,Mattche).Microencapsulationprotects the toxin from
UV light, so the toxin remains effective on plant surfaces for
up to eight days (or two to four times longer than regular Bt-
k). Bt-k genes have also been inserted into a bacterium,
Clavibacter xyli, which lives in plant xylem (Fig 4.21). Inocu-
Figure 4.21 : Xylem vessel colonized by Clavibacterxyli, a late seeds with C. xyli, or hand-vaccinate plants to treat ECBs
bioengineered Bt carrier (SEM XI 000, McPartland). already inside stalks.
48 Hemp Dise!ases and Pests

Bt variety aizawai (Bt-a, XenTaris, Agree@,Design@)is thompsonii overwinters as maggots in hibernating ECB lar-
a new Bt that kills armyworms, budworms, and some bor- vae. Two or three generations arise per year. The first gen-
ers. Bt-a contains a different endotoxin, so it is more effec- eration often emerges too early in the spring, and must sur-
tive against Spodoptera species and insects that have devel- vive on alternate hosts, such as Papaipenza nebris, the com-
oped resistance to Bt-k. Some products contain a mon stalk borer (Mahr 1997).
transconjugated combination of Bt-a and Bt-k (Mattchm). NOTES: L , thompsonii is susceptible to microscopic
B t variety morrisoni (Bt-m), the latest Bt, being mar- biocontrols, such as Beauveria bassiana and Nosema pyrausta
keted for use against lepidopterae with high gut pHs, such (Mahr 1997).
as armyworms and cabbage loopers.
NOTES: Bt toxin is normally harmless to plants and ben- Nosema pyrausta
eficial insects, though some people may develop allergic BIOLOGY:A single-celled microsporidial protozoan that
reactions. The Bt toxin is safely pyrolysed. Unfortunately, infects ECBs. It is presumably native to Europe but now lives
heavy reliance on Bt has created Bt-resistant caterpillars, in North America.
which first appeared around 1985 (Gould 1991). DEVELOPMENT: N. pyrausta rarely kills caterpillars, but
The workof genetic engineers may accelerate resistance. causes debilitating disease in both larvae and adults, which
Bioengineers have transferred Bt genes to crop plants. Every reduces ECB longevity and fecundity. Hence N. pyrausta
cell in these transgenic plants produces its own cache of Bt- serves as a long-term population suppressor. It produces tiny
a
the equivalent of permanent systemic insecticide. This input ovoid spores (4.2 x 2 km) which spread in larval faeces, and
trait creates constant, global pressure for the selection of spreads by transovarial transmission in eggs.
resistant pests. Organic farmers, who have relied on Bt for APPLICATION: Researchers have mass-produced N.
decades, may lose their best weapon against caterpillars. To pyratlsta as spores, sprayed on foliage while young ECBs are
delay the evolution of Bt-resistant pests, farmers who plant still feeding on leaves. Spores persist on foliage for one to
Bt crops are required to plant a percentage of their acreage two weeks. ECB predators, such as green lacewings, can eat
in non-Bt crops. The idea behind this "refuge" strategy is N. pyrat~sta-infectedECB eggs and pass the spores harmlessly
that Bt-resistant pests arising in transgenic Bt-corn will mate in their faeces. These spores remain infective. Unfortunately,
with Bt-susceptible pests from neighbouring nontransgenic pamsitoids that develop within ECBs, such as Macrocentrl~s
crops. Assuming Bt resistance is recessive, and assuming grandii, may acquire the infection. N, pyratista can be used
susceptible adults are homozygous, the offspring of these with Bt-the treatments have an additive effect.
matiGs should continue to be susceptible to Bt. The prob- NOTES: A second species, Nosema furnacalis, is cur-
lems with this strategy, however, are multifarious. The cor- rently under investigation. A related microsporidian,
rect size of "refuges" has not been determined. The spatial Vairimorpha necatrix, is also being investigated. V necatrix
proximity of transgenic plants to nontoxic plants is critical- kills ECBs and various armyworms much quicker than
close enough to allow mating between resistant and Nosema species.
nonresistant moths, but far enough apart to prevent the
migration of larvae from transgenic plants to nontoxic plants. CHEMICAL CONTROL (see Chapter 11)
Convincing farmers to create pest havens (aka, refuges) may Young ECBs (those still feeding on leaves) are repelled
be difficult, and there is no enforcement mechanism. Fur- or killed with neem, nicotine, rotenone, and ryania. Once
thermore, the basic premise may be flawed: Bt resistance in ECBs bore into stalks, nothing sprayed on plant surfaces will
ECBs may not be recessive (Huang et al. 1999). Pests have affect them. Frank (1988) used a large-bore syringe to inject
alread appeared with resistance to Bt-transgenic plants stalk galls with plain mineral oil. Clarke (pers. commun.
(Gould 1998). 1995) retrofitted the spray nozzle and tube from a can of
To complicate the situation, transgenic Bt is expressed WD-40@onto an aerosol can of pyrethrin, and sprayed the
in a modified, truncated form. Hilbeck et al. (1998) pyrethrin into borer holes. After treatment, wipe away all
demonstrated that truncated Bt may harm predators, such frass from stems. New excrement indicates a need for re-
as lacewings (Ckrysoperla carnea) thatkat ~t-cdnsumingECBs. peated treatment.
The effects of transgenic Bt upon nontarget organisms cannot Chemicals and microbial biocontrols work better when
be predicted; pollen from Bt-corn, for instance, contains combined with a feeding stimulant. One comrnercially-avail-
enough Bt to kill larvae of the monarch butterfly, Danatls able stimulant, Coax@,has been used against ECBs. For more
plexippzis. Bt-toxic corn pollen blows onto milkweed plants information, see the section on Budworms.
(~sclepiasczlrassavica) within 60 m of corn fields; cateriil~ars Baiting with synthetic pheromones can lure male moths
that eat pollen-dusted milkweed plants stop feeding A d die into traps. When many moths appear in pheromone traps, a
(Losey et al. 1999). new generation of young larvae is only a week away-get
your Trichogramma and Bt ready. In North America, two
Lydella thompsonii strains of ECBs respond to different pheromone blends of
BIOLOGY: This tachinid fly is native to Europe. The tetradecenyl acetate (Howard et al. 1994), so growers will
USDA released L. thompsonii in the USA between 1920 and need two traps for the two strains. Alternatively, small rub-
1938. It became firmly established and now can be found ber septums can be impregnated with pheromones and
from New York to South Carolina and west to the corn belt. spread around hemp fields to confuse male moths and in-
L. thonlpsonii parasitizes up to 75% of ECBs in the midwest hibit mating (Nagy 1979).
(Hoffmann & Frodsham 1993).
APPEARANCE &DEVELOPMENT:L, thompsonii adults re-
semble large, bristly houseflies. Female flies run up and down HEMP BORERS
hemp stalks, looking for ECB entrance holes. Females de- Hemp borers are also called hemp leaf rollers and hemp
posit living larvae, which wriggle into entrance holes. The seed eaters. Besides boring into stems, Kryachko et al. (1965)
maggots bite ECBs and bore into them (Mahr 1997). Mature described hemp borers destroying 80% of flowering tops in
maggots leave dead ECB larvae to pupate within stalks, then Russia. Bes (1978) reported 41% seed losses in unprotected
emerge as adults. The life cycle can be as short as 20 days. L. Yugoslavian hemp. Each larva consumes an average of 16
 Chapter 4: Insects and Mites 49

Cannabis seeds (Smith & Haney 1973). In addition to culti- Life History & Host Range
vated hemp, hemp borers also infest marijuana, feral hemp Hemp borers overwinter as last-instar larvae in crop
(Cannabis ruderalis), and hops. Anti-marijuana researchers stubble, weeds, and sometimes stored seed (Shutova &
considered the hemp borer "an excellent biocontrol weapon" Strygina 1969).They pupate in April, in soil under plant de-
(Mushtaque et al. 1973, Baloch et al. 1974, Scheibelreiter 1976). bris. Moths emerge in early May and migrate at night to new
Forty larvae can kill a seedling that is 15-25 cm tall in ten hemp fields. Moths are not strong fliers; Nagy (1979) calcu-
days (Baloch et a/. 1974).Ten larvae per plant cripple growth lated flight speeds of 3.2-4.7 km hour-1 in a wind chamber.
and seed production. A Pakistani strain is host-specific on Upon finding a hemp field, females land quickly, usually
Cannabis; the larvae do not feed on hops like European hemp within 3 m of the field's edge. After mating, females lay be-
borers (Mushtaque et al. 1973).Hemp borers arrived in North tween 350-500 eggs. Adults live less than two weeks.
America around 1943 (Miller 1982). Eggs hatch in five to six days at 22-25OC, or three to
four days at 26-28°C. Out of 350-500 eggs, Smith & Haney
SIGNS & SYMPTOMS (1973) estimated only 17 larvae survived to first instar. Lar-
Hemp borers feed on leaves in the spring and early sum- vae skeletonize leaves for several days before boring into
mer. Subsequently they bore into petioles, branches, and stems. Borers pupate within stems. The Primorsk region of
stalks. Feeding within stalks causes fusiform-shaped galls Russia is evidently the northern limit of G. delineana in Eura-
(Plate 22). Stalks may break at galls, although the length of sia. Further south, two generations occur in Hungary (Nagy
tunnels within galls averages only 1 cm (Miller 1982). 1979) and the Ukraine (Kryachko et a/. 1965).A partial third
Borers that hatch in late summer and autumn spin loose generation arises in Armenia (Shutova & Strygina 1969)and
webs around terminal buds and feed on flowers and seeds. four generations overlap in Pakistan (Mushtaque et al. 1973).
Senchenko & Timonina (1978) provided an illustration of Larvae go into hibernation in September and October.
seed damage. Late season larvae pupate in curled leaves Day length under 14 hours induces diapause (Saringer &
within buds, bound together by strands of silk. Nagy 1971). Temperature also influences diapause-warm
weather slows photoperiodic effects.
TAXONOMY & DESCRIPTION
Grapkolita deli~zeana(Walker) 1863, Lepidoptera; Olethreutidae. DIFFERENTIAL DIAGNOSIS
= Cydia delineatla (Walker), = Laspeyvesia delineana (Walker), Hemp borer damage often arises in the top 1 / 3 of plants
= Gvaplzolitn sinana Felder 1874, = Cydia sinana (Felder) (Nagy 1967),while European corn borers usually form galls
G ,delineana and G. sinana were considered different species until in the lower 3/4ths of plants (Nagy 1959). European corn
1968 (Miller 1982). Smith & Hanev (1973) and Hanev & Kutscheid
2 , borers and other boring caterpillars drill longer tunnels than
(1975) report Grapholita tristrigana (dlemens) iAesting hemp.
G , delineana larvae. Weevils, curculios, and gall midges also
- .
Miller (1982) reexamined G. tvistripana specimens from Minnesota.
Iowa, Missouri, Wisconsin, Illinois, Kentucky and New York-all bore into stems and form galls. Late-season hemp borers that
were G. delineana, not G. tristvigana. A report of Grapholita infest buds may be confused with late-season budworms.
interstictana on hemp (Dempsey 1975)probably represents another
m~sldentification.These species are differentiated by their genita- CULTURAL & MECHANICAL CONTROL
lia, summarized by Miller (1982). (method numbers refer to Chapter 9)
Description: Eggs are white to pale yellow, oval, 0.4 mm wide, Methods 1 (sanitation) and 2a (deep autumn plough-
and laid singly on stems and undersides of leaves. Larvae are pink- ing) greatly reduce overwintering borer populations. Elimi-
ish-white to pale brown, up to 9-10 mm long (Fig 4.22). Several nate "sanctuary" stands of feral hemp, hops, and other weeds
pale bristles per segment are barely visible. Their heads are dark (method 3). Early crop harvest (method 4) is helpful. Since
yellow-brown, with black ocelli, averaging 0.91 mm wide. Larvae
pupate in silken cocoons covered with bits of hemp leaf. Adults are
full-grown larvae survive in harvested stems, stems should
tiny moths, with greyish- to rusty-brown bodies and brown, fringed not be transported into uninfested areas. Larvae may
wings. Body length and wingspan average 5 mm and 9-13 mm re- overwinter in seed, so method 11 is important. Methods 12d
spectively in males, and 6-7 and 10-15 mm respectively in females. (nocturnal light traps) and 13 (screening) help. Breeding
Forewings exhibit white stripes along the anterior edge with four dwarf Cannabis may decrease hemp borer infestation-Smith
chevron-like stripes near the centre (Fig 4.22, Plate 23). & Haney (1973)rarely found larvae attacking plants less than
30 cm tall, even when plants were flowering.

BIOCONTROL (see Chapter 10)

Native organisms heavily parasitize G. delineana larvae
in the USA, which may explain why hemp borers rarely harm
feral hemp. Smith & Haney (1973) found 75% of larvae in-
fested by Lixophaga variablis (Coquillett), a tachinid fly, and
Macrocentrlis delicat~~s Cresson (a braconid wasp, described
below). Gonioziis species attack G. delineana larvae in Paki-
stan (Mushtaque et a/. 1973);Scambi~sspecies parasitize 30%
of hemp borers in Hungary (Scheibelreiter 1976).
Camprag et al. (1996) used Trichogramma wasps to con-
trol the first generation infestation with "51-68% efficiency."
He released 75,000-100,000 wasps per ha, and repeated the
release one week later. He did not report which species he
used. Trichogramma species are described in the previous sec-
tion on European corn borers.
Figure 4.22: Larva, pupa and female moth of Grapholita Peteanu (1980) experimented with Trichogramma
delineana (A) compared to larger Ostrinia nubilalis (B). evanescens. He released 80,000,100,000,or 120,000wasps per
Both about x2 actual size. G. delineana from Senchenko & ha, four times per season (presumably two releases against
Timonina 1978, 0. nubilalis from Ceapoiu 1958. the first generation and two against the second generation).
50 Hemp Diseases and Pests

T,eelanescens worked best at the highest release rate, and thetic pyrethroid (see Table 4.8). Spraying pesticides only
worked better against second generation larvae than first works before hemp borers burrow into stalks. Once inside
generation larvae. Peteanu combined T, evanescens with Bt stems, no surface sprays will affect borers. Nagy (1979) de-
and pesticides, with interesting results (see Table 4.8). scribed an "edge effect" in fields infested by G, delineana.
Smith (1996) controlled a related Grapholita pest, G. Weakly-flying female moths land quickly to lay eggs after
molesta, with Tricho,qrnrnma dendrolimi released at a rate of discovering a hemp field; most infestations occur in the first
600,000 wasps per ha, repeated every five days while moths 3 m around a hemp field. Spray this edge zone with pesti-
were laying eggs. cides as moths arrive. In severe infestations, the edge zone
Bako & Nitre (1977) successfully controlled young hemp should be cut down and buried or burned.
borers with aerial applications
.-
of Bt (described in the previ- Nagy (1979) used female sex hormones to attract and
ous section). Podistls macllliventris preys on young hemp bor- trap male moths, preventing reproduction. Fenoxycarb, a
ers feeding in the leaf canopy (described in the section on synthetic juvenoid, kills eggs of the related pest Grapholita
leaf-eating caterpillars). Consider injecting stem galls with filnebrana (Godfrey 1995).
beneficialnematodes such as Steinernemu carpocapsae (de-
scribed under cutworms).

Mncrocentrus ancylivorus
OTHER BORING
BIOLOGY: Several Macrocentrt~sspecies have been mass- CATERPILLARS
reared for field use. M . ancylivorl~sis a braconid wasp native At least five other moth larvae damage Cannabis stalks.
to New Jersey. It attacks many fruitworms, leafrollers, and These borers generally d o not form galls. Infested plants
stem borers. In the 1930s M . ancylivorlls was reared in large appear stunted and unhealthy. Inspection of stalks reveals
numbers to control Grapholita nzolesta. The parasitoid does entrance holes, often exuding sawdust-like frass. Hollowed
best between Massachusetts and Georgia, west to the Mis- stalks may collapse and fall over.
sissippi river.
APPEARANCE & DEVELOPMENT: Adults are slender 1. GOAT MOTH
wasps, 3-5 mm long, amber-yellow to reddish brown in col- Cossus C O S S ~ L S(Linnaeus) 1758, Lepidoptera; Cossidae.
our, with antennae and ovipositors longer than their bodies. Description: C. cossus caterpillars are said to smell like goats (as
Female wasps are nocturnal, most active at 18-27°C and will stalks, even after larvae exit to pupate in soil). Caterpillars are
>40% RH, and lay up to 50 eggs, one egg per borer (Mahr red-violet and grow 90 mm long. Pupae are reddish-brown, 38-50
mm long, covered with pieces of wood and debris. Moths are robust,
1998).The wasps go after borers already within branches (sec-
brown-bodied, with olive-grey variegated wings reaching a span
ond and third instars preferred). M . ancylivorl~slarvae ini- of 90 mm.
tially feed within caterpillars, then emerge to feed externally,
and pupate in silken cocoons next to the body of their hosts. Life History & Host Range
One generation arises per generation of the host. They Ferri (1959) described goat moth larvae attacking Ital-
overwinter as larvae in hibernating hosts. ian hemp. Moths emerge from pupae in June and July. C.
NOTES: In orchards, Grapholita molesta is controlled by cocct~stakes three or four years to complete its life cycle. The
releasing three to six M . ancylivorotls females per tree (Mahr pest normally infests trees in Europe, central Asia, and north-
1998). A related species, Macrocenfrt~sdelicatl~s,attacks G. ern Africa. A related American species, Cosslls redtenbachi,
delinenna i n the midwest. Macrocentrus grandii a n d ("the worm in the bottle") infests Agave plants, the source of
Mncrocentrlls gifilensis are widespread biocontrols of Euro- tequila.
pean corn borers.
2. MANANDHAR MOTH
CHEMICAL CONTROL (see Chapter 11) Zenzera multistrigata Moore 1881, Lepidoptera; Cossidae.
Many botanicals that control European corn borers also = Zeuzera indica Walker 1856, nec Z. indicn Herrich-Schaffer1854
work against hemp borers-neem, nicotine, rotenone, and Description:Adults of this species resemble Leopard moths (Zeuzera
ryania. Peteanu (1980) killed borers with sumithrin, a syn- pyrina L.), with predominantly white bodies, three pairs of steel-
blue spots on the thorax, and seven black bands across the abdomen.
Wings are white with steel-blue spots and streaks, veins have an
Table 4.8: Effectiveness of biological and chemical control ochreous tinge, wings span 85 mm in females, 65 mm in males.
against Grapholita delineana.1 Heads and antennae are black, lower legs blue, femora white. Lawae
are predominantly white, with black heads and dark spots.

2.. Trichogramma
3. Trichogramma
4. Bt (Dipel)
;; ; :::;
74.4
Life History & Host Range
Baloch & Ghani (1972) described this species causing
serious damage in Pakistan. Z . multistrigata takes two or three
years to complete its life cycle. According to Sorauer (1958),
this pest normally feeds within coffee tree trunks, as does
its difficult-to-distinguish cousin, Zeuzera coffeea! Nietn. Z .
nzllltistrigata occurs across the Himalayan foothills from
5, sumithrin 750 g a.i. 86.1 Dharmsala to Darjeeling.
6. diazinon 5%G 25 kg 1 55.8
7. #2 + #3 100,000+2kg 2+2 84.9 3. COMMON STALK BORER
8. #2 + #4 100,000+750g 2+2 89.2 Papaipenla nebris (Guenke) 1852, Lepidoptera; Noctuidae.
=Papaipemanitela (Guenee) 1852
9. #2 + #5 100.000+25ka 3+1 76.7 Description: Eggs are white, globular, ridged. Young larvae are red-
dish-brown with a pair of white dorsal stripes. Side stripes are in-
'Data from Peteanu (1980) terrupted near the middle of the body, lending a "bruised" appear-
2Dose per individual treatment ance (Metcalf et a / . 1962). When larvae approach maturity they lose
 Chapter 4: Insects and Mites 51

their stripes and turn a dirty-grey colour. Heads and anal shields DIFFERENTIAL DIAGNOSIS
are dark brown, mandibles and true legs are black. Mature larvae European corn borers and hemp borers usually cause
reach 45 mm in length and bore large exit holes (7 mm diametre). swellings at their feeding sites, Gall midges and boring bee-
Moths have robust brown bodies and greyish brown wings with tles must also be differentiated. Split the stem and see what
small white dots, wingspan 30 mm.
you find.
Life History & Host Range
CONTROL
Tietz (1972) listed Cannabis among this borer's 68 her-
Follow recommendations for European corn borers and
baceous hosts. Young larvae bore in monocots (e.g.,
hemp borers. To reduce P. nebris and other omnivorous pests,
quackgrass, giant foxtail) until they outgrow these hosts; by
mow down grass and weeds around crops. Grass and weeds
July they migrate to larger-stemmed plants. Maize and gi-
need to be mowed again in August before moths emerge to
ant ragweed are two common targets. Larvae usually enter
lay overwintering eggs. Biocontrol researchers are testing
stems near the ground, bore for 25-50 cm, then exit in search
of a new host. This restless habit multiplies crop losses. Lar-
Scambus pterophori, a wasp (Ichneumonidae) that parasitizes
caterpillar borers and beetle grubs. Lydella thompsoni may
vae enter soil to pupate in August; moths emerge in Sep-
attack P. nebris larvae (this biocontrol is described under
tember. Females deposit eggs on monocots between curled
leaves. Eggs hatch in spring. One generation arises per year. European corn borers). All lepidoptera are killed by Bt (de-
scribed under European corn borers). The two Noctuids (P.
I? iiebris lives in North America east of the Rocky Mountains.
nebris and P, ataphracta) might also be sensitive to NPV (de-
scribed under cutworms).yo control borers with Bt and NPV
4. BURDOCK BORER
requires individual injection of each plant stem. Consider
Papalpenza cataphracta (Grote) 1864, Lepidoptera; Noctuidae.
Description: Metcalf et al. (1962) said burdock borers resem- injecting stems with beneficial nematodes (Steinernema car-
ble common stalk borers, except I? cataphracta's side stripes are not pocapsae, described under cutworms). Synthetic pheromone
Interrupted and they are smaller. Adult moths resemble those of lures are available for more common pests, such as C. cosst~s.
common stalk borers, except for different genitalia.

Life History & Host Range

Tietz (1972)listed 31 herbaceous plants attacked by this
BUDWORMS
borer, including Cannabis. Frequent hosts include rhubarb, Four budworms appear in the Cannabis literature.
burdock, and thistle. Since P. cataphracta larvae are smaller Budworms specialize in destroying plant parts high in ni-
than those of P. nebrts, they inhabit smaller stems and trogen, namely flowers, fruits, and seeds. Some species also
branches. Larvae pupate in stems, not soil. One generation skeletonize leaves. Budworms spin loose webs around flow-
arises per year. The species is sympatric (same geographic ering tops and feed (and frass) therein. Sometimes they feed
range) with P, nebris. inside floral clusters where damage is not visible until flow-
ers are ruined. Wounded buds and frass provide a starting
5. HEMP GHOST MOTH point for grey mould infection.
Endocylyta escrescens (Butler) 1877, Lepidoptera; Hepialidae.
Takahashi (1919), Clausen (1931), and Shiraki (1952) 'OTTON
cited larvae of this moth attacking Japanese hemp, Larvae Helicoveya armigera (Hiibner) 1809, ~ e ~ i d o ~ t e~octuidae.
ra;
feed on roots and possibly stems. Adults are large (81-90 =Heliothis avmigem Hiibner, =Heliothis obsoleta (Fabricius),
mm wingspan) and fly swiftly. Coloration of moth forew- =Chlovidea obsoleta (Fabricius),= Bombyx obsoleta Fabricius 1775,
nec B. obsoleta Fabricius 1793
ings varies from greenish grey to flecks' Description: Eggs are hemispherical,shiny,with ridgesthat radiate
from the apex like wheel spokes, white when newlv laid but dark-
ening to tan with a reddish-brown ring, 0.5 mm in diameter (Fig
4.23).Newly-hatched bollworms are pale yellow with dark longitu-
dinal stripes, 1.5mm in length. They grow into stout caterpillars up
to 45 mm long. Mature caterpillars vary in colour from green to
brown to almost black, with alternating light and dark longitudinal
stripes along their bodies, pale-coloured undersides, with yellow-
green heads and black legs (Fig 4.23 & Plate 24). Tiny spines cover
most of their body surface (visible through a 10x hand lens), in
addition to the dozen or so longer bristles found on each segment.
Shiny brown pupae are found 5 cm or more below the soil surface.
Moths are stout-bodied and brown; wings are yellow-brown with
irregular lines and dark brown markings near the margins,wingspan
up to 40 mm (Plate 25).

Life History & Host Range

H . armigera lives in Eurasia and Australia (Hill 1983),so
we use the name preferred by Eurasian entomologists-Heli-
coverpa, rather than Heliothis. This pest does its worst dam-
age in the tropics, where its life cycle can be as short as 28
days (Hill 1983).H. armigera commonly infests cotton, maize,
tobacco, and chickpeas.-1t sometimes turns up in canned to-
matoes. The pest has been reported on hemp by Goureau
(1866), Riley (1885), Vinokurov (1927), Shiraki (1952), Rataj
(1957), Ceapoiu (1958), Dempsey (1975), Khamukov &
Figure 4.23: Life cycle of Helicoverpa armigera. Kolotilina (1987), and Dippenaar et al. (1996). It is cited on
A. Egg viewed from side; B. Egg viewed from above; marijuana by Rao (1928) and Nair & Ponnappa (1974). In
C. Larva; D. Pupa; E & F. Adult (from Holland 1937). southern India, Cherian (1932) considered H,arnzigera sec-
52 Hemp Dis eases and Pests

ond only to spider mites in destructive capacity. He found lars are spindle-shaped or conical, 20-23 mm long and 6-7 mm wide
that 100 bollworms could eat a pound of Cannabis (0.45 kg) (Kozhanchikov 1956). Therein bagworms feed and eventually pu-
per day! pate. Doumere (1860)described moth forewings as grey with brown
spots, bordered posteriorly with grey fringe; hindwings are greyish
H. armigera and the next species (H. zea) share similar
white with silver-white fringe. Head, thorax and abdomen are grey-
life cycles. The pests produce one to six generations per year, brown. Doumere said female moths resembled males, except for
depending on the latitude (six generations in the tropics, less smaller antennae. But according to Kozhanchikov (describing
away from the equator). Populations in temperate regions Sterrhoptenjxfusca, see next paragraph), only males develop wings,
overwinter as pupae in soil. Tropical populations d o not hi- while females are wingless, milky-white, and wormlike, with only
bernate. Moths emerge from pupae as late as June in north- rudimentary legs.
ern Russia (Vinokurov 1927). Female moths, which are noc- The identity of P. cannabinella is questionable.
turnal, lay over 1000 eggs, one at a time, on upper leaves of Kozhanchikov (1956) placed the species under synonymy
crops and weeds. According to Young (1997), H. zea moths with Sterrhopteryx fi~sca Haworth 1829, family Psychidae.
lay more eggs on yellowed leaves than green leaves. Eggs Paclt (1976) considered P. cannabinella identical to Aglaope
hatch in three to five days. Larvae eat leaves, flowers, or infatisfa (Linnaeus 1758), family Pyrom6rphidae
seeds. The larval period lasts 14-51 days, depending on lati- (=Zygaenidae). Marshall (1917) suggested the insect was a
tude (Hill 1983). Caterpillars in the tropics may seek shelter Hymenopteron!
in soil during the heat of the day (Cherian 1932). Budworms
are cannibalistic and feed on their brethren if suitable veg- Life History & Host Range
etation is unavailable. DoumPre (1860)originally found P. cannabinella in Lux-
embourg on hemp growing from birdseed. Larvae hatch in
2. BOLLWORM July and usually infest female flowers, not male flowers. Ac-
Helicozlerpa zea (Boddie) 1850, Lepidoptera; Noctuidae. cording to Kozhanchikov (describing Sterrhopteryx fiisca),
=Heliothis zen (Boddie), =Bombyx obsoleta Fabricius 1793, nec B. mating takes place without females leaving their bags. Fe-
obsoleta Fabricius 1775 males lay eggs in place, then die and fall out of old bags.
Description: H. zea closely resembles H . armigern; (compare Fig 4.24
to Fig 4.23), only a close look at genitalia tells them apart. Gill & DIFFERENTIAL DIAGNOSIS
Sanderson (1998) describe the adults as having distinct green eyes. Other caterpillars may infest flowering tops during part
of their life cycles, such as European corn borers, hemp bor-
Life History & Host Range ers, cutworms, and armyworms.
A plethora of common names reflects the wide host
range of H. zea-cotton bollworm (confused with H,armig-
era), tobacco budworm, tomato fruitworm, corn eanvorm, Table 4.9: Infestation Severity Index for budworms
and vetchworm. Tietz (1972) cited H. zea on North American
Catznab~s.It may be the "bollworm" that Comstock (1879)
reported "devouring heads of hemp." H. zea is not a perma-
nent resident above 39"N (it cannot overwinter), but every
summer the adults migrate as far north as Ontario (Howard
I Light
any budworms or webs or bud damage
seen

et al. 1994).H. zea is native to the Americas, while H. armigera Moderate signs or symptoms at more than one
is a Eurasian species (Hill 1983).H. zea is easily mistaken for location, moths present
Heliotlzis virescens (F.), also confusingly known as the to-
bacco budworm-a horrific pest of tobacco, cotton, and other Heavy many worms and webbings seen,
solanaceous plants. H. zea shares the same life cycle as H. most buds in crop infested
armigera, described above.
Critical every plant with bud damage, and 10 or
3. FLAX NOCTUID more budworms/plant, moths common
Heliothis viriplaca Hufanagel 1766, Lepidoptera; Noctuidae.
=Heliotlzis dipsncea Linnaeus 1767
Description: Larvae have greyish green to dark brown bodies up to
14-22 mm long, with green to yellow-green heads. Dorsal stripes CULTURAL & MECHANICAL CONTROL
are yellow, narrow, wit11 dark borders described as a rust colour (method numbers refer to Chapter 9)
(Kirchner 1906).Moths and eggs resemble those of the aforemen- Methods 2a (deep ploughing) and 3 (weeding) are very
tioned Noctuids. important. Cherian (1932)used method 9-he shook infested
plants every ten days and collected all caterpillars that fell
Life History & Host Range to the ground. Method 12d (nocturnal light traps) works
The Flax noctuid attacks flax, lucerne, soyabean, and at against moths. Tomato breeders have conferred some resist-
least 20 other plant hosts (Vinokurov 1927).H. viriplaca been ance against H. zea by selecting plants that produce methyl
found on flowering hemp by Kirchner (1906),Blunck (1920), ketones (Kashyap et al. 1991). Methyl ketones are produced
Vinokurov (1927), Clausen (1931), Shiraki (1952), and by Cannabis (Turner et al. 1980).
Ceapoiu (1958).H. viriplaca lives in cooler Eurasian climates
than H. arnzigera, and produces only one to two generations BIOCONTROL (see Chapter 10)
per year. Other than that, its life cycle is the same as that of Khamukov & Kolotilina (1987) controlled hemp
H. arlnigera (see above). budworms with Trichogramma pretiosz~mand T. nzinzlt~im(de-
scribed under European corn borers). They did not cite re-
4. HEMP BAGWORM lease rates. Smith (1996) said corn growers release a million
Psyche cannabinella Doumere 1860, Lepidoptera; Psychidae. T pretiosum wasps per ha every two or three days while
Description: Larvae slender and pale, pinkish white, with light moths are laying eggs, for a total of 11-18 releases per sea-
brown heads and legs. Shortly after hatching, larvae weave a co- son. T. brassicae and T. dendrolimi have also controlled
coon and cover it with flower fragments. Bags of mature caterpil- budworms, at release rates of 200,000-600,000 wasps per ha
 Chapter 4: Insects and Mites 53

(Li 1994).Less commonly used parasitoids include Ckelontls tive to the southern USA. Each fly lays up to 3000 eggs dur-
insularis, Eucelatoria byani, Microplitis croceipes, and Arckytas ing her 50-70 day life span. Eggs are deposited on leaves of
marmoratus (discussedbelow), and Cotesia marginiventris (de- plants, not caterpillars, so the hatching maggots must con-
scribed under cutworms and armyworms). nect with their hosts.
Predators work well indoors. The larvae of big-eyed
bugs (Geocoris species)consume an average of 77 first-instar Nuclear Polyhedrosis Virus "NPV"
budworms or 151budworm eggs before pupating into adults BIOLOGY Several strains of NPV infest budworms. Three
(which continue to feed!).Yet these statistics pale compared are described below.
to those of adult ladybeetles (62 budworm eggs per day), Heliothis zea NPV (HzNPV) specifically kills Heliothis
larval ladybeetles (34 eggs per day), and larvae of green and Helicoverpa species. Sandoz, via IMCC, registered
lacewings (28 eggs per day). Pirate bugs (Orius species) also HzNPV for use on tobacco in 1975 (as Elcar@),but discontin-
eat eggs and young larvae. Podisus maculiventris preys on ued the product in 1983. A new strain (Gemstara) is avail-
young budworm larvae (described in the section on leaf- able. According to Hunter-Fujiata et al. (1998), the recom-
eating caterpillars). mended dose of HzNPV is 6 x 1011OBs per ha, which can be
Effective microbial agents include Bt (see European corn obtained by grinding up the cadavers of 25-50 infected
borers) and NPV (discussed below). Researchers have killed budworms, or using 700 g of the 0.64% liquid formulation.
budworms with the fungus Nomuraea rileyi. For budworms In cotton, three applications are made at three- to seven-day
pupating in soil, consider the soil nematodes Steinernema car- intervals, beginning when pheromone traps start catching
pocapsae (see cutworms) and Steinernema riobravis (see flea egg-laying moths.
beetles). A u t o g r a p h a californica NPV (AcNPV, AcMNPV,
Gusanoa) produces mixed results against budworms-H.
Microplitis croceipes virescens is more susceptible to AcNPV than HzNPV, but H.
BIOLOGY: A parasitic braconid wasp that lays eggs in zea is less susceptible to AcNPV (Hunter-Fujita et al. 1998).
second through fourth instar larvae of H. zea and H. virescens Mainestra brassicae NPV (MbNPV, Mamestrins,
(Fig 4.24).The wasp is attracted to caryophyllene (Lewis & Virin@)kills some Heliothis-Helicoverpa species. For more in-
Sheehan 1997), a volatile oil produced by Cannabis plants formation on AcNPV and MbNPV, including development
(Mediavilla & Steinemann 1997). Economic mass-rearing and application, see the next section on cutworms.
methods are currently being developed.
Helicoverpa armigera Stunt Virus "HaSV"
BIOLOGY An RNA virus that infests H. armigera. Young
caterpillars are killed within three days of eating the virus.
The viral genome of HaSV has been genetically engineered
into tobacco plants to kill H. armigera (Anonymous 1996).

Nomuraea rileyi
BIOLOGY: A fungus that infects and kills many Noctuids,
including H , zea. The fungus does best in moderate humid-
ity and warm temperatures.
APPEARANCE & DEVELOPMENT:Spores germinate and
directly infect budworms through their skins. Death occurs
in about a week. Under optimal conditions, "mummified"
cadavers become covered with phialidic conidiophores and
oval, single-celled, greenish conidia. The cycle is repeated
when conidia contact new hosts. Blastospores are yeast-like
and produced in liquid media, but commercial formulations
require further research.

CHEMICAL CONTROL (see Chapter 11)

Figure 4.24: A parasitic wasp, Microplitis croceipes,
laying eggs in a budworm, Helicoverpa zea (courtesy Synthetic pheromone traps are available for H. armigera
and H. zea. The active ingredients are (Z)-11-hexadecenaland
USDA).
several related aldehydes and alcohols (Howse et al. 1998).
Pheromone traps are particularly important for monitoring
the annual immigration of H. zea into regions north of 39ON.
Chelonus insularis and C. texanus In tobacco, ten budworms per plant justifies breaking
BIOLOGY Two closely-relatedparasitic braconid wasps out the spray gun. Rotenone kills budworms, albeit slowly.
(some entomologists say they are the same species) that at- Neem also kills slowly, but it acts as an antifeedant, repel-
tack many Noctuids (budworms, armyworms, cutworms, ling budworms from sprayed surfaces (Mordue & Blackwell
etc.). Females lay eggs in budworm eggs or young larvae. 1993). Some budworms are highly resistant to pyrethrum
(e.g., H , armigera). Spinosad, a metabolite produced by an
Eucelatoria bryani actinomycete, controls budworms in cotton and vegetable
BIOLOGY: Aparasitic tachinid fly that lays eggs in fourth crops. Polygodial may be available soon. Back in the old days,
and fifth instar larvae of H,zea. Females deposit several eggs Cherian (1932) sprayed plants with lead arsenate; he esti-
in each host. The species is native to the southwestern USA. mated 100 g of sprayed marijuana contained 2.0 mg of ar-
senic, "allowed as a medical dose."
Archytas marmoratus Chemicals and NPV work better when combined with
BIOLOGY Another tachinid that preys on bollworms, Coax@,a feeding stimulant. It was developed for budworms
fall armyworms, and other Noctuids. A. marmoratus is na- on cotton, but also stimulates European corn borers on corn
54 Hemp Diseases and Pests

(Hunter-Fujita et al. 1998).Coax consists of cotton seed flour,

cotton seed oil, sucrose, and Tween 80 (a surfactant). Coax
has been used at 3.4 kg ha-1, mixed with NPV to kill
budworms in cotton fields (Hunter-Fujita et al. 1998). Feed-
ing stimulants work even better when they contain extracts
of the crop to which they are applied (i.e., try hemp seed
extracts). Monitor the effectiveness of controls by looking
for budworm frass-fresh frass is light green, old frass be-
comes dark green.

CUTWORMS & ARMYWORMS

Allen (1908)called cutworms "hemp's only enemy." Ten
species reportedly attack Cannabis. Some of these reports are
probably wrong, because cutworms and armyworms are
notorious migrants and may have been collected on Canna-
bis while en route to a true host plant. For instance, Alexan-
der (1984b) cited the eastern tent caterpillar, Malacosoma
anzericanL~m(Fabricius). Other questionable observations in-
clude the clover cutworm, Discestra (Scotogramma) trijolii
(Hufnagel) in North Caucasus hemp (Shchegolev 1929),and
the oriental armyworm, M y t k i m n a (Pseudaletia) separata
(Walker) infesting Indian marijuana (Sinha et al. 1979).
On the other hand, several armyworms we would ex-
pect on Cannabis do not appear in the literature. These poly-
phagous pests include the armyworm, Psezldaletia unipuncta
(Haworth), the fall armyworm, Spodopterafr~lgiperda(Smith),
and the cotton leafworm, Spodoptera littoralis Boisduval.

SYMPTOMS
Most farmers are familiar with cutworm damage, but Figure 4.25: Some cutworms. A. Adult and larva of Agrotis
few witness the culprits in action. Cutworms emerge from ipsilon; B. Spodoptera litura; C. Spodoptera exigua; D. Agrotis
soil at night to feed on stems of seedlings. The next morn- segetum (not to scale). All illustrations from Hill (1994).
ing, dead plants are found lying on the ground, severed at
the soil line. Older plants may not be completely severed-
instead, they tilt, wilt, and die. Since cutworms only eat a scribed cutworms causing heavy losses in late-sown hemp,
little from each plant, they destroy a surfeit of seedlings each but noted little damage in seedlings sown at the proper time.
evening. Cutworms burrow back into the ground shortly Robinson (1952) considered cutworms a problem only in
before dawn, usually within 25 cm of damaged plants. Some- hemp rotated after sod. A. ipsilon frequents weedy, poorly
times they drag seedlings into their burrows. Raking soil to cultivated crops.
a depth of 5 cm will uncover them. Cutworms roll into tight A. ipsilon overwinters in soil as pupae or mature larvae.
spirals (Plate 26) and "play 'possum" when exposed to light Moths emerge in April or early May and lay eggs on plants
(Metcalf et al. 1962). in low spots or poorly drained land (Metcalf et al. 1962).Each
Larvae hatching later in the season, lacking seedlings moth lays up to 1800 eggs. Eggs hatch in five to ten days.
to cut, will climb up plants to feed on leaves and flowers. Larvae feed for ten to 30 days before pupating, repeating
Some species accumulate in large numbers and crawl en their life cycle until a heavy frost. Two generations per year
rrlasse across fields, devouring everything in their path, earn- arise in Canada, three in most of the USA, and four genera-
ing their new name, armyworms. Armyworms are also called tions in southern California and Florida. A. ipsilon is found
"climbing cutworms." worldwide and despised for its pernicious habit of cutting
down many seedlings to satisfy its appetite. It feeds on al-
1. BLACK CUTWORM most any herbaceous plant.
Agrotis ipsilon (Hufnagel) 1765, Lepidoptera; Noctuidae.
= Euxon ypsilon Rottemburg 1761 2. PADDY CUTWORM
Description: Larvae are plump, greasy, grey-brown in colour, with Spodoptera littlra (Fabricius) 1775, Lepidoptera; Noctuidae.
dark lateral stripes and a pale grey line down their back, growing = Prodenin litura Fabricius
to a length of 45 mm (Fig 4.25). They have a dark head with two Description: Larvae reach 50 mm in length. They are stout, smooth-
~vhitespots. A magnifying glass reveals many convex granules on skinned, with black heads and dull grey to grey-green bodies par-
their skin, resembling rough sandpaper.Pupae are brown, spindle- tially covered with short setae. Mature cutworms exhibit yellow
shaped, with flexible abdominal segments. They are found 5 cm stripes down their back and sides. The dorsal stripes are bordered
down in soil near host plants. Moths are small, brown, with indis- by a series of semilunar black spots (Fig 4.25). Paddy cutworms
tinct Y-shaped markings on their dark forewing (hindwings nearly ciosely resemble cotton leafworms p podo opt era littornli; Boisduval).
white), and difficult to distinguish from other Noctuids. Eggs are Pupae are brown and spindle-shaped. Moths have robust brown
round to conical, ribbed, yellow, laid either singularly
- or in 'lusters bodies and wings coloured buff to brown (forewings are darker,
of two to 30 on plants c~dseto the ground. with buff markings), reaching a span of 38 mm. Eggs are laid in
clusters of 200-500 and covered with moth hairs.
Life History & Host Range
Black cutworms attack marijuana in India (Sinha et al. Life History & Host Range
1979)and hemp in Europe (Ragazzi 1954).Dewey (1914) de- S. litilra lives in warm climates and rarely overwinters.
 Chapter 4: Insects and Mites 55

In the tropics its life cycle takes 30 days; as many as eight Noctuidae. =Euxon segetum Denis & Schiffermuller
generations arise per year (Hill 1983).Female moths lay egg Description: Mature larvae are plump, greasy, cinnamon-grey
to grey-brown with dark heads and faint dark dorsal stripes, reach-
clusters under leaves of host plants. Eggs hatch in three or
ing 45 mm in length (Fig 4.25). Pupae are smooth, shiny brown, 12-
four days. Caterpillars destroy crops for 20 days before p u - 22 mm long and found in the soil. Moths are distinguished by black
pating in the soil. Pupation takes six or seven days, then kidney-shaped markings on their dark forewings (hindwings are
adult moths emerge once again. white). They are smaller than black cutworm moths, with a wing-
Also called the tobacco caterpillar and the fall army- span of 3242 mm. Females lay light yellow eggs haphazardly, on
worm, S . litura should not b e confused with Spodoptera plants or soil, singly or in groups of two to 20.
frugiperda (Smith) 1797, the North American fall armyworm.
5'.litura attacks crops from Pakistan to China, and south to Life History & Host Range
eastern Australia. Main hosts are tobacco, tomato, cotton, Also called the Turnip moth, Durnovo (1933) described
rice, and maize. Cherian (1932) considered S. litura a "minor A, segeturn destroying hemp near Daghestan, Georgia. This
pest." Nevertheless, the caterpillar attracted biocontrol re- polyphagous pests cuts d o w n seedlings at soil level and
searchers, w h o described it destroying flowering tops (Nair gnaws at root crops underground. In equatorial Africa and
& Ponnappa 1974). southeast Asia u p to five generations arise per year; whereas
i n northern Europe A. segettirn is univoltine (one generation
3. BEET ARMYWORM per year), and overwinters as larvae in the soil.
Spodoptera exigua (Hiibner) 1827, Lepidoptera; Noctuidae.
=Laphygma exigua Hubner 6. BERTHA ARMYWORM
Description: Newly-hatched larvae are light green with black heads. Mamestra corlfg~irataWalker, Lepidoptera, Noctuidae.
Full grown larvae vary in colour, grey-green to brown, with wavy Description: Young larvae are pale green with a yellowish stripe
lines down the back and pale lateral stripes along each side, skin on each side. Mature larvae darken to a brown-black colour, with
colour between the stripes often dotted and mottled, total length broad yellow-orange stripes on each side, and three narrow, broken
25-35 mm (Fig 4.25). Larvae have dark heads and often have a dark white lines down their backs. Some larvae, however, remain green.
spot on each side of the body above the second true leg. Pupae are Mature larvae have light brown heads and measure 40 mm in length.
dark brown, and found in the upper 10 mm of soil in a cell formed Pupae are reddish brown and tapered. Adults have grey forewings
of so11and plant debris glued together with a sticky secretion. Moth flecked with black, brown, olive, and white patches. A prominent
bodies are brown, similar to those of S. litura but smaller (wingspan white kidney-shaped spot arises near the wing margin. Forewings
25-30 mm). Forewings are grey-brown with a yellow kidney-shaped are edged with white and olive-coloured irregular transverse lines,
spot in the mid-front margin, reaching a span of 30 mm; hindwings wingspan 40 mm. Eggs are white (turning darker with age), ribbed,
are white with a dark fringe. Eggs are pearly-green (changing to and laid in batches of 50-500, in geometric, single-layer rows.
grey), spherical, with radiating ribs, 0.5 mm in diameter,laid in clus-
ters of 50-300, in several layers, covered by moth hairs and scales. Life Histow & Host Range -
M , configurata overwinters as pupae, 5-16 cm deep in
Life History & Host Range soil. Adults emerge and lay eggs from May to August. Each
Moths emerge in spring and deposit 500-900 eggs in female lays a n average of 2150 eggs, which hatch within a
four to ten days, which hatch in t w o to five days. Larvae
week. Young larvae congregate o n the underside of leaves.
feed for about three weeks and pupate in half that time. In
Crop damage peaks between July and August. One or two
w a r m areas four to six generations arise per year. Beet generations arise per year. M. config~~rata is a serious pest of
armyworms can march in enormous numbers, defoliating canola (rape seed), but also feeds o n alfalfa, flax, peas, and
hundreds of hectares of sugarbeets, rice, cotton, maize, and potatoes. Weed hosts include lamb's quarters and wild mus-
vegetable crops. tard. The species is native to North America and attacks hemp
Anative of India and China, where it feeds o n Cannabis in Manitoba, chewing off all leaf blade tissues, leaving only
(Cherian 1932), S. exigua w a s accidentally introduced into
a stalk with leaf skeletons (Moes, pers. commun. 1995). The
California around 1876 (Metcalf et al. 1962). It moved east to
pest population is cyclical-M, corzfigtlrata damage becomes
Florida and north to Kansas and Nebraska. From Florida significant for two to four years, then becomes insignificant
the pest was introduced into Europe, where it attacks hemp for eight to ten years, then cycles into significance again.
(Ceapoiu 1958).
DIFFERENTIAL DIAGNOSIS
4. CLAYBACKED CUTWORM
Cutworm damage can be confused with post-emergent
Agrotis gladiaria (Morrison) 1874, Lepidoptera; Noctuidae. damping off, caused b y fungi. Bird damage can b e confused
Description: Mature larvae are grey with a distinctly paler dor-
sum, set with numerous small, flat, shining granules arranged down with cutworm damage, because early birds damage seed-
the back like pavement stones. The body tapers both anteriorly and lings before growers get out of bed and see them in action.
posteriorly, reaching 37 mm in length. Moths resemble most Armyworms must b e differentiated from other assorted leaf-
Noctuids: robust brown bodies, forewings darker than hindwings, eating caterpillars (discussed in the next section).
with variegated dark brown markings.
CULTURAL & MECHANICAL CONTROL
Life History & Host Range (method numbers refer to Chapter 9)
Overwintering larvae cause their greatest destruction Cold, wet spring weather causes cutworms to prolifer-
from May to late June. After that they pupate underground. ate, so follow method 4 (plant late). A combination of meth-
Moths emerge in September and October, mate, lay eggs, o d s 3 a n d 9 works well-hoeing, a good eye, and quick
and repeat their life cycle. Only one generation arises each hands. Destroy egg clusters and eradicate weeds (especially
year. Tietz (1972) cited this species feeding o n hemp, along grasses) from the vicinity. Observe method 6 (avoid rotating
with 20 other hosts. A, gladiaria lives east of the Rocky Moun- Cannabis after sod). Finish t h e season w i t h method 2a
tains, and becomes scarce south of Tennessee and Virginia. (ploughing). Recultivate fields in early spring, and wait two
weeks before planting. Cultivation destroys the cutworm's
5. COMMON CUTWORM food source (weed seedlings), and a two week fallow starves
Agrotis segetlltn (Denis & Schiffermiiller) 1775, Lepidoptera; them.
56 Hemp Diseases and Pests

Light traps (method 12d) work against moths. Method applying about 50,000 nematodes to soil around each plant.
13 (mechanical barriers) is labour-intensive but effective. S, carpocapsae has also been injected into borer holes and
Place cardboard or tin collars around seedlings. Be sure to sprayed onto foliage to control above-ground insects, but
push 10-cm-tall collars at least 3 cm into the soil. Rings of nematode survival is short. Survival can be lengthened by
sand, wood ashes or eggs shells also repel larvae. Placing coating nematodes with antidesiccants (e.g., wax, Folicote,
toothpicks alongside stems, despite old wives' tales, will not ~ios~s-627). Repeat applications every three to six weeks for
stop cutworms. For armyworms, apply Tanglefoot@as de- the duration of infestations.
scribed in method 13. Armyworms may bridge the sticky NOTES: S. carpocapsae lives near the soil surface. Its "am-
barrier by marching over their stuck brethren. bush" strategy works best against active, surface-feeding
insects, like cutworms, armyworms, and fungus gnat mag-
BIOCONTROL (refer to Chapter 10) gots. Steinernema species are hardier than Heterorhabditis
Cutworms and armyworms have been controlled by nematodes, easier to mass produce, and last longer in stor-
Trichogramma species (T. dendolimi, T. brassicae, T. evanescens, age. But they do not penetrate hosts as well as Heterorhabditis
described under European corn borers).In sugar beets, Smith species. Nematodes living near the soil surface suffer higher
(1996) released 100,000-200,000 wasps per ha every one to predation rates from nematophagous collembolans such as
two months. Chelonlis species have killed cutworms and Folsomia candida. Beneficial nematodes are compatible with
armyworms (see budworms). A new parasitic wasp, Cotesia Bt and NPV, but biocontrol insects that live in the soil are
marginiventris, is described below. Cool, rainy weather sup- incompatible(e.g., pupating Aphidoletes aphidimyza).Accord-
presses all parasitic wasps. In these conditions, switch to ing to some experts, wasp parasitoids that pupate within
Steinernema carpocapsae, a parasitic nematode (described be- silken cocoons are not killed.
low). Steinernema riobravis kills caterpillarsin warmer, drier soil
(see flea beetles). Nuclear polyhedrosis virus "NPV"
Turning to microbials-NPV kills cutworms and BIOLOGY A rod-shaped baculovirus (Fig6.4)which con-
armyworms, albeit slowly (described below). Bt (described tains dsDNA. Individual NPV rods are only 300 nm long,
under European corn borers) works well against armyworms but they may clump by the hundreds into occlusion bodies
(except S. exigua). Bt is less successful against cutworms. (OBs) that grow to 15 ym in diameter (easily seen under a
Nomliraea rileyi is a fungus that infests Noctuids (described light microscope). NPV kills larvae of Noctuids (e.g., cut-
under budworms). worms, armyworms, budworms, borers, and leaf-eating cat-
Some predatory insects (e.g., pirate bugs) eat the eggs erpillars). Of course not all caterpillars are from the Noctuid
of cutworms and armyworms. Podisus maculiventris preys family so you must properly identify your pests. Many
on cutworms and armyworms (described in the section on strains of NPV are available.
leaf-eating caterpillars). Many toads, birds, small mammals, Mamestra brassicae NPV (MbNPV, Mamestrina, Virin-
and other insects relish cutworms. Make their work easier EKS@) kills at least 25 species of Noctuids, including
by frequently tilling the soil. Yepsen (1976) repelled cut- Mamestra, Spodoptera, and Heliothis-Helicoverpa species.
worms with tansy, Tanaceturn vulgare (described in the next Mamestrin Plus@combines MbNPV with a synthetic
section on leaf eating caterpillars). pyrethroid, cypermethrin.

Steinernema (Neoapectana) carpocapsae

BIOLOGY Of the >20 known species of Steinernema, this
species is the best known and most widely marketed para-
sitic nematode (Biosafem, Exhibits, EcoMask@,Exhibit SC@,
Sanoplantm).S. carpocapsaekills cutworms, armyworms, and
other caterpillars, as well as beetle grubs, root maggots, and
some thrips and bugs in soil. Steinernemanematodes are cos-
mopolitan, and do best in soil temperatures between 15-32°C
(but the Scanmasks strain of S. carpocapsaeis native to north-
ern Sweden and remains active down to 10°C).
DEVELOPMENT S. carpocapsaehunts for hosts with a "sit-
and-wait" or "ambush" strategy. It stands upright on its tail,
near the soil surface, and waits for mobile, surface-feeding
insects to pass by. Steinernema species must enter hosts via
natural openings; they cannot directly penetrate insect skin.
Within hosts, Steinernema species release bacteria
(Xenorhabdus nernatophilz~sand subspecies X. bovienii and X.
poinarii). The bacteria ooze insect-destroying enzymes, and
do the actual killing. Bacteria also produce antibioticswhich
prevent putrefaction of dead insects, allowing nematodes
to reproduce in the cadavers (Plate 31). Cutworms begin
dying within 24 hours of infection, and the next generations
emerges within ten days. Steinernema-infectedhosts become
limp and turn a cream, yellow, or brown colour.
APPLICATION: Supplied as third-stage larvae in
polyethylene sponge packs or bottles. Sealed containers of
Steinernema species remain viable for six to 12 months at 2-
6°C. To apply to soil, follow instructions for Heterorhabditis
bacteriophora (see the section on white root grubs). Leslie Figure 4.26: Dead armyworm killed by a nuclear polyhe-
(1994) reduced black cutworms by 94% with S. carpocapsae, drosis virus, MbNPV (courtesy USDA).
 Chapter 4: Insects and Mites 57

Azitographa californica NPV (AcNPV, VPN-808, APPEARANCE & DEVELOPMENT: Adults are sturdy
GusanoL), has the widest host range of any NPV AcNPV wasps, 7-8 mm long, dark coloured, with long, curved an-
kills at least 45 species of caterpillars, Including most cut- tennae. The egg-laying wasps track down volatiles emitted
bvorms A genetically engineered version of AcNPV ex- by caterpillar-damaged plants. Amazingly, leaves damaged
presses a neurotoxin from the venom of A n d r ~ c t ~ tai~~tralis,
l~~s by wind or other mechanisms do not attract parasitoids, un-
a scorpion (currently under investigation). less oral secretions from leaf-eating insects are added
Spodopteva exigua NPV (SeNPV, Spod-XE, Instar@, (Turlings et al. 1990).Females lay 150-200 eggs, inserting 20-
Ness-A;) has a restricted host range-the beet armyworm. 60 eggs per caterpillar. C. nzarglrliventr~slarvae kill their host
The Yc~derBrothers developed SeNPV in Florida. and then pupate externally, in a yellow, silken cocoon at-
Heliotlzis zea NPV (HzNPV) was formallv marketed tached to host larvae or the undersides of leaves. The l~fecy-
as Elcar; and now sells as Gem-Starc (described under cle takes 22-30 days in optimal conditions.
Budn orms) APPLICATION: C. nlargirliventris is supplied as adult
Spodoptera littovalis NPV (SINPV, Spodopterinc) kills wasps. Release 800-2000 wasps per ha, weekly vvhile inotl~s
cotton leafk7orrns. It is not available at the time. of pests are flying and laying eggs (at least three releases).
DEVELOPMENT: Caterpillars that eat plant material
sprayed with NPV stop feeding, become flaccid, darken, then CHEMICAL CONTROL (see Chapter 11)
die. Unfortunately, death may take a week; slow lethality Yepsen (1976)described a cutworm bait: mix hardwood
has been a limiting factor in the popularity of NPV. NPV- sawdust, wheat bran, molasses, and water. Scatter the sticky
killed caterpillars, unlike Bt victims, generally do not smell bait at dusk. It immobilizes larva and exposes them to their
(unless infected b7! secondary saprophytic bacteria). The car- enemies. Lace the bait with an insecticide to insure results
casses hang from plants and spread more viruses (Fig 3.26). (Metcalf et al. 1962). For spray-gun enthusiasts, Yepsen sug-
APPLIC24TION:NPV is supplied as a liquid concentrate gested a repellent spray of garlic, hot peppers, and horse-
or ivettable povvder. Since NPV must be eaten to kill cater- radish. Neem acts as an repellent, antifeedant, and it kills
p~llars,spray all foliage uniformly and completely. NPV on Spodoptera species (Mordue & Blackwell 1993). Spinosad is
foliage is degraded by LTV light, so spray outdoors in late particularly useful against Spodoptera species. Rotenone con-
afternoon or on cloudy (not rainy) days. To slow degrada- trols armyworms in cotton and 1,egetable crops.
tion, combine NPV with PlieastB or Coax", which are lepi- Diflubenzuron is the synthetic form of a natural chitin in-
dopteran feeding stimulants. hibitor that kills armyworms. Synthetic pheromone lures are
Shapiro & Dougherty (1993) increased the potency of commercially available for A, ipsilotz, A. segetlcrri, S.liturn, S.
NPV by adding lniinrlry briglltetlers. Many brands 'worked, exipia, and M. cunfigilrata.
especially stilbene brighteners (e.g., Leucophor BS,
Leucophor BSB, Phorwite AR, Tinopal LPW). They found
that O.lol,,of Tinopal LPW added to NPV worked the best.
Brighteners protect SPV from UV light; brighteners also LEAF-EATING CATERPILLARS
decrease caterpillar gut pH, which enhances NPV activity. The literature reports dozens of leaf-eating caterpillars
NOTES: Caterpillars gain resistance to NPV as they age. on Cannabis, including many improbable pests. Some improb-
For instance, the LD,(, for M. brassicae jumps from seven OBs able pests are incidental migrants, such as the American dag-
in the first instar to 240,000 OBs in the fifth instar (Hunter- ger (Acronictaamericana Harris) found on feral hemp in Michi-
Fujita r t al. 1998).Get those cutworms while they're young! gan (McPartland, unpublished data 1995). A. anzericana is
NPV is compatible with Bt and all other biocontrols. kTPV arboreal and probably fell from trees overhanging the hemp.
works well against two cutworms that are Bt-resistant, M. Hanson (1963) cited three improbable pests on leaves and
bri7ssicne and S. exiglln. "pods" of Vietnamese marijuana-the long-tailed blue s>val-
lowtail, Lan~pidesboctic~is(Linnaeus), Jllnzl~fesliochils (Stoll),
4grvtzs segeturi~granulosis virus and a Papilionides species.
BIOLOGY AsGV 1s a new OB-forming DNA 1lrus used Surprisingly, some improbable pests are cited more than
by European \\ orkers (Agrox ir") It is \ ery host-specific, once: Am!lnn octo Guen. was reported on Indian marijuana
kiillng only A segi7tniriand A rpsilun AgGV has been ap- (Rao 1928, Cherian 1932) and European hemp (Sorauer 1958).
plied as a sprav or a bran bait, at a rate of 4 x 10110Bs per ha Vice versa, several leaf-feeding caterpillars we'd expect to
(Huntei-Fuj~taet a1 1998) Young larbae are the most sus- find on Cannabis have not been reported. Examples include
ceptible, use light traps or pheromone traps to determine the pale tussock (Callitenrn p~ldiblindnLinnaeus), the red ad-
i\ hen Agrotis moths are flylng, and apply AsGV a week later miral (Vanessa atalarlfa Linnaeus), the comma or hops mer-
chant (Pol~jgunin cornina Harris), and the hops looper (Hypeiia
Agrotis segett~incytoplasmic polyhedrosis virus !~lltnilliHarris).
BIOLOGY: AsCPV is a n OB-forming RNA virus, Baloch & Ghani (1972)collected loopers from Pak~stani
discovered in the UK.Infected caterpillars lighten to a milky- plants, the polyphagous pests Plllsia nigrlsigiza Walker, lJ,
white colour before dying. iDiacizrisini orichalcea Fabricius and P.l~orficola.Loopers have
been observed on feral hemp in Illinois but not identified
Sacc!invopolyspoucla spinosn (Bush Doctor, unpublished data 1983).
BIOLOGY: A bacterium that kills S. exig~ra(a pest some- Cherian (1932)found larvae of two tussock moths (farn-
~vhatimpei~iousto Bt), cabbage loopers, and other Noctuids. ily Liparidae) causing minor damage in Indian marijuana
fields, E1~prucfisj?ater17aMoore and E,scitztillar~sWalker. These
Cvtesia lnarginiventris species occupy sympatric ranges from India to Malaysia.
BIOLOGY: A tiny braconid parasitic wasp that lays eggs Goidanich (1928)and Schaefer et al. (1987a)cited another (and
in young Soctuid caterpillars. C, n~al;oiiziilerifrislives in tem- infamous) member of the tussock family feeding on Catlila-
perate and semitropical regions around the world. It con- bis-the gypsy moth, L!/lna~lfriadispar (Linnaeus)
trols cabbage loopers, cutworms, armyworms, a n d Sl~iraki(1952)reported several odd species on Japanese
budworms in cotton, corn, and vegetable crops. hemp, including some improbable pests: Eurilctn i&x~i~,ca
58 Hemp Disslases and Pests

Heylaerts (Psychidae), Plafaplecta c o n s a n g u i s Butler A. gamma a major pest in temperate Eurasia, and now it lives
(Noctuidae) and five Nymphalids, including the Painted North America. The moths can migrate hundreds of miles.
Lady (Cynthia cardtti Linnaeus), Vanessa itrticae connexa But- Hosts include sugarbeets, assorted vegetables, Prunils, Rubits,
ler, Arasclznia btlrejana Bremer, Araschnia levana Linnaeus and and Sambuclts.
Pyranzeis indica Herbst. Anonymous (1974) also cited Pyrameis
indica inChina. Clausen (1931) added another odd east Asian 2. DOT MOTH
nymphalid, Polygoizia c-aztreum (Linnaeus). Melanchra persicariae (Linnaeus) 1761, Lepidoptera; Noctuidae.
=Polia persicariae (Linnaeus), =Mamestra persicariae (Linnaeus)
SIGNS & SYMPTOMS Description: Larvae colours vary from pale grey-green to dark green
Leaf-eating caterpillars act as leaf rollers (weaving leaves or purple-brown. They have a light green head, a pale dorsal line
together for protection while they feed), as leaf skeletonizers down their backs, a series of V-shaped markings, and a dorsal hump
(selectively eating leaf tissue between leaf veins, leaving a on the eighth abdominal segment. Setae (1-1.3 mm) cover the body
giving them a velvety texture. Fully-grown larvae reach 40 mm in
skeleton of veins), or they simply chew big holes in leaves.
length. Moths are grey to bluish-black, with darker forewings
Shaking a plant, especially in the early morning, will dis- marked by two kidney-shaped white dots, wingspan 38-50 mm (Fig
lodge all leaf-eating caterpillars except leaf rollers. For esti- 4.27).Eggs are hemispherical, ribbed, whitish green becoming pink-
mating the level of infestation, see the budworm chart (Ta- ish brown, laid on undersides of leaves, singly or in small masses.
ble 4.9). For estimating the degree of damage, see Fig 1.3.
Only seven leaf-eating caterpillars are cited often Life History & Host Range
enough to be considered serious pests: Pupae overwinter in soil. Moths emerge in June or July
and promptly lay eggs. Eggs hatch in a week and larvae feed
1. SILVER Y-MOTH or GAMMA MOTH for up to 90 days before moving underground in autumn.
Atltogvapha gamma (Linnaeus) 1758, Lepidoptera; Noctuidae. Only one generation arises per year. Blunk (1920) rarely
=Plusin gnmmn (Linnaeus), =Phytometra gamma Linnaeus found M. persicariae in hemp, while Dempsey (1975) consid-
Description: Larvae are 12-footed "semiloopers" with only three ered the species a major pest. Kaltenbach (1874), Kirchner
palrs of prolegs. They are dark green, pinched towards the head,
(1906), GoidBnich (1928), and Rataj (1957) also report it in
marked by a white dorsal stripe and two lateral yellow lines, and
reach 20-30 mm fully grown. Head and legs have black markings. central Europe. M. persicariae prefers feeding on Urtica, Platz-
They pupate on plants inside a silken cocoon. Moths are brown bod- tago, Salix, and Sambl~citsspecies, but is polyphagous.
ied with forewings sporting a silvery "Y" mark (or Greek ganzma)
against a grey-brown background, wingspan 30-45 mm (Fig 4.27). 3. CABBAGE MOTH
Eggs are white, round, and ribbed. Mamestra brassicae (Linnaeus) 1758, Lepidoptera, Noctuidae.
=Barnthra brassicne Limaeus
Description: Young larvae are yellow-green, their dorsum darken-
ing to a green-brown colour by the fourth instar. They have yellow
heads and side stripes resembling a chain of dark semilunar spots;
rusty brown setae cover the body. Full-grown larvae measure 40-50
mm in length and have a small hump on abdominal segment eight.
Adults are brown Noctuid moths, nearly indistinguishable from
other species except for a curved dorsal spur on the tibia of the fore-
leg (Fig 4.27).Eggs are transparent (turning light yellow to brown),
ribbed, and laid in batches of 50 or more, in single-layered geomet-
ric rows.

Life History & Host Range

M. brassicae overwinters as pupae in soil. Adults emerge
and lay eggs in May-June. Young larvae act as cutworms;
later they eat leaves. Two generations arise per year. The
species is highly polyphagous and a serious pest of vegeta-
ble crops. It attacks hemp in Europe (Martelli 1940, Rataj
1957, Ferri 1959) and Japan (Takahashi 1919, Harada 1930,
Clausen 1931, Shiraki 1952).The Japanese report that it also
feeds like a budworm. Yepsen (1976) suggested Cannabis
could repel this pest from other crops. M.brassicae ranges
across the Old World, and is somewhat tolerant to Bt.
Figure 4.27: Adult moths of some leaf-eating caterpillars. 4. GARDEN TIGER MOTH
A. Autographa gamma; B. Melanchra persicariae; C. Arctia caja (L.) 1758, Lepidoptera; Arctiidae.
Mamestra brassicae (not to scale). All illustrations by Hill Description: Larvae are black woollybears with white spots
(1994). on their flanks. The woolly hairs contain high concentrations ofhis-
tamine (Rothschild et a/. 1979).Moths have variegated forewings of
beige and brown, hindwings of bright orange with iridescent blue
Life History & Host Range
spots, wingspan 50 mm. Moth bodies are brown dorsally, orange
Females lay 500-1000 eggs on undersides of leaves, sin- underneath. The stiff spines growing from moths can inflict a se-
gly or in small groups. The life cycle takes 45-60 days. Two vere sting (Rothschild et al. 1979).
cycles arise per year in Europe, but up to four cycles occur in
Israel (Hill 1983).Caterpillars skeletonize leaves when young, Life History & Host Range
but later eat the entire leaf lamina. Larvae attack seedlings This species is distributed worldwide north of the equa-
(Kaltenbach 1874, Kirchner 1906), and leaves or flowers of tor. Harada (1930) and Shiraki (1952) reported this pest in
mature plants (Blunk 1920, Goidanich 1928, Martelli 1940, Japan; Sorauer (1958) reported it in Europe. But when Roth-
Ceapoiu 1958, Spaar et al. 1990). Dempsey (1975)considered schild et al. (1977) fed A. caja a pure Canrzabis diet, the larvae
 Chapter 4: Insects and Mites 59

became stunted and died before maturity. Given a choice, Ghani 1972). Deshmukh et al. (1979) offered S. obliqua larvae
A, caja larvae preferred eating high-THC plants to low-THC, 154 different plant species, and found Cannabis among the
high-CBD varieties. Yet their survival was higher when they six most preferred hosts. Yet, when fed a diet of pure Caiina-
fed upon the latter. bis, 50% of the larvae died after 24 days.
A. caja is aposematic - the moths are conspicuously Lago & Stanford (1989) reported another yellow
coloured, and the colours senre as a warning to predatory woollybear, Spilosoma (Diacrisia) virginica (F.),feeding on
birds and rodents. Moths store plant poisons in their bodies Mississippi marijuana (Fig 4.28).
which repel predators, just like their famous cousin-the
monarch butterfly. Birds will eat A. caja without hesitation 6. BEET WEBWORM
on the first occasion, but subsequently refuse to eat the dis- Loxostege sticticalis (Linnaeus) 1761, Lepidoptera;Pyraustinae.
tasteful moths (Rothschild et al. 1979).A. caja larvae arepoly- =Phlyctaenodes sticticalis Linnaeus
phagous, but show a predilection for poisonous plants, in- Description: Larvae are yellowish-green, darkening with age to
cluding Acoizifzlnz, Senecio, Convalaria, Digitalis, Solanurn, and nearly black, with a black stripe down their backs, reaching 30 mm
Urtica (Rothschild et al. 1979). One A. caja larva feeding on in length. Moths are mottled brown, with straw-coloured markings
and a dark margin on the underside of their hindwings.
C a n n a b i s can store e n o u g h THC in its cuticle to be
pharmacologically active to a mouse (Rothschild et al. 1977).
Life History & Host Range
Berenji (pers. commun. 1997) reported another Arctiid,
the Fall webworm (Hyphantria cunea Drury), infesting hemp Larvae overwinter in silk-lined cells in the soil, pupat-
in Yugoslavia. H. cilizea larvae web leaves and flowers into ing there in the spring. Moths emerge from March to June.
Moths are strong night fliers, migrating hundreds of miles
an unsightly nest. Greatest damage arises from August to
October. Larvae are black and white with long, whitish hairs. along river valleys with prevailing winds. Females lay 150
Two generations arise per year. H. cunea normally infests eggs, in groups of two to 20, on undersides of leaves. The
trees and shrubs. It is a gift from America to the Old World. entire life cycle can be as short as 35 days, with up to three
generations arising each year.
5. COMMON HAIRY CATERPILLAR This polyphagous pest is also known as the meadow
Spilosoma obliqua (Walker) 1862, Lepidoptera;Arctiidae. moth or steppe caterpillar. It infests leaves and flowering tops
=Diacrisia obliqua (Walker) throughout eastern Europe (Kulagin 1915, Ceapoiu 1958,
Description: S. obliqua woollybears have yellow bodies, orange Camprag et 01. 1996). L. sticticnlis ranges across temperate
stripes, dark heads, and long, light-coloured hairs. Moths are Asia and has been introduced into the USA and Canada
aposematic, with orange abdomens and yellow wings that are lightly (Metcalf et al. 1962).Young larvae skeletonize hemp leaves,
spattered with black spots, wingspan 40 mm. often webbing several leaves together, while older larvae eat
entire leaves. They occasionally march e n nlasse as
Life History & Host Range armyworms.
This species overwinters as pupae just beneath the soil
surface. Young larvae skeletonize leaves; later they com- 7. HEMP DAGGER MOTH
pletely consume leaves. S. obliqi~asometimes infests flower- Plataplecta coiisaizgzlis (Butler) 1879, Lepidoptera;Noctuidae.
ing tops. Females lay eggs in large conspicuous masses, up = Acronycla consanguis Butler
to 1000 at a time. In warm climates the life cycle takes 30 Description: Butler originally placed this species in the
days; three to eight generations arise per year. Bombycoidea. He only described the adult moths, which are grey
S. obliqua ranges from the Himalaya foothills of India to with lavender and brown wing markings, with 35 mm wingspans.
Japan.The larvae are polyphagous, but like many aposematic
insects, they prefer eating poisonous plants (Rothschild et Life History & Host Range
al. 1979).S . obliquu caught the attention of biocontrol research- Shiraki (1952)called this species the hemp dagger moth,
ers in India (Nair & Ponnappa 1974) and Pakistan (Baloch & found only in Japan. We know little about its life history and
host range.

8. CHRYSANTHEMUM WEB WORM

Cnephasin interjectana (Haworth) 1829, Lepidoptera;
Tortricidae. =Cnephsia virgaureana Pierce & Metcalfe 1922
Description: Larvae vary from brownish-cream to greyish-green in
colour. They are darker dorsally with yellow-brown heads and legs.
Pupae are brown, found in webbed leaves and at the bases of stems.
Moth forewings are whitish-grey with brown markings.

Life History & Host Range

Larvae overwinter in soil or crop debris. Moths emerge
in late summer. They deposit eggs on plants, singly or in
batches of two to three eggs. Fritzsche (1959) observed C .
interjectana feeding on hemp in the Halle-Magdeburg region
of Germany. First-instar larvae feed as leafminers. Older lar-
vae roll and web leaves (around mid-May), and move to flow-
ering tops in autumn. The species occurs across Europe into
Siberia. It has established a North American foothold in New-
foundland. C. interjectana feeds polyphagously upon over
130 herbaceous species, notably C k ysanthenzllnz, buttercups
Figure 4.28: The woollybear, Spilosoma virginica. (Ranilncillzls species), sorrel (Rllntex species), and plantain
A. Female moth; B. Pupa; C. Dark form of larva; (Plantago species). Goidanich (1928) and Spaar et al. (1990)
D. Light form of larva (courtesy of USDA). cited the related pest Cnephasia zuahlbomiana L. on Euro-
pean hemp.
60 Hemp Dise!ases and Pests

9. DEATH'S HEAD MOTH with which they stab their prey. Young nymphs are red with
Acllerontia atropos (Linnaeus) 1758, Lepidoptera; Sphingidae. black markings, older nymphs develop red, black, yellow-
= Splzinx atropos Linnaeus orange, and cream-white markings on their abdomen. Eggs
Description: Larvae are "hornworms," initially light green, dark- vary from yellow to copper in colour, oval, spined, 0.8 mm
ening with age to dark green or brown, with yellow lateral stripes tall, and deposited in clusters of 20-25 eggs.
and dorsal spots in the final instar, up to 130 mm in length; the dor- DEVELOPMENT: Adults and nymphs search for prey,
sal "horn" grows from the back of the eight abdominal segment in
an s-shape. Pupae are smooth and glossy, dark brown, 75-80 mm
stab them, then suck them dry. They may kill 100 prey dur-
long, with transverse ridges and two short apical spines. Moths are ing their lifespan. Recorded hosts include leaf-eating cater-
heavy-bodied ("the weight of a mouse" according to Young 1997), pillars, cutworms, armyworms, budworms, European corn
with striped, spindle-shaped abdomens tapering to a pointed pos- borers, grubs of bean beetles, and grubs of flea beetles. Adults
terior; dorsal surface of the thorax with a black-and-yellow skull- overwinter, each female lays hundreds of eggs. Adults may
like marking. Forewings are long and narrow, dark brown with live two or three months, and two or three generations arise
mottled white and tan markings, wingspan 90-130 mm; hind wings per year.
vellow with brown-blackbands. When disturbed, the moths can for-
APPLICATION: Supplied as older nymphs mixed with
cibly expel air from their pharynx, to produce a hissing or shrieking
sound. Larvae click their mandibles if molested, and large ones can loose filler. Store a maximum of one to two days at 8-10°C.
inflict a painful bite. For light infestations, release one predator per m2; for heavy
infestations, release five predators per m*. Repeat releases
Life History & Host Range at two-week intervals.
A. atropos overwinters as larvae or pupae in Mediterra- NOTES: Adults have wings and may fly from the intro-
nean Africa and the Middle East. Pupation occurs 15-40 cm duction site. I? macl~liventrisis compatible with Bt and NPV.
deep in the ground. Multiple overlapping generations arise It is very susceptible to organophosphate and carbamate
every year. Moths migrate north to Europe, beginning in June, pesticides, and less susceptible to synthetic pyrethroids and
and produce one European generation which rarely survives synthetic growth hormones.
past October. Larvae are polyphagous, feeding on many crop
plants, especially potato, tomato, and tobacco. Larvae cause Tanacetum vulgare Common tansy
minor damage on hemp (Kaltenbach 1874, Kirchner 1906, BIOLOGY: An aromatic perennial cousin of chrysanthe-
Goidanich 1928, Dempsey 1975, Gladis & Alemayehu 1995). mums, also known as bitter buttons or golden buttons.
Adult moths are night fliers and dive into hives to rob honey. APPLICATION: Yepsen (1976) repelled caterpillars from
crops by planting this herb nearby. T.vlllgare grows over 1.5
DIFFERENTIAL DIAGNOSIS m tall (the ornamental cultivar is shorter, T. vulgare var.
Stem borers feed on leaves before they begin boring. crispllm). The thujone in tansy drives away many insects.
Budworms occasionally act as leaf eaters. Early leafmining
damage may be confused w i t h skeletonized leaves. Hypl~antriacunea NPV HcNPV
Armyworms eat leaves, but are discussed separately due to BIOLOGY: Different strains of this DNAvirus have been
their marching habits. Some beetles feed on leaves, but gen- used in Japan and the eastern Adriatic region (Hunter-Fujita
erally make smaller holes. Slugs make ragged holes. Holes et al. 1998).For more information on NPV, see the section on
from brown leaf spot (a fungus disease) and holes from bac- cutworms.
terial blight can be confused with holes from leaf-eating in-
sects. CHEMICAL CONTROL (see Chapter 11)
Spraying plants with neem repels caterpillars. Or try a
CULTURAL & MECHANICAL CONTROL repellent solution of garlic, hot peppers, and horseradish.
(method numbers refer to Chapter 9) Aqueous extracts of Cannabis repelled moths of Pieris brassicae
Follow recommendations for cutworms a n d L. trying to lay eggs on cabbage plants (Rothschild &
armyworms, especially methods 2a (deep ploughing), 3 Fairbairn 1980).To kill caterpillars, combine pyrethrum and
(weeding), 6 (crop rotation), 8 (optimizing soil structure and rotenone. Sabadilla dust also works. Spinosad kills caterpil-
nutrients), 12d (nocturnal moth traps-especially against lars in cotton crops. Synthetic pheromones are available for
Nocturids) and 13 (barriers coated with Tanglefoot). A. gamma, C. interjectatza, L. sticticalis, and M . brassicae.

BIOCONTROL (see Chapter 10)

Follow suggestions in the previous section-spray Bt
when caterpillars are less than 10-15 mm long (unless you THRIPS
have M. brassicae, which is resistant). Spray with NPV (espe- Thrips (the word is singular and plural) are tiny, slen-
cially if your pest is a Nocturid), a NPV strain specific to H. der insects. Adults have wings but fly poorly. They prefer to
cunea is described below. Release Trichogramma wasps (de- jump, and they spring quickly to safety when confronted.
scribed under European corn borers), and Steinernema nema- Reports indicate at least five genera of thrips attack Cantza-
todes (described under cutworms). A general predator, bis. They pose a problem in modern glasshouses that use
Podisz~sn~act~liventris,is described below. Consider planting rockwool and hydroponics. In old soil-floored glasshouses,
tansy, described below. watering with a hose kept floors damp, which encouraged
the fungus Entomophthora thripidiim. E. thripidlim infects
Podisus maculivent~isPodibugO thrips when they drop to the ground to pupate. With no
BIOLOGY:Apredatory stink bug with a wide host range; damp soil, there is no fungus, and no natural biocontrol.
it is especially fond of caterpillars and beetle grubs in foliage
of many vegetable crops. P. rnacl~liventrisdoes best in tem- SYMPTOMS
perate climates. Immature and adult thrips puncture or rasp plant sur-
APPEARANCE:Adults are called "spined soldier bugs"- faces, then suck up the exuded sap. This gives rise to symp-
they are shield-shaped, khaki coloured, 8-12 mm long, with toms of white specks or streaks (sometimes coloured silver
prominent spurs behind their heads, and a swordlike snout or yellow, Plate 27), appearing first on the undersides of
 Chapter 4: Insects and Mites 61

leaves or hidden within flowers. Infested plants become cov- ing in glasshouses. Each female lays about 45 eggs. The H.
ered with tiny black specks of thrips faeces. Leaves may curl haemorrhoidalis life cycle turns in as little as 30-33 days at 26-
up. In extreme infestations, plants wither, brown, and die. 28°C (Loomans & van Lenteren 1995). This species pupates
on foliage of host plants (Hill 1994).The rectal fluid secreted
1. ONION (TOBACCO) THRIPS by larvae gums up parasitoids and repels predators. Larvae
Thrips tabaci Lindeman 1888, Thysanoptera; Thripidae. and adults are rather sluggish, compared to T.tabaci.
Description; Adults vary in colour from pale yellow to dark
brown, are pointed at both ends, and grow to 1.2 mm in length. 3. MARIJUANA THRIPS
Female's wings are slender and fringed with delicatehairs (Fig4.29). Oxythrips cannabensis Knechtel1923, Thysanoptera; Thripidae.
Eggs are white and kidney-shaped, oviposited in stems or leaves, Description:Adults are yellow, turning brown near their tails.
and all but invisible to the naked eye. Larvae are small, pale, wing- Females reach 1.6 mm in length, and males are 1.3 mm. Larvae are
less versions of the adults, barely visible without a hand lens. This smaller, lighter in colour, and wingless (Fig 4.30).
species pupates in the soil; pupae look like larvae with wing buds.
Life History & Host Range
Life History & Host Range This species appears to be host-specific on Cannabis. It
The life histories of many thrips are nearly identical. has been isolated from hemp in Romania, the Czech Repub-
Outdoor thrips overwinter in soil and plant debris. Glass- lic, Siberia, and the USA. 0. cannabensis infests leaves and
house thrips do not hibernate. Most thrips stir into activity female flowers of feral hemp growing in Illinois (Stannard et
when the temperature reaches 16°C. Thereafter, the warmer al. 1970) and Kansas (Hartowicz et al. 1971).Hartowicz et al.
the temperature, the worse their damage. Females lacking considered it a possible biocontrol agent against marijuana.
ovipositors usually lay eggs in cracks and crevices. Those They noted that 0. cannabensis is a potential vector of plant
with ovipositors insert their eggs into leaf and stem tissues, pathogens. Little is known of its life history.
well protected from biocontrols. Eggs hatch in three to ten
days under favourable conditions. First and second instars
feed voraciously, and become full grown in less than a month.
Third and fourth instars (called pre-pupae and pupae) gen-
erally become inactive, do not feed, and most go under-
ground. During pupation, thrips acquire wings (in some
species males do not have wings). Males are rare and repro-
duction is usually parthenogenic. Both mated and unmated
females produce eggs; virgin females generally produce only
females. In the field, four generations arise per year, and up
to eight generations arise indoors.
Onion thrips attack a wide variety of vegetable and field
crops. They are very active insects, especially when dis-
turbed. Indoors, T. tabaci has pestered afghanica cultivars in
glasshouses (Potter, pers. commun. 1999).Outdoors, the pest
attacks fibre hemp, where it transmits hemp streak virus
(CeapOiu 1958) and the pathogen sun- Figure 4-30;Adult female marijuana thips, OxythrjW.
flower virus (Traversi 1949).
cannabensis (McPartland, LM x48).

4. INDIAN BEAN THRIPS

Caliothrips indicus (Bagnall) 1913, Thysanoptera; Thripidae.
= Heliothrips indictls Bagnall
Description: Eggs are oval, white, and oviposited in the upper sur-
faces of leaves near main veins. Larvae are small, pale, wingless
versions of adults; they feed along veins. Adults are blackish brown
with brown and white banded forewings. Females reach 1.2 mm in
length. Males are shorter (0.9 mm) and lighter brown.
Figure 4.29: Egg, larvae, and adult of the onion thrips,
Thrips tabaci (about x25, courtesy USDA). Life History & Host Range
Indian bean thrips attack flax and various legumes. The
species is limited to the Indian subcontinent. C. indials closely
2. GREENHOUSE THRIPS resembles the North American bean thrips, Caliothrips
Heliothrips haemorrhoidalis (BouchC)1833,Thysanoptera;Thripi- fasciatus (Pergande).Cherian (1932) considered C. indict~sa
dae. minor pest in Madras. Mated females produce an average of
Description: Larvae are wingless, slender, and light yellow- 65 eggs, unmated females average 51 eggs. In tropical zones
brown to almost white in colour. They waddle around with a glob- C. indicus completes its life cycle in 11-14 days. Indian bean
ule of black faecal fluid protruding from their anus; the globule thrips are very active insects. Even the pupae can jump when
grows until it falls off and a new one begins to form. Adults have disturbed.
dark brown to black bodies ornamented by a reticular pattern, light-
coloured legs, and glassy-appearingwings which look white against 5. WESTERN FLOWER THRIPS
the black body. Adults are up to 1.3 mm long. Frankliniella occidentalis (Pergande) 1895, Thysanoptera;
Thripidae.
Life History & Host Range Description: Eggs are kidney-shaped, 0.25 x 0.5 mm, and
Greenhouse thrips are tropical, and found in warm oviposited in leaves near main veins. Larvae are cigar-shaped,white
glasshouses everywhere. They attack almost all plants grow- to pale yellow, red-eyed, and rather slow moving. They feed along
62 Hemp Dis;eases and Pests

veins of leaflets. Adult females vary in colour from pale yellow to doors (Aeolotlzrips intermedit~s,Franklinotllrips vespformis), all
dark brown, with darker bands across the abdomen, light-coloured discussed below. Hypoaspis miles (discussed under fungus
heads, and dark eyes. Forewings are light-coloured,with two longi- gnats) c a ~ oprovide
t thrips control by itself, but can reduce
tudinal veins bearing two rows of delicate hairs. Females reach 1.3- populations of soil-pupating thrips by 50%. Less effective
1.7 mm in length. Males are shorter (0.9-1.1 mm) and uniformly
\,ellow. thrips predators include lacewings (Chryosopa carnea) and
ladybeetles (Hippodanzia convergens, Coccinella undecim-
Life History & Host Range pt~nctata).Most thrips predators cannot be mixed together,
F. occidetltalis is California's gift to glasshouses around except for a combination of Neoseiulzls, Orius, and Hypoaspis.
the world. It invaded Europe around 1983 (in Holland) and Parasitoid wasps include Thripobius semilutetis, Cerani-
has spread throughout the continent. Surprisingly it has not slis menes, and Goetheana shakespearei, described below. The
yet been cited on Cannabis, probably for lack of recognition. life cycles of these parasitoids are longer than their hosts, so
Lago & Stanford (1989) described a relative, Frankliniella they work best as preventatives.
fusca (Hinds), on Mississippi marijuana. Beneficial soil nematodes control soil-pupating thrips.
F. occidentalis infests at least 250 species of plants, and Heterorkabditis bacteriophora (described under white root
vectors many viruses. Females of the species preferentially grubs) works better against thrips than Steitzernemafeltiae or
infest flowers and buds. At optimal temperatures (26-2Y°C), Steinernema riobravis (Cloyd et al. 1998). Nematodes will not
the life cycle turns in seven to 13 days. Adult females live reproduce in thrips cadavers, so they must be reintroduced
another 30-45 days and lay 150-300 eggs. Mated females are several times during the season.
more fecund than unmated ones. Females feeding on leaves Turning to biocontrol with fungi, Cloyd et al. (1998) re-
and pollen are more fecund than females feeding solely on ported better control of F, occidetztalis with Metarhizit~m
leaves. This species pupates in flowers or in soil (usually 1.5- anisopliae than with Verticillit~nzlecanii. Both fungi are de-
2.0 cm deep). scribed under aphids. Hussey & Scopes (1985)killed T. tabaci
and other thrips with the Mycotaln strain of Verticillit~m
DIFFERENTIAL DIAGNOSIS lecatzii. Hussey & Scopes also controlled T, tabaci with
Immature thrips nymphs are practically impossible to Beaziveria bassiana, spraying twice at ten-day intervals with
differentiate by species. Even the adults can be hard to tell a 0.25% water suspension of Boverinn. B. bassiana is described
apart. Thrips damage resembles that of other sap-sucking under whiteflies. Entornopktlzora tlzripidum and E. pawispora
insects, especially spider mites. But mites cause tiny, round are described below. Fungi kill more adults than larvae, pos-
lesions on the upper leaf surface, whereas thrips leave ir- sibly because many adults prefer feeding in flowers (where
regular-shaped (often elongate) lesions on undersides of humidity is higher and more favourable for fungi), and be-
leaves, lesions often fill the spaces between leaf veins. Thrips cause larvae moult periodically, shedding spores attached to
nymphs can be confused with leafhopper nymphs. their cuticle (Cloyd et al. 1998).
Onions (Allium cepa) and garlic (Alliz~nzsatiot~m)serve
as trap crops for T. tabaci. Infested trap crops can be sprayed
Table 4.10: Infestation Severity Index for thrips with pesticides or bagged and removed from the growing area.

Neoseiulus (Amblyseius) cucumeris

Light any thrips damage seen BIOLOGY: N. cl~cunzeriswas the first predatory mite ob-
served to attack thrips, in 1939. It feeds onyoung (first instar)
Moderate thrips damage and occasional larvae and eggs; in the absence of thrips it survives on alter-
thrips seen native food sources-pollen and mites. N. c~~ct~nzeris does
best in moderate temperatures (19-27°C) and moderately
Heavy thrips damage on many plants but high humidity (70-80% RH). It was originally sold for
confined to lower leaves OR control of T. tabaci, but works well against E occidetztalis and
2-1 0 thrips per leaf other thrips.
APPEARANCE &DEVELOPMENTN. cuct~merisadults are
Critical thrips damage on growing shoots elongate-oval in outline, 0.5 mm long, white to beige to pink,
OR >10 thrips per leaf with legs shorter than those of Plzytoseiult~spersimilis (Plate
28). The life cycle takes six to nine days at 25°C. Adults live
another month. Adults consume up to six thrips per day and
CULTURAL & MECHANICAL CONTROL lay two or three eggs per day (Houten et al. 1995). Most
(method numbers refer to Chapter 9) strains of N. cz~ct~meris enter diapause under short
Sanitation is important, so observe methods l a and l b . photoperiods (Gill & Sanderson 1998), but some new com-
In thrips-infested glasshouses, remove all plant residues af- mercial strains do not (Ravensberg, pers. commun. 1998).
ter harvest and heat the house for several days, to starve any APPLICATION: Supplied as adults in shaker bottles or
remaining thrips. Water-stressed plants are particularly sus- supplied as all stages in sachets. Sachets are small paper bags
ceptible to thrips damage, so follow method 7a. Thrips dis- filled with 300-1000 mites (all stages), a food source, and a
like getting wet, so frequent misting may slow them down. tiny exit hole. Hang sachets on plants to deliver a "slow re-
Method 12a (repellent mulch) works for young, low-grow- lease" of mites. When using sachets, avoid very high hu-
lng plants, and method 12c (sticky traps, blue or pink) works midity (>YO%RH), which causes sachets to mould (Cherim
for all plants. Glasshouses can be barricaded with thrips 1998). Store shaker bottles up to one to two days in a cool
microscreens (method 13). (8-15"C), dark place. Mites in sachets can be stored a little
longer. N. cticumeris can be sprinkled directly on plants, sprin-
BIOCONTROL (see Chapter 10) kled into distribution boxes (Plate 85), or heaped on rockwool
Predators have been released indoors (Neoseit~lt~s blocks. Used preventatively, release 100 mites per m2 per
clicumeris, lplziseitis degenerans, Neoseilllus barkeri, El~seil~s month (Reuveni 1995). For light to moderate infestations,
hibisci, Deraeocoris brez~is,and several Orills species) and out- release 100-500 mites per plant every two to three weeks
 Chapter 4: 11?sects and Mites 63

(Thomson 1992), or 200-300 mites per m2 per week until the per day and lay one to two eggs per day (Houten et al.
infestation subsides. 1995).Each mite eats about 85 thrips larvae during its life-
NOTES: N. ctictimeris survives well without prey, which time (Riudavets 1995). Unfortunately N. barkeri enters
makes it a good preventative biocontrol. N,ctictimeris prefers diapause with short photoperiods.
living on the undersides of leaves, rather than flowers (Cloyd APPLICATION: Supplied as adults in pour-top bottles.
et al. 1998). This trait, along with its need for high humidity Store a maximum of up to two or three days in a cool (8-
and its propensity to diapause, make N,ctictimeris a subopti- 1O0C),dark place. To control T. tabaci, researchers released
mal choice for flowering plants. Results have been poor in glass- 50,000 predators per ha every 14 days, or 15-20 per m2
house Cannabis (Watson, pers. commun. 1998). glasshouse, or 1200 per plant, or three per leaf (Riudavets
N.ctictimeris is compatible with Orit~stristicolor and 0 . in- 1995).
sidioslis. When released with other predatory mites, such as P. NOTES: N.barkeri does not work as well as N. czictinleris
persimilis and I. degenerans, the predatory mites prey upon each (lower predation and oviposition rates), and commercial
other (in the absence of pests), but the predators can coexist strains diapause under short days (Houten et al. 1995).
(Hussey & Scopes 1985). Some N.cltczlmeris strains are suscep- Nevertheless, mass rearing of N. barkeri is relatively easy,
tible to the biocontrol fungi Verticillizim lecanii and Beatlueria and the species is somewhat resistant to pesticides. N.
bassiai~a,SO they are not compatible in conditions of high hu- barkeri may be compatible with other predatory mites. This
midity (Gilkeson 1997).N,cucllnzeris tolerates most insecticides predator is therefore best suited for preventive control on
better than I. degenerans and other predatory mites, but it is vegetative plants. N.barkeri, like N.clicllrneris, has been
very sensitive to pyrethroids and organophosphates. Some plagued by protozoan infections that depress its effective-
companies mass-rearing N.cticlimeris were plagued by proto- ness (Gilkeson 1997).
zoan infections in the recent past. Protozoans cause sublethal
infections of N,cuclinzeris, so the infected biocontrols are sold Euseius (Amblyseius) hibisci
and exported to release sites (Gilkeson 1997). Infected BIOLOGY: A mite that preys on young larvae of E
biocontrols are less effective, and they may spread infection to occidentalis and other thrips,
.
spider mites, citrus mites, and
-

other biocontrol mites. immature whiteflies. It lives in southern California avocado

and citrus orchards, and loves dry, subtropical conditions.
Ipkiseitis (Amblyseius) degenerans APPEARANCE &DEVELOPMENTEggs are oblong and
BIOLOGY Apredatory mite that attacks the first larval stage almost transparent. Larvae are also transparent. ~ d u l t ares
of T.tabaci and F. occidentalis. In the absence of thrips, I. pear-shaped, very shiny, and smaller than other predatory
degeneraizs preys on spider mites. In the absence of any prey, I. mites. Their colour depends on what they are eating-
degeilerans survives and reproduces on pollen. The predator is yellow when feeding on thrips larvae, white when feed-
native to Eurasian and African subtropical regions, and pre- ing on pollen, and red when feeding on citrus mites. The
fers moderate humidity (55-80% RH), and warm temperatures life cycle takes nine to 12 days at 26°C. Adults live another
(21-32°C). 25-30 days. Adults consume three or four thrips per day
APPEARANCE:Adults are dark-brown, oval, 0.7 mm long, and lay one to two eggs per day (Houten et al. 1995). E.
and mobile. Larvae have a brown X-mark on their back. Eggs kibisci does not diapause (Houten et al. 1995).
are transparent and turn brown before hatching, laid along NOTES: E. hibisci eats and reproduces slower than N.
veins on undersides of leaves. cuctinzeris and I. degenerans. In theabsence of pests, E. hibisci
DEVELOPMENT: I. degenerans does not diapause under actually reproduces faster on pollen. It tolerates drought
short daylengths. The life cycle takes ten days. Adults live an- better than any other predatory mites (Houtenet al. 1995).
other month. Adults consume four or five thrips per day and Drought tolerance and lack of diapause make this mite a
lay one to two eggs per day (Houten et al. 1995). good preventative biocontrol in flowering plants. Natural
APPLICATION: Supplied as adults in shaker bottles, or a populations of E , hibisci have developed resistance to many
mixture of adults, nymphs, and eggs on bean leaves in plastic organophosphate insecticides, but are still sensitive to
tubs. They are stored the same as N.aiainzeris. Used preventatively, carbarnates and pyrethroids. Unfortunately, this biocontrol
release two to four mites per m2 per month. or light infesta- must be reared on leaves, which hampers its commercial
tions, release ten mites per m2 every two weeks. For moderate availability.
infestations, release 20 mites per m2 weekly (Cherim 1998).
NOTES: I. degenerans works better in flowering crops than Orius species "Pirate bugs"
N. ci~cz~rneris-it tolerates lower humidity, does not diapause, BIOLOGY: Six different species of Orius are available
migrates well in tall plants, and prefers living in flowersiather (described below). These predators are polyphagous; af-
than leaves (Cloyd et al. 1998). Nevertheless, results have been ter thrips are gone, pirate bugs eat other pests (aphids,
poor in sinsemilla Cannabis, because of the lack of pollen as an mites, scales, whiteflies), then other biocontrols, then each
alternative food source (Watson, pers. commun. 1998). 1. other. They do best in temperate climates, optimally 60-
clegenerans is compatible with Oritis but may feed on N. 85% RH and 20-30°C.
czlctrmeris. It tolerates neem and many fungicides,
- but is more APPEARANCE:Adults are yellow-brown to black with
sensitive to insecticides and miticides than N.cucumeris, espe- white wing patches, 1.5-3 mm long (Plate 29). Nymphs
cially soap, sulphur, and pirimicarb. are teardrop shaped, bright yellow to brown, with red eyes.
DEVELOPMENT: Adults overwinter; females lay 30-
Neoseiulus (Amblyseius) barkeri (=Amblyseius mackenziei) 130 eggs in plant tissue. The life cycle takes as little as two
BIOLOGY A mite that preys on T. tabaci, F. occidentalis, and weeks. Adults live another month. Usually two to four
spider mites (Tetranychlis urticae). It occurs naturally around generations arise every year. Oritis species move quickly
the world, and does best at 65-72% RH and moderate tem- and catch prey with their forelegs. Young nymphs feed
peratures (19-27°C). primarily on pollen or young thrips. Older nymphs and
APPEARANCE & DEVELOPMENT: Adults resemble N . adults feed on all stages of thrips; adults eat five to 20 thrips
cilc~irneris,perhaps a deeper tan colour. N.barkeri can repro- per day. Unfortunately, many Orills species diapause un-
duce every 6.2 days at 25°C.Adults consume two or three thrips der short day lengths.
64 Hemp Diseases and Pests

APPLICATION:Supplied as adults and nymphs in shaker China (Fig 4.39). The species associates with many plants,
bottles. Store for up to three days in a cool (10-15"C), dark including Urtica species. 0 . minuttis prefers T. tabaci but will
place. Used preventatively, release one predator per plant eat spider mites (Tetranychusspecies), eggs of European corn
(Ellis & Bradley 1992) or one predator per m2 (Reuveni 1995), borers (Ostrinia ntibilalis), and aphids (e.g., Aphis gossypii).
make two releases two weeks apart. Moderate infestations Anthocoris nemorum is a minute pirate bug (Fig 4.39)
may require five to ten predators per m2 per week (Riudav- used against thrips in England (van Lenteren 1995). It is re-
ets 1995). Heavy pest infestations cannot be controlled by lated to Oritis species.
Orius species, the biocontrols reproduce too slowly. Koppert
(1998) recommended releasing Orius in groups of 15-20 in- Deraeocoris brevis
sects to encourage mating. BIOLOGY: This predatory mirid bug eats all stages of
NOTES: Adults can fly and find remote infestations, and thrips. It also eats aphids, mites, whiteflies, lygus bugs, and
they like to inhabit flowers. In mixed pest infestations, pi- psyllids. Cherim (1998) said some D. brevis populations ac-
rate bugs will eat the largest pests first. So even if the preda- tually prefer aphids over thrips. D. brevis occurs naturally in
tors are swimming in baby aphids, they would rather eat a temperate climates (optimally 30-70% RH, 18-30°C). A re-
large thrips larva. In the absence of pests, Oritis species eat lated species, Deraeocoris nebtllostls, may also be commercially
predatory mites (but not subterranean Hypoaspis miles).Young available.
Orills nymphs have a hard time with hairy-trichomed plants, APPEARANCE & DEVELOPMENT: Adults are mottled,
adults do better. All Ori~isspecies are sensitive to pesticides. brown bugs 5 mm long. Nymphs are lighter-coloured ver-
0, tristicolor is native to western North America (Fig sions of adults. Adults overwinter; females lay up to 200 eggs
4.39) and widely available. It feeds on F.occidentalis, T. tabaci, in plant tissue. Nymphs begin preying on pests as second
Caliothripsfasciatus, spider mites (Tetranychtls species), aphids instars. The life cycle takes 30 days. Adults live another 21 days.
(Myztis persicae), and eggs of budworms (Helicoverpa zea). 0. APPLICATION: Supplied as a mix of adults and nymphs
tristicolor populations parallel thrips populations; i.e., adult in vials and bottles. Used preventatively, release one preda-
longevity and fecundity increases as the thrips population tor per 10 m2 every two weeks. For light infestations, re-
increases (Riudavets 1995).The predator is reportedly com- lease one to two predators per m2 every week. For heavy
patible with Neosei~iltiscticumeris, although 0 . tristicolor eats infestations, release five predators per m* every week.
other beneficial mites. Cherim (1998) recommended releasing a "critical number"
0.insidiosus is native to the midwestern USA (Fig 4.39). of individuals (2300)to establish a population. In the absence
The predator successfully controlled F.occidentalis in Euro- of prey, adults survive on pollen. Young nymphs move with
peanglasshouses (~iudaiiets1995). It also eats spider mites difficulty across surfaces with glandular trichomes; adults
(Tetranychtis tirtlcae), aphids (Aplzis species), and the eggs of do better. Unfortunately, adults enter diapause with short days.
budworms (Hellcozlerpa zea, H. vlrescens). It is compatible with
Neoseitill~scuclinzerls but eats the aphid predate; Aphidoletes Aeolothrips intermedius
npizidii?iyza (Riudavets 1995). 0. insidiostls diapauses in BIOLOGY: Seczkowska (1969) found this banded thrips
photoperiods under 11 hours, but warm temperatures may preying on T. fabaci in Polish Cannabis fields. It also feeds on
postpone diapause. Frankliniella species, H.hernorroidales, and spider mites
0 , albidipennis ranges across southern Europe east to (Tetratzychus cinnabarinus, T.tirticae). The predator is native
Iran. It eats F.occidentalis, T. tabaci, spider mites (Tefranychlls to Europe, the Middle East, and India.
species), aphids (Aphis gossypii), whiteflies (Bernisia tabaci), APPEARANCE: Adults are yellowish to dark brown,
budworms (Heliothis arinigera), and cutworms (Spodoptera about 1.6 mm long, with relatively broad, banded wings.
lltzira). It coexists w i t h ladybeetles (Coccinella Larvae are lighter coloured than adults.
~inidecinzpunctata), lacewings (Chysopa carnea), and 0. laevi- DEVELOPMENT Females lay 30-70 eggs, with three or
gattis. Riudavets (1995) considered it a promising biocontrol four generations arising each year. Each A, intermedills larva
agent. 0. albidipennis prefers warmer temperatures (up to consumes about 25 T. tabaci larvae. Adults feed on plant pol-
35°C) and does not diapause-which makes it perfect for len and may pierce leaves and stems for water (Riudavets
flowering crops in hot greenhouses (Cloyd et al. 1998). 1995). A. intermeditis is eaten by other predators, so it is not
0. laevigatus hails from the Mediterranean region and compatible with pirate bugs (Oritis species), lacewings
western Europe. It feeds primarily on F.occidentalis and T. (Chysopa species), and ladybeetles (Coccinellaseptenzpunctata).
tabaci, but also eats spider mites (Tetranyclztlsspecies), Euro-
pean corn borers (Ostrinia i~~lbilalis), aphids (e.g., Aphis Franklinothrips vespiformis
gossypii), and cutworms (e.g., Spodoptera litura). It coexists BIOLOGY: Another predatory thrips, related to the pre-
with ladybeetles (Coccinella ~lnidecimptinctata),lacewings vious predator, this species is found in California avocado
(Clzrysopa carnea) and 0 . albidipennis. Riudavets (1995) noted orchards. F. uespiforrnis also feeds on immature aphids,
0. laeuigatus adapts well to glasshouse conditions on many whiteflies, scales, mites, and moth eggs.
crops, and survives without thrips for a long time. 0. laeui- APPEARANCE &DEVELOPMENT:Adults are black with
gatks diapauses in photoperiodLunder 11 hours, but warm white-banded legs; their round heads and thin waists make
temperatures may postpone diapause (Ravensberg, pers. them look like wasps or ants. Larvae have a wide, red band
commun. 1999). covering their abdomen. These predators are less effective
0. majusctllus is common in central Europe (Fig 4.39). than predatory mites and ~hripoiiolissenziluteus, below.
Good control of F. occidentalis was obtained in cucumber
glasshouses using a ratio of one predator per 100 thrips. 0 . Thripobius semiluteus
nzaltisct~lusproduces more eggs than any other Orilis spe- BIOLOGY: A parasitic eulophid wasp that parasitizes H.
cies. But 0 . nzaj~iscz~lt~s
is more sensitive to photoperiod, and lzaemorrhoidalis, is native to Africa, Australia, Brazil, and South
diapauses in days shorter than 16 hours, although warm tem- Africa, and does best at 23°C and 50% RH.
peratures may postpone diapause (Ravensberg, pers. DEVELOPMENT Adults crabwalk across leaf surfaces in
commun. 19991. search of first and second stage larvae. Females lay at least
0,minutus ranges across western Europe to Siberia and 40 eggs. Parasitized hosts turn into black, slightly swollen
 Chapter 4: Insects and Mites 65

mummies. Adults emerge from thrips cadavers to repeat the sprays. Yepsen (1976)concocted a thrips-repellent spray from
lifecycle. T.semilzitetls takes 22 days to develop (Loomans & field larkspur (Consolida regalis or Delyllinitim consolida). Met-
van Lenteren 1995). calf et al. (1962) eliminated thrips with sabadilla dust, or a
APPLICATION: Supplied as pupae in vials; attach vials mix of diatomaceous earth and pyrethrum. Spraying a slurry
to plants horizontally, as high in the plant canopy as possi- of clay microparticles may work. Sprays and dusts gener-
ble (or tape to stakes if plants are small). In orchards, the ally require two or three applications at weekly intervals.
standard release rate is 5000 pupae per ha or 250 pupae per Apply at mid-day, when thrips are most active. Be sure to
tree (Loomans & van Lenteren 1995). T.semilntetis has been treat the undersides of leaves. Imidacloprid (a synthetic nico-
introduced into California and Hawai'i to control H. lzaem- tine) and spinosad (a fermentation product) kill thrips. Meier
orrlloidalis. Do not use botanical insecticides within one week & Madiavilla (1998) eliminated thrips with Cymbush
of release, or other pesticides within 30 days of release. (permethrin). ThripsticksG (polybutenes mixed with
deltamethrin, a synthetic pythrethrum) can be applied to
Ceranisus menes plastic around the bases of plants, so when thrips drop to
BIOLOGY: A tiny Eulophid wasp that parasitizes F. the ground to pupate, they die.
occidentalis and T. tabaci larvae, and lives around the world.
DEVELOPMENT: Female wasps insert ovipositors into
young thrips larvae and lift struggling larvae-into the air;
the larvae soon become paralysed and a single egg is depos-
FLEA BEETLES
ited (Loomans & van Lenteren 1995). C . menes larvae even- Beetles are the most common animals on earth, and the
tually kill their hosts, then exit to pupate in soil. C. nzenes most common beetles on Cannabis are flea beetles (family
wasps also kill thrips by poking them to feed off body fluids. Chrysomelidae). Flea beetles damage plants as adults and
APPLICATION: C . nzenes was used in a large scale effort larvae (grubs). Adults feed on foliage, flowers, and unripe
to eliminate T.tabaci from Hawai'i, but has not been com- seeds. Grubs feed on roots or occasionallv act as leafminers.
mercially available since then. Dense leaf hairs interfere with Many flea beetles are polyphagous and attack a wide vari-
the parasitoid's ability to locate prey. ety of plants. Others specialize, infesting only one or two
plant species. One such specialist is the hemp flea beetle.
Goetheana shakespearei Rataj (1957) considered the hemp flea beetle insignificant,
BIOLOGY:Another Eulophid wasp (sometimes called G. but Ragazzi (1954), Dempsey (1975), and Bosca & Karus
parvipennis), smaller than the aforementioned C . menes. This (1997) cited it as a major pest. Durnovo (1933) considered it
species is native to south Asia and Australia. It was released the number one problem in the North Caucasus. Biocontrol
in California to control young larvae of H. haemorrhoidalis. It researchers thought the species would be an excellent can-
also controls C. indicus. The wasp can complete its life cycle didate for eradicating illegal marijuana (Mohyuddin &
on T.tabaci, but attempts to control T. tabaci have failed Scheibelreiter 1973).
(Loomans & van Lenteren 1995).
APPLICATION: G. shakespearei is more successful as a SIGNS & SYMPTOMS
biocontrol in tropical areas and less effective in temperate The hind legs of adult beetles are large and they leap
climates (Loomans & van Lenteren 1995). Its minute size like fleas when disturbed. Since flea beetles are small and
makes the parasitoid difficult to rear. leap frequently, they do not eat much in one place. Damage
consists of many small, round to irregular holes, formed be-
Entomophthora thripidutn and Neozygites parvispora tween leaf veins (Plate 30). Leaves of heavily infested plants
BIOLOGY:Two closely-related soil fungi that infect thrips may be completely skeletonized. Young plants are killed.
pupating underground. Both species occur naturally in damp Grubs of most species feed on roots, usually at depths of 4-
soil in Europe and North America. N. parvispora is an obli- 8 cm underground. They feed on cambium, making longitu-
gate pathogen of thrips, known from T.tabaci, F.occidentalis, dinal or tortuous galleries in roots. Grubs of one species act
and Linzotlzrips species (Carris & Humber 1998). E. thripidtiin as leafminers, mining beneath the upper surface of leaves,
is not so specialized-it also infects thrips predators. tracking in a tight spiral that ends in a brown blotch.

CHEMICAL CONTROL (see Chapter 11) 1. HEMP FLEA BEETLE

Thwart thrips with tobacco tea or nicotine. Cherian Psylliodes attentiata (Koch) 1803, Coleoptera; Chrysomelidae.
= Halticn attenuata Koch, = Psylliodes japonica Jacoby 1885,
(1932) repelled thrips with a mix of fish emulsion and soap
= Psylliodes apicalis (Steph.)apud Spaar et a/. 1990
Description: Eggs are pale yellow, deposited singly around plant
roots near the soil line. Grubs are cylindrical, white with tiny spots
and bristles, with six minute legs, a brownish head, and reach 4.5
mm in length. Pupae are initially pearly white but slowly darken,
beginning with the eyes, followed by other parts of the head, legs,
and finally the elytra. Adult are oval in outline, black with minute
bronze-grey hairs, and average 1.3-2.6 mm long (Plate 30). Wing
covers are striated and punctuated. Heads exhibit sharp outgrowths
crossed in an "x" at the eyes, and sprout antennae reaching half
their body length (Flachs1936).See Fig 4.31.

Life History & Host Range

Adults overwinter in soil and emerge at the end of March
to feed on young seedlings. Adults mate in April and begin
Figure 4.31 : Larvae, pupae, and adults of the hemp flea laying eggs ten days later. Females lay an average of 2.6 eggs
beetle, Psylliodes attenuata (line drawing on left from per day for a total of 55 eggs (Angelova 1968). Egg laying
Flachs 1936 except side view by McPartland, shaded lasts from late April to the end of July. Larvae hatch in five to
version on right from Senchenko & Timonina 1978). 16 days, and grubs feed within roots when young. By June,
66 Hemp Diseases and Pests

grubs exit roots and live in soil. Pupation occurs from mid 1957). P. malvae normally infests kenaf or marsh mallows.
June to August, 4-15 cm underground. Adults emerge from Hill (1983)reported a Podagrica species chewing holes in Cannabis
pupae from late June to September. Adults feed on plant tops leaves in Thailand, which he called the "cotton flea beetle."
until autumn, then burrow into soil to depths of 20 cm (An- Hartowicz et al. (1971) found a n unspeciated
gelova 1968). Only one generation arises per year. Chaetocnema species infesting feral hemp in Kansas. Kulagin
Adults cause more damage than grubs. Damage peaks (1915) described Chaetocnema hortensis Geoffr. 1785 attack-
twice--on seedlings when adults emerge in early spring, and ing plants near Moscow; the species normally infests grasses.
again in late summer when adults emerge from pupae. Late- Borodin (1915) and Kovacevic (1929) cited Chaetocnema
season beetles may selectively infest female tops and feed concinna Marsh 1802 on hemp. But as Sorauer (1900)pointed
on seeds (Silantyev 1897).I? attenlrata crop damage increases out, this beet-eating beetle is easily confused with the hemp
in warm, dry weather (Flachs 1936).Adults are most active flea beetle. Lago & Stanford (1989) collected Chaetocizema
on warm sunny days; in overcast or wet weather they re- denticulata (Illiger) and Chaetocnema pulicaria Melsheimer
treat to soil cracks, beneath clods of earth, or on the lower from Mississippi marijuana. C. detlticl~latais a tiny (2.5 mm
surface of leaves. Hemp flea beetles also feed on hops and long), oval, black beetle with bronze highlights. C. plilicaria
nettles, especially in the spring (Silantyev 1897). P. attenuata looks similar but is even smaller.
ranges from Great Britain and France to eastern Siberia, Bantra (1976) described an "Altica" species on Indian
northern China, and Japan (where it was called P, japonica). marijuana. Takahashi (1919)cited "Halticaflavicornis Baly"
on Japanese hemp. Heikertinger & Csiki (Coleopterorurn
2. OTHER FLEA BEETLES Catologus Part 116) question whether this species is a mem-
The hops flea beetle, Psylliodes punctulata Melsheimer ber of the subfamily Alticinae (the genus Haltica is no longer
1847, attacks hemp in Canada (Glendenning 1927).I? punc- recognized). Goidanich (1928) cited a number of additional
tlilata is a North American version of P. attenliata. Adults are Chrysomelidae on Cannabis-Chysomela rossia Ill. 1802,
small (2-2.5 m m long), oval, and greenish-black. The bee- Crepidodera ferruginea Scop. 1763, Gastroidea polygoni L.
tles develop bronze highlights after several days in sunshine 1758, Gynandrophthlma cyanea F. 1775, and Ltrperusflavipes
(Fig 4.32).Adults feed on leaves of seedlings, larvae feed on (L.) 1776.
roots. P. pl~nctlilatafeeds on hops, nettles, canola, sugar beets,
rhubarb, and many cruciferous crops. For more details on DIFFERENTIAL DIAGNOSIS
morphology and life cycle, see Chittenden (1909) and Parker Leaf damage from flea beetles can be confused with
(1910). damage caused by leaf-eating caterpillars, weevils, and other
Phyllotreta nemorum (Linnaeus) 1758 attacks hemp in beetles. No other beetles, however, leap like fleas. Grub dam-
Europe (Kirchner 1906, Borodin 1915, Goidanich 1928, age on roots resembles that of maggots or other root-eating
Kovacevic 1929). The grubs are yellow-bodied, brown- beetles. Leafmines by P. nenzorunz must be differentiated from
headed, and small enough (5-6 mm long) to bore into stems leafmines by fly larvae. Some planthopper nymphs resem-
or feed as leafminers (Fig 4.39). Adults are metallic black, ble adult flea beetles, as d o short, black bugs. Grubs of wee-
sport a broad yellow stripe down each wing cover, and av- vils are indistinguishable from grubs of flea beetles, espe-
erage 3.5 mm in length (Fig 4.32). Borodin (1915) cited a re- cially when young.
lated species, Phyllotreta atra (Fabricius) 1775. This small
black flea beetle attacks many vegetable crops in Europe. CULTURAL & MECHANICAL CONTROL
Dempsey (1975) considered Podagrica aerata Marsh (method numbers refer to Chapter 9)
1802 a serious pest of hemp; Kaltenbach (1874)mentioned it Follow methods 1(sanitation), 2a (deep ploughing), and
in Germany. P, aerata usually feeds on brambles (Rubus spe- 3 (especially eliminate nettles and wild hops). Chinese farm-
cles) in central Europe. The closely-related species ~ o d a ~ r i c a
ers use method 4 by harvesting crops early to escape late-
lnalvae Illiger 1807 also attacks hemp in central Europe (Rataj
season flea beetles (Clarke 1995). European farmers use
method 4 by sowing seeds early, resulting in seedlings that
are older, hardier, and able to withstand flea beetles emerg-
ing in the spring (Camprag et al. 1996). Avoid susceptible
cultivars, such as 'USO-9' ('YUSO-9') from the Ukraine. P.
attenuata-resistant lines include southern Russian landraces
and Chinese landraces (Bocsa 1999). For crop rotations,
Camprag et al. (1996) suggested planting hemp crops at least
0.5-1 km away from the previous year's hemp crops.
For horticultural situations, use method 12c (white
sticky traps) and method 13 (protecting seedlings with fine
mesh or screens). Since flea beetles prefer bright sunlight,
consider planting in partial shade. Parker (1913b) stopped
I? ptinctulata from crawling up stems by applying 5-cm bands
of Tanglefoots.
To catch adults on plants, fashion two "flea beetle cones"
out of cardboard or metal. Make them 60 cm across and 20
cm deep. Coat the inner surface with Tanglefoota or other
adhesives. Hold the cones on either side of an infested plant
Figure 4.32: Assorted leaf beetles (all x5). A. Phyllotreta
nemorum (from USDA); B. Diabrotica undecimpunctata
and bring them together quickly, like cymbals, to surround
howardi (from Westcott 1964); C. Popillia japonica (from the plant. Darkness disturbs the flea beetles and they leap
Westcott 1964); D. Chaetocnema concinna (from Hill onto the sticky surface. This treatment should be repeated
1983); E. Psylliodes punctulata (from Senchenko & at least once a week. Larger plants can be treated one limb at a
Timonina 1978). time.
68 Hemp Diseases and Pests

plants in Europe. Durnovo (1933)said damage peaks in May, eaten away by grubs. Activity by moles, sod-digging skunks,
especially in hemp fields adjacent to weedy wastelands. 0. and raccoons indicates grubs are in the soil.
sabulosum is related to Tribolitlm confnstlm, a beetle occasion-
ally eating seeds in marijuana (Smith & Olson 1982). 1. JAPANESE BEETLE
Popillia japonica Newman 1838, Coleoptera; Scarabaeidae.
DIFFERENTIAL DIAGNOSIS Description: Adults of Japanesebeetles feed on Cannabis leaves
Leaf beetles can be confused with weevils and curculios and are described in the previous section. Grubs are plump and
(see two sections below). Flea beetles are small leaf beetles white with a brown head and six dark legs, 15-25 mm long. They
lie curled in a C-shaped position in the soil. Their life cycle is de-
with their own idiosyncracies and control methods (see the scribed in the previous section on leaf beetles.
section above).
2. EUROPEAN CHAFERS
CULTURAL & MECHANICAL CONTROL a. Melolonth hippocastani Fabricius 1801, Coleoptera;
. .
(method numbers refer to Chapter 9) Scarabaeidae.
Use methods 2a (deep ploughing), 3 (weeding), 9 (shake b. Melolonth melolonth Linnaeus 1775, Coleoptera;
plants early in the morning while beetles are stiff with cold), Scarabaeidae.
12c (white sticky traps), and 13 (mechanical barriers). Heavy c. Melolonth vulgaris Linnaeus 1775, Coleoptera;
-
mulching around seedlings offers some protection. Several Scarabaeidae.
Japanese beetle traps are commercially available. The best Description: These species are related to "Junebugs," the familiar
traps use "dual luresr'-food and sex attractants. The sex at- nocturnal beetles attracted to lights in the early summer. Adults have
tractant is a pheromone, Japonilure@.Food attractants include robust brown bodies with reddish-brown ridged elytra and long
eugenol and geraniol. According to Turner et al. (1980), Can- slender spiny legs, averaging 20 mm in length (Fig 4.33). Larvae
nabis produces eugenol and geraniol, so it is no surprise that resemble those of Japanesebeetles but are larger (reaching 44 mm
in length) and their posterior abdomens have smooth, thin skins
the plant is attractive to Japanese beetles. Baited traps work
with dark body contents showing through.
best in sunny spots during the early summer, set away from
desirable plants. Life History & Host Range
Grubs hibernate in soil deep beneath the frost line (Met-
BIOCONTROL (see Chapter 10) calf et al. 1962 found grubs 1.5m underground). In the spring,
In humid conditions, adult beetles are susceptible to the
grubs return from the deep to feed on shallow roots, spend
mycoinsecticide Metarkizil~nianisopliae (described under
the summer close to the surface, and then return to their
aphids). The tenebrionis strain of Bt kills Japanese beetles and
deep winter cells. Grubs may run this cycle a third year, de-
other leaf beetles (Leslie 1994). Beetle larvae in soil can be
pending on the species. Eventually they pupate. Adults
controlled with Bacillus popilliae (described in the next sec-
emerge from the soil in spring, and feed at night on tree fo-
tion). Larvae in soil can also be killed with entomophagous
liage. At dawn they return to soil where females lay pearly
nematodes, especially "cruiser" species, such as
white globular eggs in batches of 12-30, under sod or weedy
Heterorliabditis bacteriopkora and Steinerlzenza glaseri (described
patches of grass. Eggs hatch in three weeks and young grubs
in the next section). "Ambusher" species, such as Steinernemu
feed on roots for the summer. Melolontlla species range across
carpocapsae, work less well.
Eurasia from Ireland to Japan, but have only been reported
Yepsen (1976)repelled Japanese beetles by planting gar-
on Cannabis in Germany and Italy (Kirchner 1906, Goidanich
lic (Allium sntivz~m),tansy (Taliaceturn zlulgare), and geranium
1928, Gutberlet & Karus 1995). These species sometimes
(Pelargonillm hortorum). Or reverse your strategy by plant-
develop synchronous life cycles-in Switzerland M.
ing a trap crop-white zinnias (Zinnia elega~zs)attract Japa-
melolontha adults arise every three years.
nese beetles; lure the pests onto zinnias and eliminate them.
Yepsen also claimed Japanese beetles have a fatal attraction
for larkspur (Delplzinil~n~ species); they eat the foliage and die.

CHEMICAL CONTROL (Chapter 11)

Rotenone, nicotine, and spinosad kill leaf-eating beetles.
Neem repels some beetles. A Cannabis extract weakly repelled
Japanesebeetles (Metzger &Grant 1932).Yepsen (1976)killed
cucumber beetles with wild buffalo gourd, Clicurbita
foetzdissinza. He ground gourd roots into powder, suspended
the powder in soapy water, and sprayed plants.

Figure 4.33: Chafers. A. Adult and grub of Melolontha

melolontha; B. Adult of Melolontha hippocastani (from
WHITE ROOT GRUBS Sorauer 1958).
Of all underground insects, white grubs are the most
destructive. Larvae of four Scarab beetles cause Cannabis
losses. All four pests are polyphagous, and commonly at-
tack grasses. Dewey (1914) noted that white grubs cause the 3. BLACK SCARAB
greatest damage in hemp rotated after sod. The grubs often Maladera holosericea Scopoli 1772, Coleoptera; Scarabaeidae.
Description & Life History Adults are dark brown, 7-9 mm
attract moles which become a secondary problem. in length, and their elytra are marked by rows of shallow puctations.
Both adults and larvae overwinter in soil. Adults emerge in spring
SIGNS & SYMPTOMS and are active at night. Sorauer (1958)said larvae normally feed on
Seedlings grow 30-60 cm tall before showing symptoms. beets and hops. The pest has been observed in hemp fields in Yugo-
Then they wilt, yellow, and die. Damage arises in patches slavia (Camprag1961)and Korea (Anonymous 1919).Shiraki (1952)
across infested fields. Roots are slightly gnawed or entirely cited a related species, Maladera orientalis, on hemp in Japan.
 Chapter 4: Insects and Mites 69

4. AFRICAN BLACK BEETLE ooze protein-destroying enzymes, and do the actual killing.
Heteronychus arator (Fabricius) 1794, Coleoptera;Scarabaeidae. Bacteria also produce antibiotics which prevent putrefaction
Description: Adults are black, rounded scarabs between 15- of dead insects, allowing nematodes to reproduce in the ca-
20 mm long. Larvae and soft-bodied and fleshy, curved in a c-shape, davers. H , bacteriophora does best in soil temperatures be-
wit11 well-developed thoracic legs, 35 mm long at maturity. tween15-32°C.
DEVELOPMENT: H. bncterioz?hora uses a classic "cruiser"
Life History & Host Range strategy to find insects-thirdlstage larvae actively track
Adults overwinter, emerging in the spring to chew on down hosts, following trails of C 0 2 and excretory products.
young seedlings near the soil line. Females lay eggs in moist Larvae penetrate hosts through the cuticle (body wall), as
soil at the base of plants. Larvae feed on roots, but their dam- well as through natural openings (the mouth, spiracles, or
age is rarely significant. Usually one generation arises per anus). Cuticular penetration is important regarding white
year. H . arator occurs throughout tropical Africa and south- root grubs, because scarabs have sieve plates protecting their
ern Africa (Hill 1983). It is also found in Australia, where it spiracles, and they defaecate so often that most nematodes
is called the African Black Beetle. H. nrator fre~uentlvinfests entering the anus are pushed back out. Once nematodes pen-
sugarcane, maize, and wheat, and occasionally infests hemp etrate their hosts, they release toxins which inhibit the insect
in Western Australia (Ditchfield, pers. commun. 1997). immune system, then release bacteria. The bacteria kill in-
sects and liquefy host organs within 48 hours. Nematodes
DIFFERENTIAL DIAGNOSIS then use the cadaver to reproduce and multiply (Plate 31).
Above-ground symptoms caused by white root grubs Insects killed by H . bacteriophora turn brick red, and fresh
(i.e.,wilting) can be confused with symptoms caused by root cadavers may glow dimly in the dark. A new generation of
maggots, nematodes, root-rot fungi, or drought. Dig up roots nematodes emerges from the carcass in two weeks.
for inspection. Antennae of adult scarab beetles end in flat- APPLICATION: Supplied as third-stage larvae in
tened, palmate segments which unfurl like a hand-fan; this polyethylene sponge packs or bottles. Sealed containers of
feature distinguishes them from all other beetles. The hard H , bacteriophora remain viable for up to three months at 2-
part is differentiating root grubs from each other-they look 6"C, but d o not store as well as Steinernerna species. Gently
quite similar (compare Fig 4.33 with Fig 4.34). Grubs can be stir contents of package into 5 1 water (15-20°C), let stand for
differentiated by their rasters (the ends of their abdomens), five minutes, then transfer to watering-can or spray tank.
see Olkowski et al. (1991). Some gel preparations must be mixed with warm water
(<32"C) and allowed to dissolve for 30 minutes before they
CULTURAL & MECHANICAL CONTROL are released. Release within three hours, otherwise the nema-
(method numbers refer to Chapter 9) todes drown. Apply in early evening, to avoid direct sun-
Use methods 2a (deep ploughing), 3 (especially grassy light. To apply with a sprayer, remove all filters, use a spray
weeds), and 6 (do not rotate after grass or cereal crops). Ex- nozzle at least 0.5 mm wide, spray with a pressure under 5
pect greater damage during damp summers. Dry soil de- Bars (<73pounds/in2), and agitate the spra; tank frequently
stroys many eggs and newly-hatched grubs. Ploughing fields so nematodes remain suspended. The usual recommended
midsummer kills many pupae. In horticultural situations, dose is 50,000 nematodes per plant or 500,000-1,000,000 per
walk around plants wearing Lawn Aerator Sandals@.The m2. Water the soil before and after application (at least 0.64
6 cm-long spikes puncture and kill larvae. cm of irrigation), or apply during a rainstorm, and keep the
soil moist for at least two weeks after application.
BIOCONTROL (see Chapter 10) NOTES: Nematodes kills insects quicker than most bio-
Nematodes kill white root grubs, especially "cruisers" logical control agents. Heterorkabditis species live deeper in
like Heterorkabditis species and Steinernema glaseri (described the soil than ~ t e h e r n e m aspecies (down to 15 cm), and have
below). Microbial biocontrol of white root grubs began 50 superior host-penetrating abilities. H. bacteriophora is com-
years ago with Bacillus popilliae, described below. Gutberlet patible with Bt, Bacillus popilliae, and predatory mites (in-
& Karus (1995)promoted a European variety, Bacillus popil- &ding Hypoaspis miles). ~ u biocontrdl t insect; that live in
liae var. melolonthae, for use against chafers. Bacill~ls the soil are incompatible (e.g., pupating Aplzidoletes
tlii~ringiensisjaponica is a new Bt variety released against Japa- aphidimyza). According to some experts, wasp parasitoids that
nese beetles (for more on Bt, see European corn borers). Some pupate within silken coccoons are not killed.
people have success with the microinsecticides Beauveria H , bacteriophora is more sensitive to physical stress than
brongniartii, Nosema melololztlzae (described below), and other beneficial nematodes, so avoid most insecticides while
Metnrlziziunz anisopliae (described under aphids). utilizing this species. Cherim (1998) noted that many nema-
The USDA has imported several Tipkia species from Ja- todes packaged in gels and granules are raised on artificial
pan, described below. Consider pasturing big biocontrols (i.e., nema chow. These products contain a high percentage of dead
hogs) in infested fields during summer and early autumn. nematodes, cornpired to nematodes raised on insect hosts.
Hogs happily dig up grubs and eat them. Birds also eat grubs.
Flocks follow tractors, eating grubs turned up by the plough. Heterorhabditis megidis
This species was discovered in Japanese beetle grubs
Heterorhabditis bacteriophora (=Heterorhabditis keliotkidis) from Ohio. Nevertheless it is not currently registered in the
BIOLOGY: These tiny nematodes, 1-1.5 m m long, USA. Third-stage larvae are sold Europe, in a dispersible clay
parasitize many insects in soil, such as caterpillars, beetle formulation (Lavanem@,Nemasys-H@),and used against
grubs, root maggots, a n d thrips. Of the eight known vine weevil larvae. H . megidis is a large species (which makes
Heterorkabditis species, H. bacteriophora is probably the favour- it expensive to raise), and it has a short shelf life, like H. bac-
ite (Cruiserm, HeteromaskB, Nema-BIT@,Nema-green@,Lawn teriopkora.
Patrol@,Otinema). The species was discovered in Australia
but also lives in North America, South America, and Eu- Steinernema glaseri
rope. Heterorliabditis nematodes contain gram-negative BIOLOGY: This Steinernema nematode is a cruiser (Vec-
anaerobic bacteria (Photorkabdus lllrninescens). The bacteria tor@),unlike its better-known cousin, Steinernerna carpocap-
70 Hemp Diseases and Pests

iar (described under cutworms).It is highly mobile in sandy Tiphia species

soil (less so in clay soil), and highly responsive to host BIOLOGY: White root grubs are parasitized by small
chelnoattractants. S. glasc1.i works best against sedentary root- wasps from the superfamily Scoliidae. The USDA has irn-
feeders, such as white root grubs. Glaser and Fox (1930) dis- ported several wasps to control Japanese beetles. Tiphia
covered 5.gln~erikilling Japanese beetle grubs in New Jer- popilliavora was first, imported from Japan in 1921. It be-
he)-. Si11cethen, S. glaseri has been found across the southern came established in the Mid-Atlantic states. T i p l ~ i az1enzalis
USA and also in Brazil. Besides scarab grubs, S. glaseri was imported from China and Korea in 1925. It spread from
parasitizes soil-dwelling leaf beetles, weevils, and even some New Jersey south to North Carolina and west to Indiana.
caterpillars and grasshoppers. DEVELOPMENT: Adults are shiny black wasps, 5 mm
NOTES: 5,glaseri is a big nematode (twice the lengtliand long. Female Tiplzia wasps burrow into soil to locate grubs,
eight times the volume of S,cnrpocapsae, see Plate 31). This paralyze them with a sting, then insert a single egg in a spe-
makes S. glaserl expensive to produce. cific location on the grubs' bodies (Fig 4.34).T. z1ernalls adults
emerge in the spring (May and early June) and sting third-
Bncillus popilline instar grubs in the soil. T.popilliavora adults emerge in au-
BTOLOGY &DEVELOPMENT:Asoil bacterium that infests tumn (August and early September), and sting third-instar
scarab grubs and possibly other beetles. It is native to New larvae. Females of both Tipilia species lay up to 25-50 eggs
lerse); and does best in moderate temperatures (>16"C). in 25-30 days. Wasp larvae eventually consume their entire
Grubs eat spores, which germinate in the larvae's guts; the host (except the head and legs), and pupate in the soil cell
bacteria proliferate a n d eventually penetrate the built by their victim. One generation arises per year.
haemolympl~.Grubs gradually turn milky-white and die of NOTES: The wasps are poor fliers. Tiphia adults must
"milky spore disease." As grub cadavers decompose, they have a food source nearby. Adult T, vernalis feeds on aphid
release millions of spores into the soil. honeydew, T. popilliauora feeds on the nectar of wild carrot
APPLICATION: B.popilliae was the first bacterial insecti- (Dalicus carota) and other umbelliferous flowers. Tipllzn spe-
cide available in the USA, registered in 1948, and marketed cies are compatible with Bacillils species.
as Dooms and Japidemic'" Supplied as spores in a talc pow-
der (100 million spores per g). It can be stored for months in CHEMICAL CONTROL
a dry cool 8-10°C) place. Broadcast soil with spores, about 5 Drenching soil around roots with rotenone kills white
g m-2, and follow with 20 minutes of watering. Unlike Bt root grubs.
products, ti.popilliae products are viable, and released bacte-
ria can replicate and become established in stable soils, pro- ,Tiphia koreana
v i d ~ n glong-term control. Temperature is a critical factor-
5,pop71llineworks only in soils warmer than 16°C.This limits
its effectiveness above 40" latitude in the USA. Unfortunately,
A. p~opilline is slow-acting and requires a month or more to
kill its host.
UOTE: Much of the USA supply (produced by Ringer
Corporation as Grub Attackp) was tainted with other bacte-
ria in 1992 and temporarily withdrawn from the market. Cur-
rently no product is registered in the USA. Arelated species, someris annulata
Bacillus lerztii?zorbtis, is considered a strain of B. popilliae by
s o ~ n eexperts.

Beaziueriu brongglzinrtii (=B, tenella)

BIOLOGY: This fungus is marketed for control of
sugarcane root grubs (Betel&)and Melolotltl~aspecies and
other scarabs (Engerlingspilz"). The Japanese use this spe-
cies to control longhorn beetles and other species (Thomson
1998). B. broilg~ziartiiis native to central Europe, and does
best in moderate temperatures. Its morphology and life cycle Figure 4.34. Parasitized grub of Popillia japonica, showing
resembles that of Bealiveria b a s s i a m (described under the egg locations of 4 different Scoliidae wasps (from
ivhiteflies). Clausen & King 1927).
APPLICATTOA: Formulated as liquid blastospores or in
clay granules. Blastospores are sprayed on trees, so night-
feeding adults become infected and then return to the soil to WEEVILS & CURCULIOS
die. B. brongnlnrtii then sporulates on cadavers underground, Weevils are beetles with long curved snouts. They are
spreading spores throughout the field (Roberts & Hajek 1992).
also known as snout beetles. Curculios are weevils with even
In years when Melolor~thaadults are not flying, the clay gran-
longer snouts. These snouts sprout antennae and chexving
ules are drilled into the ground or applied as a soil drench
mouthparts. About ten members of the family Curculionidae
(Hajek 1993). Benlci>rrznspecies persist in soil, but organic
reportedly attack Camzabis; the cabbage curculio causes the
materlal llrlz~ierssurvi\ral and nitrogen fertilizer kills them most damage.
[Leslie 1994).
SIGNS & SYMPTOMS
Noseina gl~ielolonthae Adults chew small holes in leaves or notch leaf mar-
BIOLOGY: A microscopic protozoan lethal to Melolonthn
gins. The holes and notches often become surrounded by
grubs. This biocontrol is not yet commercially available. See chlorotic halos. Most larvae (grubs) feed on pith within
the section on grasshoppers for discussion of a related stems, causing a slight swelling at feeding sites. Some grubs
species, ~Voscinaloclistae.
feed on roots.
 Chapter 4: Insects and Mites 71

1. CABBAGE CURCULIO
C ~ ~ i t o r h y l z c i zrapae
~ i s Gyllenhal1837, Coleoptera; Curculionidae.
Description: Young lawae have cylindrical white bodies and
bro~vnheads. Older larvae slightly darken in coloul; become some-
~vhatplump, and reach 4-6 mm in length. They assume a C-shape
whei~exposed. Adults are oval to oblong, grey to black, and cov-
ered rvith yellow to grey hairlike scales. The curved snout is slightly
longer than the head and thorax; the snout is slender, c)rlindrical,
finely punctuated with striae on the basal half, and antennae arise
near ~ t middle.
s Wing covers are marked by fine longitudiilal ridges
and rows of shallorv punctuations. Length 3.5-5.0 mm (Fig 4.35).
When d~sturbed,adult cabbage curculios, like most weevils, draw
in the~rlegs and antennae and drop to grouild and play dead. Figure 4.35A: Adult
and grub of Rhinocus
Life History & Host Range pericarpius (from
Adults overwinter a n d emerge in April-May to mate. Harada 1930).
Females insert eggs into hemp stems w h e n seedlings are 3-
8 cm tall. As a result, only the lower 50 c m of mature plants
contain grubs (Nagy et a1. 1982). Grubs leave exit holes in Polygonl~nzspecies. R. pericnrpiiis lives throughout temperate
June to pupate in small coccoons just beneath the soil sur- North America, Europe and Asia. Grubs feed within stems
face. Adults emerge in late JuneJuly and feed on leaves. One and cause small galls. The galls are weak and may snap
generation arises per year. Grubs pupate in stems. Adult beetles feed on leave.; and
C. rnpae infests hemp in the Czech Republic (Rataj 1957), overwinter in the soil. One generation arises annually.
Hungary (Nagy et al. 1982),Yugoslavia (Camprag et al. 1996),
and Italy (Goidanich 1928, Martelli 1940, Ferri 1959, 1 9 6 1 ~ ) . 3. OTHER WEEVILS A N D CURCULIOS
Nagy et al. (1982) reported u p to 40% crop losses; they called Tremblay & Bianco (1978) found the cauliflower wee-
C. rhy?ne the "hemp curculio." The pest ranges across tem- vil, Ceutorhynchus pallidactyl~is(Marsham) 1802, feeding
perate Europe and was accidentally introduced into North o n hemp in Italy. The life cycle of C , pallidactyll~smimics that
America about 1855. In America, C. rapae primarily infests of C. rapae except C. pallidactyllrs eggs are laid o n leaves. C.
cruc~ferouscrops. It has attacked d r u g cultivars in Vermont pallidactyllls beetles are slightly smaller than C. rapae but oth-
(Bush Doctor, pers cornmun. 1994). erwise identical; the grubs cannot be differentiated. Tremblap
(1968) also reported Ceutorlzynclzus pleurostigma (Marsham)
1802, Cetitorhynclitrs qliadridens ( P a n z e r ) 1795, a n d
Ceutorhynchus roberti Gyllenhal 1837 on Italian hemp. He
illustrated the morphological differences between C. iapae,
C. roberti, C. q~ladrideilsand C. pleurostiglila.
Mohyuddin & Scheibelreiter (1973) cited Cetrtorhynchus
macula-alba Herbst 1795 o n Romanian hemp. They tested
the species as a biocontrol agent against marijuana (the pest
usually infests opium pods). Goidanich (1928) cited selren
other Curculionidae o n Italian hemp-Ceutorlzyncl~l~s
strleicollis (Payk.) 1800, Gymnetron labile (Herbst) 1795,
Gymnetron pascltoruin (Gyllenhal) 1813, Polydrost~ssericelis
(Schall. 1783), Sitorla humeralis (Steph.) 1831, Sitona lineattrs
(L.) 1758, and Sitona sulcifrons (Thumb.) 1798.
The gold dust weevil, Hypomeces squamous (Fabricius)
1794, is a pest in Thailand (Hill 1983).Adults notch leaf mar-
gins, and grubs live in the soil and feed on roots. These broad-
Figure 4.35: Cabbage curculio, Ceutorhynchus rapae. nosed weevils attack many crops in southeast Asia. Adults
A. Adult from above; B. Adult from side; C. Grub; D. Head are grey, 10-15 m m long, and covered with a fine golden-
of grub from front; E. Pupa (from Blatchley 1916). green "dust" of hairs. Their wings fuse together in midline,
rendering the adults flightless.
O t h e r r e p o r t s i n c l u d e t h e Asiatic o a k weex.11,
2. HEMP WEEVIL
Cyretepistonrnus castaneus (Rolofs) o n hlississippi marl-
R ~ z ~ ? ~ operic11rpills
~ I c I ~ s Linnaeus 1758, Coleoptera;Curculionidae.
Description: Larvae are white with dark brown-black heads;
juana ( L a g o & S t a n f o r d 1989), t h e Ber weevil,
they sooil grow plump, reaching 4-6 mm in length (Fig 4.35A). Xantlzoprochilis faunus Olivier 1807 (=Xnnthoclzellrs
Adults are broadly oval, dark reddish-brown to black, thinly cov- s ~ ~ p e r c i l i o s l lGyllenhal
s 1834) in India (Cherian 1932),
ered w ~ t hgreyish-yellow hairs, length 3 . 5 4 inm. The scutellum Corigetus mandarinus Fairmaire 1888 in Vietnam (Hanson
sports a conspicuous triangular spot, the beak is slightly longer than 1963), and Apion species in India (Bantra 1976) and Italy
the head and slightly ridged (Fig4.35A).Eggs are oval, white, less than 1 (Goidanich 1928). Apio~zspecies are often called weevils but
mm long. belong to the family Apicinidae, not the Curculionidae.

Life History & Host Range DIFFERENTIAL DIAGNOSIS

Harada (1930) called R. pericnrpi~lsthe most injurious Adult weevils and curculios may be confused with other
pest of hemp in Japan. Wang et al. (1995) labelled it a major beetles until you spot their unmistakable snouts. Grubs are
pest in Anhui, China. Clausen (1931) considered it less con- harder to separate from other stem- and root-inhabiting bee-
sequential. Shiraki (1952) and Wang et al. (1995) called R. tles, especially the grubs of flea beetles. Stem gall symptoms
perlcarpills "the hemp weevil," although R. pericarpius also must b e differentiated from those caused by caterpillars and
attacks bouncing bet (Snporznrin officinalis L.), Euphorbia, and gall midges.
72 Hemp Dise,sses and Pests

CULTURAL & MECHANICAL CONTROL (1931) considered it less consequential. Besides hemp, the
(method numbers refer to Chapter 9) adults tumble in umbelliferous tops, such as parsley and car-
Observe methods 1 (sanitation), 2a (deep ploughing), 3 rot (Anonymous 1974).
(eliminate cruciferous weeds), 6 (avoid vegetable crops), and
9 (hand removal of adults). Consider using crucifers as trap
crops for C. rapae. Destroy trap crops after eggs are depos-
ited on them. Collect adults by laying a tarp under infested
plants and shaking the plants. Weevils and curculios fall and
remain motionless, long enough to be squashed or gathered
for soup. Or let chickens and geese at them.

BIO & CHEMICAL CONTROL

Biocontrol researchers are testing Scambus pterophori,
a parasitoid of stem-boring beetle grubs (especially weevils).
For grubs or adults in the ground, utilize bacteria and para-
sitic nematodes described in the sections on flea beetles and
white root grubs. Rotenone and cryolite kill weevils and
curculios. Try injecting stem galls with a large-bore syringe
to kill pith-feeding larvae.

ASSORTED BORING BEETLES

In this section w e grouped together a few tumbling
flower beetles, longhorn beetles, and a marsh beetle. Grubs
of all these species bore into Cannabis stems and roots.

1. TUMBLING FLOWER BEETLES

Only adult Mordellistena beetles tumble in flowers to
elude predators. The grubs hide in stems. Flachs (1936)called
the pests "barbed beetles," Lindeman (1882) and Shiraki
(1552) called them "hemp beetles," and Anonymous (1974)
called them "hemp flour beetles." Some Mordellistena spe- Figure 4.36: Mordellistena micans (=Mordellistena
cies have attracted attention as potential biocontrol of illicit cannabis/).A. Adult, side view; B. Adult, dorsal view; C
Cannabis crops (Baloch & Ghani 1972, Mushtaque et 01.1973). Larva; D. Pupa; E. Damage in stems and roots (from
Anonymous 1974).
Signs & Symptoms
Grubs of tumbling flower beetles feed on stem pith. b. Mordellistena parvula Gyllenhal 1827, Coleoptera;
They bore into the central stem and replace pith with fine Mordellidae.
borings and frass (Plate 32). Feeding sites often swell. The Description: Adults resemble those of M. micnns, except they
sites are structurally weak and may snap, causing a wilt of are shorter (2 mm long) and more slender. The larvae also look alike,
distal plant parts. Damage peaks in late July and early Au- except M. pavz~ulngrubs lack barbs.
gust. One species, M. nzicans, feeds in the lower ends of stems
(Kirchner 1906) or roots (Anonymous 1974) where its tun- Life History & Host Range
nel can be up to 6 cm long (Lindeman 1882). Another spe- In the north Caucasus and Kirgizia, M. parvula has in-
cies, M, parvtlla, feeds in upper parts of plants, within nar- fested 95% of hemp sown in April or May, and 66-78% of
row branches, petioles, and even central leaf veins (Sorauer 1958). crops sown later (Dumovo 1933). Krustev (1557) counted
ten grubs per plant on severely damaged crops. In Pakistan
a. Mordellistena micans Germar 1817, Coleoptera;Mordellidae. M. parvula attacked marijuana and sunflowers (Baloch &
=Moudellistena cnnnabisi Matsumura 1919 Ghani 1972).
Description: Adults are wedge-shaped, humpbacked, black on the In addition to M. micans and M. parvula, Baloch & Ghani
upper surface and covered with a dense pubescence of grey-brown (1972) identified Mordellistena gurdneri in Pakistan, which
hairs, brown on the underside, 2.5-3.5 mm long (Fig 4.36). They normally infests alder (Alnus species) and absinthe (Artemi-
have long strong hind legs like flea beetles. The abdomen tapers to sia species). Goidanich (1928)reported Mordellistena reichei
a barb which projects beyond the wings. Larvae are legless, yellow, Em. 1876 in Italian hemp, and Kyokai (1965) cited Mordel-
covered with bristles, reach 3 4 mm in length, and sport a dark red
listena comes Marseul in Japanese hemp.
barb on their posterior end (Fig 4.36).Pupae have a red-brown dor-
sum and a yellow abdomen.
2. LONGHORN BEETLES
Life History & Host Range Grubs of most longhorn beetles (family Cerambycidae)
Grubs of M. n~icansoverwinter in stems and crop stub- feed within tree trunks. They are also called roundheaded
ble. They pupate there in spring. Adults emerge by May and wood borers. The antennae of adult males may grow sev-
lap eggs within a month (Krustev 1957).M. micans has been eral times their body length, hence their longhorn name.
cited on hemp in Europe (Lindeman 1882, Kirchner 1906, Some longhorn beetles live for several years as larvae.
Flachs 1936, Krustev 1557, Sorauer 1958, Camprag et al. 1996), Clarke (unpublished research, 1996) found Anoplohora
Pakistan (Baloch & Ghani 1972, Mushtaque et al. 1973), and glabripennis (Motschulsky) on hemp stalks in China. Adult
North Africa and Syria (Sorauer 1958).Sorauer said the Japa- beetles have black bodies speckled with white spots. The
nese species M. cannabisi is identical to M , micans. Takahashi Chinese call them "starry night beetles." The grubs normally
(1919)called M ,cannabisi a serious pest in Japan, but Clausen bore into hardwood trees. Vermont entomologists call this
 Chapter 4: Insects and Mites 73

species the Asian longhorned beetle, and fear its accidental CULTURAL & MECHANICAL CONTROL
introduction into the USA wdl destroy the sugar maple industry (method numbers refer to Chapter 9)
Three other longhorns have been cited on Cannabis. An Stem-boring beetles can be reduced by methods 1
unspeciated A g a p a n t h i a was collected in Turkey (Clausen recommended burning crop debris to kill T.geblerz),
(Mohyuddin & Scheibelreiter 1973); Rataj (1957) cited 2a (deep ploughing), and 4 (plant late to avoid M.parvula).
Agapanthia cynarae Germar 1817 on hemp in the Czech Avoid swampy ground and you avoid marsh beetles.
Republic. However, the most common longhorn pest is
Tliyestes gebleri (see below). BIOCONTROL (see Chapter 10)
In the future, mordellids may be controlled by
3. HEMP LONGHORN BEETLE parasitoids. Baloch & Ghani (1972)found several Tetrastickus
Tllyestes gebleri Faldermann 1835, Coleoptera; Cerambycidae. species, R h a c o n o t t ~ sspecies, a n d a B u r e s i u n z species
Description: Beetles are black with white stripes down the parasitizing grubs of tumbling flower beetles. The Japanese
prothorax, elongate, cylindrical, u p to 15 mm long, with striped an- use Beauveria brongniartii to control longhorn beetles (de-
tennae nearly as long as their bodies. Grubs are robust, cylindrical, scribed in the section on white root grubs). Try injecting sterns
creamy white, with prominent heads capped by a black dot, up to with Bacillus popilliae or beneficial nematodes, described
20 mm long. Pupae are intermediate in size between grubs and under white root grubs.
adults, light brown.
CHEMICAL CONTROL (see Chapter 11)
Signs & Symptoms
Treating plants with rotenone kills boring beetles. Stems
Grubs feed within larger stems and stalks, ejecting ex-
must be individually injected to kill the grubs. After treat-
crement at intervals through frass holes. Their tunnels are
ment, remove all frass from around plants. Any subsequent
much longer than those of tumbling flower beetles. Infested
sign of excrement indicates a need to repeat the treatment.
stems may snap at tunnel sites.

Life History & Host Range

T. gebleri grubs overwinter in plant debris. They pupate PLANT BUGS
in roots at the ends of tunnels, which they plug with fibrous People call a variety of animals "bugs," including bee-
hemp fragments. Adults emerge in May or June. Females tles (e.g., June bugs) and non-insect arthropods (e.g., sow-
deposit one to two eggs at a time into hemp stems, "usually bugs). True bugs are only found in the insect order Hemi-
5 inches below the first joint," according to Takahashi (1919). ptera. They are distinguished by their wings, which are half
One generation arises per year. Shiraki (1952) called T.ge- (hemi-)membranous, and half thickened and leathery Many
bleri the hemp longhorn beetle. Anonymous (1974) called it bugs are polyphagous and move from host to host. Pest
the hemp beetle. Takahashi (1919)and Clausen (1931)considered populations build up in weeds and wild plants before the
T,gebleri the most destructive pest of hemp in Japan. T, gebleri pests disperse to cultivated hosts, and vice versa (Panizzi
lives in eastern Siberia, northern China, Korea, and Japan. 1997). Pest populations often follow a regular sequence of
host plants as the seasons change (Fig 4.38).
About a dozen Hernipterans are regularly found on Can-
nabis, some are serious pests. One species acts symbiotically:
Vavilov (1926) described a red bug (Pyrrhocoris apterus
Linnaeus 1758) associating with Cannabis ruderalis in the
former USSR. P, aptertis is attracted to a fat pad at the base of
the seed, not the seed itself. In the process of fat pad feeding,
the bug carries seeds "far distances" and facilitates the spread
of Cannabis ruderalis.

SIGNS & SYMPTOMS

True bugs, like their Homopteran cousins (aphids,
leafhoppers, whiteflies), have piercing-sucking mouthparts
and feed on plant sap. They feed on succulent parts of
plants-unripe seeds, flowering tops, leaves, and even young
branches and stems. Bugs feeding on flowers cause bud abor-
tion, which delays maturity and reduces yields. Some bugs
Figure 4.37: The hemp longhorn beetle, Thyestes gebleri, inject a toxic saliva as they feed, turning plant tissues brown
A. Adult; B. Grub (from Takahashi 1919). and lumpy.

1. SOUTHERN GREEN STINK BUG

4. MARSH BEETLE Nezara viridula (Linnaeus) 1758, Hemiptera; Pentatomidae.
Scirtes japoniclls Kirsenwetter, Coleoptera; Helodidae. Description: Adults are easily recognized by their flattened
shield-like shapes, 15 mm long and 8 mm wide (Fig 4.39). Green
Shiraki (1952)cited this marsh beetle infesting Japanese stink bugs are not always green; they may turn a russet colour be-
hemp. Most helodid beetles live in marshes and other fore diapause. McPartland (unpublished data 1982) found white
swampy places. Adults are black-bodied, oval, and 2-4 mm colourmorphs on feral hemp in Illinois (Plate 33). Adults make a
long. Helodid grubs are aquatic. They are characterized by stink when handled. Females lay barrel-shaped, pale yellow eggs in
long slender antennae. Not much is known about S.japonicus. dense batches of 50-60 on undersides of leaves. The egg masses re-
semble tiny honeycombs. Nymphs are oval, bluish-green with red
DIFFERENTIAL DIAGNOSIS markings.
Grubs of M. micans and T.gebleri can be confused. For
further differentials, see the previous section concerning wee- Life History & Host Range
vils and curculios. Adults overwinter in secluded places in deciduous
74 Hemp Diseases and Pests

G = Generation of Nezara viridula

Castor Bean
(Ricinus communis)

SUMMER I AUTUMN I WINTER I SPRING

= host for feed~ng& reproduction = host for feeding, = host for feeding,
no reproduction unknown reproduction

Figure 4.38: Sequence of wild and cultivated plant hosts used by successive generations of the southern green stink bug,
Nezara virjdula, in a semitropical climate where 6 generations arise per year (McPartland adapted from Panizzi 1997).

woods. They overwinter in above-ground debris, so their generation of L. lineolaris arises per year.
range is limited by winter temperatures. Females lay up to L , lineolaris is abundant on feral hemp in Illinois
250 eggs. Total development time averages 22-32 days in (McPartland, unpublished data 1981), where it feeds on new
warm weather (depending on host plant) and adults live for soft tissue near the apical meristem. Feeding sites become
20-60 days (Panizzi 1977). Three generations may arise per brown lesions and young plants grow malformed. L. lineola-
year in temperate zones, but in tropical regions they breed ris also attacks cultivated hemp in Manitoba (Moes, pers.
continuously (Fig 4.38). N. viridula probably originated in commun. 1995),and Mississippi marijuana (Lago & Stanford
Ethiopia but now lives worldwide. It is very polyphagous, 1989). Protective colouring and retiring habits make L. line-
a .
with preference for brassicas and legumes (especially
- -
olaris an underestimated threat. The pest infests hundreds
soybeans). N. viridula feeds on hemp seeds (Ferri 1959a, of wild and cultivated plants, primarily alfalfa, canola, cot-
Hartowicz et al. 1971),hemp leaves (Rataj 1957, Sorauer 1958, ton, vegetable crops, and peach trees. L, litzeolaris damage
Dlppenaar et al. 1996), and flowering tops of marijuana (Rao worsens in hot, dry weather. The bugs are very mobile and
1928, Cherian 1932, Nair & Ponnappa 1974). The bug also move quickly between hosts. Watch for sudden infestations
vectors diseases. Paulsen (1971)
~, cultured the brown blight
" after neighbburing alfalfa is cut for hay.
fungus, Alternaria alternata, from N . virid~ilafeeding on feral L. lineolaris is native to North America east of the Rockies.
hemp in Kansas. N.v i r i d ~ ~ can
l a be confused with a slightly
- Its nearly-identical homologue, Lygus lzesperus Knight, is
smaller green stink bug, Acrosternum hilave (Say). found zuest of the Rockies. L, lineolaris has also been reported
on Cannabis in Europe (Goidinich 1928, Rataj 1957, Sorauer
2. TARNISHED PLANT BUG 1958);its native European counterpart is L y p s rugulipennis
L y p s lineolaris (Palisot de Beauvois) 1818, Hemiptera; Miridae. Poppius.
Description: Adults are flattened and oval in outline, 4-7 mm Other Lygz~sspecies cited on Cannabis include Lygus
in length, mostly greenish-brown but irregularly mottled by red- apicalis How. and Lygus pubescens Reut. in Italy (Martelli
d~sh-browncolouring, with a distinct yellowish triangle or "V" lo-
cated on their backs. Their triangular wingtips are characteristically 1940), and Lygus arboreus Taylor and L y p s nairobiensis
yellow, tipped by a black dot (Fig 4.39 & Plate 34). Young nymphs Poppius in Uganda (Sorauer 1958). Clarke (pers. commun.
look like yellow-green aphids, but lack abdominal cornicles are more 1990) photographed unidentified Lygus bugs in Hungarian
active than aphids. Final-instar nymphs have green bodies marked and Chinese hemp (Plate 34).
by four black dots on the thorax, one dot on the abdomen, long an-
tennae, and red-brown legs with indistinct stripes. Eggs are elon- 3. FALSE CHINCH BUG
gate and curved N y s i ~ i sericae Schilling 1895, Hemiptera; Rhopalidae.
Description: Adults are small, 3-4 mm long, with light to dark
Life History & Host Range grey bodies and lighter coloured wings (Fig 4.39).Nymphs are tiny
Adults overwinter in soil, weeds, and crop stubble. Fe- reddish-brown versions of adults, darkening as they grow older.
males emerge in spring and insert eggs into stems of crop
plants and weeds. Nymphs moult five times, gradually tak- Life History & Host Range
ing on an adult appearance. The life cycle takes only three or Adults fly south to winter quarters for hibernation. They
four weeks under optimal conditions, permitting three to five migrate north in the spring. Females lay several hundred
generations each year. But in northern extremes only one eggs on sheaths of grasses. Nymphs limit their damage to
 Chapter 4: Insects and Mites 75

grasses and weedy hosts. Adults feed on a wide variety of Life History & Host Range
crops. N . ericae infests hemp in Europe (Gilyarov 1945, C, norvegicus overwinters as an egg in stem tissues of
Sorauer 1958). Two broods arise per year. Species of the ge- herbaceous plants. Sorauer (1958)called C. norvegicl~sa "po-
nus Nysi~rsare not easily differentiated. N.ericae is limited to tato bug," but it normally feeds on legumes, mustards, and
Europe a n d citations of it elsewhere are probably grasses. It infests leaves and flowers of hemp in Italy
misidentifications. (Goidanich 1928, Ragazzi 1954, Ferri 1959a) and Germany
(Flachs 1936, Gutberlet & Karus 1995).The species originated
4. POTATO BUG near the Mediterranean, but now lives in North America.
Calocoris norvegicus (Gmelin) 1788, Hemiptera; Miridae.
=Calocoris bipunctntus F.
Description: Adults are elongate, dull greenish-yellow in colour, 5. OTHER PLANT BUGS
thinly covered by fine yellow and black hairs, with two small black Cherian (1932) considered the Indian stink b u g
dots marking their pronotums, and reach 6-7 mm in length (Fig4.39). Dolycoris indicus St51 (family Pentatomidae) as destructive

Figure 4.39: Bugs-the good, the bad, and the ugly (all x6): A. Lygus lineolaris (from Illinois Natural History Survey);
B. Anthocoris species (from Kelton 1978); C. Calocoris norvegicus (from Southwood 1959); D. Dolycoris species (from
Southwood 1959); E. Geocoris species (from I.N.H.S.); F. Nezara viridula (redrawn from Hill 1994); G. Nysius ericae (from
I.N.H.S.); H . Nabis rugosus (from Southwood 1959); J. Liocoris tripustulatus (from Southwood 1959); K. Orius majusculus,
L. 0. minutus; M. 0. insidiosus; N. 0. tristicolor; D. (K-N from Kelton 1978); 0. Pyrrhocoris apterus (from Sorauer 1958).
76 Hemp Diseases and Pests

as N. viridula on Indian ganja (Fig 4.39). It also infests Euro- APPEARANCE:Adults are very tiny (wingspan 1-2 mm),
pean hemp (Rataj 1957, Sorauer 1958).D. indicus usually in- black or very dark brown, with brown antennae, and light-
fests jute, corn, and alfalfa. coloured bands along their legs.
Bantra (1976)recovered six bugs from C. indica in north- DEVELOPMENT Female wasps lays eggs in Lygus eggs.
ern India: two members of the Miridae (Paracalocoris spe- Development time from egg to adult can be as short as ten
cies and Rhopalus species), a cotton stainer, Dysdercus days. Adults emerge from L y g eggs
~ and live up to 65 days;
cingulatus ( ~ a b r i c i i s )(family Pyrrhocoridae), a n d a females lay 150 eggs under optimum conditions. Nearly two
pentatomid (Canthecoides species). Two bugs were benefi- generations of A. iole develop for each Lygus generation.
cial insect predators-damsel bugs, Nabis species (family APPLICATION: Supplied as adults in vials. Releases of
Nabidae, F& 4.39) and assassin bugs, sycan& collaris Fab- 37,500 wasps per ha (15,000 wasps per acre) per weekworked
ricius (family Reduviidae). Clarke photographed an uniden- well in California strawberry fields, especially when used
tified pentatomid infesting hemp flowers in Hungary, and preventatively.
he collected an unknown shield bug in China (subfamily
Scutellerinae, possibly an Elasmostethus species). Trissolcus basalis
Watson & Clarke (pers. commun. 1994) identified the BIOLOGY: A tiny scelionid wasp that parasitizes eggs of
inirid Liocoris tripusttilatus (Fabricius) 1781 feeding on Can- N , viridula. It is found around the world in temperate regions,
nabis pollen in Dutch greenhouses. L. trip~istlilaft~s adults are and works best at 22°C.
light yellow-brown and less than 5 m m long (Fig 4.39). They APPEARANCE & DEVELOPMENT: Adults are minute,
look like tarnished plant bugs, and frequently infest nettles black wasps (wingspan 1-2 mm), with short antennae that
in Europe. L. t r i p ~ ~ s t i ~ ladults
a t ~ ~ scan be confused with Orius bend downward. Adults mate immediately after emerging
biocontrols from host eggs. Female wasps lay up to 300 eggs; they insert
one egg into each Nezara egg. Larvae develop and pupate in
DIFFERENTIAL DIAGNOSIS Nezara eggs and emerge as adult wasps. The life cycle takes
Bugs (Hemipterans) can be confused with Homopterans about 23 days in optimal conditions.
such as leafhoppers, planthoppers, and treehoppers. Very APPLICATION: Strains from southern Europe were in-
young bugs look like aphids but lack cornicles and move troduced into California in 1987, becoming established in
faster than aphids. Short, black bugs can be confused with cotton, soyabean, and vegetable-growing areas. Unfortu-
beetles. Bug damage is nearly identical to that of other in- nately, when T. basalis was released in Hawai'i, it moved to
sects with sucking mouth parts. Bud abortion caused by tar- several native nontarget hosts and nearly caused their ex-
nished plant bugs can be confused with late flowering asso- tinction (Howarth 1991). T.basalis tolerates permethrin, but
ciated with late-maturing plants (i.e., Thai landraces). Ob- is susceptible to organophosphate insecticides.
sewation with a hand lens will detect damaged flowers.
Trichopoda peunipes
CULTURAL & MECHANICAL CONTROL BIOLOGY:A tachinid parasitoid of N . viridula and squash
(method numbers refer to Chapter 9) bugs, native to North and South America. A related species
Control bugs with methods 3 (weeding), 1(sanitation), from Argentina, Trichopoda giacomellii, has been released
2a (deep ploughing), and 9 (some heavier bugs can be shaken against N . viridula in Australia.
off plants like beetles, best done early in the morning before APPEARANCE &DEVELOPMENT:Adults look like house-
they "limber up"). Tarnished plant bugs are attracted to white flies with orange highlights, and a fringe of feather-like black
or yellow sticky traps (method 12c).They can be vacuumed bristles on their hind legs. Female flies produce up to 100
off plants, leaving behind beneficial insects. The bugs see eggs, and lay one egg per bug (on nymphs or adults). Mag-
the nozzle coming and try to fly off. Many bugs migrate from gots feed within bugs for two weeks, then pupate in soil.
host to host. So learn what sequence of hosts are infested, The life cycle takes about five weeks, three generations may
and monitor these hosts before bugs spread to crops (see arise per season, maggots overwinter in hibernating hosts.
Panizzi 1997). NOTES: T, pennipes has been released in Hawai'i and
Italy, but is not currently available.
BIOCONTROL (see Chapter 10)
Five parasitoids infest bugs. Anaphes iole, Trissolct~sbasa- Peristenus digoneutis
hs, Tvichopoda pennipes, and Peristenus digonelitis are described BIOLOGY: A braconid parasitoid of Lyglis ri~gulipeni~is,
below. Ooencyrtus submetallicus, an egg parasitoid of stink discovered in northern Europe. The USDA established a
bugs, works well in laboratory studies but zero in the field. population in New Jersey that controls L, lineolaris. It has
Bugs are eaten by their predatory brethren: the mirid spread north to the Canadian border, but has not moved
Deraeocoris brevis (described under thrips), and the big-eyed south of New York City (41" latitude).
bug, Geocoris puncfipes (described under whiteflies). Bantra APPEARANCE & DEVELOPMENT: Adults are small,
(1976)discovered two predatory bugs on Indian Cannabis - brown-black wasps, 3 mm long. Female wasps lay a single
S>/canl~s collaris and a Nabis species. egg into each L, lineolaris nymph. Eggs hatch in seven days
Biocontrol of bugs with fungi like Beauveria bassiana is and larvae take another ten days to kill their host and drop
inconsistent at best. Hill (1983) estimated that 15 chickens to the ground to pupate in soil. The second generation of I?
will keep 0.1 ha free of cotton stainers. Soyabean (Glycine digoneutis emerges in synchrony with the second generation
i?zns) serves as a trap crop for N,viridula. Strips of alfalfa of Lygus bugs. A third generation may arise, pupae overwinter.
(Medicago sativa) are interplanted in cotton fields at 100-150 NOTES: This biocontrol is not commercially available,
m intervals to trap L y g ~ i sbugs (Hokkanen 1991). Alfalfa is but is slowly spreading to the Midwest. A related species,
also excellent habitat for many biocontrol insects. Peristenus howardi, infests L. hespertis west of the Rocky
Mountains in the USA.
Annphes iole
BIOLOGY: A "fairyfly" mymarid wasp that parasitizes CHEMICAL CONTROL (see Chapter 11)
eggs of tarnished plant bugs ( L . hespevus and L. lineolaris). It Dusting plants with rotenone, ryania, or sabadilla con-
is native to Arizona and works in a range of 13-35°C. trols green stink bugs, cinch bugs, tarnished plant bug (lygus
 Chapter 4: Insects and Mites 77

bugs), and other hemipterans (Metcalf et al. 1962).Young bugs leaf epidermis. Other species drop to the ground to pupate.
are killed by nicotine, some species are susceptible to soaps Pupation takes one week to several months, depending on
and pyrethroids (tarnished plant bugs infesting cotton have the species and season. Outdoors, two to six generations arise
become resistant to pyrethroids). Researchers are trying to per year; in glasshouses the generations overlap so all stages
find the pheromone for L. Iineolaris, to develop synthetic can be found at any time.
pheromone traps. Six Agromyzid species appear in the Cannabis literature.
Surprisingly, Liriomyza trifolii, the serpentine leafminer, is
not one of them. L, trifolii is native to Florida and now lives
LEAFMINERS in glasshouses worldwide, attacking a wide variety of plants.
Many less-famous leafminers do not have common names.
Leafminers tunnel through tissues zoitllin leaves, like
miniature coal miners. This way they avoid THC and other 1. Liriomyza (Agromyza) strigata (Meigen) 1830, Diptera;
insecticidal chemicals on the leaf surface. Most leafminers
~~romyzidae.
are maggots-members of the fly family Agromyzidae (see Description:Maggots are pale, legless, seemingly headless, up
exceptions listed under "differential diagnosis" below). Leaf- to 2 mm in length. They mine slender light green serpentine tun-
miners can become serious pests in Dutch glasshouses if al- nels that end in brown blotches. Mines begin near the leaf margin
lowed to get out of control. and end near the midrib. Larvae pupate externally. Adults resem-
ble tiny houseflies. They are shiny black with yellow marlungs along
SIGNS & SYMPTOMS the sides and femora, averaging 1.7-2.1 mm in length, with brown
Leafminer tunnels can be seen through the leaf surface. legs and white antennae. Females lay eggs near leaf edges.
They appear pale green or white, from light reflecting off air This pest occurs onGerman hemp (Kirchner 1906, Blunk
in the mines. Each species makes tunnels with the same iden- 1920, Flachs 1936, Hering 1937, Spaar et al. 1990), in Italy
tifiable "signatureu-either linear, serpentine, or blotch-like (Goidanich 1928, Martelli 1940, Ciferri & Brizi 1955, Ferri
(Fig 4.40 & Plate 35). Most mines are made on the upper sides 1959a), and the Czech Republic (Rataj 1957). The species is
limited to Europe. Kirchner & Boltshauser (1898) provided
a fine illustration of L. strigata mines.

2. Phytomyza horticola Goureau 1851, Diptera;Agromyzidae.

= ~ h y t o & ~ zntricornis
a Meigen 1838, = ~ h y t o r n ~ zEhysnnthenli
a
Kowarz 1892
Description: Maggots are greenish-white and 3 4 mm long.
They mine long, serpentine tunnels according to Hill (1983) and
Spaar et al. (1990), but Hering (1937) said the tunnels are short and
straight. Tunnels contain small, widely-spaced pellets of frass. Lar-
vae pupate inside leaves, visible as brown puparia at the wide end
of the mine. Flies have black bodies marked by yellow lateral lines,
2-3 mm long, with pale heads and yellow "knees" (Fig4.41).Phyto-
nzyzlzn flies, unlike most other Agromyzids, lack crossveins in the
posterior portions of their wings.
Known as the pea leafminer, polyphagous P. korticola
has been cited under its different synonyms by Goidanich
(1928), Hendel (1932), Hering (1937), Martelli (1940), Rataj
(1957), Ferri (1959a), and Spaar et al. (1990). Pea leafminers
live throughout Europe, Asia, and parts of Africa, but not
(yet) North America. I? korticola should not be confused with
Figure 4.40: Leafmines by 3 maggots and a beetle grub. the American species Phytomyza syngenesiae Hardy (chry-
A. By Phytomyza horticola; B. By Liriomyza cannabis; santhemum leaf miner), which was also called P, atricornis
C. By Agromyza strigata; D. By Phyllotreta nemorum in earlier literature.
(McPartland redrawn from Hering 1937).
3. Agromyza reptans Fallen 1823, Diptera; Agromyzidae.
=Agronzyza hplacnze Steyskal 1972
of leaves. Tunnels increase in width as the maggots grow. Description: Maggots are similar to those of P. horticoln in size and
Frass may be expelled in continuous strips, in widely-spaced shape, but they mine linear tunnels along leaf margins, which ex-
pellets, or in one big dump at the end of the mine. Plants pand into brown blotches, filled with frass. Flies are brownish black,
rarely die from leafminer damage, but heavy infestations legs black with yellowish tibiae and tarsi, and the puparia are red-
cause leaf wilting and reduce crop yields. Mines also provide dish brown.
entrances for fungi and other pathogens. Mines in fan leaves
of flowers make the flowers unattractive to consumers,
especially when filled with frass.

LIFE HISTORY
Most Agromyzids have identical life cycles. They
overwinter outdoors as pupae. Adults emerge in spring to
mate. In warm glasshouses, leafminers do not hibernate, they
breed continuously. Females drill eggs into leaves, one at a
time but often clustered together. Flies feed on sap oozing
from ovipositor wounds. Some species lay up to 350 eggs
(Hill 1983). Eggs hatch in two to six days. Maggots of most
species undergo four moults, then pupate. Some species Figure 4.41 : Pea leafminer, Phytomyza horticola.
pupate in mines, projecting two spiracular "horns" through A. Adult fly; B. Larva leafminer; both x 10 (from Hill 1994).
78 Hemp Diseases and Pests

Buhr (1937),Hering (1937,1951) and Hendel(1936) re- and 12a&b (repel flies with reflectant material, trap them
ported A.reptans in northern Europe. Bush Doctor (unpub- with yellow sticky tape) Van Lenteren (1995) noted more
lished data 1986) found this species on marijuana in Illinois leafminer damage in glasshouses using rockwool.
(Plate 35). A. reptans has been reported on nettles and hops
in temperate Europe, and in Michigan. BIOLOGICAL CONTROL (see Chapter 10)
Ferri (1959a) described several parasitoids attacking
4. Lin'omyza enpatorii (Kaltenbach) 1874, Diptera; Agromyzidae. hemp leafminers. Van Lenteren (1995)controlled vegetable-
= Liriomyza pusilla Meigen 1830 subspecies eupatorii Kaltenbach mining flies (Liriomyza bryoniae and L. trifolii) with a mix of
Description: Mines often commence in a spiral pattern, a charac- two parasitoids, Dacnilsa sibirica and Diglyphils isaea (de-
teristic this species shares with L. cannabis (the next species). The scribed below). Success with these parasitoids varies with
spiral, however, may be minute or even lacking. The rest of the mine the leafminer species and the host plant. A new parasite,
is linear. Adults exhibit black bands on the lower halves of their Opiils pallipes, is described below.
abdomens, legs are yellow, usually with brownish-black striations;
Hara et al. (1993) used a soil nematode, Steinernenza car-
total length 1.5 mm, wing length 1.6-2.2 mm.
pocapsae, to kill leafminers in leaves, by spraying nematodes
Hering (1921),Buhr (1937),Rataj (1957),and Spaar et al.
onto foliage at night. The nematodes entered leaf mines
(1990) cited L, eupatorii Kaltenbach on hemp. Hering noted
through oviposition holes and killed >65% of L, trifolii lar-
high mortality in Cannabis; he blamed "poisonous alkaloids"
vae (S. carpocapsae is described under cutworms). Leafmin-
present in leaves. L. eupatorii frequently attacks Aster spe-
ers pupating in soil can also be controlled with beneficial
cies, goldenrod (Solidago species), and hemp agrimony (Ellpa-
nematodes.
torium cannabinum). It less frequently infests hemp nettle
(Galeopsis tetrahit) and Cannabis. Buhr (1937) transplanted
Dacnusa sibirica
qaggots between Cannabis and Galeopsis; the pests mined
BIOLOGY: A braconid wasp that parasitizes Liriomyza
either plant a n d p u p a t e d in soil. L. eupatorii lives
larvae. It is native to northern Eurasia and does best in cool
sympatrically with A. ueptans in North America and Europe.
to moderate temperatures (optimally 14-24°C) and moderate
humidity (70% RH).
5. Liriomyza cannabis Hendel 1932, Diptera; Agromyzidae.
APPEARANCE: Adult wasvs are dark brown-black, 2-3
= Lirionzyza e~lpatorii
Hering 1927 [non Kaltenbach 18741
Description: Maggots form tight spiral mines as illustrated by mm in length, and have long antennae.
Hering (1937). He described frass scattered along tunnels "like DEVELOPMENT Wasps lay eggs in all ages of leafminer
strings of pearls." Hendel(1932) said adults resemble L. strigata ex- larvae, but prefer first and second instars. D. sibirica para-
cept for sharply defined abdominal stripes, pointed warts on the sites develop and pupate within leafminer larvae. The D.
heads, and smaller size (1.5 mm long). sibirica lifecycle takes 16 days under optimal conditions;
L. cannabis infests hemp in Germany (Hering 1937, Spaar adults live another seven days, females lay 60-90 eggs.
et al. 1990), Finland (Hendel 1932), Italy (Ferri 1959a), and APPLICATION: Supplied as adults in shaker bottles. Store
Japan (Kyokai 1965). Mushtaque et al. (1973) reported the a maximum of two days at 8-10°C. Release in the early morn-
pest in Afghan marijuana. ing or evening. Hussey & Scopes (1985)released three wasps
per 1000 small plants, weekly. For light infestations, Koppert
DIFFERENTIAL DIAGNOSIS Leafmining is unique and (1998)suggested releasing two wasps per m2 crop area, with
not easily confused with other symptoms. Maggot leafmin- a minimum of three weekly introductions. For moderate in-
ers must be differentiated from caterpillar leafminers or bee- festations, release five wasps per m2 weekly.
tle grub leafminers. One caterpillar, Grapholita delinearza, NOTES: D. sibirica is the most popular leafminer
mines within leaves when young (Mushtaque et al. 1973). biocontrol, but it only works as a preventative. It reproduces
After it outgrows this niche G. delineana assumes its primary too slowly to stop even a moderate infestation. D. sibirica is
role as the hemp borer. The flea beetle grub Phyllotreta compatible with D. isaea. Mixtures of the two species are
nenzort~nzmakes tunnels in tight spirals which end in blotches. available. Avoid insecticides. Cherim (1998) suggested D.
Differentiating maggots from caterpillars and grubs requires sibirica may be attracted to yellow sticky traps.
close inspection; only the maggots are legless. Leafminer eggs
can be confused with thrips eggs; see what emerges. Diglyphus isaea
BIOLOGY: Apredatory/parasitic chalcid wasp that feeds
CULTURAL & MECHANICAL CONTROL on leafminers. It is native to Eurasia and does best in mod-
(method numbers refer to Chapter 9) erate humidity and temperatures (80% RH, 24-32°C).
Perform methods 1 (sanitation), 2a (deep ploughing) APPEARANCE: Adults are black with a metallic green-
2b&c (sterilizing or pasteurizing soil), 3 (for species pupat- ish tinge, 2-4 mm long, with short antennae (Plate 36). Pu-
ing in soil), 6 (crop rotation against Liriomyza cannabis), 9 pae can be seen in leafminer mines as tiny black spots.
(pick off infested leaves and compost them or burn them), DEVELOPMENT Wasps kill leafminer larvae, then lay
eggs next to them. D, isaea larvae feed on carcasses and pu-
pate in mines. The lifecycle takes 17 days under optimal con-
Table 4.11: Infestation Severity Index for leafminers. ditions; adults live another three weeks. Females kill up to
I Light few mines seen I 360 leafminers.
APPLICATION: Supplied as adults. Store one or two days

I Moderate many mines on a few plants

I at 8-10°C. Release in the early morning or evening. Used
preventively, Hussey & Scopes (1985) introduced three wasps
per 1000 small plants per week. For light infestations,
Heavy leaves starting to curl on plants Thomson (1992) released one or two wasps per 10 m2 per
OR 10-99 mines per plant week for at least three introductions. Koppert (1998) sug-
gested 1wasp per m2, with a minimum of three weekly in-
Critical plants losing vigour and wilting troductions. Cherim (1998) suggested two to four wasps per
OR >I00 mines per plant m2 every two weeks.
 Chapter 4: Insects and Mites 79

NOTES: D. isaea works well at higher leafminer densi- SIGNS & SYMPTOMS
ties than D. sibirica, so it senres as a preventative and a cura- Leafhoppers and their relatives are sap suckers. Most
tive biocontrol. D. isaea prefers warmer temperatures than are phloem feeders, but some suck xylem sap (e.g., cicadas,
Dnclzlisa sibirica. It is compatible with D , sibirica and with spittlebugs, many planthoppers, and some leafhoppers).
beneficial nematodes. Avoid using insecticides at least a Xylem sap is an extremely dilute food source (see Table 3.2),
month prior to release. Cherim (1998) suggested D , isaea may so these pests must ingest great quantities of sap. Spittlebug
be attracted to yellow sticky traps. nymphs can suck xylem sap at a rate greater than ten times
their body weight per hour. They digest what little is avail-
Opius pallipes able, then excrete >99% of the fluid. They surround them-
BIOLOGY: A parasitic wasp that feeds on leafminers. It selves with a froth of excreted spittle.
is native to temperate regions and does best in moderate Leafhoppers and their relatives cause symptoms simi-
humidity and moderate temperatures. lar to those caused by aphids and whiteflies-wilting, a light
APPEARANCE & DEVELOPMENT: Adults are 2-3 mm coloured stippling of leaves, and sooty mould from honey-
long and resemble Dacniisa sibirica. Wasps lay eggs in young dew. Xylem feeders also cause leaf veins to become swollen
leafininer maggots. The eggs hatch and wasp larvae slowly and lumpy, because their stylets are thick and stout com-
kill leafminer lanlae. pared to the fine flexible stylets of phloem feeders (Press &
APPLICATION:Supplied as adults. Store one or two days Whittaker 1993).
at 8-10°C. Release the same as Dacnusa sibirica.
1. GLASSHOUSE LEAFHOPPER
CHEMICAL CONTROL (see Chapter 11) Zygina (Erythronel~raipallidifvorzs (Edwards 1924),Homoptera;
Catch egg-laying flies with synthetic pheromone traps. Cicadellidae.
Repel them with a foliar spray of neem or oil. Description: Adults are small (3-4 mm long), pale yellow-
Larvae are protected within mines, so surface-sprayed green, with iridescent wings extending beyond the rear ends of their
insecticides may not reach them. Adding surfactants may bodies. Between the eyes are two small dark spots, and behind the
improve the leaf-penetration of surface sprays. ~eafminerk eyes (near the bases of the forewings) are two larger dark spots (Fig
4.42). Legs and antennae are long. When disturbed the adults hover
in other crops have been controlled with abamectin and over plants, like whiteflies. Nymphs are small, white, and initially
spinosad (two natural fermentation products), permethrin almost transparent; wing-pads appear during the third instar and
(a synthetic pyrethrin), methoprene a n d c y r o k z i n e (syn- enlarge during the fourth (final) instars. Cast skins from moults re-
thetic insect growth hormones) and imidacloprid (a synthetic main attached to leaves, are translucent, and are commonly called
nicotine). Try watering plants with a 0.4% neem solution. "ghost flies" (Wilson 1938). Eggs are elongated, slightly curved,
The solution is absorbed systenzically, with activity against white, 0.5-0.7 mm long.
larvae for about three weeks.
Control species which pupate underground by cover- Life History & Host Range
ing the soil with plastic. Coat plastic with Thripsticka, which Adults overwinter on weeds outdoors. In warm glass-
kills leafminers as well as thrips. This can be integrated with houses they do not hibernate, so overlapping- generations
-
biocontrol by Diglyplzus isaea,-which does not in the arise all year long. Eggs are inserted into leaf veins, usually
soil (Hussey & Scopes 1985). singly. Young nymphs feed on undersides of leaves. In opti-
mal conditions (26"C), the life cycle takes 42 days. Adults
live another four months. Z. pallidifrons poses a problem in
English and Dutch glasshouses (Wilson 1938).It attacks many
LEAFHOPPERS & crops (especially tomato and mint) and many weeds (espe-
THEIR RELATIVES cially chickweed, Stellaria media).
Leafhoppers, planthoppers, treehoppers, spittlebugs,
2. REDBANDEDLEAFHOPPER
and cicadas are grouped in a
Auchenorrhyncha. Many have been collected on Cannabis.
the Graphocephnla coccinea (Foerster) l77l.Homoptera;Cicadellidae.
Description: Adults 8-10 mm in length, slender, with yellow,
Few cause serious feeding but may inflict set- pointed heads; wings reflect alternate bands of magenta and green
ondary losses by spreading plant viruses. A dozen species with yellow margins. Nymphs are yellow to green.
are described below, in order of occurrence. They rarely cause
serious economic damage. Life History & Host Range
Adults overwinter in leaf trash on the ground. Eggs are
thrust into soft plant tissues in early spring. One or two gen-
erations arise per year. Hartowicz et al. (1971)
and DeWitt (unpublished data) collected G.
coccirlea from feral hemp in Kansas and
Illinois. Frank & Rosenthal (1978) described
them in California. This leafhopper infests
many hosts, but injury is rarely serious. Its
size and striking colour make it a high-profile
pest. Lago & Stanford (1989) described a
relative, Graphocephala versiita (Say) on
A B Mississippi marijuana.
3. POTATO LEAFHOPPER
Figure 4.42: Leafhoppers and their cousins (all x7): A. Empoasca E m ~ o a s c afabne (Harris) 1841, Homoptera;
flavescens (from USDA); B. The potato leafhopper, Empoasca fabae Cicadellidae.
(from I.N.H.S.); C. The glasshouse leafhopper, Zygina pallidifrons; =Enzponscn mnli LeBaron 1853
D. The buffalo treehopper, Stictocephala bubalus; E. The spittlebug, Description: Adults are green with faint white
Philaenus spumarius (C, D, E by McPartland). spots on the head and thorax, reaching 4 mm in
80 Hemp Diseases and Pests

length. Seen from above they appear wedge-shaped, with 1 mm I? sp~~marilis

attacks at least 400 species of plants. The spe-
wide heads tapering to pointed wingtips (Fig 4.42). Nymphs are cies accumulates in areas of high humidity across North
similar in shape but even smaller, nearly impossible to see as they America.
feed on undersides of leaves. Nymphs and adults have strong hind
legs and readily jump or "crab-walk to safety when disturbed. 6. BUFFALO TREEHOPPER
Stictocepkala bubahis (Fabricius) 1794, Homoptera;Membracidae.
Life History & Host Range Description:Adults are green and 6 mm long, their blunt heads
E.fabae ranges across the eastern half of North America, support two short horns. Seen from above, adults appear triangu-
but cannot overwinter north of the Gulf states. Each spring, lar, pointed at the rear (Fig4.42). Adults are shy and fly away with
adults hitch rides on weather fronts in Louisiana. They mi- a loud buzzing noise. Nymphs are tiny, pale green, humpbacked
grate north to Manitoba, Ontario and Quebec, and east to and covered by spines. Eggs are yellow.
New England. Some years they are blown as far as New-
foundland. Their numbers decrease in the west where rain- Life History & Host Range
fall falls below 65 cm per year. Nymphs hatch late in the spring from overwintering
Upon landing, females lay an average of 35 eggs in peti- eggs laid in tree branches. They drop to the ground and suck
oles and leaf veins. Up to four generations arise each year. sap from herbaceous plants. Adults appear by August.
Potato leafhoppers feed from both phloem and xylem (Press Nymphs feed on Cannabis, adults feed on trees. The real dam-
& Whittaker 1993), and frequently plug these vessels, which age is done by egg-laying females-not feeding, but slicing
impairs fluid movement in leaves, causing "hopperburn." stems with their knifelike ovipositors to lay eggs. They per-
Hopperburn symptoms begin as a browning of leaf mar- forate rings around branches, causing branches to break off.
gins and leaf tips. Affected leaves become deformed, lumpy, One generation arises per year.
and curly. In heavy infestations the leaves become badly Buffalo treehoppers infest marijuana near Buffalo, NY
scorched with only rnidveins remaining green. Dudley (1920) (Hilligpers. commun. 1993) and in southern Indiana (Clarke
described hopperburned hemp in New Hampshire. E.fabae pers. commun. 1985). Hartowicz et al. (1971) reported
attacks more than 100 plant species, especially alfalfa, clo- "Membracidae treehoppers" on feral hemp in Kansas.
ver, apple, and many vegetable crops. In potato crops, the DeWitt (unpublished) found S . btiballis on feral hemp in Illi-
presence of ten nymphs per 100 mid-plant leaves is the nois. The species lives throughout North America and causes
threshold for control measures (Howard et al. 1994). economic damage in orchards and tree nurseries.

4. OTHER CICADELLIDAE LEAFHOPPERS 7. PLANTHOPPERS

The "flavescent" leafhopper, Empoasca flavescens F., Description: Planthoppers (superfamily Fulgoroidea) can be
infests European hemp (Spaar et al. 1990). This species also separated from leafhoppers, treehoppers, and spittlebugs by the
lives in the USA; it resembles a pale (nearly white) version movable spur on their hind tibia, and by the location of their anten-
of E.fnbae (Fig 4.42 & Plate 37), and like E.fabae, it feeds on nae and ocelli. Planthoppers seldom occur as abundantly as
leafhoppers or spittlebugs.
xvlem and phloem (Press & Whittaker 1993). An unidenti-
fied Enlpoasca species infests marijuana leaves in Thailand Life History & Host Range
(Hill 1983).Anew species, E~npoascauniprossicae Sohi, was All Cannabis planthoppers are reported from Asia.
described on Calznabis in India (Sohi 1977).
Euybrachys tomentosa Fabricius (family Fulgoridae) is a
Frank & Rosenthal (1978) claimed Rose leafhoppers
minor pest of marijuana in India (Cherian 1932). Geisha
(Edwardsiana rosae L.) attack marijuana in the San Fran-
distinctissirna Walker 1858 (family Flatidae) attacks hemp
cisco area. Bantra (1976) reported Iassus indicus (Lethierry)
in China, Korea and Japan (Takahashi 1919, Clausen 1931,
1892 [=]assus clzlorophanus (Melichar) 19051 as the most
Shiraki 1952, Sorauer 1958). The Japanese broadwing plan-
common cicadellid on Indian marijuana. DeWitt (unpub-
thopper, Ricania japonica Melichar 1898 (family Ricaniidae)
lished data) collected a Gyponann species from feral hemp in
is also a frequent offender (Takahashi 1919, Clausen 1931,
Illinois. Lago & Stanford (1989)cited Gyponana octolineata
Shiraki 1952, Sorauer 1958). Kuoh (1980) collected a small
(Say) on ~ i s s i s s i ~marijuana,
pi along with Agallia contricta
planthopper from Chinese hemp, Stenocranus qiandainus
Van Duzee and about fifteen other uncommon cicadellids.
Kuoh (family Delphacidae).
Wei & Cai (1998) described a new cicadellid, Macropsis can-
nabis Wei & Cai, attacking hemp in Henan Province, China.
8. CICADAS
Botkrogoniaferrtiginea (Fabricius) 1787 (family Cicadidae)
5. SPITTLEBUG
infests hemp in Eurasia (Clausen 1931, Shiraki 1952, Sorauer
Piiilnen~~s spi~lnarii~s(Linnaeus) 1758, Homoptera; Cercopidae.
Description: Spittlebugnymphs are easily spotted-they usu- 1958). Cicadas are recognized by their large size and the
ally occupy the crotches of small branches and surround themselves males' summer mating songs. Some cicadas have life cycles
with froth (see Plate 84).Nymphs resemble tiny pale green frogs, 6 as long as 17 years. Most of their damage is inflicted on trees.
mm in length. Adult spittlebugs are the same size and shape as fi-
nal-instar nymphs, but turn a mottled brown colour (ranging from DIFFERENTIAL DIAGNOSIS
straw-coloured to almost black, Fig 4.42),and no longer hide in spit- Leafhopper nymphs and their damage may resemble
tle. Eggs are ovoid, 1mm long, yellow to white, and laid in groups other Homoptera such as aphids. Some planthopper nymphs
of two to 30 in hardened spittle,on stems and leaves near the ground.
(Rickanin species) look like flea beetles, and planthopper
adults can be mistaken for small moths. Hopperburn symp-
Life History & Host Range
toms can be confused with sun scald or nutrient imbalances,
Eggs overwinter. Nymphs feed on xylem sap in the
except for the irregular lumps on leaf veins.
spring, and metamorphize into adults from late May to late
June. Females lay eggs in August and September, then die.
CULTURAL & MECHANICAL CONTROL
One generation arises per year. We have seen spittlebugs on
(method numbers refer to Chapter 9)
feral hemp and marijuana in Oregon, Illinois, New Jersey,
A11 Auchenorrhyncha are controlled by methods 1 (sani-
and Vermont. Goidanich (1928) reported the species in Italy
tation), 2a (deep ploughing), and 3 (weeding). Leafhoppers
Spittlebugs rarely kill plants but they decrease crop yields.
 Chapter 4: Insects and Mites 81

are controlled with methods 12a&b (mechanical trapping Life History & Host Range
and repelling). Potato leafhoppers are attracted to blacklights Long-tailed mealybugs feed on a wide range of hosts
(method 12d).For spittlebugs, use methods 6 (crop rotation, and are found worldwide. They live out-of-doors in warm
and plant away from alfalfa and clover fields). Avoid plant- climates (including Florida, Texas, and southern California).
ing in shaded, protected areas preferred by spittlebugs. To In northern latitudes P. longispinils becomes a serious glass-
avoid treehoppers, avoid elm trees and orchards. house pest (Frank & Rosenthal 1978, Frank 1988). Unlike
other mealybug species, I? longispinils births live larvae. New
BIOCONTROL (see Chapter 10) generations may emerge every month.
Fungi w o r k best against sap-sucking insects:
Metarhizillm anisopliae (brand name Metaquino'a) is a 2. COTTONYCUSHION SCALE
mycoinsecticide that kills spittlebugs (described under lee y a ptlrchasi Maskell 1878, Homoptera; Margarodidae.
aphids). Paecilonzyces f i ~ i n o s o r o s e i ~ is
s used against Description: Mature scales are reddish brown, with black legs
planthoppers in the Philippines (described under whiteflies). and antennae.Females are partly or wholly covered by egg sacs that
are large, cottony-white,and longitudinally ridged (Fig 4.43).Total
Hiustitella fhompsonii (Mycar@)was previously registered for
length is 8-15 mm, two-thirds of which is egg sac. Sacs contaln 300-
use against planthoppers in the USA (described under hemp 1000 oblong red eggs. The red-bodied nymphs have black legs,
russet mites). Erynia (Zoophthora) radicans has been used prominent antennae, and congregate under leaves along the midribs.
experimentally to control potato leafhoppers (Samson et al.
1988). It arises naturally in the midwestern USA and kills Life History & Host Range
leafhoppers in two or three days. I, purchasi overwinters in all stages. Depending on the
The rnirid bug Deraeocoris brevis preys on young psyllids climate, eggs hatch in a few days to two months. Larvae re-
(described under thrips). Amynarid wasp, Anagvus atomus main mobile, unlike most scales. Adults are gregarious and
(=Anagui~sepos?) parasitizes eggs of Z. pallidifrons and other often suck sap in groups. Up to three generations arise per
leafhoppers. Commercial development is hampered by egg year in California. I. purchasi is native to Australia, unwit-
desiccation and the short lifespan of the adult wasp. Frank tingly introduced into California around 1868. The pest posed
& Rosenthal(1978) repelled leafhoppers by planting gerani- a serious threat to the citrus industry until a natural enemy,
ums around crops. Yepsen (1976) repelled leafhoppers with the vedalia beetle, was imported. Bodenheimer (1944) and
geraniums and petunias. Sorauer (1958) reported cottonycushion scales infesting
hemp. Cottonycushion scales are very polyphagous and live
CHEMICAL CONTROL (see Chapter 11) throughout the warmer parts of the world.
Damage is rarely bad enough to justify chemicals. Frank
& Rosenthal(1978) used insecticidal soap, especially against
leafhopper nymphs (spray on undersides of leaves). Neem
and diatomaceous earth (or better, diatomaceous earth mixed
with pyrethrum) kill most hoppers. Frank (1988) killed
leafhoppers with pyrethrum and synthetic pesticides. Bor-
deaux mixture repels adults and serves as a feeding deter-
rent (Metcalf et al. 1962). Clay microparticles may also work
this way.
Z . pallid2frons is controlled with nicotine sprays (two or
three applications at ten-day intervals); rotenone and pyre-
thrum are less satisfactory (Wilson 1938).Potato leafhoppers
succumb to sabadilla dust, but new waves of migrating
leafhoppers may require five to eight applications per sea-
son. Spittlebugs are susceptible to rotenone, especially when
nymphs are less than a month old. Alfalfa growers start
spraying fields when spittlebug concentrations reach one
pest per plant. Imidacloprid (a synthetic nicotine) kills all Figure .4.43: Mealybugs and scales (all x4).
insects with sucking mouthparts. A. Pseudococcus longispinus (from Comstock 1904);
B. Palthenolecanium corni (McPartland); C. lcerya
purchasi (McPartland redrawn from Sorauer 1958).

MEALYBUGS & SCALES

Mealybugs a n d scales suck plant sap, as d o their 3. EUROPEAN FRUIT LECANIUM
Homopteran cousins, the aphids and whiteflies, in the sub- Parthenolecaniurn corni (Bouche) 1844, Homoptera; Coccidae.
order Sternorrhyncha. They also gum-up plant surfaces with =Lecaniurn corni Bouche, =Lecnnium robinnrium Douglas 1890
honeydew. Honeydew attracts ants and supports the growth Description: First and second instar nymphs are tiny (<0.5mm),
green, and active. Later instars turn brown and settle on twigs and
of sooty mould. stems. Adults are oval, brown with dark brown mottling, fluted or
ridged near the edges of their bodies, 3-5 mm long. Egg-bearing
1. LONG-TAILED MEALYBUG females become quite convex, nearly hemispherical (Fig 4.43).Eggs
Psetldococc~~s longispinus Targioni-Tozzetti 1867, Homoptera; look like a woolly white ring appearing under the edges of the fe-
Pseudococcidae. male's body.
=Pseudococcusndonidtlnz (Linnaeus)Zimmerman 1948,nec Coc-
cus ndonidum L i ~ a e u 1767
s Life History & Host Range
Description:Young nymphs have tiny yellow bodies. With age,
In warm climates, nymphs and adults overwinter on
they become covered with waxy secretions and turn white; late-
instar females grow four elongated wax filaments from their cau- twigs and branches. In cool climates only adult females
dal ends. Adult females are broadly oval, segmented, 2 4 mm long overwinter. Females lay 200-300 eggs in May-June and die
and 1-2 mm wide. Tails nearly equal bodies in length (Fig 4.43). (the body provides a protective covering). Eggs hatch in June-
82 Hemp Diseases and Pests

July, and young nymphs feed on leaves near midveins. As (described below). Harmonia axyridis and Lindoris lophanthae
winter approaches they move to twigs and stems. One gen- prey on many scales and mealybugs (described below). Pre-
eration arises per year. dation of scales by Chysoperla carnea seems less effective (de-
Under different Latin names this pest has been recorded scribed under aphids).
o n Ca~irlabisin Germany (Kovacevic 1929), Russia Turning to parasites, Encarsia formosa attacks some soft
(Borchsenius 1957) and central Asia (Mostafa & Messenger scales but much prefers whiteflies. Scales are parasitized by
1972). P. corni usually infests fruit trees and cane crops in Aphytis melirtlls and Metaphycl~shelvolus, Encytus lecaniort~ni
Europe, north Africa, south Asia, Australia, and the west (described below), and Microterys flavus and Encytus
coast of North America. It may be the unnamed small brown lecaniorurn (wasps currently u n d e r investigation).
scale that Frank (1988) described on stems of Californian Leptolnastix dactyloiii and ~ n a ~ y rpsetldococci
t~s arepopular
marijuana. biocontrols of the citrus mealybug (Planococcus citri Risso).
Although polyphagous P, citri does not appear in the Can-
4. HEMISPHERICAL SCALE nabis literature, we describe the biocontrols below. The fun-
Snissetia coffeae (Walker) 1852, Homoptera; Coccidae. gus Verticilli~imlecanii kills scales and mealybugs, but works
=Saissetia Izerrzispimerica (Targioni) 1867 better against aphids and whiteflies (described under the latter).
Description: Adults look like the aforementioned P.corni-hemi-
spherical in shape, like miniature army helmets, light brown be- Rodolia cardinalis "Vedalia"
coming dark brown, 2-3 mm diameter. Eggs are tiny, oval, pink- BIOLOGY: The ladybeetle that saved agricultural Cali-
beige, in clusters of 400-900 under the shell of dead females. First-
instar nymphs (crawlers) are flat, pink-beige, 0.7 mm long, with
fornia from I, pi~rchasi-entomologists imported only 129
two red eye spots. Older nymphs develop a pattern of ridges in an vedalias from Australia, but within 18 months their offspring
H-shape on their dorsal surface, and begin to dome. nearly eradicated I, pt~rchasifrom Californian orange groves
(Metcalf et al. 1962).
Life History & Host Range APPEARANCE & DEVELOPMENT Adult vedalias have
First and second instar nymphs crawl and feed on round, convex bodies, wing covers red, densely pubescent,
leaves. Later instars settle on branches and stems. They tend marked with irregular black spots, 3-5 mm long (Fig 4.4).
to group in clusters, and secrete a lot of honeydew. Several Larvae are pink, wrinkled, spindle-shaped, covered with soft
overlapping generation arise per year in warm glasshouses. spines, and reach 8 mm long. Eggs are red and oval. Females
Bush Doctor (pers. commun. 1992) observed S. coffeae lay 150-190 eggs near I. ptrchasi egg masses. Young larvae
infesting branches of glasshouse marijuana in Becker, Michi- feed on eggs. Mature larvae and adults feed on all scale
gan. S. cofleae lives in the tropics, subtropics, and warm en- stages. Up to 12 generations arise per year in hot, dry, in-
closures. It probably came from South America. S. coffeae at- land valleys. Pupae overwinter in cooler coastal climates.
tacks a wide range of woody and herbaceous hosts, from APPLICATION: Supplied as adults. Unfortunately the
Ficus to ferns. pest I. pt~rchasiabsorbs plant poisons from certain crops, ren-
dering it unpalatable to vedalias. R. cardinalis is susceptible
5. WHITE PEACH SCALE to many insecticides (especially juvenile growth hormones);
Pseildnl~lacaspis
pentagona (Targioni-Tozzetti)1886, Homoptera; DDT almost wiped the species out during the 1950s.
Coccidae.
Description: Adults are broadly oval with slightly notched Cryptolaemtrs montrouzieri "Mealybug destroyer"
sides, white to reddish pink, and usually less than 10 mm long. BIOLOGY: "Crypts" eat many mealybug species, as well
as young scales and aphids. Californian entomologists im-
Life History & Host Range ported these ladybeetles from Australia in 1891, as biocontrol
White peach scales are polyphagous, but prefer the pioneers against citrus mealybugs. They do best at 27OC
branches and trunks of woody plants. Shiraki (1952) cited P. (range 17-32OC) and 70-80% RH.
pentagona on Japanese hemp. The species originated in the APPEARANCE: Adults are 4 mm long, with red-orange
far East but now infests all of subtropical Asia, southern heads and thoraxes, and shiny, black-brown wing covers
Europe, and the southern USA (from Maryland down to the (Plate 38). Larvae are long (up to 13 mm) and covered with
Gulf of Mexico). woolly projections of wax-they resemble monstrous, di-
shevelled, mobile mealybugs. Eggs are white or yellow, ob-
DIFFERENTIAL DIAGNOSIS long, and found in the cottony egg-masses of their prey.
The long tails of P. longispinlis and the fluted egg sacs of DEVELOPMENT Adults overwinter in moderate climates
I. p~~rchilsl
make these pests easy to identify, at least as adults. but cannot survive cold winters. Adults feed on mealybug
Nymph stages can be confused with whitefly nymphs, eggs and young larvae; females lay up to 500 eggs, one per
aphids, or mealybug destroyers. P. corni and S. coffeae can be host egg-sack. Young larvae eat mealybug eggs and young
confused with other small, sessile, brown, barnacle-like pests, larvae; older C. montrot~zierilarvae attack all mealybug . -
such as the black scale Saissetia oleae (Bernard), and the stages. Larvae eat up to 250 mealybugs and eggs, then pu-
California red scale Aonidiella aurantii (Maskell). pate in sheltered places on plant stems or greenhouse struc-
tures. The life cycle takes 29-38 days at 27°C. Adults live
CULTURAL & MECHANICAL CONTROL another 50-60 days.
(method numbers refer to Chapter 9) APPLICATION: Supplied as adult beetles in trays or bot-
Methods 1 (sanitation), 2b (deep ploughing), and 9 tles. Store up to one or two days in a cool (10-15°C), dark
(hand removal) are the most important cultural controls of place. Release in the early morning or late at night to de-
mealybugs and scales. crease dispersal. Release five beetles per plant or 20-50 per
m2 in infested areas (Thomson 1992).
BIOCONTROL (see Chapter 10) NOTES: Short days may send this species into diapause.
Controlling cottonycushion scales with Rodolia cardinalis C. molztrouzieri works best-in heavy infestations. It tends to
is an international success story. Other useful ladybeetles fly away before its job is done, like most ladybeetles. To in-
include Cryptolaemus montrouzieri and Rhizobius ventralis duce it to stay, use strategies described under Hippodamia
 Chapter 4: Insects and Mites 83

conilrrgeizs, in the section on aphids. C. nzontrouzieri must lay and Psel~dal~lncnspisspecies). It is native to India and Paki-
its eggs in host egg masses, so it cannot reproduce in pests stan and tolerates a wide range of temperatures (13-33°C)
that give birth to lixte larvae, such as I? longispinus (Cherim and moderate humidity (40-609/0 RH).
1998). This biocontrol is compatible with Lepton~astix APPEARANCE & DEVELOPMENT: Females are tiny
dactylopii. Avoid most insecticideshhile utilizing ladybeetles, yellow-brown wasps, 1.2 mm long. They are attracted to
especially diazinon. Cherim (1998) exercised caution when pheromones released by female scales when females are
~isingwhite sticky traps-the beetles are attracted to white. ready to reproduce. Ready females loosen their attachments
Mass rearing of C. n~oiztro~~zieriwas hampered for years by a to plants, allowing males to fertilize them. A. ine11111isuses
parasitoid of its own that proved difficult to eradicate this window of opportunity to lay eggs in about 30 scales.
(Gilkeson 1997). Eggs hatch into larvae which kill female scales, pupate there-
in, then emerge as adults. The life cycle takes two or three
Hnrmoilia asyridis "Multicolored Asian ladybeetle" weeks in optimal conditions. Adults live another three weeks.
BIOLOGY:This predator was imported from Japan by APPLICATION: Supplied as adults in large vials and ice-
the USDA as a successful biocontrol against scales and cooled boxes. A. nzelilzl~sshould be released immediately,
aphids. H. nsyridis also feeds on whiteflies, mealybugs, and during the mating flight of male scales. In orchards, they are
even some mites. The species does best in moderate tem- released 25,000-250,000 per ha (= 2.5-25 per m2). Indoors,
peratures and humidity (optimally 21-3O0C, 70Y0 RH). Cherim (1998)controlled light infestations with 16-24 wasps
APPEARANCE &DEVELOPMENT:Adults are 8 mm long, per rn' every three weeks, and treated heavy infestations
orange w ~ t hblack spots (they are also called Halloween with 24-48 wasps per m2 every week. A. wleliizus tolerates
ladvbeetles) Adults o.i erwinter; females lay up to 700 eggs, Bt, sabadilla, and abamectin (Van Driesche & Bellows 1996).
lar~-.ae
resemble 10 rnm alligators with orange spots. ~a&;ae
take 25 days to reach adulthood; adults then live up to three Leptomastix dactylopii
years (Cherim 1998). BIOLOGY: A parasitic chalcid wasp that lays eggs in
APPLICATION: Supplied as adults. Store for a week at third-stage larvae (and sometimes adults) of Platlococct~scitrz.
8-10°C. Koppert (1998) recommended releasing one preda- It is native to Brazil and does best at 2S°C and 60-65'% RH.
tor per 50 pests. Other sources recommend four predators APPEARANCE & DEVELOPMENT: Females are ~ e l l o t \ ~ -
per m' crop area, or eight predators per m3 foliage volume, brown, 3 mm long; male wasps are smaller and have halry
repeat~ngevery three weeks, two or three times. antennae. When searching for mealybugs, females prefer
NOTES: In Japan the adults overwinter in cliffs, but in walking to flying, so they-do not disperse very efficiently.
the USA, the next best thing is a house. The adults aggre- Females lay up to 200 eggs, one per mealybug. Parasitized
gate in large numbers on vertical surfaces, typically swarm- mealybugs turn into dark, swollen mummies. Wasps emerge
ing on light-coloured walls with southern exposures. They from small, round holes. The life cycle takes about three
enter houses through cracks and become a nuisance. Like weeks in optimal conditions.
all ladybeetles, H. asyridis can be "flighty." See comments APPLICATION: Supplied as adults in shaker bottles or
regarding Cr!lptolaetrlus ~?zo~ltrol~zie~i,
above. tubes. They must be dispersed the day of receipt, either in
the early morning or evening. Release two wasps per rnz as
Liildovis (Rhyzobius) lopl~alztlzae a preventative o r five wasps per plant if pests a r e evident.
BIOLOGY: Yet another ladybeetle that feeds on imma- The wasps work well at low pest densities. L. liactylopii can
t~lresoft scales, including brown and black scales (Saissetia be combined with predators such as C. nzontro~~zieri.
and Pnrasaissctia species), and red scales (Aonidiella species).
In the absence of soft scales, L. loplmnthae eats immature hard Anagyrus pseudococci
scales and mealybugs. The species does best in moderate BIOLOGY: A parasitic Encyrtidid wasp that lays eggs in
tern-peratures (15-25°C) and a wide humidity range (20- third-stage larvae (and sometimes second- and fourth-stage
90';'o RH). larvae) of Platzococclrs citri. A related species, Ailagyrtls
APPEARANCE & DEVELOPMENT: Adults are small, pu- fiisciventris, is also available.
bescent, black beetles with red-orange highlights, 2.5 mm APPEARANCE & DE\'ELOPMENT: Females are bro~vn,
long. Fernales lay over 100 eggs, one at a time, on plants 1.5-2 mm long, with black and white banded antennae. Male
among scales. Larvae are grey, reach 3 mm long, and prey wasps are smaller, without the distinctive antenna bands.
on young scales and eggs. The life cycle takes three weeks. Wasps lay eggs in mealybug larvae, which turn into swol-
Adults live another six to eight weeks, eating constantly. The len, yellow-brown (striped) mummies. Wasps emerge frnrn
final generation of the season overwinters as adults. mummies via small, ragged holes. The entire lifecycle may
APPLICATION: Supplied as adult beetles in trays or bot- take only three weeks.
tles. Store up to one or two days in a cool (10-15"C), dark APPLICATION: Supplied as adults in shaker bottles or
place. Release in early morning or late at night. For light tubes. Store up to one or two days in a cool (8-1O0C), dark
infestations, release three to six beetles per m2, repeat every place. Koppert (1998)recommended releasing 0.25 wasp per
three weeks, two or three times. Double the rate for heavier mz every 13 days as a preventative. For infestations, release
infestations (albeit an expensive proposition). L. lophantlzae one wasp per mz per week.
is probably compatible with parasitic biocontrols.
Metaphyctis helvolus
Rlzyzobills ventvnlis BIOLOGY: A parasitic and predatory wasp that controls
BIOLOGY: A ladybeetle related to the aforementioned many species of soft scales, especially black and brown scales
species; it also preys on mealybugs and scales. Adults are (Snissetia,Parasnissetia, and perhaps Cocc~~s species). It is na-
shiny \ elvety black with reddish abdomens. tive to semitropical regions and does best in warm, relatil~ely
dry glasshouses (24-32", RH 50%). A related species from
Apllytis inelinus Australia, Metaplzycus alberti, was released in California to
BIOLOGY: A parasitic chalcid (Eulophidae) wasp that control Cocct~shesperidil~in.
feeds on hard scales (Aonidiella,Aspidiotus, Ql~nclraspinioti~s, APPEARANCE & DEVELOPMENT Wasps are tin); yellow
84 Hemp Disealses and Pests

and black, 1.3 mm long. Female wasps lay 100400 eggs, one termites infesting marijuana in the southern USA. The first
at a time, under immature scales. Larvae burrow into scales, report was from a Florida grower who invited disaster by
kill them, pupate therein, and emerge as adults. The life cy- mulching his soil with wood chips. Clarke (pers. commun.
cle takes 24 days. Adults live another two or three months. 1996)found termites infesting marijuana in Mexico and equa-
Adults poke holes in scales and feed on body fluids. torial Africa. Termites hollowed out the main stem and
APPLICATION: Supplied as adults, which should be re- branches, up to the levels of flowers, and then plants col-
leased immediately. Used preventively, release one or two lapsed.
wasps per m2 per month. For light to moderate infestations,
release five to ten wasps per plant or ten wasps per m2, every DIFFERENTIAL DIAGNOSIS
two weeks until controlled. M.helvolus is not useful in heavy Symptoms from ants and termites are similar to those
infestations. According to Cherim (1998), the wasps are at- caused by white root grubs, root maggots, nematodes, root
tracted to bright lights and yellow sticky traps. The wasps rots from assorted fungi, and drought.
live longer if supplied with honeydew or nectar.
CULTURAL & MECHANICAL CONTROL
CHEMICAL CONTROL (see Chapter 11) (method numbers refer to Chapter 9)
Synthetic pheromone lures are available for some Observe methods 1 (remove stumps, roots and other
mealybugs and scales (Olkowski et al. 1991). Puritch (1982) termite-attractive debris), 2a (deep ploughing), 7a (termites
killed 96.9% of mealybugs with Safer's insecticidal soap attack water-stressed plants), and 8 (optimize soil structure
spray. Neem repels mealybugs and young scales, oil kills and nutrition). Avoid mulching with woody materials ex-
mealybugs and young scales. Frank (1988)suggested spray- cept for cedar chips. Compost, humus, and manures exacer-
ing mealybugs with nicotine sulphate, or daubing mealybugs bate ant infestations. Frank & Rosenthal(1978)repelled ants
with cotton swabs dipped in rubbing alcohol. Buprofezin and termites by flooding soil near infested plants. Clarke
and kinoprene are synthetic insect growth hormones toxic (pers. commun. 1996) located nests and flooded them with
to many mealybugs and scales. Imidacloprid (a synthetic water. Frank (1988), a proponent of hydroponics, noted that
nicotine) kills all insects with sucking mouthparts. ants and termites only cause problems in soil-grown plants,
not hydroponically-grown plants.

BIOCONTROL
ANTS & TERMITES Fire ants have been biocontrolled experimentally with
Ants and white ants (termites) often curse Caizizabis in Pyemotes tritici, Pseiidacteorz species (described below),
semitropical climates. Although they look somewhat simi- Thelohania solerzopsae, and Steinernema nematodes. The ter-
lar, they are from unrelated orders. mite 0. obestls is parasitized by Termitomyces striattls (Belli)
Heim, but this fungus is not commercially available. Other
SIGNS & SYMPTOMS termites are controlled with Mefarhiziilm anisopliae (see un-
Ants and termites tunnel into taproots and stems (Plate der aphids) and Steinernema carpocapsae (see under cutworms).
39). Plants wilt and sometimes collapse. They are easily
pulled from the ground. Ants also colonize aphid-infested Pseudoacteon species
plants. Ants protect aphids from parasitoids and predators, BIOLOGY: About 20 species of parasitic humpbacked
and the aphids supply ants with honeydew. Leafcutter ants flies (family Phoridae) are known to infest ant nests. Related
cut pieces of leaves and carry them off to underground nests. flies infest termite nests. Pseildoacteon species from North
In severe cases, leafcutter ants devour entire marijuana plants, America parasitize S.geminata, and Pseudoacteorz species from
leaving only the roots. Leafcutter ants are nocturnal and can South America parasitize S , invicta and S . richteui. Adult flies
destroy several plants a night. are tiny, half the size of their hosts. Females dive-bomb the
back of ants and quickly insert a single egg, using their hy-
1. ANTS podermic-like ovipositor. Females repeat this aerial attack
Cherian (1932) cited the red fire ant Solenopsisgeminata on dozens of ants. Eggs hatch and maggots migrate into the
(Fabricius) 1804 (Hymenoptera; Formicidae) chewing seed- head of ants. Maggots do not eat vital parts, so ants remain
lings and tunnelling into roots of mature marijuana plants. alive until maggots are fully mature. At that point, maggots
Siege1 (1989) also reported "red ants" attacking marijuana release an enzyme that causes the ant's head to fall off. Pu-
in Mississippi. S. geminata workers are three to six mm long, pation occurs in the decapitated head; the adult fly emerges
and can inflict painful bites and stings. The species is native from the mouth. The life cycle takes four to six weeks.
to Central America and the Caribbean, north to Texas and
Florida, and has been accidentally introduced into India, Pyemotes tritici
Africa, and some Pacific islands. S. geminata is the only crea- BIOLOGY: These predatory mites are used against fire
ture nasty enough to slow the spread of imported fire ants, ants and other pests in stored grain. Unfortunately, I? trifici
Solenopsis invicta and Solenopsis riclzteri. is known as the "straw itch mite," because it bites people
Frank & Rosenthal (1978) described unidentified ants who handle infested materials.
damaging roots of marijuana in California. Clarke (pers.
commun. 1996) encountered leafcutter ants (Atta species) CHEMICAL CONTROL
in Mexico. Overnight, the ants cut entire plants into little Cherian (1932) irrigated infested soil with oil to drive
pieces and carried the pieces to their underground nests. away termites and ants. Drench the soil with pyrethroids
(especially tralomethrin), neem oil, or abamectin. Frank &
2. TERMITES Rosenthal (1978) repelled ants from plants by dusting the
Odontotermes obesus (Rambur) 1842 (Isoptera; surrounding soil with cream of tartar. Clarke (pers. commun.
Termitidae) has been a major pest of Indian marijuana 1996) stabbed an opening into nests and drowned ants and
(Cherian 1932). 0.obesils occurs across the Indian subconti- termites by pouring water into the nests. More effectively
nent, east into Burma. The species builds spectacular mounds but less ecologically, he poured a kerosene-gasoline mix into
up to 3 m tall. Alexander (1984c, 1985)cited two instances of nests and burned them.
 Chapter 4: Insects and Mites 85

GALL MIDGES & roots often rot from invading soil fungi. Plants, if they
survive, become yellow and stunted. They wilt during the
ROOT MAGGOTS day but recover at night. D. platlira pupates 2-4 cm under
You'd think a midge named Chortophila cannabina Stein the soil near damaged plants; two to five generations arise
would infest Cannabis, but no. Its host is the hemp linnet, per year.
Carduelis cannnbina (a bird named for its love of hemp seeds). D.platura infests hemp in Japan (Harukawa & Kondo
Some gall midges are carnivorous and predatory-for instance, 1930, Shiraki 1952), and Europe (Rataj 1957). A Delia spe-
Aplzidoletes aphidimyza larvae devour aphids. They are used for cies has also been reported in India (Bantra 1976).D. platura
biocontrol. Other gall midges attack plants. They are related is cosmopolitan. It usually feeds on decaying vegetable
to root maggots. matter, but attacks a wide range of crops (Hill 1983).Ger-
minating seeds of corn and bean are particularly vulner-
1.NETTLE MIDGE able to attack, especially in cold, wet springs when seed
Melarlogromyza ~lrticivoraSpencer 1966, Diptera; Agromyzidae. germination is slow (Howard et al. 1994).
Description: Adults are green and black flies with 2.4 -3.0 mm
wingspans. Larvae are small, white maggots without legs or distinct 4. CABBAGE MAGGOT
heads. Delia radicum (Linnaeus) 1758, Diptera; Anthomyiidae
=Chortophila brassicae Wiedemann 1817,=Chortophila brassicae
Life History & Host Range Bouche 1833
This species probably overwinters as pupae in soil. Adults Description: Female flies have dark grey bristly bodies, 7
emerge in spring and lay eggs on host plants. Maggots hatch mm long, with black stripes on their thoraxes. They lay fusiform-
and form galls in stems of Pakistani marijuana (Mushtaque et shaped eggs covered by longitudinal striations. Cabbage mag-
al. 1973).Baloch et al. (1974)reported M . urticivora only attacks gots are spike-shaped, with pointed heads and wide flat posteri-
Cannabis growing near nettles (Urtica species, its primary host). ors, and the posteriors have paired tubercles that are forked at
their apex.
M , llrticivora lives in the western Himalaya.
Life History & Host Range
2. OTHER GALL MIDGES
In early spring adults emerge from overwintering
Mushtaque et al. (1973) described an Asplzondylia species
pupae 2-10 cm underground. Females lay eggs in soil abut-
(family Cecidomyiidae) forming galls in male flowers in Paki-
ting plant roots. Larvae hatch within a week and burrow
stan. Baloch et al. (1974) reported, "It does not appear to be a
into roots. After feeding for several weeks, larvae pupate
promising biocontrol agent [of Cannabis],because pollination
in the plant or in nearby soil. By late June the next genera-
is almost complete by the time the infestation starts."
tion of flies emerge to lay eggs. A third generation may
emerge in autumn. Because maggots thrive in cool moist
3. SEEDCORN MAGGOT
weather, only the first generationcauses much damage.
Delia platllra Meigen 1826, Diptera; Anthomyiidae.
=Chortoplzila cilicr~ltaRondani 1859 According to Goriainov (1914), D. radicum destroyed
Description: Female adults look like half-sized house flies, 5 mm long, 40% of a hemp crop in the former USSR. The cabbage mag-
with grey pointed abdomens (Fig 4.44). They lay fusiform-shaped eggs got normally attacks Crucifers; Yepsen (1976), in fact, sug-
covered by net-like surface ornamations. Pupae are spindle-shaped gested using Cannabis to repel D. radicum. This Old World
and brown. Maggots are 5-7 mm long, white, with pointy heads and pest was introduced into North America about 100 years
blunt posteriors, and the posteriors sport a pair of unforked tubercles. ago. Cabbage maggots can kill 95% of unprotected cab-
bage crops north of the Mason-Dixon Line (39'40' latitude).
Life History & Host Range
D. platl~raoverwinters as pupae; adults emerge in early DIFFERENTIAL DIAGNOSIS
spring. Females lay 100 eggs, a few at a time, in disturbed soil. Galls can be caused by European corn borers, hemp
Hatching maggots burrow into ungerminated seeds and young borers, and assorted stem-boring beetles and weevils. Some
seedlings. Seeds are often completely destroyed. Seedlings fungi and bacteria also cause swellings confused with in-
become honeycombed with slimy brown tunnels. Damaged sect galls (e.g.,Fusarium canker). Root maggots cause wilt
symptoms similar to those caused by white grubs, nema-
todes, and root-rotting fungi. Their symptoms may also
be confused with those caused by damping-off fungi. Dig-
ging up plants or seeds reveals the tiny maggots.

CULTURAL & MECHANICAL CONTROL

(method numbers refer to Chapter 9)
To discourage root maggots, observe methods 1 (sani-
tation), 2a (deep ploughing), 2b&c (sterilizing or pasteur-
izing soil), 3 (eliminate cabbage-family weeds), 4 (plant
late to avoid the first wave of egg-laying D , radicllrn flies,
about two or three weeks after mustard weeds flower), 7a
(maintain proper soil moisture), 12c (light traps), and 13
(mechanical barriers). Protect seedlings from pregnant flies
with a tent of gauze or finely-woven screen (20-30 threads
to the inch). When plants outgrow tents, fit a 15 cm
diametre disk (made of felt, foam-rubber, or tar paper)
around the base of each stem. Plastic sheeting mulch also
works if the sheets are pulled tight around the base of
Figure 4.44: Delia platura, enlarged x6. A. Female adult; stems. This prevents flies from laying eggs in soil next to
B. Reticulate egg; C. Puparium; D. Larva; E. Close up of plants. Disks and plastic mulch also provide a humid
larval head; F. Close up of larval posterior (from Hill 1994). microhabitat for predatory ground beetles, and the con-
86 Hemp Diseases and Pests

served soil moisture permits plants to better tolerate minor Canada, grasshoppers destroyed apical meristems, stunting
infestations without wilting. Organic growers pour a ring of plants and severely reducing seed production (Scheifele
wood ashes around seedlings, but this must be repeated af- 1998). Crickets feed on young seedlings, causing cutworm-
ter every rainstorm. Bait sticky tape and water traps with like damage. Mole crickets feed on roots.
allylisothiocyanate to lure flies to their deaths. Although these
traps will not provide complete control, growers may moni- 1. TWO-STRIPED GRASSHOPPER
tor traps to determine when other controls are needed. Melatzopltis bivittattis (Say) 1825, Orthoptera; Acrididae
Description: These short-horned grasshoppers are members
BIOCONTROL (see Chapter 10) of the "spur-throated" subfamily of Acridids, they sport a short
Parasitic nematodes control root maggots; D. radiclinz tubercle protruding from between their front legs. Females coloured
olive-brown to brownish-yellow above, pale yellow below, without
was controlled by applying Steinernema feltiae (described
spots but with a distinct pale stripe on each side of the head, begin-
under fungus gnats), or Heterorhabditis bacteriophora and ning behind the eyes and extending down the back. Hind tibiae
Steinernema riobravis (see white root grubs and flea beetles). often red or purple-brown with black spines. Females average 29-
The rove beetle Aleochara bilineata has been used against Delia 40 mm long, males average 23-29 mm.
species (described below). Not-yet-commercially available
controls include maggot predators (Strongwellsea castrans Life History & Host Range
and Trybligrapha rapae), and the fungus Entoinophthora M . bioittatus overwinters underground in the egg stage.
muscae, which infects many types of flies (see Plate 40). A Each female lays one or two clusters of eggs. Southern
companion crop of basil repels adult flies (Israel 1981).Mari- populations reach maturity by early June, northern
golds, parsnips and onions also deter the maggots. populations appear by late July. The species ranges across
southern Canada from Newfoundland to British Columbia,
A'leochara bilineata south to Mexico in the west and North Carolina in the east;
BIOLOGY: Apredatory rove beetle whose larvae are para- M , bivittattis predominates in the Great Plains, where it has
sitic. The species occurs in North America and Eurasia in infested hemp in Manitoba (Moes, pers. commun. 1999).
temperate regions. Two-striped grasshoppers frequent grass pastures and prai-
APPEARANCE & DEVELOPMENT: Adults have elongate, ries, as well as clover fields. The pest occasionally migrates
glossy black bodies 3-6 m m long, with small reddish-brown in masses, "eating the choicest of everything." Arelated spe-
forewings. They look like earwigs without rear pinchers. Fe- cies, the migratory grasshopper (Melanoplus sanguinipes
males lay up to 700 eggs in soil around root maggot-infested Fabricius), also infests hemp in Manitoba.
plants. Laniae hatch in five to ten days, and actively search
for root maggot pupae. Larvae chew into maggot pupae and 2. SPRINKLED LOCUST
complete their life cycle therein, emerging as adults after 30- Chloealtis conspersa Harris 1841, Orthoptera; Acrididae
40 days. Adults eat up to five maggots per day. Two genera- ~ e s c r i ~ t i d These
n: short-horned grasshoppers are members
tions usually arise per year. of the "slant-faced" subfamily of Acridids, their heads look stream-
APPLICATION: A. bilineata has been mass reared in the lined. Females vary in colour from dull yellow to dark brown, with
former Soviet Union, Europe, and Canada (Hoffmann & front wings speckled by small black spots. They average 20-28 mm
Frodsham 1993). Some strains are tolerant of insecticides. in length. Males are 15-20 mm long, light brown with a black bar
across their thorax, and lack sprinkled tegmina. Both sexes sport
CHEMICAL CONTROL (see Chapter 11) yellow to red hind tibiae, and antennae 10-12 mm long.
Eliminate root maggots with neem and horticultural oil,
applied as soil drenches. Life History & Host Range
C. conspersa overwinters in the egg stage, underground,
in masses of 15-50 eggs encased by a hardened gummy sub-
stance. Sprinkled locusts make their home in grassy thick-
GRASSHOPPERS & ets alongside pastures, fields, and streams. They live east of
THEIR ALLIES the Mississippi, from the Canadian border down to the
The order Orthoptera includes grasshoppers, locusts, mountainous regions of Virginia and North Carolina. Males
crickets, and cockroaches. Differentiating grasshoppers from produce a familiar tsikk-tssikk-tssikk song. Bush Doctor (un-
locusts is like separating mushrooms from toadstools. These published data 1990)found this species on marijuana in New
moderately large and elongated insects have prominent eyes, York, and DeWitt (unpublished) collected C. conspersa from
jaws, and large hind legs (Fig 3.5 & Plate 42). They may be feral hemp in Illinois.
winged or wingless. Not all Orthoptera are evil-Praying
mantids (family Mantidae) are beneficial carnivorous insects. 3. CLEARWINGED GRASSHOPPER
As a corollary, not all evil pests infest Cannabis; Rothschild et Camntila pellucida (Scudder) 1862, Orthoptera; Acrididae
al. (1977)fed a swarm of locusts (Schistocerca gregaria Forsk.) Description: These small short-horned grasshoppers are mem-
bers of the third Acridid subfamily, the "band-winged" grasshop-
nothing but Cannabis, and the diet proved lethal.
pers-they do not have spurred throats or slanted faces. This par-
ticular species doesn't even have banded wings. Females are col-
SIGNS & SYMPTOMS oured brownish-grey with black markings until they become sexu-
Grasshoppers, locusts, and crickets eat large, round, ally active, and turn bright yellow. Forewings are mottled with light
smooth-edged holes in leaves. In heavy infestations, Lassen stripes along their angles, hindwings are transparent. Females av-
(1988) reported plants being stripped to stalks in a matter of erage 21-25 mm long; males average 17-21 mm. Nymphs are strik-
days. Swarms of locusts in western Africa d o the same in a ingly coloured black with white to tan markings. Eggs are 4 mm
matter of minutes (Clarke, pers. commun. 1994, see Plate 41). long, laid in clusters of ten to 30.
Clarke also reported large grasshoppers in Mexico biting
through stems of young seedlings to topple them so the pests Life History & Host Range
could easily feed on leaves. Grasshoppers in Indiana chewed C , pellucida overwinters in the egg stage, underground,
through stems and branches, causing limb tips to topple. In usually in bare ground, often along roadsides. This species
 Chapter 4: Insects and Mites 87

emerges and matures earlier than the two aforementioned

species. C. pellilcida ranges across western North America
from northern Mexico to northern Alberta; east of the Great
Plains the species inhabits land along the USA-Canadian
border. It has infested hemp in Manitoba (Moes, pers.
commun. 1999).C. pellz~cidafrequents grassy pastures on high
ground. The nymphs sometimes migrate in swarms, caus-
ing trouble. This small species has a big appetite: populations
greater than 25 per m2 can completely denude rangeland
forage grasses.

4. STINK GRASSHOPPER
Zonoceriis elegans (Thunberg) 1773, Orthoptera; Acrididae
Description: Adults are handsome hoppers, mostly dark green
with bold stripes of black, yellow, and orange, 35-55 mm long.
Nymphs are black with appendages ringed with yellow or white.
Eggs are sausage shaped, 6 mm long, and laid in the soil within
spongelike masses of dried froth about 2.5 cm wide.

Life History & Host Range

Rothschild et a2. (1977) fed Z . elega~zsnymphs a diet of
pure Cannabis for six weeks, and most survived. Rothschild
demonstrated that Z. elegans sequestered some cannabinoids
in body tissues (as a predator deterrent?), and excreted the
rest in frass. This colourful, aposematic grasshopper exudes
an unpleasant odour when handled. Hill (1983) called it the
skink grasshopper, Sorauer (1958) gave it the sinister mon-
icker Stinkschrecke. The pest ranges across all of Africa below Figure 4.45: Adult crickets (all approximately life size).
the Sahara, and feeds on many dicot crops in the seedling A. Gryllotalpa hexadactyla (from Comstock 1904);
stage, especially cassava and finger millet (Hill 1983). One B. Acheta domesticus (from Ill. Nat. Hist. Surv.);
generation arises per year. C. Gryllus deseiius; D. Gryllus chinensis.
(C & D from Anonymous 1940)
5. CITRUS LOCUST
Clzorldracris rosea (DeGeer) 1773, Orthoptera; Acrididae black and produce familiar nocturnal chirping sounds.
Description: Adults are yellow to greenish brown, with rosy- Selgnij (1982)described unidentified crickets attacking mari-
v~oletwings. They have red hind femurs with yellow spines. Body juana seedlings. Rataj (1957) cited the house cricket Acheta
length averages 64-85 mm (females) and 49-60 mm (males). domesticus (Linnaeus) 1758 infesting Czech hemp. A.
C. rosea attacked mature hemp crops in Taiwan (Sonan donzestict~sis a European species introduced into North
1940). Citrus locusts damage many crops, and range from America by early settlers. Adults are shiny with dark cross-
northern India to Japan. bars and light-coloured heads, 10-20 mm long (Fig 4.45).
House crickets live outdoors until the weather turns cold in
6. OTHER GRASSHOPPERS & LOCUSTS the autumn.
Lassen (1988) described unidentified grasshoppers at- Gryllus desertus Pall 1771 and Gryllus chinensis We-
tacking marijuana in California. Siege1 (1989) cited unidenti- ber 1801 are reported in Italy (Goidanich 1928, Martelli 1940,
fied grasshoppers attacking feral hemp in the Midwest. In Ragazzi 1954,Ferri 1959)and the Czech Republic (Rataj 1957).
South America, Dichroplus maculipennis Blanch 1851 (Or- These species are the same size as house crickets (14-18 mm
thoptera; Acrididae) caused "intense" damage on hemp long), but darker in colour (Fig 4.45).
(Liebermann 1944), despite predation by birds and flesh flies The mole cricket, Gryllotalpa gryllotalpa (L.), attacks
(Diptera; Sarcophagidae). hemp in Europe (Barna et al. 1982, Gutberlet & Karus 1995).
Atractomorpha crenulata Fabricius 1793 (Orthoptera; Clarke (pers. commun. 1994) reports mole crickets in Africa
Acrididae) was the most common grasshopper pest in India (Gryllotalpa gryllotalpa or Gryllotalpa afircana Pal.) and
(Bantra 1976). Sorauer (1958) called A. crenulata the Tobacco California (Gyllotalpa hexadactyla Perty). Mole crickets are
grasshopper, which ranges from India to Malay. Adults are distinguished by their broad, shovel-like front legs (Fig 4.45).
spotted green with reddish wings and abdomens, 24-35 mm They burrow underground and eat roots and tubers.
long. Other Asian pests include Atractotnorpha bedeli The tree cricket Oecanthus indicus Saussure 1878 has
Bolivar 1884 in J a p a n (Shiraki 1952), H i e r o g l y p h u s been reported in India (Bantra 1976). Tree crickets are pale
nigrorepletus Bolivar in India (Roonwal 1945), a n d green. Females damage plants by splitting open stems with
Chrotogonus saussurei Bolivar 1884 in India (Cherian 1932). their long, strong ovipositors. 0. indictis ranges from India
In Italy, Goidanich (1928) cited Tettigonia cantans to Java to Taiwan. A related species, Oecanthus celerinictus
(Fuessly) 1775 and two uncommon Tettigoniidae, Tettigonia Walker, damaged Mississippi marijuana in late summer and
ferrtrgitzea and Tettigonia orientalis. These long-horned early autumn (Lago & Stanford 1989).
grasshoppers are related to katydids. Clarke (pers. commun.
1994) reported minor damage by katydids in California, In- 5. COCKROACHES
diana, and western Africa. Frank & Rosenthal(1978) described unidentified cock-
roaches attacking marijuana. They were probably Blattella
4. CRICKETS gevmanica (L.) or Supella longipalpa (F.). Dass Baba (1984)
Crickets (family Gryllidae) have shorter legs and longer described unidentified Hawaiian cockroaches eating Can-
antennae than grasshoppers and locusts. Most species are nabis seeds in peat pots.
88 Hemp Disealses and Pests

DIFFERENTIAL DIAGNOSIS APPEARANCE & DEVELOPMENT: M. flavoviride is a

Grasshopper and locust damage can be confused with hyphomycete with branching, phialidic conidiophores and
damage caused by caterpillarsor flea beetles. Damage caused simple oval spores (conidia).Conidia in contact with insects
by crickets may be confused with cutworms or damping-off quickly germinate and grow into their hosts. Infected insects
fungi. Root damage caused by mole crickets may be con- stop feeding, then die in four to ten days, depending on the
fused with root maggots or root grubs. temperature. In humid conditions, M. flavoviride reemerges
from dead hosts to sprout more green conidia and repeat the
CULTURAL & MECHANICAL CONTROL life cycle.
(method numbers refer to Chapter 9)
Observe methods 2a (deep ploughing), 3 (eliminate Entomophthora grylii
grassy weeds), 6 (do not rotate after pasture), 7a (avoid BIOLOGY:A fungus that infests grasshoppers and locusts.
drought), and 9 (hand removal). Use method 13 outdoors- It is native to North America and Africa, and does well in
screened tents and row covers will keep hoppers away from moderate temperatures. This fungus produces sticky, mu-
tender seedlings. cus-covered ballistospores, which are forcibly discharged
from dead insects. Resting spores arise within cadavers and
BIOCONTROL are mass-produced in liquid media fermenters. A related
Noserna locustae is commercially available, Metarhizium pathotype from Australia, Entomophthora praxibuli, has a
flavoviride is registered in Africa, and Entornophthora gylli is broader host range. E. praxibtili was released in North Da-
being developed (all described below). Beauveria bassiana also kota to suppress rangeland grasshoppers,but the fungus died
kills grasshoppers (described under whiteflies). Biocontrol out after several winters (Bidochka et al. 1996).
organisms must tolerate high temperatures, because sick APPLICATION: Entomophthora spores directly penetrate
grasshoppers instinctively cling to the tops of plants, seek- grasshopper skin-spores do not have to be eaten, just
ing bright sunshine ("summit disease syndrome"), and can touched. Hoppers infected with the fungus climb to the tops
raise their temperatures to 40°C. To control species that live of elevated surfaces, instinctively seeking sunshine. They
in the soil, such as mole crickets, try beneficial nematodes hang on for dear life and usually expire in the late afternoon,
(Steinernema and Heterorhabditis species, described under seven to ten days after infection (Fig 4.46).
cutworms and white root grubs, and Steinernema scapterisci,
described below). Melanoplus sanguinipes entomopoxvirus
Lassen (1988) knocked grasshoppers to the ground by BIOLOGY: The MsEPV is a DNA virus being tested
spraying Cannabis with water two or three times a day, then against grasshoppers in the western USA. The cadavers of
his chickens finished the job. Chickens also scratch eggs out 35 infected grasshoppers are required to produce a dose suf-
of the ground and eat them. Almost all birds feed on hop- ficient for 1ha of crop area (Hunter-Fujita et al. 1998).
pers (except vegetarians such as doves and pigeons). Other
natural enemies include spiders, flesh flies, snakes, toads,
and assorted rodents (field mice, ground squirrels).

Nosema locustae
BIOLOGY: A protozoan that parasitizes grasshoppers,
Mormon crickets, black field crickets, and pygmy locusts
(Locucide", SemisporeBaits; Evans discontinued Nolo Bait@
in 1993). N. loclistae does best in temperatures above 15°C.
According to Hunter-Fujita et al. (1998), the spores from a
single grasshopper cadaver can be used to treat 4 ha.
APPLICATION: Supplied as 7.5% concentrate, 0.05%
powder, and in a wheat bran bait. It can be stored for 13
weeks in a dry, cool (8-10°C), dark place. Cherim (1998) rec-
ommended N. loclistae when grasshopper populations ex-
ceed 40 m-2.He spread bait around plants at a rate of 25 bran
flakes per square foot ( ~ 2 7 0flakes m-2). Distribute bait in
the morning after the temperature reaches 16"C, and when
no rain is forecasted (rain or heavy dew makes the bran un-
palatable to pests).
NOTES: After ingesting N. loctlstae, grasshoppers stop
feeding, develop discoloured cuticles, and die a slow death.
Because of this slow death, the parasite is more effective for
long-term suppression of grasshopper populations than
stopping sudden outbreaks. Once established in a grasshop-
per population, N. loctistae spreads via cannibalism of in-
fected individuals. N. loclistae is compatible with all other
biocontrols. A related species, Nosema acridophagus, kills
grasshoppers quicker and the USDA is developing it.

Metarhizium flavoviride
BIOLOGY: This soil fungus (Green Musclea) has been
used against grasshoppers and locusts in Africa and the trop-
ics. It is closely related to Metarhizitlm anisopliae (described Figure 4.46: Grasshopper infected with the biocontrol
under aphids). fungus Entomophthora praxibuli (courtesy USDA).
b Chapter 4: Insects and Mites 89

Steinernemu scapterisci 1. CRANE FLIES

BIOLOGY: This species of Steinernema (Otinem S@, Adult crane flies resemble gigantic mosquitoes or
Proacta) was discovered in the 1980s. It parasitizes mole crick- "daddy-longlegs with wings." The adult's feeding habits are
ets (Scapterlscus species), house crickets (Acketa domesticus), unknown, but they d o not bite humans. Bantra (1976)caught
and field crickets (Gylllrs species). The gut of S. scapterisci an unidentified Tiptila species in marijuana. Bovien (1945)
contains mutualistic bacteria (Xenorkabdzrs species). S. reported the European crane fly, Tiplula paludosa Meigen
scapterisci lives in the USA and temperate South America, 1830 (Diptera; Tipulidae), attacking Danish hemp crops.
and does best in soil temperatures between 15-33°C. Watson (pers. commun. 1998) found T.paltrdosa infesting a
N O E S : Infective juveniles of S,scapterisci move through glasshouse in the Netherlands. T.paltrdosa larvae usually feed
the soil (although they are also described as sedentary on grasses. The maggots, known as leatherjackets, are pink
ambushers), and enter hosts through mouths or spiracles. to greyish-black, with black heads, 35-40 mm long. Adults
The life cycle is similar to that of other Steiizernerna species. have grey-brown bodies (up to 25 mm long) and delicate
Its commercial availability varies. milky-white wings (Fig 4.47). The species was introduced
into the Province of British Columbia and Washington State
CHEMICAL CONTROL around 1955.
Dusting plants with diatomaceous earth or spraying
plants with neem deters most grasshoppers and crickets.
Yepsen (1976) used sabadilla dust against grasshoppers and
pyrethrum against crickets. Hydroprene is a synthetic growth
hormone targeted against cockroaches. Boric acid kills crick-
ets and cockroaches. Sticky baits made from wheat bran, simi-
lar to cutworm baits, attract all Orthopterans. Poisoning baits
with pesticides works even better.

FLOWER FLIES
Black and yellow-striped syrphid flies, also called hover
flies, resemble wasps and bees. They hover over flowers to
sup nectar and serve as pollinators (second in importance
behind bees). They do not sting or bite. Larvae of some flower
flies feed voraciously on aphids, leafhoppers, and mealybugs.
Schmidt (1929) described large numbers of syrphid larvae
infesting stored hemp seed in Brandenbug. He noted the
preceding summer's crop was heavily infested with aphids,
which led to a great syrphid boom until autumn, when the
aphid population crashed. Remaining syrphid larvae, un-
able to pupate, hibernated in Cannabis flowers, ending up in
seed during threshing.
A few s y r p h i d species feed o n plants. Datta & Figure 4.47: An adult crane fly (Tiplula species) emerging
Chakraborti (1983) collected flower flies from Cannabis in from its pupal case (courtesy USDA).
northern India: Metasyrphus latifasciatus (Macquart) 1829,
Episyrpktrs balteatus (DeGeer) 1776, Ischiodon scutellaris
2. FUNGUS GNATS
(Fabricius) 1805, S y r i t t a pipiens (Linnaeus) 1758,
Fungus gnats come from t w o fly families, the
Splzaerophoria scripta (Linnaeus) 1758, and unidentified
Mycetophilidae and Sciaridae. Bantra (1976) caught a
adults of Ischyrosyphus species and Melanostoma species.
Bradysia species (Diptera; Sciaridae) buzzing around Indian
Cannabis.Arnaud (1974)found Bradysia species infesting con-
fiscated stocks of midwestern marijuana. An anonymous
author reported fungus gnats attacking a drug cultivar
('William's Wonder') in Ontario, Canada (High Tinzes No. 233,
CRANE FLIES & p. 77, 1994). For some reason, plants growing in rockwool
FUNGUS GNATS and sterile potting media are more susceptible than plants
Larvae of crane flies and fungus gnats infest plant roots growing in soil (Howard et al. 1994).
close to the soil surface. Infested plants lose vigour and col- Bradysia larvae are slender maggots with translucent
our. Close inspection reveals small surface scars on roots, bodies and black heads, growing to 4-5 mm in length. They
and fine root hairs are eaten away. Root wounds may serve pupate in soil. Bradysia adults are small, delicate, hump-
as portholes for pathogenic fungi, and the pests may trans- backed flies, with long gangly legs and beaded antennae,
mit soilborne pathogens, notably Pytkiunz and Ftisarizln~spe- grey to black in colour, and 2-4 mm long. They are poor fli-
cies (Howard et al. 1994). ers and usually come to one's attention when plants are
Crane fly and fungus gnat larvae attack plants that are moved or shaken. Fungus gnats can be confused with adult
stressed by nutrient imbalances or waterlogging. They pre- Aphidoletes apkidinzyza (aphid biocontrols). Females lay up
fer feeding on decaying vegetation and fungi in damp soil. to 200 eggs on soil or rockwool in seven to ten days. New
Fungus gnat larvae also feed on the green algae that covers generations arise on a monthly basis and overlap in warm
damp rockwool. The adults d o not feed on plants, but may climates and warm glasshouses.
become trapped in the sticky resin of mature flowers and
are aesthetically displeasing. Crane flies are common out- CULTURAL & MECHANICAL CONTROL
doors, fungus gnats dominate indoors. Observe methods 2b&c (sterilize or pasteurize soil), 3
 Chapter 4: Insects and Mites 91

Life History & Host Range CONTROL

S . pilicornis infests Cannabis in Mississippi, where it Gutberlet & Karus (1995) recommended biocontrol with
occurs abundantly from early May to the middle of August Metarhiziunz anisopliae. Beneficial nematodes may work
(Lago &Stanford 1989).The larvae act as stem borers (Sands (Heterorhabditis and Sfeitlernema species, described under
et 01. 1987). Previously, S . pilicornis was described as a host- white root grubs). Wheat is used as a trap crop for wireworms
specific pest of common purslane (Portulaca oleracea L.), in Switzerland (Hokkanen 1991). Yepsen (1976) suggested
where it acts as a leafminer (Gorske et al. 1976).At 24"C, the trapping wireworms with potatoes. Skewer potatoes on a
egg stage lasts 80-100 hours, larvae mature in 130 hours, stick, then bury them 10 cm underground, leaving the stick
pupation takes 200 hours, and the adult wasps live 25 hours as a marker. Set traps 1-3 m apart. After a couple of weeks,
(Gorske et al. 1977). Female wasps deposit eggs in slits at pull up potatoes full of wireworms. Carrot slices, untreated
edges of leaves, generally one egg per leaf, a total of 20 eggs. corn seed, and mesh bags filled with 30 g of whole wheat
flour also work.
CONTROL
In China, 15-30% of T. cannabis are killed by ladvbeetles,
lacewings, and parasitoids. Neodiprion sertijer ~ P v ' ( ~ i r o x @ ,
Sertanm) controls pine sawflies, but these pests are not in the
SPRINGTAILS
same family as T. cannabis (for more on NPV, see the section Springtails are small, simple, and surprisingly abundant
on budworms). Wang et al. (1987) killed 90-100% of young insects. They are seldom seen because of their tiny size and
T. cannabis larvae with a variety of synthetic pesticides. retiring habits in concealed soil habitats. Most springtails feed
Yepsen (1976) controlled other sawflies with rotenone and on decaying plant material, fungi, and bacteria. A few spe-
hellbore. Neem also works. cies, such as the garden springtail, Bourletiella hortensis
In the USA, wild populations of S . pilicornis are culled (Fitch), damage plants in greenhouses. Another species, con-
by a microsporidium, Nosenla pilicornis Gorske & Maddox. fusingly called a "flea," causes stippling of leaves, much like
S. pilicornis larvae are completely killed by malathion and spider mites.
carbaryl, and Bt causes 50% mortality (Gorske et al. 1976).
LUCERNE FLEA
Sminthl~rllsviridis Linnaeus 1758, Collembola; Sminthuridae.
Description: Adults are yellow-green, wingless, globular in
WIREWORMS shape, up to 3 mm long, with a large head in relation to their body.
Wireworms are the grubs of click beetles. In German,
Saatsschnellkafer. Grubs feed on roots and freshly-sown seeds. Life History
Wireworms attack many plants, especially grasses (corn, This European species has recently invaded Australia,
lawn, etc.) in poorly-drained soil. where it infests hemp (Ditchfield, pers. cornmun. 1997).Early-
sown crops suffered serious infestations, whereas later
LINED CLICK BEETLE sowings were unaffected. S. viridis is a serious pest of alfalfa
Agriotes lineatus Linnaeus 1758, Coleoptera; Elateridae. (lucerne), where up to 6000 pests may be found per square
Description: Adults are light brown with lined electra, cylin- foot (Hill 1983). The pest also infests clover ("clover
drical, 7.5-11 mm long. Grubs are hard, jointed, yellow brown, cov- springtail") and various cereals.
ered by minute hairs, with three pairs of legs behind the head, up to
25 mm long (Fig 4.49). CONTROL
Biocontrol can be achieved with Hypoaspis miles, a mite
Life History that preys on small soil insects (described under fungus
This European species also lives in north Africa and the gnats). Chemical controls have not been elucidated.
Americas. Gutberlet & Karus (1995) reported A.lineatus on
German hemp. The species lives its life cycle underground,
taking about four years. A related European species, Agriotes
obsclrrus (L.), has become established in British Columbia EARWIGS
(Howard et al. 1994). Wireworms can be confused with Beetle-like in appearance, these insects are distinguished
millipedes. by a pair of prominent forceps-like cerci at their tail end. Al-
though beneficial (they eat aphids and other pests), omnivo-
rous earwigs can become garden pests. Large males can in-
flict a painful pinch with their cerci. They are nocturnal.

EUROPEAN EARWIG
Forficula allricularia Linnaeus 1758, Dermaptera; Forficulidae.
Description: Adults are dark reddish-brown, 10-15 mm long,
and backed by the aforementioned cerci (Fig4.50). Females lay round,
pearly-white eggs in underground masses. Nymphs are pale brown,
and their wings and cerci are much reduced or absent.
Earwigs overwinter as adults in small underground
nests. Eggs are laid in late winter and females carefully guard
their brood until they hatch around May. Earwigs infest all
ages of Cannabis and all parts of Cannabis, including seed-
lings in California (Frank 1988), and flowering tops in India
(Bantra 1976).Earwigs have been found in the stem galleries
of European corn borers in Italy (Goidanich 1928). Earwigs
Figure 4.49: Agriotes lineatus. A. Adult click beetle; can fly, but not far. To travel distances they must take off
B. Larval wireworm (from Hill 1994). from a high place with a good tail wind. They are frequent
pests of urban gardens.
92 Hemp Diseases and Pests

CONTROL
Culpepper (1814) repelled earwigs with the juice of
crushed Cannabis seeds. Frank & Rosenthal (1978) trapped
earwigs by placing pieces of cardboard around the garden.
During the day, earwigs gather in cool, moist, dark areas-
find them under the cardboard. Frank (1988) eliminated
earwigs with a rotenone soil drench. Puritch (1982) reported
100% mortality of earwigs with insecticidal soap sprays.
Poisoned baits, the kind described for cutworms, work
against earwigs if poisoned with boric acid.

Figure 4.50: Female Forficula auricularia in winter

quarters with her eggs (from Fulton 1924).
 "Achlya species (water molds) exhibit a sexual ambivalence in which maleness and femaleness
are determined in each mating by common consent of the mated."
-J. R. Raper

Chapter 5: Fungal Diseases

At least 8000 species of fungi attack plants (Cook & Qualset The grey mould fungus tends to attack Cannabis in
1996). They cause more crop losses than the rest of Earth's two places-flowering tops and stalks. Flower infestations
organisms combined. How many species cause Cannabis tend to arise in drug cultivars and seed cultivars with large,
diseases? The scientific literature lists 420 Latin names of fungi moisture-retaining female buds. Scheifele (1998) reported
associated with Cannabis. Many names are taxonomic 30-40%incidence of "head blightwinfields of early-matur-
synonyms. The fungus causing grey mould, for instance, ing seed hemp; 'Fasamo' and 'FIN-314' also suffered spo-
masquerades under seven different Latin names (McPartland radic infections. Stalk rot seems more common in fibre va-
1995e). Other species cited in the literature are misidentifica- rieties (Patschke et al. 1998). The grey mould fungus can
tions (McPartland 1995a).Yet more names in the literature de- also infest seeds, destroy seedlings (see Damping off), and
scribe obligate saprophytes. Saprophytic "hemp retters and attack plants after harvest (see Chapter 8).
rotters" unable to attack living plants are excluded here. (See
Chapter 8 concerning post-harvest problems.) After a name- SYMPTOMS
by-name review, McPartland (1992) determined the 420 taxa Flower infestations begin within buds, so initial symp-
in the literature actually represented 88 species of Cannabis toms are not visible. Fan leaflets turn yellow and wilt, then
pathogens. . pistils begin to brown. In high humidity, whole inflores-
The fungal diseases described here are sequenced by their cences become enveloped in a grey fuzz, then degrade into
economic importance. Grey mould tops the list. We use grey-brown slime (Plate 45). The grey fuzz is a mass of
common names of diseases approved by the American microscopic conidia (Fig 5.1). In low humidity, the grey
Phytopathological Society (McPartland 1989, 1991). Latin fuzz does not emerge; infested flowers turn brown, wither,
names of all spore states (teleomorphs and anamorphs) are and die (Plate 46).
included for each species, with the most common spore state Stalk rot begins as a chlorotic discolouration of in-
listed first. We have included synonyms for the sake of fected tissues. Chlorotic sections turn into soft shredded
continuity and reference to earlier literature. cankers. Stalks may snap at cankers. Cankers may encir-
cle and girdle stalks, wilting everything above them. In
high humidity, cankers become covered by conidia (Plate
GREY MOULD 47). Conidia are liberated in a grey cloud by the slightest
Grey mould has become the most common disease of Can- breeze. Small, black sclerotia may form within stalks.
nabis (Plate 44). It afflicts fibre and drug cultivars, outdoors
and indoors (glasshouses and growrooms). The fungus caus- CAUSAL ORGANISM & TqXONOMY
ing grey mould attacks hundreds of other crops around the Botrytis cinerea Persoon: Fries, Systema mycologicum 3:396,
world. It is so pervasive in vineyards that some wines deliber- 1832.
ately use grey-mouldy grapes, such as Sauterne (in France), =Botrytis infestans (Hazslinszky) Saccardo 1887, =Polyactis
Trokenbeerenauslesen (in Germany), and Tokay (in Hungary). infestans Hazslinszky 1877; =Botytis felisiana Massalongo 1899;
The fungus thrives in high humidity and cool to moderate tem- =Botytis vulgaris Link: Fries 1824.
peratures. Disease peaks in drizzly, maritime climates (e.g., the teleomorph: Botyotinia fuckeliana (deBary)Whetzel, Mycolo-
Netherlands, the Pacific Northwest). In these climates grey gia 37:679 1945; =Sclerotiniaftlckeliana (deBary)Fuckel 1870.
mould reaches epidemic proportions and can destroy a Can- Research reveals these seven names refer to the same organ-
nabis crop in a week (Barloy & Pelhate 1962, Frank 1988). ism, susceptible to the same control methods. There is some dif-

Figure 5.1 : Conidiophores and conidia of Botrytis cinerea. A. Stereopair: view with a stereoscope or hold about 20 cm
away, cross your eyes and align double vision into a single middle 3-D image, SEM x1250 (courtesy Merton Brown and
Harold Brotzman, Brown & Brotzman, 1979); B. LM x260 (courtesy Bud Uecker).
93
94 Hemp Diseases and Pests

ference of opinion: Ferraris (1935) considered B. citlerea and B. vtrl- easily tells them apart, the conidia are very different-see
pnns d~fferentspecies. Gitman & Boytchenko (1934) and Barloy & Fig 3.2. Powdery mildew looks like a white dust, not a gre?
Pelhate (1962) separated B. cinevea from B. infestans and B. felisiana. mould, and powdery mildews do not cause cankers. Sev-
Spaar et '11. (1990) confused the teleomorph name. eral fungi cause cankers, such as anthracnose fungi and
Description: Conidiophores upright, grey-brown, branching Fusariunz species, but none grow as fast as grey mould.
near the apex, 5-22 pm m diameter. Conidia borne on conidiophore
Sclerotia formed by B , cirierea can be confused with scle-
apex in botryose clusters, hyaline to yellow-grey (grey en masse),
aseptate, round to ovoid, 8-14 x 6-10 pm (Fig 5.1). Microconidia rotia formed by Sclerotinia sclerotiorl~m,the cause of hemp
rare, Myr~oconiurn-like,arising from phialides, hyaline, oval, asep- canker (see next section). Generally, B. cirieren sclerotia arise
tate, 2.0-2.5 ym in diameter. Apothecia rare, arising from sclerotia within stalks, whereas S . sclerotiorllln sclerotia appear within
on a 3-10 mm stalk, topped by a yellow-brown disc; disc flat to stalks or upon the exterior surface. B. ciizerea sclerotia are
slightly convex, 1.5-5mm in diameter, capped by a single layer of smaller than S. sclerotior~lnzsclerotia, and they have different
nsci and paraphyses (Flg 5.2). ASCIcylindrical, w ~ t hlong tapered cross sections (Fig 5 . 3 ) . If sclerotia produce apothecia, the
stalks, eight-spored, 120-140 x 8 Llm. Paraphyses hyaline, septate, species are easily distinguished under a microscope.
hl~form.Ascospores aseptate, unlguttulate, uniseriate, hyaline, ovoid
to ellipsoid, 8.5-12 x 3.5-6 pm. Sclerotia hard, black, rough, plano-
convex, irregularly round to elongate, 1-15mm long (averaging 5mm DISEASE CYCLE & EPIDEMIOLOGY
on Cannabis, Flachs 1936), sometimes m chains; cross-section of B. cinerea can overwinter in infected seeds (Fig 5.28).
sclerotlum reveals a t h ~ nblack rind with a hyaline interior (Fig 5.4). Infected seeds may mould in storage or give rise to seed-
borne infections of seedlings the following sprlng
DIFFERENTIAL DIAGNOSIS (Pietkiewicz 1958, Noble & Richardson 1968). B. cllzerea also
In flowering tops, this disease may be confused with overwinters within stalk debris as sclerotia or dormant hy-
brown blight caused by Alternaria nlternata. A microscope phae (Fig 5.2A).Sclerotia may persist for years in soil. Spring

Figure 5.2: Disease cycle of gray mold; for description see text (McPartland redrawn from Agrios 1997, insert from Barloy
& Pelhate 1962).
 Chapter 5: Fungal Diseases 95

rains induce sclerotial germination, producing conidia (Fig plants) rarely suffer from bud rot, whereas the dense, tightly-
5.2C) or rarely producing apothecia with ascospores (Fig packed buds of Cannabis nfglmnica tend to hold moisture and
5.2D). Hyphae from sclerotia may also penetrate plants di- rot easily (Clarke 1987).Afghan biotypes evolved in very arid
rectly (Fig 5.2E). conditions and have no resistance to grey mould. This
Conidia (rarely ascospores) are blown or splashed onto unfavourable trait even appears in hybrids with a small
seedlings. High humidity or dew is needed for conidial percentage of afglzanicn heritage. What marijuana breeders
germination. The optimal temperature for conidial gained from the introduction of nfgharlica (potency, short
germination is around 20°C. Conidia germinate and directly stature, early maturity), they paid for with extreme
penetrate thin young epidermis. On older plants the fungus susceptibility to bud rot.
can only penetrate epidermis damaged by insects, rough Susceptibility to grey mould has no correlation with
handling, frost, or improper fertilization. After infection, the plant THC or CBD levels (De Meijer et al. 1992, Mediavilla et
fungus grows within the host, at an optimum of 24°C. al. 1997).Cannabinoids inhibit some fungi (Elsohly et al. 1982,
In humid conditions, the fungus forms new conidia, McPartland 1984), but apparently not B. cilzeren. Perhaps B .
which spread to sites of secondary infections (Fig 5.2F). ciizerea evolved the ability to metabolize THC, as have other
Conldlal production requires ultraviolet (UV) light, fungi (Robertson et al. 1975, Binder 1976).
specifically at wavelengths shorter than 345 nm (Sasaki & Mediavilla et al. (1997) reported that 87% of 'Swihtco'
Honda 1985). Microconidia may spread the disease (Fig plants [a drug cultivar with afglinnica heritage] suffered grey
5.2G),but many experts consider microconidia noninfective. mould-more than any fibre cultivars, including 'Felina 34'
Cycles of secondary infections eventually build to epidemics (75% diseased), 'Fedora 19' (63%),'Uniko-B' (55?/0),'Kompolti
in mature plants (Fig 5.2H). Hibrid TC' (50%), 'Futura 77' (47%), 'Secuiemi' (37%), and
Grey mould flourishes in high-density stands of hemp. the most resistant, 'Livoniae' (25%).Van der Werf et al. (1993)
De Meijer et nl. (1995)seeded four fields at rates of 20,40,80, said Hungarian 'Kompolti Hibrid TC' and French 'Fedrina
and 110 kg seed per ha. These rates initially produced 104, 74' are more susceptible to grey mould than other fibre
186, 381 and 823 seedlings per mz. Grey mould decimated cultivars. Dempsey (1975) reported good resistance in 'JES-
the high-density stands. By harvest, all fields yielded about 1' and 'JUS-7, but these cultivars may no longer be available
100 plants per m'. Crop yield (stalk biomass) was actually (see de Meijer 1995).
greatest in crops sown at the lowest seedling rate. Several corporations produce monoclonal antibody-
Fibre varieties become more susceptible after canopy based assays. These detection kits can identify B. cinere17 in
closure. Drug varieties become most susceptible during flow- plant tissues before symptoms appear.
ering. B. cinerea often colonizes senescent leaves and flow-
ers, and from these footholds it invades the rest of the plant. BIOCONTROL (see Chapter 10)
bJatch for grey mould epidemics during periods of high hu- Foliage, flowers, and stems can be sprayed with Gliocln-
midity (>60(Y0 RH) and cooling temperatures, especially at diunz rosennt and related Trichodernza species (described be-
d e ~ vpoint when plants cannot dry out. low). Bees have successfully delivered these fungi to flow-
ers of other crops, by dustingbees with spores as they emerge
CULTURAL & MECHANICAL CONTROL from hives (Sutton et al. 1997).Unfortunately, Cannabis is not
(numbers refer to methods listed in Chapter 9) an attractive nectar source for this unique bud biocontrol de-
The key to grey mould control is method 7c-keeping livery system.
relative humidity below 50%. Also use methods 1 Damping off caused by B. cinerea can be prevented by
(sanitation), 2a (deep ploughing), 2b&c (sterilizing or mixing Glioclacliz~inand Triclzoderlnn species into the soil
pasteurizing soil), 2d (flooding soil), 3 (weeding), 4 (described under damping off). Other soil biocontrols include
(harvesting early), 5 (genetic resistance-see below), 8 (avoid Strepton~ycesgriseouiridis (described under damping off) and
excess nitrogen and phosphorus, add calcium, neutralize Coniothyri~~ln nzinitans (described under hemp canker).
acid soils to enhance calcium absorption), 10 (careful Post-harvest disease by B. czneven has been controlled with
pruning), and 11 (avoid seedborne infection). yeasts (Picliiag~~illiernzotidii
and Cntzdida oleopiziln) and the bac-
In glasshouses, keep light intensity high and the tem- terium Psez~domonassyringae, described near the end of
perature warm (>25"C) to inhibit conidial germination. Chapter 8.
Outdoors, avoid planting in shade in humid climates. Do
not crowd plants-sow seed at low densities. Avoid Gliocladium roseum (=Clonostaclzys rosea)
\'\rounding plants during susceptible periods, except to prune BIOLOGY: This cosmopolitan fungus lives in a wide
away ~njuredor infected branches. Trunoff (1936) noted B. range of soils, from tropical rainforests to subartic deserts
czizcreiz attacks males first (males lose vigour before females), (Sutton et al. 1997).G. roseunl grows best between 20-28°C. It
then spreads to female plants. Rogue males if possible. has been isolated from hemp stems in the Czech Republic
Flowers should be harvested while resin glands are white (Ondrej 1991). The fungus protects plants in many ways: it
or amber, not brown. Brown, wilted pistils are prime B. ciilerea aggressively colonizes all surfaces of plants-above and
fodder. Harvested material should be cured and dried in dark below ground-which prevents pathogens from gaining a
rooms with good air circulation. foothold; it grows within senescent plant tissues and
Sasaki & ~ o n d (1985)
a slowed grey mould in glasshouse competes with pathogens for nutrients and space; it induces
tomatoes by covering windows with sheets of UV-absorbent resistance in host plants against other fungi; it oozes
vinyl. Since the grey mould fungus requires UV light to pro- metabolites that inhibit pathogens; and it directly parasitizes
duce conidia, epidemics are prevented. Sasaki & Honda other fungi, including B. cirierea, Sclerotiilin sclerotior~~nz,
(1985) filtered UV-B radiation with Hi-S vinyl by Nippon Rhizoctotzin solani, Phyinatotriclz~~m
onznivori~nl,and Vert~c~lliiinz
Carbide Industries; Lydon et al. (1987) used Mylar Type S species. It also parasitizes some nematodes (Heterodern and
film. Reducing UV-B also decreases production of THC Globodera species).
(Lydon et 171. 1987). APPLICATION: Spore suspensions (lob-108 conidia per
Breeding resistant plants is the ultimate solution for grey ml) mixed with a surfactant such as Triton X-100 have been
mould. Tall indica biotypes (Mexican, Colombian, and Thai sprayed on flowers to prevent bud rot, or sprayed on conifer
96 Hemp Diseases and Pests

stalks at canopy closure to prevent stalk rot, or daubed on America, the disease hampered production of hemp in Wis-
clone-pruning wounds of mother plants to prevent wound consin (Buchholtz, USDA archives 1943),in Ontario (Conners
infections. Protect seedlings by dressing seeds with spores 1961, Scheifele 1998), and caused 40% losses in Nova Scotia
rnlxed ~ 1 1 tah sticking agent, or by quick-dipping clones in (Hockey 1927). Hemp canker has infested Indian ganja
spore slurries, or by drenching seedbed soil with spore (Bilgrami et al. 1981), Finnish seed-oil crops (Callaway &
suspensions. Laakkonen 1996),and Australian hemp (Synnott 1941, Lisson
Treatments should be applied shortly before nightfall, & Mendham 1995).
because dew and darkness facilitate spore survival and
activity (Sutton et al. 1997). A few hours of dampness are SYMPTOMS
needed for germination, which can be scary to growers Symptoms usually begin in late summer, on full-,arown
familiar with grey mould. Heavy rainfall is not good, because plants. The disease may arise near the soil line (Holliday
it washes spores off plants before spores can germinate. Some 1980) or in the upper 2/3rds of the plant (Tichomirov 1868).
researchers report a single G. rosellm treatment can last eight Watersoaked lesions appear first, on stalks and branches.
to 12 weeks or more. Others reapply G. roseuin weekly during Cortical tissues beneath the lesions collapse, creating pale,
peak grey mould seasons (Sutton et al. 1997). light-brown cankers. In humid conditions, the surface of the
NOTES: Spores stored in sealed containers at 3-5'C stalk becomes enveloped in white mycelium (Plate 48). Black
remain effective for over a year. G. roseuin is compatible with spots of sclerotic tissue emerge on the surface of cankers,
Tricl~ociernzal z a r z i a n l ~ ~T.
~ zviride,
, T,koningii, and mycorrhizal usually by September. Larger sclerotia form within the hol-
fungi. Its compatibility with other biocontrol fungi has not low of stalks. Flowering, if it has begun, often ceases (Flachs
been evaluated. It can probably be mixed with BT, NPV, and 1936). Plants remain in this condition, or wilt and fall over.
biocontrol insects, but these combinations remain untested. A secondary fungus may overgrow hemp canker, see "red
Unlike fiicl~odernza species, G. roseuin can reproduce on the boot disease."
surface of plants, and the fungus bears conidia within 72
hours of initial treatment (Suttonet al. 1997). G. rosezrnl is not CAUSAL ORGANISM & TAXONOMY
harmful to bees a n d does not grow at h u m a n body Sclerotinia sclerotiort~m(Libert) deBary, Vergh. Morpil. Biol.
temperatures. Because of its unique morphology, ecology, Piize, Mycet., Bact. p.236, 1884.
and DNA, G. rosewrz will be moved to the genus Clotzostacl~ys (Libert)Korf & Dumont 1972,-Pezlza
- =Witetzeiinia sclevotrorr~n~
and renamed C . rosea (Samuels 1996). scievotiovunl Libert 1837;=Scleuotinin iibevtiann Fuckel1870; =Sclerotinin
kauffilnniliana (Tichomirot,)Saccardo 1889, =Pexzn kaz~ffnla?~ninirn
Tichomirov 1868 (see McPartland 1995e).
Trichodenna species No anamorph has been named for S.sclevotiorurri, but
BIOLOGY: Four cosinopolitan fungi, Trichodeuma har- Brandenburger (1985)described one on Cnn?~ilbls, albeit namelessly.
zianun~,Triclzoderma polysporl~m,Trichoderma viuens, and Ferraris (1915) erroneously called Botnjtis o~ilgarrsthe anamorph of
Trichodernzn viride, control B . cinerea and other pathogens. S . sclerotiorum. His mistake followed Kirchiler (1906), ~,iyl~ucalled B.
These species are discussed further under Rhizoctonia sore fuckeliana the hemp canker orgaillsm (B. fiiuckelinnn causes grey
shin disease. mould).
APPLICATION: These fungi are marketed in a variety of
biocontrol products. Most products are designed for seed and
soil treatments. When applied as afoliar spray, the T-39 strain
(Trichodexa) controlled B. cineyea in grape
of T , llnrziarz~~i11
vineyards and glasshouse vegetables, using a spray
concentration of lo8 conidia ml-' (Reuveni 1995). Foliar
application inay need repeating, because Trichoderma species
do not survive long on leaf surfaces (Samuels 1996).Pruning
wounds can be painted with a spore suspension of the T-39
strain. Foliar application of the T.harziaizl~in-T,p o l ~ j s p o r ~ ~ m
product (Binab TQ)does n o t control B. cinerea in flowering
tops (Watson, unpublished data 1991).

CHEMICAL CONTROL (see Chapter 11)

Bordeaux mixture knocks back grey mould in early
stages, and pre\.ents infections until the next rainfall washes
the mixture off plants. Doctor Indoors (1988) painted prun-
ing ~voundswith alcohol or sodium hypochlorite (bleach).
Mixing calcium cyanamid into the soil kills overwintering
sclerotia. B , cinerea develops resistance to most synthetic fun-
gicide sprays. Fungicides may still be useful as seed treat-
ments (Patschke et nl. 1997). Seed dressings have a low im-
pact on the environment (see Chapters 11 and appendix).

HEMP CANKER Figure 5.3: Sclerotinia sclerot~orum.A. Sclerotia and

This disease goes by many names around the world, mycelium in pith (xl); B. Sclerotium with apothecia (XI);
including cottony soft rot, watery soft rot, stem rot, white C. Cross section of apothecium ( X I50); D. Asci,
mould, and grey rot. Europeans call it hemp canker. Some paraphyses, and ascospores ( ~ 3 0 0 )E.
; Close-up of three
consider it the #1 scourge of hemp cultivation (Rataj 1957), ascospores (~430).All from Tichomirov (1868) except B
or #2 behind grey mould (Termorshuizen 1991). In North by DeBary (1887).
 Chapter 5: Fungal D i s e a s e s 97

Description: Sclerotia within stalks are hard, yellow becom- midity), 8 (add calcium to soil, neutralize acid soils to enhance
111gblack, smooth, oblong, 5-13 x 3-7 mm, cross section of sclerotia calcium absorption, Ferraris (1935) suggested adding
re\ eals a dark rind surrounding a soft, white centre (Fig5.4):Apoth- phosphate), 10 (careful pruning), and 11 (avoid seedborne
ecla conslst of brown'cylindrical stalks up to 25 mm long, topped infection). Deep ploughing of debris is particularly effec-
1~1th by yellow-brown cups, concave, 2-8 mm wide, filled with a tive-S. sclerotiorunz sclerotia d o not germinate if buried
slngle layer of asci and parapheses (Figs 3.3 & 5.3).Asci cylindrical,
e~gl-it-spored, bluing with Melzer's reagent, 110-140 x 6-10 pm. Para- deeper than 6 cm underground (Lucas 1975). Rotation with
pheses flllform, liyaline, same length as asci. Ascospores aseptate, nonhost crops must last three or four years to stanTelong-
elliptical, Ilyaline, usually biguttalate, 9-13 x 4-6 pm (Fig 5.3). lived sclerotia. Avoid high-density stands; seed at low rates
(see discussion in grey mould section). Do not use seeds from
infected plants (Serzane 1962).
Soil suspected of harbouring pathogens can be tested
with biotechnology-based assays. A kit detecting S.
sclerotior~rnzis commercially available (though expensive,
US$148 for a set of six tests). Handling plants infected with
S. sclerotiorlrm may cause dermatitis. The fungus produces
two compounds, xanthotoxin and bergapten, which sensitize
skin to sunlight (Centers for Disease Control 1985).

BIOCONTROL (see Chapter 10)

Infected glasshouse soil can be dried for several days,
and then soaked for two or three weeks. This encourages the
Figure 5.4: A. Thin section through a sclerotium of Botrytis growth of soil organisms whith parasitize and kill S. sclero-
cinerea ( ~ 3 1 0 )B.
; Section through a sclerotium of t i o r ~ r n isclerotia. A parasitic fungus, S p o r i d e s ~ n i u m
Sclerotinia sclerotiorum (x300), from DeBary (1887). sclerotivorum, attacks sclerotia of S,sclerotiorliilz and is be-
ing investigated (Cook et al. 1996). Commercially-available
biocontrols of S. sclerotiorurn include Trzchoder~nahar-~ l a ~ l l ~ i n
DIFFERENTIAL DIAGNOSIS (see the section on Rhizoctonia sore shin), Bacillus sirbtilis (de-
Symptoms of hemp canker can be confused with stalk scribed under damping off), and Colziothyriunz mi~litaiis.
rot caused by the grey mould fungus, B. cinerea. In humid
conditions, B. cinerea forms its characteristic blanket of grey Coniothyri~imnzinitans
conidia. In dry conditions, you can differentiate sclerotia. BIOLOGY: A fungus (ContansWGa, KoniB) that
Sclerotia on stalk exteriors are usually caused by S. sclerotio- parasitizes sclerotia of S. sclerotior~rnz,B. ci~zerea,and a few
ruiil. 5.s c l e r o t i o r ~ ~sclerotia
~r~ are larger than B. cinerea sclero- other pathogens. C. rr~initnnslives in temperate zones worldwide.
tia. The rind covering S. sclerotiorl~~i~ sclerotia consists of three APPLICATION: C. nlinitans is formulated as water dis-
or four layers of cells, and the cells have thick black walls. bursable granules, applied as a soil drench to sclerotia-
The r ~ n dof B,ci~lerenis thinner, usually one or two cells thick infested soil. It has also been sprayed directly on susceptible
(Fig 5.4). The internal medulla of both species is white, but plants. Application rates range from 50-3000 kg ha-1 (Whipps
the medulla of B. cirlerea is denser and more gelatinous. & Gerlagh 1992).C. ~ninita~zs has protected canola, sunflower,
Hemp canker can also be confused with southern blight peanut, soyabean, and vegetable crops.
or Rhizoctonia sore shin. Both of these diseases, however,
tend to cause symptoms adjacent to soil. The fungi causing CHEMICAL CONTROL (see Chapter 11)
these diseases form tiny sclerotia, smaller than S. sclerotior~llrl Preventative methods are more effective than curative
sclerotia. Hemp canker is common in temperate climates; treatments. The only effective chemicals are synthetic; they
southern blight predominates in warmer regions. are best applied as seed dressings (see Appendix 1 and
Patschke et al. 1997).
DISEASE CYCLE & EPIDEMIOLOGY
S . sclerotioruin overwinters as sclerotia in plant debris
or in soil. Springtime moisture and warm temperatures (15-
20°C) initiate germination. Each sclerotium sprouts two to DAMPING OFF
selren apothecia. Apothecia forcibly eject asci into the air. At least two "water moulds," Pythiunz species, cause
Ascospores germinate in the presence of moisture and di- damping off disease of Cannabis seedlings. Pytilillni species
rectly penetrate host epidermis, or penetrate via wounds. occasionally attack mature plants in field soils (Frezzi 1956)
The fungus grows inter- and intracellularly through host pa- and hydroponic systems (McEno 1990). Curiously, no
renchyrna and cortex. S,sclerotiorunz invades the seeds of Plzytoplztlzora species (closely related to Pytlliurn) parasitize
many plants, but this has not been documented in Cannabis. Cannabis. Hemp in fact serves as a biocontrol against
S. sclerotioruliz attacks over 360 species of crops and Plzytophthora infestatzs (Israel 1981);aqueous extracts of hemp
xveeds, mostly herbaceous dicots. Strains isolated from inhibit the growth of I? iilfestails (Krebs & Jaggi 1999).
sunflowel; Jerusalem artichoke, potato, safflower, flax, and
colza also attack hemp (Antonokolskaya 1932). S. sclerotio- SYMPTOMS
run1 thrives in cool, moist conditions, just like grey mould. Damping off presents itself in two scenarios: In pre-
Both diseases accelerate after canopy closure, especially in emergent damping off, seeds or seedlings die bqfore they
high-density stands (De Meijer et nl. 1995). emerge from the soil (Plate 49). Post-emergent damping off
hits after seedlings have emerged from the soil (Plate 50).
CULTURAL & MECHANICAL CONTROL (see Chapter 9) Seedlings with post-emergent damping off usually develop
Use methods 1 (especially burying), 2a (deep a brown rot at the soil line, then wilt and topple over. In older
ploughing), 2b&c (sterilizing or pasteurizing soil), 3 seedlings (with up to eight pairs of true leaves) growth ceases,
(xveeding), 5 (genetic resistance), 6 (rotate with grains and leaves turn pale yellow, then seedlings wilt and topple over
grasses and keep free of dicot weeds), 7c (avoid excess hu- (Kirchner 1906).
98 Hemp Diseases and Pests

Figure 5.5: Ten organisms that cause damping off in Cannabis seedlings. Not drawn to scale (McPartland).

CAUSAL ORGANISMS & TAXONOMY germinate as germ tubes or (rarely) they produce an elongate emls-
1. Pytlliuln n y k n l z i ~ e r t n a t t l(Edson)
m Fitzpatrick, M?lco/ogin sion tube, 5-135 pm long, with a vesicle at its end. Vesicles clea\'e
15:168, 1923. into eight to 15 zoospores (Fig 5.5).
Description: S p o r a ~ i g ~mature
a illto elongated irregular
s~\~ellings,branched or unbranched, 50-1000 x 2-20 pm, sometimes 3. Other causal orgarzisms
scarcely d~stinguishedfrom the mycelium. Sporangia germinate into Europeans cite a third Pytliiiinz species attacking hemp,
em~ssiontubes of variable lengths, 2-5 pm diameter, and form vesi- Pythiutn debaryanum Hesse (Kirchner 1906, Serzane 1962,
cles at their ends. Vesicles cleave into 100 or more zoospores.
Kirchner 1966, Vakhrusheva 1979, Barna e f 01.1982, Gutberlet
Zoospores reniform, laterally biflagellate, 7.5 x 12 pm. Oogonia
spher~cal,formed terminally (rarely intercalary), 22-27 ym diam- & Karus 1995, Bbsca & Karus 1997). P.debaryntzllriz is not a
eter. Each oogonium is fertilized by one or two antheridia. Antheridia true species; its original description was based on a mixture
are tormed on hyphae adjacent to oogonia, terminally or intercalary, of fungi (McPartland 1995a). "P.debauyan~ini"citations usu-
doliiform to broadly clalrate, 9-11 x 10-15 pm. Each fertilized ally indicate misidentfications of l? iiltinzilnz (Plaats-Nitennk 1981).
oogonium forms a single oospore. Oospores spherical, containing a Besides Pyfhiiim, several fungi also cause damping off-
central \.acuole, wall 2 l m thick, smooth, and d o not fill oogonia, Bot~ytiscinerea, Mncrop!wtnina pkaseolina, Rhizocfonia solani,
17-19 pm d~ameter(Fig 5.5). and several Fiisariiiin species (Fig 5.5). These pathogens
usually attack mature plants. Their taxonomic descriptions
2 . Pytlzizrin trltimtlm Trow, Annals of Botany 15:300,1901. are presented elsewhere.
Description: Sporangia spherical to doliiform, 20-29 x 14-28
p m , and germinate only by germ tubes. Oogonia smooth, spherical,
DISEASE CYCLE & EPIDEMIOLOGY
formed term~nall!; 19-23 ym in diameter. Each oogonium is ferti-
llzed by one antheridium (rarely two or three antheridia).Antheridia Pyflzi~itnspecies produce two types of spores, zoospores
swollen, sausage-shaped, curved, originating immediately below and oospores. Zoospores swim using flagella, and spread
oogoiua. Each fert~lizedoogonlum forms a single oospore. Oospores disease fromplant to plant Oospores are sexually produced.
are spherical, containing a single central vacuole and eccentric re- They do not migrate, but spread the disease over time (from
frillgent body, thick walled, smooth, 14-18 pm diameter. Oospores season to season) as overwintering spores.
 Chapter 5: Fungal Diseases 99

P liltlnlllnz attacks hemp seedlings in central Europe

(Marquart 1919, Schultz 1939) P u l t ~ n z l ~ nalso z causes a root
lot of inat~ireplants (Frezzi 1956) Root rot begins at the root-
tip, e~~entually causing above-ground parts to wilt. P. ~ ~ l t i n z ~ ~ n z
lives in temperate regions worldwide. This pathogen attacks
many crops, especially at cooler temperatures (12-20°C).
P, aplza~zidernlatllnlis the "warm weather pythium," with
optimum growth at 32°C.Although the specie; arises world-
wide, it has only been reported on C a n n a b i s in India
(Galloway 1937).
B o t y t i s ciizerea causes damping off because it can spread
by seedborne infection (Pietkiewicz 1958, Noble &
Richardson 1968, Patschke et nl. 1997). Expect epidemics in
seeds harvested from females infested by grey mould. B.
iiitrrea conidia disperse by wind (not water like most
damping off fungi).
Macropko~riinapliaseolilla causes charcoal rot in older
plants, as well as damping off in seedlings. It is common in
the midwest, especially on maize. Epidemics caused by M.
pl~aseoliiiapeak in warm weather (optimal temperature 37°C).
Rlzizoctonia solani causes sore shin and root rot in mature
plants. It tends to damage seedlings at a later stage than
P!/tliilriiz species. Mishra (1987)reported R. solani causing 78%
mortality in Cannabis seedlings (44% as post-emergence, 34%
as preemergence). Dippenaar et 121. (1996) reported heavy
seedling losses by R , solani despite spraying with a fungi- Figure 5.6: Signs and symptoms of post-emergent
cide (Rhizolexa). R . solnni does not require excess moisture, damping off caused by two Fusarium species.
inl like most darnping off organisms. A. F: solani; B. F: oxysporum (from Barloy & Pelhate
Several Ftlsarililiz species cause damping off, including 1962).
E solaizl, E oxyspor~cm(Patschke et al. 1997),and less frequently
E sldpiz~~relini, F,at1eizace1117?,and E ~ r a n ~ i i z e a r ~Some
~ n z . of these many different temperatures, pH ranges, lighting conditions,
fungi also cause cotyledon drop (Rataj 1957).In mature plants and soil types. Their coinmon denominator is excess
these pathogens cause Fusarium foot rot, root rot, stem moisture, so avoid overwatering-follow methods 7b and
cankei', and Wilt. F L L S ~ Y conidia ~ L I ~ I spread via splashed rain 7c. Irrigate soil before planting; watering afterward will pack
and water runoff. They generally prefer warm climates. If soil around seeds. Damping off increases in seeds planted
introduced into sterilized soil, F ~ i s a r i u msoecies run wild. deeper than 2 cm. Some horticulturists germinate seeds
They cause greatest losses in damp soil, but cannot survive between layers of lightly moistened paper towels before
in zunterlogged land.
planting them in soil. Right out of the plastic wrap, paper
towels are nearly sterile.
DIFFERENTIAL DIAGNOSIS
Do not plant in heavy, wet, poorly draining soils that
Differentiating between Pythillnz species is difficult, and
tend to puddle. On questionable sites, try planting in raised
requires laboratory cultivation. Differentiating Pythiunz spe-
beds (>I0 cm l~igh)or lighten soil with perlite. (Vermiculite
cies from other organisms can be easier: Dig up a stricken
is a poor second choice; it eventually decomposes into sludge,
seedling and lightly grasp the root between your thumb and
which exacerbates soil heaviness.) Some authors suggest
forefinger. Then pull the root away from the stem. If the outer
using a sandy soil to suppress damping off. Others say a
layer of the root (epidermis and cortex) slips away leaving
perlite and peat mix is best. The key characteristic is good
only a thin inner cylinder (endodermis and stele), you have
drainage.
a P!/tllil~nzproblem. If tiny bits of soil dangle from the root of
Observe method 4-do not plant too early in the spring.
a seedling pulled froin the ground, you have a R. solalzi prob-
Most damping off pathogens thrive in cold soil. Vetter (1985)
le~n-this diagnostic sign is attributed to the pathogen's
coarse, clinging hyphae. Barloy & Pelhate (1962) differenti- described a rule of "thumb" for Calllzabis: "I walked into the
ated symptoms caused by F. solani and F. o x y s p o r ~ ~ m - middle of my unplanted garden plot, dropped my pants and
seedlings with F, solnili develop red discoloured roots (Fig sat ... if the soil is too cold for your bare ass, it's too cold for
5.6 A ) , whereas seedlings with F. o x y s p o r u n z become the seeds you're getting ready to plant." Indoors, use an
enveloped in a pink mass of hyphae (Fig 5.6 B). insulated horticulture table to provide "bottom heat" for
Pre-eii~erge~it damping off can be erroneously attributed seedlings and clone cuttings. Unfortunately, heat may
to outdated, dead seed. A lack of emerging seedlings may encourage M. phaseolina and P. a p h a n i i i e r m a t u n ~ .Full-
also be due to underground insects, such as crickets and root spectrum lighting greater than 1000 Lumens inhibits
maggots, which kill germinating seeds. Post-enzergei~t damping off. Etiolation predisposes plants to damping off,
symptoms can be confused with damage from insects and is prevented by high light intensity (Smith et a / . 1981).
(cutworms) or environmental causes (frost, hail, or heavy Follow method 11-do not plant seeds harvested from
rains). Leaf spots are not damping off. For instance, Tri- diseased females-B. cinerea and many Flisariliin species can
c l ~ o t l ~ e c i ~roseliin
~ i ~ z and Septoria cannabis attack cotyledons. spread by seedborne infection. Robust seedling growth in-
Slnce stems remain untouched and seedlings stay upright, hibits damping off, so follow method 8. But excess soil elec-
this is not damping off. trolytes of any type (N, P, K, Ca, Fe, Etc.) increase damping
off-so avoid chemical fertilizers and concentrated organics
CULTURAL & MECHANICAL CONTROL (see Chapter 9) (e.g.,blood meal, chicken manure). Aim for a pH of 7.0-7.5.
The many patl~ogensthat cause damping off prefer Hydroponic operators should take note of method l c
 Chapter 5: Fungal Diseases 101

YELLOW LEAF SPOT and dried lesions rarely flake away. Olive spot produces more
symptoms on the undersides of leaves. Fungi causing these
This disease is ubiquitous. It frequents fibre crops, but
diseases look quite different under the microscope.
also arises on drug plants (Mushtaque et al. 1973, Ghani &
S. cannabis-and S. neocannabina are distinguished by sev-
Basit 1975,Ghani et al. 1978)and Cannabis rtlderalis (Szembel
eral characteristics (McPartland 1995d). Spots caused by S.
1927, Gamalitskaia 1964). Since leaf photosynthesis is the
neocannabina have a dark reddish-brown border (Peck 1884,
engine driving crop yields, yellow leaf spot can reduce yields
Kirchner 1906, Kirchner 1966). Microscopically, S. cannabis
of fibre, flowers, and seeds.
pycnidia are larger and have thicker walls than S.
neocannabina pycnidia
-. (Fig
- 5.7). Ostioles in S. cannabis
SYMPTOMS
pycnidia are relatively small, whereas S. neocannabina ostioles
Small lesions first appear on lower leaves in early June.
may open to nearly half the diameter of pycnidia. Conidia
Lesion colour is variously described as white, yellow, ochre,
of S. cannabis are larger and contain more septa than those of
or grey-brown (Plate 51). Spots may remain small and round
S. neocannabina.
but usually enlarge to irregularly polygonal shapes, their
edges partially delineated by leaf veins. Spots sometimes
DISEASE CYCLE & EPIDEMIOLOGY
have reddish-brown perimeters (Peck 1884, Kirchner 1906).
Septoria species overwinter as pycnidia in crop residue
Tiny pycnidia arise within leaf spots. These fruiting
near the soil surface. Seedlings and young plants become
bodies form on the upper surfaces of leaves, not on
infected in early spring. Conidia of S. cannabis germinate rap-
undersides as reported by Flachs (1936). Eventually, leaf
idly and infect plants by penetrating stomates (Ferri 1959b).
spots dry out and fragment, leaving ragged holes in leaves.
Optimal growth of S , cannabis occurs at 25'C. The incuba-
In severe infections the leaves curl, wither, and fall off
tion period-between inoculation and first symptoms-is six
prematurely, defoliating the lower part of the plant (Ferraris
1935, Watanabe & Takesawa 1936, McCurry & Hicks 1925, or seven days (Watanabe & Takesawa 1936).Pycnidia on in-
Barloy & Pelhate 1962, Ghani & Basit 1975). fected plants spew copious amounts of conidia (Fig 5.8).
Conidia spread by splashed rain and give rise to epidemics
Kirchner & Boltshauser (1898) illustrated yellow leaf
in the summer.
spot in a classic lithograph. Many subsequent illustrations
are copied from this engraving (see Flachs 1936, Ceapoiu
1958, Barloy & Pelhate 1962). Although yellow leaf spot
principally affects lower leaves, Bush Doctor (unpublished
data 1986) saw it on fan leaves of flowering tops in Nepal
(Plate 52). Spots may also arise on stems (Gitman &
Boytchencko 1934, Ferri 1959b, Ondrej 1991), and seedling
cotyledons (Ferri 1959b).

CAUSAL ORGANISMS & TAXONOMY

At least two species of Septoria cause yellow leaf spot.
Rataj (1957)described a third species on Cannabis-Septoria
graminum Desmarieres. This citation is probably an error
(McPartland 1995a).

1. Septoria cannabis (Lasch) Saccardo, Sylloge Fungorum 3:557,

1884. ~ A s c o c h y t acannabis Lasch 1846; =Spilosphaeria cannabis
Rabenhorst 1857; =Septoria cannabina Westendorp 1857; =Septoria
cannabis Saccardo, nomen nudum.
Description: Pycnidia epiphyllous,gregarious, immersed but
eventually erumpent, globose to flask shaped, averaging 90 pm in
diameter; peridium dark brown, thick-walled textura angularis-globu- Figure 5.7: Two Septoria species. A. S. cannabis sec-
losa; ostiole round, 15pm in diameter (see Figs 3.3 & 5.7).Conidiog- tioned pycnidium and conidium; B. S. neocannabina
enous cells subglobose to ampulliform, simple, hyaline, holoblas- sectioned pycnidium and conidia (pycnidia LM x500,
tic. Conidia hyaline, filiform, pointed at both ends, straight or conidia LM x980, McPartland).
curved, three-septate, 45-55 x 2.0-2.5 pm. Pycnidia in vitro grow to
465 pm in diameter and form long necks (Watanabe & Takesawa
1936, Ferri 195913).Conidia exude from ostioles in a slimy, ribbon- Barloy & Pelhate (1962) suggested S , cannabis spreads
like cirrhus. via seedborne infection, but did not confirm this experimen-
tally. Other studies of seedborne fungi (Pietkiewicz 1958, Ferri
2. Septoria neocannabina McPartland, Sydowia 47:46,1995. 1961b, Babu et al. 1977)have not implicated Septoria species.
Septoria cannabina Peck 1884 (non Septoria cannabina Westen- Ferraris & Massa (1912) cited Leptosphaeria cannabina as a
dorp 1857);=Septoria cannabis var. microspora Briosi & Cavara 1888. "probable" teleomorph of S. cannabis. Evidence suggests they
Description: Pycnidia epiphyllous, gregarious, immersed, glo- erred (see the section concerning L. cannabina).
bose, eventually erumpent and nearly cupulate, 66 pm in diameter;
peridium honey-brown near the ostiole to almost colourless at the
base, thin-walled textura angularis-globulosa;ostiole irregular, 20 pm CULTURAL & MECHANICAL CONTROL (see Chapter 9)
in diameter (Fig 5.7). Conidiogenous cells short ampulliform to Methods 1 (sanitation) and 10 (careful pruning) are the
lageniform, simple, hyaline, holoblastic, up to 8 ym long. Conidia cornerstone of control. Also use 2a (deep ploughing), 5
hyaline, filiform, pointed at apex with a truncate base, usually (genetic resistance-see below), 6 (crop rotation), 7c
curved, one to three septate, 20-30 x 1.0-2.0 ym. (especially avoid overhead irrigation), 8 (avoid excess
nitrogen, add phosphorus and potassium), and 11 (unproven,
DIFFERENTIAL DIAGNOSIS but hedge your bets).
Brown leaf spot causes smaller, darker spots than yel- Monitor for disease during rainy seasons. Stay out of
low leaf spot. White leaf spot is paler than yellow leaf spot, Cannabis fields when plants are wet, since conidia spread
102 Hemp Dise'ases and Pests

Description: Sclerotia deep brown to black, smooth, somewhat

flattened and irregular in shape, no differentiation between rind
and medulla, up to 6 mm in diameter. Hyphae wide (5-12 pm),
multinucleate, with dolipore septa, no clamp connections, at first
colourless but rapidly becoming brown; branches form
geometrically at 45' or 90" angles from parent hyphae. Hyphae
become slightly constricted at branching points; a septum always
forms near the base of the branch (Fig 5.5). Basidiocarps arise on
stem surface, thin, effuse, discontinuous.Basidia are barrel-shaped,
10-25 x 6-19 pm, borne on imperfectly-symmetricalracemes, with
four sterigmata per basidium. Sterigmata are 6-36 pm long and bear
spores. Basidiospores hyaline, ellipsoid-oblong, flattened on one
side, 5-14 x 4-8 pm (Fig 5.5).

2. Binucleate R h i z o c t o n i a species
Description: These fungi are morphologically similar to R.
solani, but have thinner hyphae (4-7 pm) and only two nuclei per
hyphal cell. Binucleate Rhizoctonia species, unlike R. solani, often
produce Ceratobasidium teleomorphs. See Parmeter (1970) for an
introductory discussion.
McPartland & Cubeta (1997) isolated a binucleate
Figure 5.8: Pycnidium of Septoria neocannabina erupting Rhizoctonia species from the roots of a drug cultivar ('Skunk
from the leaf surface, spewing masses of conidia (SEM No. 1') growing in Holland. The binucleate condition was
x500, McPartland). determined by staining hyphae with Safranin 0 and DAPI.
Amplification of hyphal rDNA with PCR, followed by di-
gestion with four different restriction endonucleases, dem-
easily by contact. Ferri (195913) described resistant hemp va- onstrated that the restriction phenotype of the Cannabis iso-
rieties in Italy. Conversely, B6csa (1958) said "in-bred" late was identical to Ogoshi's anastomosis group G strain.
monoecious cultivars lose resistance to Septoria infection.
DIFFERENTIAL DIAGNOSIS
BIOCONTROL & CHEMICAL CONTROL Symptoms of Rhizoctonia sore shin disease resemble
No biocontrol against yellow leaf spot fungi is available. symptoms of hemp canker and southern blight. The fungi
For spray-gun enthusiasts, the disease can be slowed with causing hemp canker and southern blight, however, produce
Bordeaux mixture (Ferri 195910). more prominent external hyphae and larger sclerotia than
Rhizoctonia species. If examined under a microscope, the
unique hyphae of Rhizoctonia species are easily separated
from other fungi (Fig 5.5).
RHIZOCTONIA SORE SHIN
& ROOT ROT DISEASE CYCLE & EPIDEMIOLOGY
R. solani overwinters as sclerotia in soil. Sclerotia ger-
This disease has been described on fibre and drug plants
minate in early spring and produce hyphae. Hyphae enter
in Europe and India. Disease severity ranges from mild to
roots near the soil line, either by direct penetration or through
severe. The two fungi causing this disease are found
wounds. After penetration, R. solani oozes cellulose-degrad-
worldwide, and they parasitize approximately 250 plant
ing enzymes (which disrupt xylem and cause wilting), and
species. They also attack fellow fungi, animals, and humans.
pectolytic enzymes (which cause cortical rot). The fungus
They can live saprophytically in the soil.
may also kill seedlings (see damping off disease).
A virulent strain of R . solani destroyed 80% of Cannabis
SYMPTOMS
in an Indian epidemic (Pandotra & Sastry 1967).This strain
Symptoms begin as discoloured roots, usually undetec-
produced neither sclerotia nor basidia-just hyphae. Most
ted (Plate 53). Then leaf chlorosis arises. followed bv wilt-
strains in Europe produce sclerotia (Ray110 1927). A
ing. A dark brown discolouration moves up from th; roots
Ukrainian strain formed basidiospores (Trunoff 1936).These
to the base of the stalk. Several cm of stalk rot away, leaving
strains probably represent different anastomosis groups. R.
a shredded "sore shin" appearance. Within six to eight weeks
solani encompasses at least 11anastomosis groups, each with
young plants (<three months old) topple over and die. Older
unique host ranges and disease cycles. Anastomosis group
plants may survive. Small black sclerotia sometimes appear
AG-4 usually causes sore shin in tobacco. But recently,
in the shredded area. Occasionallya pale mat of basidiospores
anastomosis group AG-3 has become a problem, causing leaf
forms around the base of the stilk.-
lesions instead of sore shin, because AG-3 readily forms
basidiospores which splash onto lower leaves.
CAUSAL ORGANISMS & TAXONOMY
Disease symptoms increase under cool, damp condi-
1. R h i z o c t o n i a solani Kiihn, Krankh. Kulturgew., p. 224,1858.
tions, or paradoxically, when temperatures are elevated.
=Rhizoctonia napaeae Westendorp & Wallays 1846.
teleomorph:Thanatephorus cucumeris (Frank)Donk, Reinwardtia 3:376, Beach (in Lucas 1975) simply explained that sore shin wors-
1956; aHypochnus cuctlrneris Frank 1883; =Corticium solani (Prillieux ens at temperatures and conditions not optimal for the host.
& Delacroix) Constantineanu & Dufour 1895; =Hypochnus solani
Prillieux & Delacroix 1891; =Corticium vagnum Matuo 1949; CULTURAL & MECHANICAL CONTROL (see Chapter 9)
=Cortic~umvagum Berkeley & Curtis var. solani Burt apud Rolfs; Methods 2b&c (sterilize and pasteurize the soil) are very
=Pelliculaviafilamentosa (Patouillard)D.P. Rogers 1943. useful. Also apply methods 1 (sanitation), 4 (delay planting
Teleomorph taxonomy is unstable, see Parmeter (1970). until soil warms up), 7b (avoid overwatering), and 8 (avoid
Rhizoctonia napaeae is a "probable synonym" according to Parmeter
excess nitrogen, add calcium). Acidic soils increase disease.
(1970).Ajrekar & Shaw (1915)cited an orthographic variant, "Rhizoc-
tonia napi," on Cannabis in India. Avoid planting in previously infested areas-sclerotia re-
 Chapter 5: Fungal Diseases 103

main viable in soil for up to six years. Many weeds serve as cide protects seeds as they sprout, then T. harzianltm protects
alternate hosts. R. solani viability decreases in soils amended the developing root system. On the down side, some Trichode-
with high-carbon mulches, such as straw, corn stover, and rma species produce trichothecenes and other highly toxic
even pine shavings (Lucas 1975, Windels 1997).Residues of metabolites. Heat-tolerant strains may cause infections in
cabbage and other brassicas release isothiocyanates into the humans (Samuels 1996).
soil, which are toxic to R. solani. Enhance the toxic effects of
isothiocyanates by covering soil with a plastic tarp, as de- Trichoderma (Gliocladium) virens
scribed in method 2c (Howard et al. 1994). BIOLOGY: A soil fungus native to the USA that
Dippenaar et al. (1996) tested 'Fedora-19,' 'Futura-77,' antagonizes and parasitizes R. solani, Pytlzii~multiml~m,
'Felina-34,' 'Kompolti,' and 'Secuini,' and all the cultivars Botytis cinerea, and Sclerotil~mroysii. It is marketed for use in
were equally susceptible to R. solani. Broglie et al. (1991) en- glasshouses as SoilGards (formerly Gliogardo).
hanced resistance to R. solani using genetic engineering. They APPEARANCE & DEVELOPMENT:The smaller hyphae of
transferred a tomato gene coding for chitinase (an antifun- T. virens coil around R. solani hyphae and destroy them (Fig
gal enzyme) into tobacco and canola plants, which then be- 5.9). Many strains of T.virens also produce gliotoxin, which
came resistant to R. solani. suppresses the growth of l? ultimlim and R. solani (and mam-
Root-knot nematodes (Meloidogyne species) act mals, the LD50 in rodents is 25 mg kgl).
synergistically with R. solani. These underground "land APPLICATION: Formulated as 12-20% alginate-wheat
sharks" must be eliminated before sore shin can be bran prill and granules. Store up to five weeks at 10°C with
controlled. Disease detection kits utilizing ELISA (enzyme no loss in efficacy. Mix granules into moistened soil at a
linked immunoabsorbent assay) biotechnology can identify minimum rate of 2 g 1-1. If granules are broadcast upon the
R. solani in plants or soil. The kits are sensitive enough to soil surface, use a rate of 21-30 g m-2. T. virens works best
detect fungi before symptoms arise. against damping off fungi if applied three or two days prior
to planting. Disease control lasts at least a month. T.virens is
BIOCONTROL (see Chapter 10) compatible with mycorrhizal fungi (Linderman et al. 1991).
Biocontrol bacteria include Bacillus subtilis, Burkholderia NOTES: T.virens produces conidia held in slime (Fig 5.10),
cepacia, Pse~~domonasfl~~orescens,and the actinomycete Strep- which differs from the dry, powdery conidia produced by
tomycesgriseoviridis (all described under damping off). Fungi most Trichoderma species (Samuels1996).Because of the slime,
are also effective, especially Trichoderma species (described T. virens should disperse better in wet weather.
below), and Gliocladium roseurn (described under grey
mould). The mycorrhizal fungus Glornt~sintraradices is de-
scribed under Fusarium stem canker disease. The soil fun-
gus Verticillium biguttatum has experimentally controlled
R. solani. At least 30 other species of soil fungi parasitize R.
solani (Jeffries& Young 1994).

Trichoderma harzianum
BIOLOGY: A soil fungus that parasitizes the sclerotial
fungi Rhizoctonia solani, Sclerotit~mrolfsii, Botrytis cinerea, and,
to a lesser degree, Macrophomina pkaseolina and Sclerotinia
sclerotiort~rn,as well as non-sclerotial Fllsaritlrn, Pythium, and
Colletotriclz~~m species. It is found worldwide, and grows best
at 30°C (Gams & Meyer 1998).T. lzarzianum looks a lot like T.
viride (see Fig 5.10),but with smooth-walled conidia.
APPLICATION: T.harzianum is formulated in a variety
of biocontrol products (Binabo, Binab To, Bio-Fungus@,Bio-
Trek@, Rootshieldo, Supresivito, T-220, Topshieldo,
Trichodexo, Trichopelo), sold by itself or mixed with other
Trichoderma species, such as Trichoderma polysporum.
Supplied as a wettable powder, granules, or planter box
formulation. Unopened containers can be stored up to a year
in a cool (8-10°C), dark place. Mix wettable powder with water
and apply as a soil drench; mix granules directly into Figure 5.9: Biocontrol fungus Trichoderma virens coiling
moistened soil or potting media. T. harzianl~rnis heat-tolerant, around hypha of Rhizoctonia solani (SEM x3900), courtesy
so it can be added to soil immediately after sterilization or USDA.
pasteurization. The planter box formulation is mixed with
stickers and used as a seed treatment before planting. The
T-22 strain is a hybrid, selected for effectiveness against a Trichoderma viride
variety of pathogens on a variety of hosts. The T-39 strain BIOLOGY: A soil fungus sold for control of R. solani, Py-
(Trichodexo)can be applied as a foliar spray or painted on thium, Fusarium, and other soil pathogens. It grows world-
pruning wounds, although T. harzianum is not normally a wide in temperate regions, and researchers have isolated it
phylloplane fungus (Samuels 1996). from hemp (see last page in this Chapter). T. viride prefers
NOTES: T. lzarzianllm is compatible with Gliocladi~~m cooler climates, whereas T. harzianl~mgrows better in warm
roseurn, mycorrhizal fungi, and other biocontrol fungi. Its climates (Samuels 1996).
compatibility with beneficial nematodes is unknown. Some APPLICATION: T.viride is formulated with T,harzianum
T. harzian~~rn isolates (e.g., T-22) tolerate fungicides such as as Trichopels, Trochoseals, and other products. Supplied as
benomyl, captan, and PCNB. T-22 has been applied as a seed a wettable powder or granules for mixing into soil. T, viride
coat over seeds previously treated with capian-the fungi- is compatible with other Trichoderrna species.
104 Hemp Diseases and Pests

Some researchers simply list the cause as "Phoma sp.," from

Lithuania (Brundza 1933), Iowa (Fuller & Norman 1944,
1945), Kansas (Paulsen 1971), India (Srivastara & Naithani
1979),and the Czech Republic (Ondrej 1991).Most research-
ers, however, attempt to label their pathogens with specific
names. At least 23 names appear in the Cannabis literature.
McPartland (1995~)sorted them out, and reduced the 23
names to nine species:

1.Phoma cannabis (Kirchner)McPartland, Mycologia 86:871,1994.

~Depazeacannabis Kirchner 1856,=Phyllosticta cannabis (Lasch)
Spegazzini1881,=Phyllosticta cannabis (Kirchner)Speg. apud others,
=Ascochyta cannabis (Spegazzini)Voglino 1913 [non: Ascochyta
cannabis Lasch 1846 (Septoria cannabis (Lasch) Sacc.)];=Eysiphe
communis var. urticirum Westendorp 1854; =Diplodina cannabicola
Petrak 1921,=Diplodinaparietaria Brun. f. cannabina von Hohnel1910.
teleomorph: Didymella cannabis (Winter) von Arx, Beitr.
Figure 5.10: Trichoderma species x640: A. T: viride Kyptogamanflora Schwiez 11:365, 1962; =Sphaerella cannabis Winter
(McPartland redrawn from Rifai 1969); B. T: virens 1872, =Mycosphaerella cannabis (Winter) Magnus 1905,
~Mycosphaerellacannabis (Winter)Roder 1937.
(McPartland redrawn from Samuels 1996).
Conidial morphology is variable, which explains the
plethora of synonyms. The fungus also produces ascospores
and chlamydospores. The only known host of P. cannabis is
CHEMICAL CONTROL (see Chapter 11) Cannabis, both fibre and drug cultivars. Herbarium speci-
Mishra (1987)controlled R. solani with several synthetic mens are limited to the northern hemisphere (North America,
fungicides, by soaking seeds in fungicide solutions for 15- Europe, the Indian subcontinent, Japan).The fungus infests
20 minutes before planting them in infested soil. He also root- flowering tops (Plate 55), leaves, and stems.
dipped Cannabis seedlings in fungicide solutions before trans-
planting seedlings into infested soil (see Appendix 1). 2. Ascochyta arcuata McPartland, Mycologia 86:873,1994.
teleomorph: Didymella arcuata Roder, Phytopath. Zeits. 12321-333,1939.
Like the previous species, A, arcuata produces pycnidia,
BROWN LEAF SPOT & ascocarps, and chlamydospores. It has only been collected
from European hemp, on leaves and stems. Roder (1939)
STEM CANKER erroneously connected Didyrnella arcuata with Ascochyta can-
Several species of Phoma and Ascochyta cause this disease nabis. This misconception continues to arise (Spaaret al., 1990).
all over the world. Brown leaf spot and stem canker rivals
yellow leaf spot as the most common Cannabis problem in 3. Phoma exigua Desmazieres,Annls. Sci. Nut. (Bot.)ser. 3,11:282,
Europe. Sometimes the two diseases appear on the same 1849; =Ascochyta phaseolorum Saccardo 1878; =Plenodomus cannabis
plant (Plate 54). Brown leaf spot, like yellow leaf spot, is a (Allescher)Moesz & Smarods in Moesz 1941,=Plenodonzus herbarum
"disease of attrition1'-as leaf losses increase,, crop yields f. cannabis Allescher 1899.
decrease (whether fibre, flowers, or seed). Sutton & Waterston (1966) listed Cannabis as a host of
A. phaseolorum. Ondrej (1991) described P. exigua appearing
SYMPTOMS "abundantly" on stalks in the Czech Republic. According to
Leaves low in the canopy begin developing small brown Kaushal & Paul (1989), P. e x i g t ~ ais resistant to fungitoxic
spots in late May. Rarely, spots turn brick red (Shukla & extracts of Cannabis. Domsch et al. (1980)considered P. exigtla
Pathak 1967). Spots remain circular (Plate 57), or form a the most common pycnidial fungus in the world. It attacks
straight edge along leaf veins. They average 5 mm in over 200 plant genera. Over 100 synonyms exist for this cos-
diameter, rarely up to 15 mm. Sohi & Nayar (1971) described mopolitan pathogen. No teleomorph is known. P. exigua-in-
spots coalescing into large irregular lesions but this is unu- fected Cannabis has only been found in central and eastern
sual. Spots may become peppered with tiny black pycnidia Europe, limited to stem infections (Plate 56).
(Fig 5.11).Two of the fungi causing this disease subsequently
produce tiny pseudothecia (Roder 1939, Barloy & Pelhate
1962).Old spots often break apart or drop out, leaving small
shot-holes in leaves. Flowering tops may also develop brown
spots (Plate 55).
Stem cankers begin as chlorotic spots. They quickly turn
grey or brown, and elongate along the stem axis (Plate 56).
Pycnidia form on the surface of stem lesions. Two species
subsequently develop pseudothecia on stem lesions ( ~ o d e r
1937,1939).Diseased plants are stunted, 30 cm shorter than
healthy plants by midJune (Roder 1939).

CAUSAL ORGANISMS & TAXONOMY

Many pathologists erroneously cite the cause of brown
leaf spot as "Phyllosticta cannabis."McPartland (1995~)ex-
amined 25 herbarium specimens labelled "Phyllosticta can-
nabis" from the USA and Europe. N o n e were true Phyllosticta Figure 5.11: Dense growth of Phoma cannabis pycnidia
species; most were misidentified Phorna or Ascochyta species. on a leaf with brown leaf spot (LM x135, McPartland).
 Chapter 5: Fungal Diseases 105

4. P h o ~ ~ glolnerata
za (Corda) Wollenweber & Hochapfel, 2. 8. Ascochyta catztlabina E.I. Reichardt, Btllletin lie la Station
l'ninsztkde 8:592, 1936. Rigionale Pyotectrice rfes Plants 2 Leningrad 5:46, 1925.
McPartland (1995~)collected this species from leaves Reichardt described A. calznabina causing leaf spots on
of a C. sation-C. afglza~zicahybrid in Illinois (Plate 57). The hemp near St. Petersburg. He did not report a teleomorph.
chlamydospores of I? glomeratn can be confused with conidia
produced by Alternnria nlternata (Fig 5.17). P.glo~izerata 9. P h o m a nebulosn (Persoon:Fries) Berkeley, Outlilze of British
parasitizes at least 94 plant genera worldwide (Sutton 1980). Fzlngi p. 314,1860.
No teleomorph has been found. Gruyter et al. (1993) cited this species on C. snfioa in the
Netherlands. It is a common soilborne saprophyte, isolated
5 . P h o ~ n nherbnvum Westendorp, B~llletinsAcaderrlie Royale from various herbaceous and woody plants in temperate
Bt'lgi~~ueCl. Sci 19:118, 1852. climates worldwide.
This fungus reportedly causes stem cankers on hemp
in Italy (Saccardo 1898), France (Brunard 1899), Denmark MORPHOLOGICAL DESCRIPTIONS
See McPartland (1995~)for full descriptions. The nlne specles
(Lind 1913), the Netherlands (Oudemans 1920), and Japan share many characteristics: Pycnidia dark brown, immersed then
(Kyokai 1965, Kishi 1988). Boerema (1970)considered P. Izer- erumpent, subglobose, unilocular, ostiolated, usually 90-200 pm di-
bnrlim a very weak pathogen, but Domsch et 01. (1980) re- ametre on host substrate (up to 450 pm diameter in culture). Conid-
ported it from over 35 host genera, worldwide. No iogenous cells phialidic, discrete, determinate, simple, ampulliform
teleomorph is known. to short cylindrical. Conidia hyaline, light brown en masse, ellipsoi-
dal to short cylindrical, straight or curved, usually zero or ongsep-
6 . P h o ~ n npiskorzii (Petrak) Boerema & Loerakker, Pevsoonin tum (rarely two septa), not constricted at the septum, sometimes
11:31C7,1981;-Diploplenodorntis piskorzii Petrak 1923; non: Pl~on~aacuta finely guttulate, size ranging from small (4 x 2 pm) to very large (28
auct., nomen a m b i g u u m . Teleomorph: Leytospizneria acuta x 8 pm) on host substrate; larger in culture. Chlamydospores vary
(f3offman:Fries)P.Karsten 1873. from single-celled and globose (8-12 pm diametre) to multicellular,
Saccardo & Roumeguere (1883)reported the teleomorph even alternariod in appearance (Fig5.17).Pseudothecia brown-black,
immersed or erumpent, globose on leaves and flattened on stems,
on fibre plants in Belgium. They may have erred (McPartland
130-150 pm diameter on host (up to 250 pm in culture). Ostiole sim-
1995c,e).The fungus normally attacks stinging nettles, Urtica ple. Asci bitunicate, clavate to pyrlform, with a slight pedicle, eight-
riiolcll L. spored, 50-90 x 9-15 pm. Ascospores hyaline, subovoid to oblong,
submedially one-septate with cells unequal in size, constricted at
7. Ascocliyta prnsndii Shukla & Pathak, Syrfoii~ia21:277, 1967. septum, ranging 16-22 x 4-6 pm. Pseudoparaphyses filamentous,
Shukla & Pathak isolated this fungus from leaves of a zero to three septa, ranging 23-60 x 0.5-1.5 pm.
drug plant in Udaipur, India. No teleomorph is known. Differentiating the nine anamorphs is not easy on the

Table 5.1: Characteristics of nine Phoma or Ascochyta species reported from Cannabis

SPECIES (p)
PYCNIDIA CONIDIA(p) (p)
CHLAMYDOSPORES TELEOMORPH

i? cannabis 65-(130)-180 1-2-celled, oval-ellipsoid, globose to dictyo- Didymella

guttulate, 3-8 x 2-3 form, 8-1 7

A. arcuata 90-(135)-150 2-celled, short-cylindrical, globose to oval, Didymella

usually guttulate, 9-28 x 3.5-8.0 8-1 2

?
i exigua 110-220 1-2-celled, ellipsoid-cylindrical, absent none known
guttulate, 5-1 0 x 2-3.5

P glomerata 50-300 usually 1-celled, ellipsoid-cylin- dictyoform, none known

drical, guttulate, 4.5-1 Ox15-4 30-65x15-20

P herbarum 80-260 usually 1-celled, ellipsoid, absent none known

nonguttulate, 4.5-5.5 x 1.5-2.5

P piskorzii 300-500 usually 1-celled, ellipsoid, absent Leptosphaeria

nonguttulate, 6-1 6 x 2-3

A. prasadii 55-93 usually 2-celled, oblong, absent none known

nonguttulate, 7.5-1 0 x 1.5-2.5

P nebulosa 100-250 usually 1-celled, oblong, absent none known

guttulate, 3.6-6.6 x 1.4-2.0

A. cannabina 120-1 30 usually 2-celled, oval-oblong, absent none known

nonguttulate, 7-8.8 x 3.2-4
106 Hemp Diselases and Pests

host. Teleomorphs are also difficult to tell apart. Refer to Ta- necrose and fall off. Whole plants and entire fields may
ble 5.1 for key characteristics. Most Phoma species must be follow this course.
cultured in a laboratory to confirm their identification.
In culture, the two most common species (P. cannabis CAUSAL ORGANISMS & TAXONOMY
and A. arcclata) are distinguished by conidial size, shape, and 1. Pseudoperonospora cannabina (Otth) Curzi, Riv. Pat. Veg.
septation. According to Roder (1937,1939),who cultured both Pavia 16:234, 1926; =Peronospora cannabina Otth 1869, =Perono-
fungi, the conidia of A. arctlata produce septa, and the conidia plasmopara cannabina (Otth) Peglion 1917, =Pseudoperonospora can-
of P. cannabis do not. A. arctlata conidia, i n vitro and on the nabina (Otth)Hoerner 1940.
Description: Sporangiophoreshyaline with several arising per
host, are longer than P. cannabis conidia. leaf stomate, dichotomous branching meagre, occasionally up to
the third order, often with swellings on the main axis and branches,
DIFFERENTIAL DIAGNOSIS 100-350 x4-8 pm. Sporangia ovoid to ellipsoid, grey-violet(turning
Yellow leaf spot and white leaf spot produce larger le- brown with age),with protruding apical papillae, germinating into
sions than brown leaf spot, and they are lighter in colour. zoospores or hyphae, usually 26-30 x 16-19 pm (Figs 3.2 & 5.12).
When brown leaf lesions break up and fall out, the holes may Zoospores reniform, laterally biflagillate, rounding up to produce
resemble insect damage. Dobrozrakova et al. (1956) noted hyphae. Oogonia with oospores rare, reported once within Cannabis
"stains on stems" from this disease resembled stains caused cotyledon mesophyllum (by Peglion 1917,who gave no description
or measurements).
by Dendrophoma marconii and Microdiplodia abromovii. Stem
cankers may be confused with symptoms caused by
2. Pseudoperonospora humuli (Miyabe& Takahashi)Wilson, My-
Ftlsaritlm, Coniothyrit~m,Leptodothiorella, Ftlsicoccum, or Pho-
cologia 6:194,1914; ~Peronoplasmoparahumuli Miya. & Takah. 1905;
mopsis species. These fungi are hard to distinguish without a =Pseudoperonosporaceltidis (Waite)Wilson var. humuli Davis 1910.
microscope. Description: Sporangiophores hyaline, with two to five aris-
ing per stoma, dichotomous branching abundant, occasionally up
DISEASE CYCLE & EPIDEMIOLOGY to the sixth order, 200460 x 6-7 pm. Sporangia ellipsoid, grey, with
Pycnidia and pseudothecia overwinter in plant debris. blunt apical papillae, germinating into zoospores, usually 22-26 x
P. cannabis, A. arctlata, and P. glomerata also survive as chlamy- 15-18 pm. Oogonia oval, brown, 40-49 pm in diameter.Oospores
dospores in soil. Infection occurs in early spring, but symp- spherical, brown, smooth, 25-35 pm in diameter.
toms may take a month to appear. Conidia spread in splashed Only three authors report P. hclmuli on Cannabis-
rain and wind-driven water. Epidemics build by late sum- Hoerner (1940), Ceapoiu (1958), and Glazewska (1971). The
mer. The optimal temperature for mycelial growth and spore pathogen normally infests hops. Hoemer suggested P. htlmcili
germination is 19-22°C (Roder 1939). and P. cannabina represent different physiological races of
one species. Berlese (1898) considered P. cannabina a species
CULTURAL & MECHANICAL CONTROL (see Chapter 9) dubium, identical to P. epilobii Otth. Waterhouse & Brothers
Follow all measures described under yellow leaf spot. (1981) recognized P. htlmtili and P. cannabina as different spe-
Brown spot worsens on hemp grown in peat soils (Rataj 1957); cies, based on sporangiophore branching patterns and spo-
avoid overclse of nitrogen fertilizer. P. exigtla can spread by rangia papillae.
seedborne infection (Sutton & Waterston 1966); do not har-
vest seed from infected females. Many Phoma diseases bloom DIFFERENTIAL DIAGNOSIS
in crops weakened by environmental stress. Be sure plants The English call this disease "false mildew," to differ-
are watered and well-maintained. entiate it from powdery mildew. Downy mildew can also be
confused with olive leaf spot, brown leaf spot, or gall midges.
BIOCONTROL & CHEMICAL CONTROL Ferraris (1935) said leaf contortions caused by downy mil-
Biocontrol against brown leaf spot and stem canker is dew resembled symptoms by Ditylenchys dipsaci, the stem
not available. Kirchner (1966) used copper-based chemical nematode. Careful observation with a microscope or strong
sprays such as Bordeaux mixture. magnifying lens makes these problems easy to differentiate.

DISEASE CYCLE & EPIDEMIOLOGY

Sporangia are spread by wind or water. Germination of
DOWNY MILDEW sporangia requires a wet period (a heavy dew will do). Spo-
Downy mildew infests half-grown to mature plants. A
world map outlining the range ofdowny mildew, dublished
by the Commonwealth Mycological Institute (1989), is
outdated-the disease now occurs on every continent except
Antarctica (McPartland & Cubeta 1997). Two oomycetes
cause downy mildew of Cannabis. Both pathogens attack fibre
and drug cultivars. Researchers have proposed using downy
mildew to destroy clandestine Cannabis plantations (McCain
& Noviello 1985).Zabrin (1981) claimed, "A single infected
plant introduced into Colombia or Jamaica during a wet
season could cause complete devastation."

SYMPTOMS
Downy mildew begins as yellow leaf spots of irregular
size and angular shape, limited by leaf veins. Opposite the
spots, on undersides of leaves, the organisms emerge from
stomates to sporulate. Mycelial growth on undersides of
leaves is best seen in early morning when dew turns the
mycelium a lustrous violet-grey colour. Lesions enlarge Figure 5.12: Sporangia and sporangiophores of
quickly, and affected leaves become contorted. Leaves soon Pseudoperonospora cannabina (LM x150, McPartland).
 Chapter 5: Fungal Diseases 107

rangia germinate into hyphae or cleave into zoospores. Hy- cardo 1879,=Botnjospizaeria saubiizetii (Montang~~e) Niessl1872; =Bat-
phae penetrate plant epidermis directly. Smith et 01. (1988) ryosphnerin dispersa DeNotaris 1863; =Gibberella quinilileseptntn
noted P.11lliizi~lizoospores settle directly over open stomates Sherbakoff 1928.
Description: Conidiogenous cells hyaline, cylindr~cal,aris~ng
during the day, but in darkness the zoospores settle randomly palmately from metulae, phialidic, 12-20 x 3-5 pm. Macroconld~a
over leaf surfaces. Since zoospores can only invade via fusiform, hyaline, with a curved, pointed apical cell and a marked
stomates, infection becomes dependent on daytime leaf wet- foot cell, usually three or four septa (range zero to SIX), 30-50 x 3.5-
ness. Epidemics arise w h e n w a r m humid days are followed 5.0 pm (Fig 5.5).Microconidia absent. Cl~lamydosporesrare, single
by cool wet nights. or in short chains within hyphae or macroconidia, smooth-~valled,
Downy mildew fungi may persist i n a field, and become 8-10 Fm in diameter. Perithecia superficial, scattered, blue-black,
progressirrely worse over the years. Barloy & Pelhate (1962) globose, peridium textura nnglllaris-ylob~~losa, with a distinct ostiole,
blamed oospores as the source of these perennial infections. 150-300 pm in diameter. Asci clavate, e~ght-spored,65-90 x 12-20
pm. Pseudoparaphyses present but evanescent.Ascospores ellipsoid,
hyaline, straight, slightly constricted at three septa, 20-25 x 5-7 p n ~
CULTURAL & MECHANICAL CONTROL (see Chapter 9) (Fig 5.5).
Methods 1 (sanitation), 2b&c (sterilize and pasteurize F. slllphurel~n?or its teleotnorph (commonly cited as G.
the soil), 5 (genetic resistance-see below), 6 (rotate other
s a l ~ b i n e t t i )infest h e m p in Europe (Saccardo S.F.11 1883,
cropsfor a minimum three years), 7c (important-avoid
Voglino 1919, Oudemans 1920) and the USA (Scherbakoff
glasshouse dew), 8 (optimize soil structure and nutrition),
1928, Miller et al. 1960). The teleomorph has been confused
10 (very important-completely r o g u e infected plants with G. zeae (Wollenweber & Reinking 1935, Vakhrusheva
including roots). McCain & Noviello (1985) cited two Italian
1979). McPartland (1995e) added G. q l ~ i i ~ q l ~ e s e p t to
a t nthe list
hemp cultivars, 'Superfibra' and 'Carmagnola Selezionata,' of G. cyanogena synonyms.
with resistance to a strain of P. can~znbinathat destroyed all
drug culti~,ars. 2. F u s a r i t r m g r a m i n e a r u m Schwabe, Flora Anhalttnn 2:285, 1838.
=Fllsariunl roseurit Link emended Snyder, Hansen & Oswrild
BIOCONTROL & CHEMICAL CONTROL 1957, p ~ oparte.
A unique strain of B a c i l l l ~ ssubtilis (Serenade@)is sold teleomorph: Gibberella zeae (Schweinitz:Fries) Petch, Annales
as a foliar spray for controlling downy mildew (described Mycologici 34:260,1936;-5phaeria zeae Schweinitz 1822 (non Spiznerin
under damping off). Yepsen (1976) sprayed downy mildew zeae Schweinitz 1832).
with a solution of boiled horsetail leaves. Ferraris (1935) con- Description: Co~udiogenouscells hyaline, short, dolilform,
trolled c a i ~ n a b i n nepidemics with copper sulphate. Hewitt phialidic, simple or branched dichotomously, proliferating through
(1998) treated P. l z l ~ n ~with
~ ~ l i Bordeaux mixture or copper
previous phialides, 10-14 x 3.54.5pm. Macroconidia sickle-shaped,
hyaline, with a well marked foot cell, three to seven septate, 25-30 x
oxychloride. Be sure to spray the undersides of leaves. 3-4 pm. Microcollldia absent (although some are illustrated by Booth
1971).Chlamydospores rare, single or in chains, 111 hypl~aeor rarely
in macroconidia, hyaline to brown, thick-walled, smooth or slightly
rough surface, 10-12 pm in diameter. Perithecia superficial, clus-
FUSARIUM STEM CANKER tered, blue-black, ovoid, with a rough tuberculate surfaced, 140-250
This section begins a subchapter o n several Fusarium pm in diameter. Asci clavate, usually eight-spored (rarely four to
diseases. Fusarium stem canker is caused by six F l ~ s a r i l ~ n l six), 60-85 x 8-11 pm, but rapidly deliquesce after ascospore pro-
duction. Ascospores fusoid, hyaline to light brown, curved, three-
species, most wit11 Gibberella sexual states. Fusarium root rot, septate (200'0 two-septate or less), 19-30 x 3-5 pm.
described in the next section, is caused by a F ~ ~ s n r i i species
~nl F.gvanlinenr~~rrl often confuses mycologists, even so emi-
with a N e c t r ~ nsexual state. F u s a r i u m wilt, described nent a pair as Wollenweber & Reinking (they called F.
thereafter, is caused by two F~lsarililrlspecies without sexual ~ conidial state of G. saubinetii, see McPartland
g r a n ~ i n e a r u nthe
states. F l ~ s a r i l ~ species
nl also cause damping off of seedlings 1995a). E granzinerlllrl or its teleomorph, G. zene, infest hemp
(Figs 5.5 & 5.6). in Europe (Ceapoiu 1958, Ferri 196lb). Francis & Burgess
(1977) described two groups of G,zenr: Group 1 is soilborne,
SYMPTOMS causes crown rots, and rarely produces perithecia (recently
Fusarium stem canker usually arises o n mid-season-to- renamed F u s a r i u r n p s e u d o g r a m i n e a r u m 0 8 D o n n e l l &Aoiki
mature Calzizabis. Symptoms begin as watersoaked epider- 1999). Group 2 is airborne, causes stem cankers, and readily
mal lesions, followed by epidermal chlorosis and cortical forms perithecia. McPartland & Cubeta (1997) isolated a
necrosis. Stems often swell at the site of the lesion, creating Group 2 homothallic strain from feral hemp in Illinois.
fusiform-shaped cankers that may split open (Plate 58).
Leaves on affected stems wilt a n d necrose, without falling 3. F u s a r i u m l a t e r i t i t r m Nees:Fries, Syst. Piize Scli~canzniep. 31,
off plants. Cankers rarely girdle stems but, if they do, all 1817, emended Snyder & Hansen 1945, pro parte.
upper leaves wilt and die. Slicing open cankers often reveals teleomorph: Gibberelin baccata (Wallroth) Saccardo, Sylloge
n reddish-brown discolouration. Fungorllni 2:553, 1883.
Pietkiewicz (1958) cited this species infesting hemp seeds
CAUSAL ORGANISMS & TAXONOMY in Poland, possibly erroneously. It normally infests woody
Six Fiisnrillnl species reportedly cause canker, b u t only hosts. For a description, see Booth (1971).
two have been isolated on a regular basis. F ~ ~ s n r i itaxonomy
~in
is confusing. Even the experts disagree-Wollenweber & 4. F t r s a r i u m s a m b u c i n t r m Fuckel, Syntbolae Mycologlcae p 167,
Reinking described 143 F ~ ~ s a r i u species,
ln varieties, and spe- 1869; =Ftrsantlnl sarcocl~ro~~rn (Desmazieres)Saccardo 1879,=F~isarlurrt
cial forms; Booth cited 44 species and seven varieties, but voseunz Link emended ~ n y d e r& Hansen 1945, pro parte.
according to Snyder & Hansen there are only nine species ... teleomorph: Gibberella pulicaris (Fries:Fries)Saccardo, Mlcheiin 1:43,
see Toussoun & Nelson (1975) for a n overview. 1887; =Botryospl~aeriapulicaris (Fries) Cesat~& DeNotarls ajllid
Oudemans 1920.
G. p ~ l l i c a r i s h a s b e e n described o n h e m p once, by
1. F t r s n v i u m strlplztireum Schlecl~tendahl,Flora berolinensis p.139,
Oudemans (1920).The species causes a basal canker on hops
1824.
teleomorph: Gibberella cyanogena (Desmazieres) Saccardo, and woody hosts. F. s a n ~ b l ~ c i n z ~may r i i be identical to F. s ~ l -
Sylloge Fungonlm 2:555,1883; =Gibberella sat~bi~zetii
(Montagne) Sac- pllllrellnl (Howard et al. 1994).
7 08 Hemp Diseases and Pests

5 . Flrsaritlnz nvenacel4tn (Corda:Fries)Saccardo, Sylloge Fuizgorltrr~ ploughing), 2b&c (sterilizing and pasteurizing the soil), 2d
4:713, 1886. (flooding soil), and 10 (careful pruning) are paramount for
teleomorph: Gibbei.elln nelelzncen R.J. Cooke, Phytopntlzology reducing inoculum. Other important methods include 7b
57:732-736, 1967. (avoid overwatering), 11 (avoid seedborne infections), and
G. nuenncen caused Cnizlznbis disease in host-range ex- 5 (cultivar 'Fibramulta 151' is resistant to Filsnri~~nt
species
periments (Zelenay 1960). It attacks a wide range of hosts. according to Dempsey 1975).Do not plant in heavy, wet soils
For a description, see Booth (1971). or low lying areas. Avoid excess nitrogen; increase potassium
and phospl~orus.Choose KC1 as a potassium supplement
6. Ftlsariuin c t d ~ n o r t ~ r(W.G.
n Smith) Saccardo, Sylloge Fungorunz instead of K2S04or KN03. KzSOl and KN03 may actually
11:651,1895. increase the disease (Elmer, pers. commun. 1950). since
This pathogen normally infests cereal crops. Abiusso fusaria have a wide host range and live saprophytically, crop
(1954)cited it on Argentinean Cannabis, perhaps erroneously. rotation is not effective. Diseased plants should not be
For a description, see Booth (1971). harvested for human consumption.

DIFFERENTIAL DIAGNOSIS BIOCONTROL

Separating Filsarii~nzspecies usually requires culturing Biocontrols include the bacteria Bacillus silbtilis, Burkkol-
them in lab and hoping their teleomorphs develop. F. sul- deria cepacia, Pseudon~o~zas fluorescens, and the actinomycete
p111irr1intmacroconidia are larger than E gramitiearunz mac- Strepfmzyces griseoiiiridis, all described under damping off.
roconidia. The conidiogenous cells of F. s~llplzl~reuni are cy- Piclioderma and Gliocladillin fungi are also useful (see Rhizoc-
lindrical and branch in a palm-shape, whereas F. grl~nzii?e- tonia sore shin).
~11'11111 conidiogenous cells are stubby (doliiform), branch in
palrs, and proliferate through previous conidiogenous cells. Gloinus intraradices
Fusarium foot rot (see next section) may be confused BIOLOGY: A mycorrhizal fungus (Mycori-Mixa, Nutri-
with Fusarium stem canker. The cause of foot rot (Fl~sarillin Link@)that protects roots against soil pathogens such as
solani) can be distinguished from F. sl~lph~treunz and F. Fusarii~nzand Pythiilnz species. It does best in soils with
grnintnmr~lniby its microconidia. moderate moisture and moderate temperatures.
The reddish xylem discolouration seen in Fusarium stem APPEARANCE & DEVELOPMEET: See the section on myc-
canker does not appear in grey mould, hemp canker, and orrhizae (in Chapter 5) to understand the utility of Glontus
southern blight. Stem swelling caused by Fusariunz species species.
can be confused with stem swellings caused by European APPLICATION: Supplied as spores which can be stored
corn borer, hemp borer, and assorted beetles. for months in a cool (8-1O0C), dark place. G. intraradices is
used as a soil inoculant to improve plant nutrition, and it
DISEASE CYCLE & EPIDEMIOLOGY protects plants from a variety of root pathogens.
Perithecia overwinter in crop stubble or soil. Some NOTES: Although mycorrhizal fungi have wide host
Fiisarz~iinspecies also overwinter as seedborne infections. In ranges, they also show preferences for certain soils, climates,
the spring, ascospores or conidia infect seedlings. Later in and hosts. Thus, mixtures of mycorrhizae are becoming
the season, F I L S I ~ Y ~species
I I I ? I invade roots via wounds cre- available. Mycorrhizae work well in combination with other
ated by nematodes and Orobnilche parasites. Since Fllsariunz microbial biocontrols, such as Trichoderma, Gliocladi~lni,
conidia move in water droplets, they do best in damp condi- Talaroinyces, Bacilllls, Pseu~ionzonas,Streptom!/ces, and Bacilllis
tions and heavy soil. Watch for Fusari~linepidemics during species (Linderman et nl. 1991). Other soil organisms attack
seasons wit11 above-average rainfall. Above ground, F1lsariiini mycorrhizae, including fungi (Cephalosporillnt species), nema-
conidia cause secondary infections at stem nodes and wound todes (Apkelenchoides species), and insects (Folsoniin species).
sites. Wounds are caused by stem-boring insects, wind, im- Pesticides, synthetic fertilizers, and frequent soil tillage kill
proper pruning, and from damage caused by dry, caking soil. mycorrhizae.
Hydroponic cultivators should not allow rockwool blocks
to dry out, because evaporated fertilizer salts accun~ulate CHEMICAL CONTROL
around the base of the stalk and favour infections (Howard Seeds suspected of harbouring Fl~snriilnz should be
et nl. 1994). disinfested. Booth (1971)soaked seeds in formalin (see Chap-
All six F~~siirilli~?species are distributed worldwide. F. ter 12).Spraying fungicides on plants has not proved practi-
grnnlii1enrunl and F. avenacelini predominate in cooler climates. cal against Fl~sariiliiz.Neither has soil fumigation, except to
F grm~~'ir~enrlcn~is considered the most pathogenic of the control synergistic organisms such as nematodes. Vysots'kvi
bunch (Booth 1971). Most Filsariilnl species are facultative (1962) protected pine seedlings from F~lsariilirzby mixing
parasites, existing as soil saprophytes until a parasitic hemp leaves into soil.
opportunity presents itself.
Some fusaria pose a threat to humans. F. graininearllm
produces zearalenone, a toxic metabolite. Consuming - foods FUSARIUM FOOT ROT
~nfestedwith F, grnnziiiearllnz may cause nausea, vomiting,
diarrhea, headache, chills, and convulsions (Rippon 1988).
& ROOT ROT
F granziizenr~inialso produces trichothecenes, known as "T-2 Fusarium foot rot and root rot has been reported in the
toxins." T-2 toxins cause a haemorrhagic syndrome, gaining former USSR (Gitman & Boytchenko 1934, Vakhrusheva
notoriety for their reputed use in chemical warfare ("yellow 1979), Poland (Zelenay 1960), southern France (Barloy &
rain"). Lastly, fusaria can invade human tissue and grow in Pelhate 1962), and Maryland (Miller et al. 1960). Data in
dirty wounds and in the skin of burn victims (Rippon 1988). USDA archives also described the disease affecting hemp in
Illinois a n d Virginia. The causal organism is found
CULTURAL & MECHANICAL CONTROL (see Chapter 9) worldwide. On hemp it sometimes causes stem canker
Booth (1971) characterized stem canker as a "disease of instead of root rot (Wollenweber 1926).
accumulation." Thus methods 1 (sanitation), 2a (deep
 Chapter 5: Fungal Diseases 109

SYMPTOMS Mixing dried Cannabis leaves into soil may suppress

Plant stems turn brown at the soil line; this "foot rot" is Fi~sariurnspecies (McPartland 1998). Grewal (1989) mixed 3
an extension of the root rot. Common above-ground symp- kg of dried leaves into 137 kg of wheat straw compost and
toms include partial or systemic wilting. Roots turn red, rot- suppressed F. solani growth. Pandey (1982) reported an aque-
ten, and necrotic (Plate 59).According to Zelenay (1960)and ous extract of hemp leaves inhibited F~~sarillni
in petri plates.
Barloy 6r Pelhate (1962), this disease knocks down plants in Vysots'kyi (1962) applied an aqueous extract in the field to
all stages of development, including seedlings (Fig 5.6). protect pine seedlings from a Filsarii~nzspecies. Dahiya &
Jain (1977) reported that pure THC and CBD inhibited the
CAUSAL ORGANISM & TAXONOMY growth of F. solani.
Fusarium solnni (Martius) Saccardo, Mlci~riii~
2:296, 1881,
emended Snyder & Hanson, Anl. 1. Bot. 28:740, 1911.
=Flisa~.il~ni jai~anicziniKoorders 1907.
teleomorph: Nectuia llaenlatococca Berkeley & Broome, I. Linn. Soc.
FUSARIUM WILT
iBotan!yi l l : l l h , 1873;=Hypomyceshaernatococca (Berkeley & Broome) The pathogen causing this disease is a xylem inhabitant,
Wollenweber 1926; =Hypornyces cnncri (Rutgers) Wollenweber 1914, rather than a cortical pathogen like the aforementioned
~Nectriaozncri Rutgers 1913; 11011:Hypoinyces solani Rein. &Bert. 1879. Fllsariz~nzspecies. It plugs plant water-conducting tissues and
Description: iMacuoconidiai conidiogenous cells hyaline, short, causes a wilt. Fusarium wilt is a serious disease in eastern
doli~forni,frequently branched, phialidic. Macroconidia fusoid, Europe (Dobrozrakova et al., 1956, Rataj 1957, Ceapoiu 1958,
stout, broad, thick-walled, hyahne, with a pointed, somewhat beaked Czyzewska & Zarzycka 1961, Serzane 1962, Zhalnina 1969),
ap~calcell, one to six septa, 40-100 x 5-8 pm (Fig 5.5). Micvoconidial Italy (Noviello & Snyder 1962),and southern France (Barloy
conidiogenous cells hyaline, long, cylindrical, phialidic, up to 400
& Pelhate 1962). It may occur in Pakistan (Ghani et al. 1978)
pm In length. Microconidia hyaline, ovoid to allantoid, sometimes
becom~ngone-septate, 8-16 x 2-4 Fm. Cl~lamydosporesusually in and the USA (McPartland 1983a).Gitman (196813)considered
pairs, but sonietnnes single or in chains, globose to oval, smooth to it a little-known disease in the former USSR. The causal fun-
rough walled, 9-12 x 8-10 pm. perithecia superficial upon a thin gus also causes damping-off in seedlings (Pietkiewicz 1958,
pseudostroma, scattered or clustered, pale orange to brown, globose Zelenay 1960, Barloy & Pelhate 1962).
( i ~ p o drying
n they collapse laterally, as often illustrated in textbooks), A scientist at UC-Berkeley, Arthur McCain, suggested
1 ~ 1 tah \carted surface, 110-250 pm in diameter. Asci cylindrical to to President Nixon that Fusarium wilt could destroy illegal
clavate w ~ t ha roulided apex, eight-spored, 60-80 x 8-12 pm. marijuana cultivation (Shay 1975). Subsequently, the Nixon
.4scospores ellipsold to obovoate, hyaline but becoming light brown
\ c ~ t hlongitudinal str~ationsat maturity, slightly constricted at single
administration funded research to mass-produce the wilt
septum, 11-18 x 4-7 p n ~( F ~ g 3.2 s & 5.5). fungus (Hildebrand & McCain 1978). Zubrin (1981)
interviewed McCain, who claimed, "Just introduce a couple
DISEASE CYCLE & EPIDEMIOLOGY of pounds [of the fungus] into an area, and while it wouldn't
F solnni overwinters as conidia, chlamydospores, or as- have much of an effect the first year, in several years it would
cospores in crop debris or in soil. Perithecia are abundant in spread throughout the country with devastating results." The
!'vet tropics and less common in temperate zones (Booth project was terminated by Carter administration officials
1971). The mode of infection is similar to Fllsariilin species (Zubrin 1981).
described in the previous section. Although F. solaili is char- Interest in eradicating "Cannabis satizlrls" (Sands 1991)
acterized as a weak pathogen, it acts synergistically with with the wilt fungus resumed during the Reagan/Bush ad-
nematodes and parasitic plants. It easily invades root ministration. A virulent isolate of the fungus was collected
wounds created by other organisms. Barloy & Pelhate (1962) in Russia; Ronald Collins subsequently released the patho-
considered a combination of F. solaizi and broomrape gen in a USDA field plot in Beltsville, Maryland. The fungus
(Orobnnihe ranzosa) the greatest threat to Cannabis cultiva- attacked Calztznbis plants, successfully overwintered in the
t ~ o nin southern France. soil, then killed seedlings planted in the same soil the fol-
Like other fusaria, F. solaiii may also pose a threat to lowing year (Sands 1991). More recently, 25 strains of the
humans. Ingesting F. solnizi-moulded sweet potatoes may fungus were isolated by researchers in Russia and
cause respiratory distress. F.solani also causes eye infections, Kazakhstan (Semenchenko et al. 1995).Some strains reduced
especially in agricultural workers (Rippon 1988). Carlnabis survival by up to 80% (Tiourebaev et al. 1998). The
current deployment of genetic engineering to create super-
DIFFERENTIAL DIAGNOSIS wilt Ca~i~iabis pathogens has elicited public outrage and-sci-
Initial symptoms (wilting) entific criticism (McPartland & West 1999).
. . - resemble wilts caused by
nematodes and some soil insects (e.g.,root maggots, white
root grubs). F,solnni can be distinguished from F. oxysporuni SYMPTOMS
(a related pathogen described next) b y its long Small, dark, irregular spots initially appear on lower
il~icroconidiophoresand stouter, thicker macroconidia (Fig leaves. Affected leaves suddenly become chlorotic. Wilt
5.5).Grey mould, hemp canker, and southern blight produce symptoms begin with an upward curling of leaf tips. Wilted
sclerotia on stalks. No red staining is present in root rots leaves dry to a yellow-tan colour and hang on plants with-
caused by other fungi. out falling off (Plate 60). Stems also turn yellow-tan. Cutting
into wilted stems reveals a reddish-brown discolouration of
CONTROL xylem tissue. Pulled-up roots show no external symptoms.
Follow recommendations in the previous section. Barloy & Pelhate (1962) described the fungus wilting whole
Fusarium root rot usually appears after abundant rainfall in plants. Noviello & Snyder (1962) illustrated plants with only
fields with heavy soils. Recognize disease symptoms and one side wilting. Dead plants may become partially envel-
oped in a white-pink mycelium. Surviving plants are stunted.
rogue affected plants as soon as possible. Eliminate
synergistic organisms such as nematodes and broomrape.
Pseudo~izoiznsstutzevi has controlled F. solani root rot
CAUSAL ORGANISM(S) &TAXONOMY
1. Ftlsarit~moxysportun Schlechtendal~l:Friesf.sp. vnsinfecturn
in kidney beans (Plzaseolils z~lllgaris).This bacteria is related
(Atkinson) Snyder & Hansen, Am. J. Bot. 27:66,1940.
(described under damping off).
to Pse~~~ioit~oizasf2l~oresceils
=Fusavitim oasinfectllliz Atkinson 1892.
110 Hemp Disea.ses and Pests

2. Fusarium oxysporum Sch1echtendahl:Fries f.sp. cannabis DIFFERENTIAL DIAGNOSIS

Noviello & Snyder, Phytopathology 52:1315-1317,1962. F. solani (the cause of Fusarium foot rot and root rot)
Description: The two subspecies are morphologically identi- causes a reddish-brown discolouration in roots; F. oxyspo-
cal-mycelium in culture usually floccose or felty, white to pink to rum does not (Barloy & Pelhate 1962). Microscopically, F.
purple, growing abundantly. Conidiogenous cells hyaline, short, oxysporum is distinguished from F. solani by its thin macro-
barrel-shaped, phialidic, in tufts of one to four atop metulae, 10-12
pm long. Macroconidia hyaline, three to five septa, sickle-shaped, conidia and short microconidial conidiophores (Fig 5.5). F.
ends curved inward with a hooked apex and pedicellate base, as oxysporum sometimes produces only microconidia (Gams
large as 45-55 x 3.54.5 km (Figs 3.2,5.5& 5.13).Microconidia hya- 1982). Microconidia might be confused with those of
line, aseptate (rarely one septum),oval to cylindrical, 5-16 x 2.2-3.4 Acremonium (=Cephalosporil~m), which also attack Cannabis
pm Chlamydosporeshyaline, thick walled, with a rough or smooth (Fuller & Norman 1944, Babu et al. 1977, Gzebenyuk 1984).
surface, spherical, borne singularly or in pairs, formed atop Fusaria differ from Acremonillm species by faster growth,
conldiophores or intercalary within hyphae or macroconidia, 7-13 wider hyphae, and larger microconidia (Gams 1982).
km in diameter.
Symptoms of Fusarium wilt are similar to those of
Verticillium wilt, Texas root rot, southern blight, Rhizoctonia
sore shin, early symptoms of charcoal rot, some nematode
diseases, and injury caused by soil insects (e.g.,root maggots,
white root grubs). Finding a reddish discolouration of xylem
is helpful, but observing the fungus with a microscope may
be needed for positive identification.

DISEASE CYCLE & EPIDEMIOLOGY

F. oxysporum overwinters as chlamydospores in soil or
crop debris. In the spring, chlamydospores produce hyphae
which directly penetrate roots of seedlings. In older "root-
hardened" plants, the hyphae must enter via wounds. Thus
disease worsens in fields harbouring root-wounding broom-
rape or nematodes (particularly Meloidogyne incognita).
Clarke (1993field notes, Kompolt, Hungary) reported a wet-
dry-wet summer predisposed plants to a Fusaritlm epidemic.
Lack of rain caused heavy clay soil to cake and crack, which
wounded plant roots, allowing Ft~sarit~m to invade. Clarke
(1993 field notes, Kompolt, Hungary) found plants infected
with Hemp streak virus exhibited resistance to Fusarit~mwilt.
After hyphae penetrate roots, F. oxysporum invades
water-conducting xylem tissues. Microconidia arise in these
vessels and flow upstream to establish a systemic infection.
The fungus eventually plugs xylem vessels, interrupts wa-
ter flow, and plants wilt. Wilt symptoms may appear within
two weeks of fungal inoculation (Semenchenko et al. 1995).
Fusarium wilt is a warm-weather disease. Optimal tem-
perature for fungal growth is 26°C (Noviello & Snyder 1962).
Figure 5.13: Macroconidia, microconidia, and chlamy-
Disease symptoms may not become evident until the ad-
dospores of Fusarium oxysporum f.sp. cannabis (LM x950,
vent of hot summer temperatures.
McPartland).
There is no windborne transmission. Conidia arising on
dead plants may be rain-splashed onto neighbours.
In 1962 Noviello & Snyder published a new form-spe- Chlamydospores also arise on dead plants. F. oxysporum in-
cies, F. oxysporum f.sp. cannabis. They justified this new name vades Cannabis seeds. Seedborne infections lay dormant until
saying, "...the wilt disease and its pathogen have not been seedlings sprout the following spring (Pietkiewicz 1958,
described." They apparently overlooked many descriptions Zelenay 1960). Mycoherbicide researchers discovered that
of F. oxysporum f.sp. vasinfectum wilting Cannabis (e.g., spore-coated Cannabis seeds effectively spread F. oxysporum
Dobrozrakova et al. 1956, Rataj 1957, Ceapoiu 1958, through the soil (Tiourebaev ef al. 1997).
Czyzewska & Zarzycka 1961, Serzane 1962).
Armstrong & Armstrong (1975) recognized six races of CULTURAL & MECHANICAL CONTROL (see Chapter 9)
F. oxysporum f.sp.vasinfectt~m.These races attack cotton, mung Methods 1 (sanitation) and 11 (avoid seedborne infec-
beans, pigeon peas, rubber trees, alfalfa, soyabeans, coffee, tion) provide a cornerstone for wilt control. Methods 2b&c
tobacco, and many other plants. The tobacco wilt fungus was (sterilizing or pasteurizing soil) often work. Flooding soil
originally called F. oxysporum f.sp. nicotianae, but Armstrong (method 2d) kills strongly aerobic Fl~saril~m species while
& Armstrong (1975) proved it to be a race of vasinfectt~m maintaining a beneficial bacterial population. Method 5 of-
(Lucas 1975). Similar experiments may prove "f.sp. canna- fers control-the Romanian hemp cultivar 'Fibramulta 151'
bis" to be another race of vasinfectum. is resistant to Fusarium wilt while the Italian hemp cultivar
Mycoherbicide researchers consistently misspell the 'Super Elite' is susceptible (Noviello et al. 1990). Generally,
form-species as "f.sp. cannabina" (Tiourebaev ef al. 1997, fibre plants have more resistance than drug plants (McCain
Tiourebaev ef al. 1998).Ceapoiu (1958) cited Neocosrnospora & Noviello 1985).Goebel & Vaissayre (1986) devised a rapid
vasinfecta Smith 1897 on Romanian hemp. This ascomycete method for screening cultivars for resistance to Fusarium
was erroneously described as the sexual stage of F. vasinfec- wilt in the field.
tum. Ceapoiu's citation of N. vasinfecta on Cannabis is prob- Noviello ef al. (1990) claimed different soil types affect
ably due to this enduring misconception (McPartland 1995a). plant resistance, but did not elaborate. Barloy & Pelhate
 Chapter 5: Fungal Diseases 111

(1962)reported epidemics in sandy, alluvial soils. Soils defi- SYMPTOMS

cient in calcium and potash predispose plants to wilt dis- Early signs of infection include raised humps or blisters
ease and must be corrected (see Elmer's comments regard- on upper leaf surfaces. From these areas the powdery myc-
ing optimal K supplements in the section on Fusarium stem elium arises. Mildew may remain isolated in irregular pus-
canker). Excess nitrogen increases wilt disease. Zhalnina tules or coalesce over the entire leaf. Leaves soon look like
(1969) reported that "acid fertilizers" increased wilt (he cor- they were dusted with flour or lime (Plate 61). Infected plants
rected this by adding lime). Keep soil p H near neutral. Mix remain alive, or they prematurely yellow, brown, and die. If
organic material and green manure into soil to encourage the the disease is permitted to run its course, black specks (fun-
growth of natural Fzlsarizlm antagonists (Windels 1997). gal cleistothecia) arise in the powdery mycelium.
Continuous cropping of Cannabis causes a buildup of
inoculum and the creation of "wilt-sick soil" (Czyzewska & CAUSAL ORGANISMS & TAXONOMY
Zarzycka 1961). Armstrong & Armstrong (1975) found F. 1. Sphaerotheca macularis (Wal1roth:Fries) Lind, Danish Fungi
oxysporum f.sp. vasinfecttlm growing i n roots of many p. 160,1913; =E ysiphe macularis (Wallroth)Fries 1824; =Sphaerotheca
symptomless plants. These unidentifiable hosts make crop humuli (DeCandolle)Burrill 1887.
rotation difficult. Czyzewska & Zarzycka (1961) suggested anamorph: Oidium sp.; =Acrosporiurn sp.
Description: Superficial hyphae flexous, branched, with
rotating hemp after wheat. Ceapoiu (1958) laid soil fallow
inconspicuousappressoria, cell diameters4-7 pm, lengths averaging
for five years to eliminate fusaria. (37-) 64.5 (-80)vm. Conidiophores upright, simple, hyaline, 50-100
Lucas (1975) cited interesting atmospheric studies: as pm tall. Conidia produced in chains, hyaline (turning brown with
carbon dioxide increases, F. oxysportim replication decreases. age), containing fibrosin bodies (which disappear with age), ovate
Supplementing a glasshouse with 20% C 0 2 cut F. oxysporzlm to barrel shaped, single-celled, averaging 30.2 x 14.0 pm (Figs 3.3 &
growth rate and spore production by 50%. 5.14). Cleistothecia globose, black, smooth, 60-125 pm in diametre,
with few to many hyphal appendages (Fig 3.3). Appendages hya-
BIOCONTROL line to brown, unbranched, flexous, tapering at ends, 300-500 Fm in
Czyzewska & Zarzycka (1961) mixed Trichoderma length. Asci one per cleistothecium, subglobose to broadly ellipti-
cal, indistinctly stalked, eight-spored, 50-90 x 45-75 p.Ascospores
lignorum into soil to protect their hemp crop. Commercially ellipsoidal to oval, hyaline, unicellular, 18-25 x 12-18 pm.
available biocontrols include Blirkholderia cepacia and Strep- S. mactllaris lives worldwide and commonly parasitizes
tomyces griseoviridis (described under damping off), Gliocla-
hops (Miller et al. 1960). Only the anamorph Oidizrn~state
dilirn species (see Rhizoctonia sore shin), Trichoderma har-
(subgenus Fibroidium) has been found on drug plants in the
zianzim (see Rhizoctonia sore shin), and Glomtls intraradices
USA (McPartland 1983a, McPartland & Cubeta 1997), on
(see Fusarium stem canker).The latter two work well in com-
Cannabis in South Africa (Doidge et al. 1953), and on fibre
bination against F. oxysporlim (Datnoff et al. 1995). A
varieties in Russia and Italy (Hirata 1966).
hypovirulent strain of F. oxysporum is described below.

Fusarium oxysporum (nonpathogenic)

BIOLOGY: A nonvirulent strain (Biofox C@,Fusacleano)
of the wilt fungus. It prevents virulent pathogens from at-
tacking roots. The nonpathogenic strain colonizes roots of
plants and lives as a saprophyte. It has been used on tomato,
basil, carnation, and cyclamen crops.
APPLICATION: Supplied as spores in a liquid or micro-
granule formulation. It can be stored for months in a cool
(8-10°C), dark place. F. oxysporlim can be applied as a seed
treatment, mixed with potting soil, incorporated into drip
irrigation, or applied as a soil drench.

CHEMICAL CONTROL
Disinfect seeds with fungicides; see the section on
Fusarium canker. Spraying wilted plants with fungicides is
not useful. Some farmers fumigate soil with nematicides to
reduce root wounding in Fzlsaritln?-infestedfields.

Figure 5.14: Conidia and conidiophores of

POWDERY MILDEW Sphaerotheca macularis. A. LM x390; B. SEM x390;
C. Drawing of conidiophore being parasitized by
Powdery mildew is known as "mould" in England and
Ampelomyces quisqualis (A & B by McPartland, C by
"Echter Mehltau" (true mildew) in Germany. This
DeBary 1887).
distinguishes it from downy mildew. Powdery mildew arises
in temperate and subtropical regions. It infests outdoor hemp
(Transhel et al. 1933, Gitman & Boytchenko 1934, Gitman
2. Leveillula taurica (Leveille) Arnaud, Annales ~piph~ties
1935, Doidge et al. 1953) and indoor drug cultivars (Stevens 7:92,1919. =L. taurica f. sp. cannabis Jaczewski 1927.
1975, McPartland 1983a, McPartland & Cubeta 1997). anamorph: Oidiopsis taurica (Lkveille)Salmon 1906.
Two fungi cause powdery mildew on Cannabis. West- Description: Superficialhyphae often cover whole plants; pale buff
endorp (1854) described a third "powdery mildew," Eysiphe to white, persistent, densely compacted, tomentose. Conidiophores
communis var. urticirum. But McPartland (1995a)examined often two-celled, upright, simple or occasionally branched, hyaline,
Westendorp's original specimen. The fungus was a misiden- emerging from host stomates, up to 250 pm long. Conidia borne
tified Phoma species, mixed with mite webbing (Plate 55). singularly atop conidiophores, either cylindrical or navicular in
112 Hemp Dise ases and Pests

shape, s~ngle-celled,hyaline, 50-79 x 14-20 pm. Cleistothecia rarely toes (Falk ef 01. 1995). A . quisqualis grows ubiquitously in
formed, scattered, embedded In mycelium, globose, black, smooth, temperate regions, especially in high humidity and tempera-
135-250 pm in diameter, with numerous hyphal appendages. Ap- tures between 20-30°C.
pendages hyaline to light brown, ind~stinctlybranched, less than APPEARANCE & DEVELOPMENT: In optimal conditions,
100 in length. Asci usually 20 per cleistothecium,ovate, distinctly
stalked, t.ivo-spored,70-110 x 2540 Fm. Ascospores cylindrical to parasitized mildew colonies become flattened and dull grey
p!.r~form,sometimes slightly curved, 2440 x 12-22 ym. within a week of infection. A. qzlisqllnlis produces pycnidia
L. taurica is distributed ~ r o r l d ~ v i dand
e parasitizes a wide which ooze tiny cylindrical conidia (Fig 5.14).
range of hosts, including C a r l l ~ a b ~ins southern France (Hirata APPLICATION: Supplied as dried, powdered conidia in
1966), eastern Europe (Jaczewski 1927, Transhel et al. 1933, extruded granules. It can be stored for months in a cool (8-
Gitman 19351, and Turkistan (Gitman & Boytchenko 1934). 1O0C),dark place. Conidia are mixed with water and sprayed
o n plants. Conidia require free water to infect powder!.
DIFFERENTIAL DIAGNOSIS mildews-this requirement is easily fulfilled in glasshouses
S . ii~nclilnrlsand L. talirica are differentiated by micro- but outdoors depends o n unpredictable dew or light rain.
scoplc examination of conidia or cleistothecia. Conidiophores The fungus can be applied with horticultural oil; it actually
of S, rilac~ilaris g r o w superficially, w h e r e a s L. t n u r i c a works better when mixed and sprayed with oil.
conidiophores emerge through stomates.
Downy mildew and pink rot may be confused with pow- CHEMICAL CONTROL (see Chapter 11)
dery mildew. Downy mildew produces a grey mycelium on The superficial nature of S,niacillaris and L. tnurica
the iiil~iersiiles of leaves. Pink rot commonly affects leaves renders them susceptible to fungicide sprays. A simple so-
and stems, whereas powdery mildew is normally restricted lution of sodium bicarbonate or potassium bicarbonate may
to 1eal.e~. Spider mite webbings have been misidentified as be sufficient. Some botanical insecticides and miticides, such
powdery mildew b y amateurs and experts (e.g., Westen- as Neemguardg and Cinnamites, also kill powdery mildews.
dorp). Look for mites and eggs. Horticultural oil (Sunsprays) works well, especially ~ 7 l l e n
mixed with baking soda. Sulphur kills powdery mildew in
DISEASE CYCLE & EPIDEMIOLOGY many crops, applied at two week intervals. Copper also
The fungi overwinter as dormant mycelia or cleistothe- works, but works better against downy mildews. Physcion
cia in plant debris. Young plants become infected in early extract (Milsanas), a new botanical, protects plants by in-
spring but may take weeks to show symptoms. Disease gen- ducing plant resistance; technically it is not a fungicide.
erally increases as rainfall decreases. Lolv light intensity (in-
doors) or shaded areas (outdoors) increase disease severity,
as does poor air circulation. S. n~aclllarisand L. taurica pro-
duce copious amounts of conidia, which spread by the slight-
CHARCOAL ROT
est breeze to sites of secondary infection. Losses multiply as Charcoal rot kills plants approaching matur~ty.Some
plants approach maturity. Succulent plants treated with ex- researchers call the disease "Premature wilt." The causal
cess nitrogen suffer the greatest damage. f u n g u s also kills y o u n g seedlings (see the sectlon on
L , tnlirica adapts well to xerophytic conditions; conidia Damping Off disease). Charcoal rot has been reported on
can germinate in 0% RH. Films of water actually inhibit L. fibre varieties in Italy (Goidanich 1955, DeCorato 1997),
t a ~ ~ r i cgermination.
~? Its optimal growth occurs at 25°C. In Cyprus (Georghiou & Papadopoulos 1957), Illinois (Tehon
contrast, S. tiinc~ilarisconidia germinate best at 100% RH (al- & Boewe 1939, Boewe 19631, and Y~igoslavia(Acinovic 1964).
tI.loug1-1 they tolerate RI-I d o w n to 10-30%), with optimal Charcoal rot also attacks drug cultivars (McPartland 1983a).
g r o ~ v t hat 15520°C The fungus is widespread and causes disease in more than
300 plant species. Maize is a particularly susceptible host.
CULTURAL & MECHANICAL CONTROL (see Chapter 9)
Strictly observe methods 7a (drought stress increases dis- SYMPTOMS
ease) and 7c (avoid overcrowding). Overhead irrigation may Plants develop a systemic chlorosis, then rapidly wilt,
~ n h i b i tL , taurica b u t enhances S . mnciilnris. Methods 8 necrose, and die (Fig 5.30). This can happen very quickly.
(optimizing nutrition) and 10 (careful pruning) are also very The pith inside the stalk becomes peppered with small black
important. Method 5 (genetic resistance) has controlled sclerotia (Figs 5.5 & 5.15).
mildexv In hops and other crops. Yarlvood (1973) suggested
blast~ngplants with a cold jet of water to knock back the CAUSAL ORGANISM & TAXONOMY
mycelium of powdery mildew fungi, but this may predispose Macrophomina plzaseolina (Tassi) Goidanich, Anrii~li ~feiiri
flo~veringplants to grey mould. spevirnentnsioize ayrnvzn Ro1nn N . 5 l(3):449-461, 1947.
=Macropilorna phnseoiinn Tass~1901, -Tiat.osporrlin pi~nseoiitza
(Tassi)van der Aa 1981; =Macvopi~orn~iln pi~aseoli ( bl a u b la n c )
BIOCONTROL (see Chapter 10) Ashby 1927, =Mncropi~orrinphaseoii Maublanc 1905; -Rl~i~octor~ia
On other crops, powdery mildew has been controlled batnticoin (Taubenhaus) Briton-Jones 1925, -5cler~ot1urri bntilticolri
by hyperparasitic fungi such as Anzpelon~ycesqliisqllal~s(see Taubenhaus 1913.
below), a strain of Verticillillnz lecanii (see the section o n Description: Sclerotia smooth, black, hard, oval to irregular In
whiteflies), a n d a n e x p e r i m e n t a l f u n g u s , Sporothrix shape, averaging 44 x 75 pm (up to 200 pm in diameter accord~ngto
floccztlosn (Kendrick 1985).A unique strain of Bncill~issubti- DeCorato 1997). Pycnidia solitary or gregarious (when present),
11s (Serenade) is sold as a foliar spray for powdery mildew brown to black, subglobose, immersed in host tissue but becoming
erumpent, ostiolate, axreraging180 pm in diameter. Conidiophores
(for more information o n B. sl~btilis,see damping off).
short, hyaline, simple (sometimes branched accord~ngto
Punithalingam 1982).Conidiogenous cells at f~rstholoblast~c,be-
A~tzpdornycesqzi~squalls(=Clclnnobol~lsccsatu) coming phialidic, with a minute cellarette, lageniform to dol~iform.
BlOLOGY A fungus that parasitizes powdery mlldew Conidia hyaline, asepta te, obovoid to fusiform, thin walled, smooth,
fungl (AQ-102) The tiny hyperparaslte has been used agalnst often guttulate, 5-10 x 14-30 pm (Fig 5.5). Punithalingam visual-
Spl~aerotllecafuligl~zea,S n~aclilarls,L ta~rrlcn,and other pow- ized a conidial appendage after treating conidia with Leifson's flag-
dery mlldews, on cucurbits, grapes, strawberries, and toma- ella stain. This appendage is apical, cap-like, and cone-shaped.
 Chapter 5: Fungal Diseases 113

SYMPTOMS
Spots on the upperside of the leaf are brown (Plate 62).
Flip over the leaf to see the olive colour. The fuzzy olive
material is the fungus itself, growing on the leaf surface.
Whole leaves soon wilt, curl, and drop off. Damage esca-
lates rapidly in August. Surviving plants are stunted with
reduced yields.
spots caused by one fungus, Cercospora cannabis, usu-
ally remain small and circular. Rarely, spots enlarge to mar-
gins of leaf veins, causing straight edges and somewhat rec-
tangular shapes. The mycelium on the underside of the leaf
Figure 5.15: Microsclerotia of Macrophomina phaseolina may look yellow-brown rather than olive. The other fungus,
within a Cannabis stalk (LM x35, McPartland). Pse~idocercospomcannabina, produces large irregularly-shaped
spots, vein-delimited at first but eventually coalescing with
other spots. Its mycelium is dark olivaceous brown and
DIFFERENTIAL DIAGNOSIS
spreads effusely across the underside of the leaf. It occasion-
The rapid necrosis caused by charcoal rot has been mis-
ally produces conidia on upper leaf surfaces as well.
taken for herbicide destruction. Flisarilim and Verticillilim
cause wilt symptoms but do not pepper the pith with scle-
CAUSAL ORGANISMS & TAXONOMY
rotia. Sclerotia are much smaller than those produced by
1. Pseudocercospora cannabina (Wakefield)Deighton, C.M.1.
hemp canker and southern blight organisms.
Mycological Paper No. 140,1976;=Cercospora cannabina Wakefield 1917;
=Helicomina cannabis Ponnappa 1977.
DISEASE CYCLE & EPIDEMIOLOGY Description: Conidiophores simple or occasionally branched,
M . phaseolina overwinters as sclerotia in soil or plant straight, septate,olivaceous brown, narrowing to a slender apex, up
debris. In dry conditions, sclerotia survive over a decade in to 150x 3.54.5pm. Conidia with unthickened basal scars, cylindrical,
the soil. Spring rains induce sclerotia to germinate and di- slightly or strongly curved, dilutely olivacous,usually five to seven
rectly penetrate roots of young seedlings. Hyphae grow in- septa, 30-90 x 3.55.5 pm (Vasudeva 1961 reports conidial lengths
up to 120 pm). Occasionally, conidia may form microconidia.
ter- and intracellularly, primarily in the plant cortex. M.
Microconidia ovoid to fusiform, unicellular, hyaline, 2-6 x 2.3.5 pm
plzaseolina is an opportunist and-rapidly colonizes plants (Fig 5.16).
stressed by drought and high temperatures. Optimal fungal
growth occurs at 37OC.
2. Cercospora cannabis Hara & Fukui in Hara, ZituyB sakaotu-
byBrigaku [Textbook in Plant Pathology] p. 594,1925.
CULTURAL & MECHANICAL CONTROL (see Chapter 9) -Cercospora cannabis Hara & Fukui in Shirai & Hara 1927,~ C e r -
Method 7a is important-prevent drought stress by wa- cospora cannabis (Hara)Chupp apud Green 1944, =Cercosporina can-
tering accessible plants during late-summer dry spells. Ex- nabis Hara Sakumotsu byorigaku [Pathology ofcrop plants] p. 195,1928,
periments with C a n n a b i s showed that drought stress =Cercosporina cannabis Hara et Fukui apud Hara J. Agricul. Soc.
predisposed plants to charcoal rot before plants started wilting Shizuoka-ken 32(364):45,1928; =Cercospora cannabis Teng 1936 apud
(McPartland & Schoneweiss, unpublished). Watering plants Korf 1996.
after they wilt is too late. We have not seen Hara's 1925 text to verify priority. Many re-
searchers cite Hara & Fukui in Shirai & Hara 1927.Korf (1996)incor-
Use methods 1 (sanitation), 2a (deep ploughing), 2b rectly erected "Cercospora cannabis Teng 1936 [non Hara & Fukui]."
(sterilizing soil), and 8 (optimizing soil nutrition). M. Teng (1936) was clearly describing C. cannabis Hara & Fukui-he
phaseolina is too heat-tolerant for method 2c. In the corn belt, cited these authors. The synonym Cercosporina cannabis has similar
do not plant Cannabis next to maize fields or in recently problems with dates and names. Morphology of C. cannabis closely
rotated fields. These soils often harbour a large reservoir of resembles that of C. apii Fresenius; it may be a synonym.
M. phaseolina sclerotia. Ghaffar et al. (1969) reduced M.
phaseolina in cotton by adding organic amendments to soil.
They added barley straw an; aEalfa meal to increase the
number (and activity) of soil organisms antagonistic to M.
phnseolina.

BIOCONTROL & CHEMICAL CONTROL

Trichoderma l m r z i a n u m infests and kills sclerotia of M.
plzaseolina (described under Rhizoctonia sore shin). The my-
corrhizal fungus Glomlis intraradices offers some protection
(described under Fusarium stem canker disease). No fungi-
cide stops charcoal rot once it has begun.

OLIVE LEAF SPOT

Two fungi cause slightly different symptoms of olive
leaf spot. The disease has been reported in South America
(Viegas 1961), Nepal (Bush Doctor 1987a), India, Pakistan,
Bangladesh, China, Cambodia, Japan (Vasudeva 1961),
Uganda, Mississippi, Missouri, and Wisconsin (Lentz et al.
1G4).As Lentz e't hl. (1974) demonstrated, the 'two causal
fungi are often confused in these reports. Both fungi attack Figure 5.16: Conidia of Pseudocercospora cannabina
fibre and drug plants. erupting from leaf surface (SEM x375, McPartland).
114 Hemp Diseases and Pests

Description: Conidiophores simple, straight or occasionally Brown blight occurs in Europe and Asia (Vakhrusheva 1979),
gemculate, septate, pale brown, not narrowing at the apex but broad Tasmania (Lisson & Mendham 1995), and the USA (Harto-
and flat, up to 100x 3.5-5.5 pm. Conidia bear thickened detachment wicz et al. 1971, Haney & Kutsheid 1975, McPartland 1983a).
scars at the base, not cylindrical but tapering toward their tips, Cigarettes made from Alternaria-infected tobacco
straight or slightly curved, hyaline, indistinctly multiseptate,25-90
x 3-5 pm. generate a harsh and irritating smoke (Lucas 1975). More
significantly, A. alternata produces toxins that mutate human
oesophageal epithelial cells (Liu et al. 1992). Smoking A.
DIFFERENTIAL DIAGNOSIS
alternata-infected material may cause oesophageal cancer,
During its early stages, olive leaf spot may be confused
which is the most common cancer of marijuana smokers
with yellow leaf spot or brown leaf spot. It later resembles
(McPartland 1995f, 1995g).
downy mildew. The differential diagnosis is accomplished
with a microscope.
SYMPTOMS
Brown blight usually arises on mature plants late in the
DISEASE CYCLE & EPIDEMIOLOGY
growing season. Symptoms begin as pale green or grey leaf
The two fungi share similar life cycles. They overwinter
lesions. These irregularly circular spots may or may not de-
in plant debris or in soil. Other Cercospora species are seed-
borne, but this has not been detected in Cannabis (Pietkiewicz velop chlorotic halos, depending on the Alternaria species.
Spots often coalesce into blight-like symptoms, as whole
1958, Ferri 1961b, Babu et al. 1977).Conidia form in the spring
leaves turn brown. Undersides of spots may develop con-
and spread by wind, splashing water, and farm hands. Mites
can also carry conidia to sites of secondary infection. Co- centric zonations (especially spots caused by Alternaria
nidia of Cercospora and Pseudocercospora can be blown great solani). Frail necrotic tissues break up, resulting in irregular
distances, unlike Septoria conidia, which are "sticky" and leaf perforations. Lesions may extend to leaf petioles and
disseminate only several metres (Howard et al. 1994). High stems. The fungus also infests female flowers (and seeds),
humidity or free water is required for conidial germination. turning them g$ey-brown (Ferri 1961b).
Penetration occurs via stomates or wounds.
CAUSAL ORGANISMS & TAXONOMY
C. cannabis is more widespread than P. cannabina around
the globe. P. cannabina seems to be more virulent than C. 1. Alternaria alternata (Fries:Fries) Keissler, Beih. Bat. Cent.
29:434, 1912; =Alternaria tenuis C. G. Nees 1816-7; ?=Alternaria
cannabis. Whereas P. cannabina may cause complete leaf cannabis Yu 1973.
defoliation, C. cannabis only causes leaf spots. McPartland Description: Conidiophores simple or branched, straight or
(1995a) described microconidia budding from conidia of P. curved, septate, yellow to golden brown, up to 50 pm long, 3-6 pm
cannabina (Fig 5.16).This "microcyclic conidiogenesis" may thick. Conidia borne in chains,straight or slightly curved, obyriform
explain why P. cannabina is more virulent than C. cannabis- to obclavate (rarely ellipsoid),yellow to golden brown, with three
because it produces copious amounts of conidia and to eight transverse and one or two longitudinal septa, tapering to a
microconidia. short beak (beak sometimes absent), length 20-63 pm, width 7-18
pm (Figs 3.2 & 5.17).
CULTURAL & MECHANICAL CONTROL (see Chapter 9) A. alternata has been isolated from hemp cultivars in
Follow instructions for yellow leaf spot. Calpouzos & Italy (Agostini 1927, Ferri 1961b), Poland (Zarzycka &
Stallknecht (in Lucas 1975) noted that sugar beets growing Jaranowska 1977), the Czech Republic (Ondrej 1991), and
in bright sunlight suffer small Cercospora leaf spots, but in the USA in Illinois (McPartland 1983a), Iowa (Fuller &
low light intensity the spots coalesce and damage increases. Norman 1946), and Kansas (Hartowicz et al. 1971). Drug
plants are attacked in India and the USA (McPartland 1995c
BIOCONTROL & CHEMICAL CONTROL
Dicyma may provide biocontrol (see below). A unique
strain of Bacillus subtilis (Serenade@)is sold as a foliar spray
for Cercospora leaf spot (described under damping off).
Hawksworth et al. (1995) said Gonatobotrys simplex kills
Cercospora species. This biocontrol is not yet available, al-
though it grow naturally on hemp stems in the Czech Re-
public (Ondrej 1991). Yu (1973) controlled Cercospora with
sulphur dusts and Bordeaux mixture, applied every two
weeks for a total of three to five applications.

Dicyma pulvinata (=Hansfordia ptllvinata)

BIOLOGY: A fungal mycoparasite that parasitizes hyphae
and spores of Cercospora species, and also works against
Clzaetomitim species. It has been used successfully in field
experiments (Jeffries & Young 1994).

BROWN BLIGHT
Four fungi reportedly cause brown blight. The most
ubiquitous fungus, Alternaria alternata, infests plants as an
opportunistic saprophyte in necrotic tissue, and as a primary
pathogen in female flowers (McPartland 1983a). It has de-
stroyed up to 46% of seed (Haney & Kutsheid 1975). It also Figure 5.1 7: Conidia of Alternaria alternata (A, LM x660)
rots hemp stalks after harvest (Fuller & Norman 1944), and compared with a chain of chlamydospores produced by
even destroys finished hemp products (Agostini 1927). Phoma glomerata (B, LM x490), McPartland.
 Chapter 5: Fungal Diseases 115

and unpublished data). Paulsen (1971) cultured the fungus n l f e r n n t n infects plants during warm, wet weather but
from green stink bugs (Nezanz uiriiluln) feeding on marijuana symptoms do not arise until the hot, dry summer.
seeds Yu (1973)described Alter'ilaria cannnbis from marijuana Alternuria conidia disperse by wind or splashed rain. Co-
growing at the Unix~ersityof Mississippi. Her description of nidia require a film of water to germinate. Conidia germ tubes
this i z o i ~ i r di ~i l b i i ~ i nstrongly resembles A. alferliatu. penetrate host epidermis directly, or enter via stomates and
wounds. Infection increases ten-fold when Alter?iarln conidia
2. Altevnaria solani (Ellis & Martin) Sorauer, Zeitscizri'jt are mixed with plant pollen (Norse in Lucas 1975)
i~tlin~=c~ikianki~riten6:6, 1896; Apparently the pollen serves as an energy source for conidia.
aiMizcrosnoriuni solani Ellis &Martin 1882, =Altevnnuin vorvi (Ellis)
Clferri f sp. solan1 (Ellis &Martin) Neergaard 1945; =~lte~'navia'dalic; CULTURAL & MECHANICAL CONTROL (see Chapter 9)
(Kuhn)Groves & Skolko f. sp, solnni (Ellis & Martln) Neergaard 1945.
Carefully observe methods 7c (avoid excess humidity),
Description: Coiiidiophores simple, straight or curved, septate,
pale to dark ol~ve-brown,u p to 100 pm long, 6-10 pm thick. Conidia
10 (careful pruning), 11 (avoid seedborne infection), and 8
borne singularly, straight or slightly curved, obyriform to obclavate, (fertilize with high P & K and low N; Lucas (1975) used a
light olive-bro\vn, with eight to 11transverse septa and zero to two NPK ratio of 3-18-15). Other important measures include 1
long~tudlnalsepta, tapering to a long beak which is usually equal (sanitation), 2a (deep ploughing), 2b&c (sterilizing or
111 length to the bod!; olserall length 140-280 pm, 15-19 pm at greatest pasteurizing soil), and 4 (escape cropping). Sasaki & Honda
iv~dth. (1985) decreased A. solani infection in glasshouse vegetables
Onll. Barloy & Pelhate (1962)have cited A . solni~ion Can- by covering glass with UV-absorbent vinyl sheets. Since A.
i l n b ~ s and
, the fungus they illustrated looks like A, alternata solaili requires UV light to produce conidia, epidemics are
not A. solni~i.They erroneously synonymized A . solnil1 with prevented (see the section on grey mould for more
Macrosporilinl C L ~ ~ I ~ Z L ~ (McPartland
~ ~ ~ Z I ~ I ~ I 1995a). A . solnili information).
attacks solanaceous crops (tobacco, potato, tomato, etc.) and
Br'asszca olrracea (cabbage, cauliflower, kohlrabi, etc.). BIOCONTROL & CHEMICAL CONTROL
Jeffries & Young (1994) controlled A. s o l a n i with
3. Altevnavin longipes (Ellis & Everhart) Mason, Mycological A n ~ p e l o t n y c e sql~isqualis(described under powdery mildew)
P n ~ ~ e2:19,
r 1928; A unique strain of Bacilllls subtilis (Serenade~E)is sold as a
s A l t ~ i ? ~ n rIongi~cs
ia (E111s & E~rerhart)Tlsdale & Wadkiiis 1931.
foliar spray for Alternnrzn species (descr~bedunder damping
Description: Conldiophores simple or branched, straight or curved,
septate, pale olivaceous brown, u p to 80 pm long, 3-5 pni thick. off). Dipping flovvers in a solution of Picilia gi~illlerii~oizdiz
may
Conldi'i borne ill cl~ains(rarely singularly), obclavate, rostrate, reduce postharvest rot (see Chapter 8). Disinfect Alterliarla-
smooth to verruculose, gradually tapered to a beak (1/3rd its total infested seeds by soaking them in fungicides (see Chapter
length) and rounded at the base, beige to pale brown, with five or 12).Yu (1973) controlled brown blight caused by Alternarin
six transverse and one to three longitudinal or oblique septa, 35- caiz~zabis(whatever that is) by spraying crops with Bordeaux
(69)-110 x 11-(14)-21 pm. mixture every month. Tobacco growers also use Bordeaux
A. longipes normally attacks tobacco; Lentz (1977) re- mixture (Lucas 1975). For farmers interested in alternative
ported it on Caizrrnbis. ~ o l y(1964) lumped A. loilgipes with A. pesticides, Dahiya & Jain (1977)reported that pure THC and
soinill; Lucas (1975) lumped it with A. alternntn. CBD inhibited A . nlter?znta.

4 . Alteniaria cheirlzizthi (Libert) Bolle [as "(Fr.) Bolle"], Meded.

~1/1!1tv~~nt11Lnb. 1Ylliie Contti~eiinSciloiteil. 7:55, 1924.
s H c l ~ r i ~ ~ l t i i o s ~ ~coIr~ie~~~~r.in~i iLibert
t i ~ i 1827, =Mncuosporiur~i STEMPHYLIUM LEAF
ili~ii.nrltl~i
(L~bert)Fries 1832.
Description: Conidiophores s ~ m p l eor branched, straight or
& STEM SPOT
cur\ ed, septate, pale olive to hyaline at the apex, up to 130 b ~ long,
n~ Spaar ct al. (1990)called this "brown fleck disease." Leaf
5-8 prn thick. Conidia borne siiigularly (rarely in chains of two or lesions arise as light-brown spots with dark margins, often
three), ovoid to pyriform at first, later becoming irregulal; gener- irregularly shaped and limited by leaf veins. Spots grow to
all>-tapered at the apex and rouiided at the base, light olive-golden 3-10 mm in diameter, become concentrically zonated, and
broxvn, \\rith numerous transverse and longitudinal septa, 20-100 x
coalesce. Disease may spread to leaf petioles and stems
13-33 urn.
(Gzebenyuk 1984).Conidia (rarely perithecia and ascospores)
Gzebenyuk (1984) reported isolating A,cheirantiri from
of the causal fungi arise in necrotic plant tissue.
96 4'0 of hemp stems sampled near Kiev He may have
~ n ~ s i d e n t ~Af ~alteirzatn
ed (VcPartland 1995a)
CAUSAL ORGANISMS & TAXONOMY
The literature is confused and confusing. Researchers
DIFFERENTIAL DIAGNOSIS
cite two taxa on Cntlriabis: Steinphyliunz botryosl~inand Ple-
Several microscopic characteristics separate A . solani
osporn herbnrl~nz.These names are frequently misinterpreted
trom A . nlteriiata: the former exhibits larger, darker conidia
as two stages in the life cycle of one organism. Careful re-
with longer beaks. A . solalzi produces solitary conidia
search refutes this, they are different species (Simmons 1985).
ivhereas A . nlternnta often sporulates in chains.
Brolvn blight can be confused with grey mould caused
1. Stemplzylitrm botryoszlm Wallroth, Flova Crypt. gcunl., pars
b!. B o t y t i s cincr'cn. A . altrrrintn conidia can be confused with
post., p. 300, 1833; =Stemphylil~rncarlnnbintlnl (Bakhtii~& Gutner)
chlamydospores of Plroriia gloii-ier'atn, one of the causes of Dobrozrakova et al. 1956, -Mncrosporitii~l cannabin~~rii Bakhtiii &
b r u ~ v nleaf spot (Fig 3.17). Gutlier 1933; =Tilyros{lora c(rnnnbls Ishiyania 1936.
teleomorph: 1'1~ospoi.ntnriia Simmons, Sylioicin 38:291, 1985.
DISEASE CYCLE & EPIDEMIOLOGY Description: Conidiophores arise in clumps (often ten to 15 per
Alternarin species overwinter as mycelia in host plant clump), straight or flexous, simple or occasionally branched at tlie~r
debris. A. alfer'nntn can survive indefinitely in the soil. A. al- base, one to seven septa, cylindrical except for swelling at the site ot
tcrilnta also overwinters as a seedborne infection (Pietkiewicz spore formation, light olive brown darkening at s~volienapes,
smooth becoming echinulate at sbvollen apex, 4-6 pm wide, swelling
1958, Ferri 1961b, Stepanova 1975). The optimal tempera-
to 7-10 pm at apex, usually 25 to 85 pm long, but continued
ture for A. solani, A . alterliatn, and A. longiyes is 28-30"C.A. proliferation tl~roughone to five previously-formed apices may
116 Hemp Diseases and Pests

extend conidiophore length to 120 pm. Conidia initially oblong to DISEASE CYCLE & EPIDEMIOLOGY
subspherical, hyaline and verruculose; at maturity oblong to S. botyosurn lives i n temperate and subtropical regions
subdoliiform, densely echinulate (Fig 5.18), with three (sometimes worldwide. It attacks many hosts. The fungus overwinters
four) transverse septa and two or three (sometimes four) complete in crop debris. Springtime infections begin via air-dispersed
or nearly complete longitudinal septa, with a single conspicuous ascospores and conidia. During the growing season copi-
constriction at the median transverse septum, dilute to deep olive ous amounts of conidia form o n diseased leaves. The buildup
brown, 30-35 x 20-25 pm. Pseudothecia large (700 pm in diameter),
hard, black scleroticbodies, with formation of asci occurring slowly, of conidia leads to epidemics of secondary infections. Ganter
often after eight months in culture. Asci subcylindrical, eight-spored, (1925) reported transmission of a Pleospora species b y infected
averaging 200 x 40 pm. Ascospores hyaline, oblong and constricted seed. Presley (unpublished data, USDA) also described
at three transverse septa when immature; when mature they exhibit "Tetracoccospori~~m species" infecting Cannabis seed.
a broadly rounded apexand almost flat base, turn yellowish-brown,
develop four seconda~ytransverse septa in addition to the three primary CONTROL
septa, form one or two longitudinal septa, and average 40 x 17pm. Follow suggestions for brown blight described above.
McPartland (1995e) synonymized Stemphylittm cannabi- Prevention is the key to control.
n u m , Macrosporium cannabint~ma n d Thyrospora
- . cannabis un-
der S. bot yosllm after examining leaf specimens from Russia
and Japan. A misidentified "Coniothecilim species" from Lima,
Peru, also turned out to be S. b o t y o s u m (McPartland 1995a).
Presley (unpublished 1955 data, USDA archives) described
a "Tetracoccospori~~m species" o n stems of hemp growing in
Beltsville, M D . N o s p e c i m e n s u r v i v e s , b u t Presley's
description suggests S. b o t y o s u m .
McCurry & Hicks (1925) reported a Stemphylium species
causing "considerable amounts" of leaf disease in Canadian
hemp o n Prince Edward Island. S. bot y o s l l m colonizes hemp
stems in Holland (Termorshuizen 1991), a n d the Czech
Republic (Ondrej 1991). Rataj (1957) considered S. botryosllm
a dangerous Cannabis pathogen. Conversely, Barloy & Pelhate
(1962) considered the fungus a weak pathogen, only able to
invade plants previously weakened b y Alternaria species.

2. Stemphylium herbarurn Simmons, Sydowia 38:291,1985.

teleo&orph: Pleospora herbarum (Persoon:Fries) Rabenhorst,
Herb. viv.mycol. Ed. 11, No. 547, 1856.
Description: Conidiophores straight or flexous, simple or
branching either at their base or subapically, one to four septa,
cylindrical except for apical swelling, dilute brown, smooth, 3-5 pm
wide swelling to 8-10 pm at apex, usually 15-20 pm long but
occasionally much longer. Conidia initially oblong to broadly ovoid
to subspherical, hyaline and minutely verruculose; at maturity
becoming broadly bvoid to broadly elliptical, sometimes unequally
sided, distinctly verruculose, with up to six or seven transverse septa
and two or three longitudinal septa, distinctly constricted at one to
three transverse septa, dilutely yellowish brown to deep reddish
brown, 3545 x 20-27 pm. Pseudothecia immersed then erumpent Figure 5.18: Conidia and conidiophores of Stemphylium
in host tissue (in culture, forming asci within a month), black, globose botryosum on Cannabis (SEM XI 000, McPartland).
or slightly flattened due to collapse of their thin walls, 250-300 pm
in diameter. Asci conspicuously bitunicate, tubular with parallel
walls and a broadly rounded apex, gradually narrowing near the
base, eight-spored, averaging 160 x 25 km. Ascospores hyaline, el-
lipsoid to obovoid and constricted at one transverse submedian
SOUTHERN BLIGHT
septum when immature; when mature they become medium brown Southern blight is sometimes called southern stem and
in colour, obovoid, often flattened on one side, with the upper half root rot. The causal fungus attacks many plants, from corn to
broader than the lower half, usually exhbiting seven transverse and one catnip. Crops in the tropics a n d warmer temperate regions
or two (rarely three) lol~gitudinalsepta, averaging 32-35 x 12-15 p. suffer extensive losses from southern blight, which has been
In the Ukraine, Gzebenyuk (1984) found P. herbarum o n reported o n Cannabis in India (Hector 1931, Uppal 1933,
2% of hemp stems, and S. botryosltm o n 7% of stems. In the Krishna 1995), Texas a n d South Carolina (Miller et al. 1960),
Czech Republic, Ondrej (1991) reported P. herbarum o n some Italy (Ferri 1961a), Japan (Kyokai 1965, Kishi 1988), and the
stems, whereas S. botyosllm appeared "abundantly." Ukraine (Gzebenyuk 1984). Recently the fungus was isolated
i n Oxford, Mississippi (Sands 1991); virulent mutants of this
DIFFERENTIAL DIAGNOSIS isolate yielded a 100% kill rate (Sands 1995).
Distinguishing the Stemphylium stages requires careful
measurements and a practiced eye. Generally, S. b o t y o s t ~ m SYMPTOMS
produces longer conidiophores with a series of knobby api- Disease arises during w a r m summer weather. Mature
cal proliferations; S. herbarum produces larger conidia with plants suddenly wilt a n d turn yellow. Systemic wilting and
a greater size range and greater complexity of septation. The leaf chlorosis progress to leaf necrosis a n d plant death (Plate
teleomorphs, P. herbartun and P. tarda, can best b e differenti- 63). Stalks decay at the soil line, turning brown and macer-
ated in culture. Stemphylium leaf symptoms can b e confused ated. Sunken brown stalk lesions sprout sclerotia after death
with brown blight o r early olive leaf spot. The stem symp- of the plant. A pale-brown hyphal mat sometimes radiates
toms can b e confused with Cladosporium stem canker. from the base of the stalk along the soil surface.
 Chapter 5: Fungal Diseases 117

CAUSAL ORGANISM 30°C, when Cannabis losses may reach 60% (Krishna 1995).
Sclerotium mlfsii Saccardo, Annales Mycologici 9:257,1911. Basidiospores germinate best at 28"C, but are short lived and
teleomorph:Athelia rolfsii (Curzi)Tu & Kimbrough 1978,=Cur- relatively unimportant. A root-boring maggot, Tetanops
ticium rolfsii Curzi 1932. luridipenis, may vector S. rolfsii between Cannabis plants
Description: Sclerotia smooth or pitted, near spherical (slightly (Sands et al. 1987).
flattened below), at first amber then turning brown to black, usually
0.7-lmm in diameter(Fig5.19);cross-sectionof sclerotia reveals three
CULTURAL & MECHANICAL CONTROL (see Chapter 9)
cell layers-a darkly pigmented outer rind (three or four cells thick),
a middle cortex composed of dense, hyaline cells (four to ten cells
Apply methods 1 (sanitation), 2a (ploughing at least 8
thick), and an inner medulla, comprised of loosely interwoven cells. cm), and 2b (soil sterilization).Method 2c-soil solarization-
Hyphae produce clamp connection at some septa, and branch at kills S. rolfsiisclerotia (Elmore et al. 1997).Decreasing moisture
acute angles. Basidiocarps resupinate, effused, loosely adherent to at ground level inhibits disease development, so observe
the stem surface.Basidia clavate, usually four-spored.Basidiospores methods 3 (weeding), 7b (avoid overwatering), and 7c (avoid
hyalme, smooth, teardrop-shaped, 5.5-6.5 x 3.54.5 pm (Fig 3.2). excess humidity). All weeds should be viewed as alternate
hosts-method 3 again. Method 8 (balanced soil structure)
is tricky, because too high a percentage of soil organic matter
may increase the severity of southern blight. Cyclical drying
and wetting of soil induces sclerotia to germinate.
Germinated sclerotia die within two weeks in fallow soil
(Lucas 1975). Krishna (1996) tested three ganja "cultivars"
from Uttar Pradesh (India) for genetic resistance to S. rolfsii.
Cultivar 'dwarf' [Cannabis afghanica?] showed the most
resistance, followed by cultivar 'medium' [Cannabis indica?],
and lastly cultivar 'tall' [Cannabis sativa?].

BIOCONTROL & CHEMICAL CONTROL

Kendrick (1985) reported biocontrol with the fungi Tri-
choderma harziantim and Trichoderma (Gliocladillm) virens, de-
scribed under Rhizoctonia sore shin. The mycorrhizal fun-
Figure 5.19: Sclerotia produced by Sclerotium rolfsii gus Glomt~sintraradices offers some protection (described
(LM x20, courtesy Bud Uecker). under Fusarium stem canker disease). Mixing urea into soil
at a rate of 100 kg ha-1 kills many sclerotia (Krishna 1995).
Drenching soil with formalin reduced hemp losses in Italy
DIFFERENTIAL DIAGNOSIS (Ferri l96la).
Symptoms of southern blight may be confused with
symptoms of hemp canker (caused by Sclerotinia sclerotio-
rum), charcoal rot (Macrophomina phaseolina), or Rhizoctonia
sore shin (Rhizoctonia solani). Sclerotia of S. sclerotiortim and BLACK MILDEW
R. solani are larger and less symmetrical than those of S. rolf- Black mildews, like powdery mildews, are obligate para-
sii, and the sclerotia of M. plzaseolina are smaller and irregu- sites that grow on the surfaces of leaves. They tap host re-
lar in shape. Stem cankers caused by Ftisarilim species may sources by immersing haustoria into host leaves. They should
be confused with southern blight, but sclerotia are missing. not be confused with sooty moulds, which are entirely su-
perficial and saprophytic. See "Phylloplane fungi" for a
DISEASE CYCLE & EPIDEMIOLOGY discussion of sooty moulds. Black mildew is known only
S. rolfsii overwinters as sclerotia in soil and plant de- from Nepal.
bris. Sclerotia are spread by wind, water, contaminated farm
tools, and some survive passage through sheep and cattle. SYMPTOMS
Sclerotia germinate in the spring. Germination decreases in Black mildew forms thin grey to black colonies on oth-
deep soil. Germ tubes directly penetrate roots or enter via erwise healthy leaves. Colonies arise on upper sides of leaves.
wounds. The fungus grows inter- and intracellularly, pri- They remain isolated or coalesce with neighbours to form
marily within the stem cortex. Optimal growth occurs near spots 5 mm or more in diameter.

Figure 5.20: Black mildew caused by Schiffnerula cannabis. A. Infected leaf (LM x2); 8.Leaf surface crowded with conidia
(LM x70); C. Close-up of a conidium, an ascus with 2 ascospores in focus, and 2 hyphopodia (LM ~ 1 4 5 0 )McPartland.
;
118 Hemp Dise,ases and Pests

CAUSAL ORGANISM & TAXONOMY

Schiffnerula cannabis McPartland & Hughes, Mycologia 86:868,
1995. anamorph: Sarcinella sp.
Description: Mycelium composed of brown hyphae, straight
or sinuate, septate (cells mostly about 16 Fm long), 4-5 pm thick,
branching irregular. Hyphopodia numerous, alternate, hemispheri-
cal or subglobose, brown, 9.5 x 6.5 pm. Ascomata circular in outline,
3743 Frn diameter, containing one to three asci, wall at maturity
mucilaginous and breaking up in water. Asci ellipsoid to subglobose,
nonparaphysate, eight-spored, 23 x 21 p m Ascospores massed into
a ball, at first hyaline, later turning light brown, oblong, smooth,
one septum, scarcely constricted, the cells unequal, 17.4 x 7.3 pm.
Conidiophores indistinguishable from hyphae. Conidiogenous cells
intercalary, short, cylindrical, integrated, monoblastic. Conidia
opaquely black-brown, ellipsoid to subglobose, dictyoseptate,
composed of eight to 15 cells, bullate but smooth walled, 31.9 x 27.5
pm (Fig 5.20 & Plate 64).

DIFFERENTIAL DIAGNOSIS
Black mildew m a y b e confused w i t h early stages of
brown blight, corky leaf spot, black dot, o r sooty mould.

DISEASE CYCLE & EPIDEMIOLOGY

S . cannabis produces copious Sarcinella conidia but none
have been observed to germinate. Their role in the life cycle
is uncertain. The fungus spreads b y water-transported as-
cospores. Schiffnerulaceous fungi occur worldwide but are
more abundant in tropical and subtropical climates.

CONTROL
Measures have not been elucidated. Since most Schifl-
nertlla species have limited host ranges, crop rotation should
effectively control the disease.
Figure 5.21: Symptoms of twig blight (from Charles &
Jenkins 1914).
TWIG BLIGHT
Known as nebbia in Italy, twig blight has been reported But Charles & Jenkins were wrong-their species w a s not
in Europe, Asia, North America, a n d South America. Dewey the same as Cavara's fungus. So w e have two different spe-
(1914) and Barloy & Pelhate (1962) considered the disease of cies with the same marconii epithet.
no economic significance, b u t others consider it a severe
problem (Ferraris 1935, Trunoff 1936, Ghillini 1951). Charles 1. Dendrophoma marconii Cavara, Atti dell'lstit. Bot. di Pavia,
& Jenkins (1914) reported losses as high a s 95% in Virginia. ser. 11, 1:426; Revue Mycologique 10(40):205,both 1888.
Losses multiply i n drought-stressed crops-McPartland & Description: Pycnidia few, arising in grey spots, concealed by
Schoeneweiss (1984) s h o w e d h o w Botryosphaeria species epidermis, flattened globose, ostiole slightly raised, 130-150 pm
diameter. Conidiophores unbranched or widely dichotomous
opportunistically attack stressed plants. Secondary infections branching, septate, hyaline. Conidia pleomorphic, ovate-elliptical,
often follow twig blight; a Ftisariz~mfollowed twig blight in teretiuscullis [tapering?], one-celled, hyaline, 4.5-6.5 x 2-2.5 pm.
Italy (Petri 1942)' a n d Alternaria alternata followed twig blight The description above is from Cavara (1888).Barloy & Pelhate
in Illinois (McPartland, unpublished data). (1962) cited smaller D. marconii conidia, 3 x 2 pm. McPartland (1999b)
described conidiophores as multiseptate, widely branched, with
SYMPTOMS irregular monilioid swellings, growing up to 25 pm long and
Twig blight begins with a wilting and drooping of leaves, ramifying throughout the pycnidial locule. Conidiogenesis could
usually late in the season (Fig 5.21). Wilted foliage turns not be determined. Conidia arise at restrictions along the
conidiophores. Conidia are quite pleomorphic and irregular in
brown and dies but remains attached to plants. Tips of young shape, often resembling short sections of conidiophores.
branches show symptoms first (thus "twig blight"). Within
Every D. marconii specimen we've examined has been
two weeks the entire plant may wilt and die. Diseased stems
concurrently infected b y other fungi, such as Pkomopsis
develop grey spots averaging 6-12 m m long and 2-6 m m
ganjae, Septoria species, Phoma species, and Botryosphaeria
wide (Flachs 1936). These spots darken and bear tiny, black
marconii. N o o n e h a s cultured D. marconii in isolation.
pycnidia a n d / o r pseudothecia. Spots continue to enlarge
Conceivably D. marconii could b e a hyperparasite of other
after plants are harvested (Gitman & Malikova 1933).
fungi, and not a plant pathogen at all. This may explain the
confusion this species caused Charles &Jenkins (1914) and
CAUSAL ORGANISMS & TAXONOMY
Petrak (1921).
At least t w o species cause twig blight (McPartland
199510, 1999b). The first species w a s named Dendrophoma
rnarconii in 1888 b y Cavara, w h o discovered it i n Italy. The 2. Botyosphaeria marconii Charles & Jenkins, I. Agric. Re-
second species w a s discovered b y Charles &Jenkins (1914), search 3:83,1914 [as Botyosphaeria marconii (Cav.)Charles & Jenkins].
anamorph 1: Leptodothiorella marconii McPartland, Mycotaxon
w h o thought it w a s identical to Dendrophoma marconii. When
53:422, 1995.
Charles & Jenkins's fungus produced a sexual stage, they anamorph 2: Fusicoccum marconii McPartland,Mycotaxon 53:421,
named it after Cavara's fungus ("Botryosphaeria marconii"). 1995; = Macrophoma marconii nomen nudum, various authors.
 Chapter 5: Fungal Diseases 119

Description: Pseudothecia usually unilocular, globose, im- - specimens collected in Afghanistan by Schultes, and isolated
mersed then erumpent, 130-160 pm in diameter. Outer wall pale the fungus from two-month old 'Skunk No. 1' growing in a
brown but darkening near the ostiole; inner wall hyaline and very Dutch glasshouse, where it caused considerable losses.
thin. Asci bitunicate, eight-spored, clavate, with a short stalk, 80-90
x 13-15 pm. Paraphyses filiform. Ascospores fusoid to ellipsoid,
aseptate, hyaline to pale green, 16-18 x 7-8 pm. Leptodothiorella SYMPTOMS
conidiomata identical to pseudothecia. Conidiogenous cells simple Pink rot is a bit of a misnomer. The fungus often presents
or rarely branched, lageniform to cylindrical, 3-12 pm long, up to as a white fuzz covering leaves or flowering buds (Plate 66).
3 pm wide at base tapering to 1 pm at apex, phialidic, integrated or The pink tint arises when conidia are produced. In Holland,
discrete, with minute channel. Microconidia ellipsoid, single-celled, the fungus surrounded stems and girdled them. Girdled
hyaline, occasionallybiguttulate and swollen towards each end, 3.0- plants wilted and fell over. Ghillini (1951) and Fuller & Nor-
4.0 x 0.5-2.5 pm. Fusicoccum conidiomata slightly larger than those man (1946) noted the fungus ruins hemp fibres.
of Leptodothiorella, but otherwise indistinguishable (Plate 65).
~onidio~hores arise from inner wall, hyaliie, smooth,.simple or
branched and septate near the base, 10-16 pm long, up to 4.0 pm CAUSAL ORGANISM
wide at the base tapering to 2.5-3.0 pm at the tip. Conidiogenous Trichothecium roseum (Persoon:Fries) Link, Magazin,
cells holoblastic, integrated, determinate. Macroconidia fusiform, Gesellschaft Natuiforschender Freunde zu Berlin 3:18,1809.
single-celled,hyaline to glaucous, smooth-walled,base truncate, 16- =Cephalothecium roseum Corda 1838.
22 x 5-8 pm (Plate 65). Description: Colonies quickly turn dusty pink with conidia.
Conidiophoresupright, unbranched, often with three septa near the
DIFFERENTIAL DIAGNOSIS base, up to 2 mm long, 4-5 pm wide. Conidiogenesis basipetal,
The two fungi causing twig blight are easy to differen- conidia often remain in contact with each other in zig-zag chains.
tiate microscopically. Symptoms of twig blight may be con- Conidia ellipsoidal to pyriform with truncate basal scars, two-celled
fused with symptoms of brown stem canker (Dobrozrakova with upper cell larger and rounder than lower cell, hyaline (pink en
masse), with a thick smooth wall, 12-23 x 8-10 pm (Fig 5.22).
et al. 1956) or Phomopsis stem canker. The fungi must be
examined microscopically.

DISEASE CYCLE & EPIDEMIOLOGY

Both species overwinter in host tissue and spread by
splashed rain in the spring. Because of confusion in the lit-
erature, the range of these two species can only be estimated:
D. marconii infests hemp stems in Italy (Cavara 1888),France
(Barloy & Pelhate (1962), and Chile (Mujica 1942, 1943).
McPartland (199913)found it on hemp stems in Austria, hemp
leaves in Michigan, and leaves of drug plants in Nepal.
Charles & Jenkins found B. marconii in Maryland and Vir-
ginia. Judging from other authors' descriptions, B. marconii
also occurs in Russia (Gitman & Boytchenko 1934), Germany
(Flachs 1936, Patschke et al. 1997), and Italy (Petri 1942).

CULTURAL & MECHANICAL CONTROL (see Chapter 9) Figure 5.22: Conidia of Trichothecium roseum (LM x1000,
Use methods 1 (sanitation), 5 (genetic resistance-see courtesy Bud Uecker).
below), 7a (avoid drought), and 7c (avoid overhead irriga-
tion, keep plants well-spaced). Charles & Jenkins (1914)
noted twig blight strikes males first, then moves to females. DIFFERENTIAL DIAGNOSIS
Male plants should be rouged. Dempsey (1975)listed Russian Pink rot, when it looks white or beige, can be confused
and Ukrainian hemp varieties with resistance to B. marconii- with grey mould, powdery mildew, or downy mildew. Mi-
'Monoecious Central Russia,' 'Odnodomnaja 2,' 'USO-1' croscopic inspection easily differentiates these diseases.
1 , , ~ , . ; ~ , , . - 1 . .+ --..,-I-+-A -,- /vri c n
(Illachs 1936, Patschke et al. 1997),and ltaly (Petri 1942).
i f- - I ~ T i c n I*\ dr 1 r n 7 * ..-I
"" w
CULTURAL & MECHANICALCONTROL (see chapter 9) Figure 5.22: Conidia of Trichothecium roseum (LM xl000,
Use methods 1 (sanitation), 5 (genetic resistancesee Bud Uecker).
below), 7a (avoid drought), and 7c (avoid overhead irriga-
tion, keep plants well-spaced). Charles & Jenkins (1914)
noted twig blight strikes males first, then moves to females. DIFFERENTIAL DIAGNOSIS
Male plants should be rouged. Dempsey (1975)listed Russian Pink rot, when it looks white or beige, can be confused
and Ukrainian hemp varieties with resistance to B. marconii- with grey mould, powdery mildew, or downy mildew. Mi-
'Monoecious Central Russia,' 'Odnodomnaia , 2,'. 'USO-1'
croscopic inspection easily differentiates these diseases.
/.rqr;n..rl,r trq~rl-&nrl -- I V T TCA 1 1
(Flachs 1936, Patschke et al. 1997), and Italy (Petri 1942).
--. t T T TCA 1 I\ I T T P A I . .-1

CULTURAL & MECHANICAL CONTROL (see Chapter 9) Figure 5.22: Conidia of Trichothecium roseum (LM x1000,
Use methods 1 (sanitation), 5 (genetic resistance-see courtesy Bud Uecker).
below), 7a (avoid drought), and 7c (avoid overhead irriga-
tion, keep plants well-spaced). Charles & Jenkins (1914)
noted twig blight strikes males first, then moves to females. DIFFERENTIAL DIAGNOSIS
Male plants should be rouged. Dempsey (1975)listed Russian Pink rot, when it looks white or beige, can be confused
and Ukrainian hemp varieties with resistance to B. marconii- with grey mould, powdery mildew, or downy mildew. Mi-
'Monoecious Central Russia,' 'Odnodomnaja 2,' 'USO-1' croscopic inspection easily differentiates these diseases.
f ;-, , n ~ , ,+--..-i-+,.~ IVT icn 7 -- ~ T lT c n I !\ IT T e n 7 I - .3
(Flachs 1936, Patschke et al. 1997), and Italy (Petri 1942).

CULTURAL & MECHANICAL CONTROL (see Chapter 9) Figure 5.22: Conidia of Trichothecium roseum (LM x1000,
Use methods 1 (sanitation), 5 (genetic resistance-see courtesy Bud Uecker).
. . . - . .. . . . .- . .
120 Hemp Dis:eases and Pests

BIOCONTROL & CHEMICAL CONTROL C. herbarttm also acts as a secondary invader of plants
N o biocontrol is known. T. rosetlrn, in fact, serves as a parasitized b y Sphaerotheca, Septoria, or Pkornopsis species
biocontrol a g a i n s t Sclerotinia sclerotiorl~rni n soil. N o (McPartland, unpublished data). Once established, the
fungicides are effective. Dahiya & Jain (1977) reported that fungus is difficult to eradicate. It may survive after fungicides
pure THC and CBD inhibited the growth of T. roseurn. eliminate the primary pathogen. Indeed, Durrell & Shields
(1960) found C. herbarurn growing at Ground Zero in Nevada
shortly after nuclear weapons testing. The fungus is one of
the most common facultative parasites i n the world. Farr et
CLADOSPORIUM STEM CANKER al. (1989) described C. kerbarl~rnfrom 90 host plant genera.
Two or more species of Cladosporit~rncause stem can-
kers. The most common species, C. herbarurn, also infests 2. Cladosporium cladosporioides (Fresenius) deVries, Con-
leaves and seeds. C, herbarurn produces cellulolytic enzymes tributions to the knowledge of the genus Cladosporium, pg. 57,1952.
which continue to ruin h e m p after it has been harvested. =Homodendrum cladosporioides(Fresenius) Saccardo 1880.
Description: Conidiophores straight or flexous,simple or branched,
smooth or verruculose, pale to olivaceous brown, 2-6 pm in diam-
SYMPTOMS eter and up to 350 pm long, but generally much shorter. Ramo-co-
O n stems, this disease begins as dark green spots. The nidia an extension of conidiophores, irregularly shaped, smooth or
spots elongate a n d turn velvety green-grey i n d a m p condi- rarely minutely verruculose, 2-5 pm wide, up to 30 pm long. Co-
tions (Fig 5.23). Cortical tissues beneath these lesions necrose, nidia formed in simple or branched chains, ellipsoidal to lemon-
creating cankers. Affected fibres stain a dark brown-black shaped, pale olivaceous brown, mostly smooth walled, rarely
colour and lose their tensile strength (Gitman & Malikova minutely verruculose, one-celled, 3-7 x 2 4 pm.
1933). Leaf spots grow round to irregular in shape, covered Gzebenyuk (1984) isolated C. cladosporiodes from hemp
by a green-grey mat of mycelium. The spots turn necrotic, stalks i n t h e Ukraine. O n d r e j (1991) rarely f o u n d C.
dry out, and break u p into ragged shot-holes. cladosporiodes infesting stalks in the Czech Republic, whereas
C. herbarurn w a s abundant.

3. Cladosporium tenuissimum Cooke, Grevillea 5(37):140,1877.

Description: Conidiophores flexous, branching, olive brown,
swollen at the apex, 2-5 pm in diameter, up to 800 ym long. Conidia
formed profusely in chains, ellipsoidal to oval, smooth walled or
minutely verruculose, one-celled, 3-25 x 3-6 pm.
Nair & Ponnappa (1974) cited C. tenltissirnurn o n dis-
eased marijuana leaves in India. N o one else has found C.
tenuissirnltrn in Asia. Nair & Ponnappa may have misidenti-
fied C. herbanirn.

4 . Cladosporium resinae (Lindau) deVries, Antonie van

Leeuwenhoek 21:167,1955.
Figure 5.23. Symptoms of Cladosporium stem canker =Homoconis resinae (Lindau) vonArx & deVries 1973.
teleomorph: Amorphotheca resinae Parberry 1969.
(courtesy Bud Uecker).
Description: Conidiophores straight or flexous, simple or
branched, smooth or distinctly warted, pale brown to olive green,
3-6 pm wide and up to 2 mm long. Ramo-conidia, when present,
CAUSAL ORGANISMS & TAXONOMY are clavate or cylindrical, generally smooth, 8-20 x 3-7 ym. Conidia
1. Cladosporium herbarum (Persoon) Link, Magazin Ges. nuturf. solitary or in chains, ellipsoid, smooth walled, pale to olivaceous
Freunde Berlin 7:37,1815; Dematium herbarum Persoon 1794; brown, without prominent scars, 3-12 x 2-4 ym.
=Hormodendrum herbarum auct. Gzebenyuk (1984) reported C. resinae o n hemp stems.
teleomorph: Mycosphaerella tassiana (deNotaris)Johanson 1884; C. resinae normally colonizes petrochemical substrates such
=MucosvlzuereIla tulasnei (Tanczewski)Lindau 1906.
d <
as jet fuel and facial creams; it has not been reported from
Description: Conidiophores cylindricaland straight (in culture)
or nodose and geniculate (in viva), unbranched until near the apex, herbaceous plants (McPartland 1995a).
smooth, pale to olivaceous brown, 3-6 pm in diameter and up to
250 pm long. Conidia blastic, produced sympodially in simple or DIFFERENTIAL DIAGNOSIS
branched chains, ellipsoidal or oblong, rounded at the ends, thick Cladosporium stem canker can b e confused with hemp
walled, distinctly verruculose, golden to olivaceous brown, with canker, brown blight, a n d Stemphylium stem disease.
scars at one or both ends, zero or one septum, 8-15 x 4-6 pm.
Teleomorph pseudothecia globose, black, scattered to aggregated, DISEASE CYCLE & EPIDEMIOLOGY
up to 160 ym in diameter. Asci bitunicate, subclavate, short stipitate,
eight-spored, 35-90 x 15-30 Fm. Ascospores hyaline to rarely pale
Cladosporit~rnspecies overwinter in crop debris. They
. -

brown, ellipsoid, usually one septum,~slightlyconstricted at sep- continue to cause destruction i n retted h e m p after harvest.
tum, 15-30 x 4.5-9.5 pm. Paraphyses not present. Cladosporium species also overwinter in seeds (Pietkiewicz
C. Izerbarurn attacked stalks and flowering tops of hemp 1958, Ferri 1961b, Stepanova 1975). C. herbarurn a n d C.
in Italy (Curzi & Barbaini 19271, a n d stalks i n the Ukraine cladosporioides grow best between 20-28'~, but they can grow
(Gzebenycek 1984), and the Czech Republic (Ondrej 1991). at standard refrigerator temperatures. Warm humid condi-
It caused over-retting of h e m p in Germany (Behrens 1902), tions favour conidial production. A slight breeze detaches
and ruined harvested h e m p & Russia it man & Malikova conidia and carries them for miles. peaks of airborne conidia
1933, Vakhrusheva 1979). C. herbart~rnhas infested drug plants arise in June-July and September-October (Domsch et al.
from Colombia (Bush Doctor, unpublished data) and India 1980).Airborne C. herbarurn and C. cladosporioidesconidia are
(Nair & Ponnappa 1974, Ponnappa 1977). Babu et al. (1977) major causes of "mould allergy." Opportunistic infections
isolated the fungus from seeds i n India. Lentz (1977) reported b y Cladosporiurn runs the gamut from eye ulcers to pulmo-
its teleomorph o n Cannabis stems. nary fungus balls (Rippon 1988).
 Chapter 5: Fungal Diseases 121

CONTROL as cabbage, mustard, lettuce, chrysanthemums, and many

Cornerstones of cultural control (described in Chapter weed hosts. C. coccodes attacks both leaves and stems of
9) include method 1 (sanitation-eliminate any damp organic Cannabis. Hoffman (1958,1959) reported heavy hemp losses
matter, especially wet paper products) and 7c (avoid excess in central Europe. Conversely, Gitman (196813) considered
humidity). Also follow methods 2b&c (sterilization or pas- the pathogen of little importance in the USSR. Ghani & Basit
teurization of soil), 8 (optimizing soil structure and nutri- (1975)probably collected C. coccodes on Pakistani drug plants
tion), 10 (careful pruning), and 11 (avoid seedborne infec- (described as "Colletotricht~rnspecies ...cylindrical conidia
tion). Pandey (1982) protected millet seeds from C. herbartirn with both ends rounded"). A hemp specimen collected by
with a 30 minute soak in Cannabis leaf extract.' Vera Charles in Virginia (deposited at herb. BPI) also proved
to be C. coccodes (McPartland, unpublished data).

2. Colletotrichum dematium (Persoon) Grove, 1. Botany (Lon-

ANTHRACNOSE don) 56:341,1918; =Vermiculariadematium (Persoon) Fries 1849; =Ver-
Three species reportedly cause Cannabis anthracnose. micularia dematium f. cannabis Saccardo 1880.
All are soilborne fungi and cause disease in temperate Description: Acervuli are round to elongated, up to 400 pm
climates worldwide. Two of the fungi parasitize a wide range diametre, strongly erumpent through epidermis and exuding smoky
of plants and also exist as saprophytes. Gzebenyuk (1986) grey conidial masses with divergent setae. Setae smooth, stiff, rarely
reported a third species, Colletotrichum lini, which normally curved, usually three or four septate, 4-7.5 pm wide at the base
tapering to sharpened apices, 60-200 pm long (Fig 5.25).
attacks only flax (Linurn species); his report is probably
Conidiophores hyaline, cylindrical. Conidiogenous cells phialidic.
erroneous (McPartland 1995a). Conidia hyaline, aseptate (becoming two-celled during germination),
smooth, thin walled, guttulate, falcate and curved, 18-26 pm long
SYMPTOMS and 3-3.5 wide in the middle and tapering to pointed apices
Leaf symptoms begin as light green, watersoaked, (Fig5.24).Appressoria club-shaped to circular, medium brown, edge
sunken spots. Spots enlarge to circular or irregular shapes usually entire, 8-11.5 x 6.5-8 pm, often becoming- complex.
-
with grey-white centres and brownish-black borders. Larger The fungus is a weak parasite on many hosts; Von Arx
spots may become zonate. Affected leaves soon wrinkle then (1957) listed about 80 additional synonyms. C. dernati~rrnhas
wilt (Hoffman 1959). been collected from hemp stems in Italy (Saccardo 1880,
Stem lesions initially turn white. Then black, dot-like Cavara 1889). Saccardo named his specimen a new subspe-
acervuli arise in the lesions, lending a salt-and-pepper ap- cies, but McPartland (1995e) considered it superfluous.
pearance (Plate 67). Affected stems swell slightly and de- Clarke collected a fungus infesting stalks of cultivar 'Uniko-
velop cankers. The periderm peels off easy. Stems sometimes B' growing in China, which turned out to be C. dernatitrrn
snap at cankers. Distal plant parts become stunted and of- (McPartland, unpublished data 1995).
ten wilt. Young plants die.

CAUSAL ORGANISMS & TAXONOMY

1. Colletotrichum coccodes (Wallroth)Hughes, Can. 1.Bot. 36:754,
1958; =Colletotrichumatramentarium (Berkeley & Broome) Tauben-
haus 1916, -Vermicularia atramentaria Berkeley & Broome 1850.
Description: Acervuli on stems round or elongated, reaching
300 pm in diametre, at first covered with epidermis; then dehiscing
irregularly and exuding slimy conidia and bristling setae. Setae
smooth, stiff, septate, slightly swollen and dark brown at the base
and tapering to sharpened and paler apices, up to 100 pm long.
Conidiophores hyaline, cylindrical, occasionally septate and
branched at their base. Conidiogenous cells phialidic, smooth, hya-
line, with a minute channel and periclinal thickening of the collarette.
Conidia hyaline (honey coloured to salmon-orange en masse), asep-
tate, smooth, thin walled, guttulate, fusiform and straight, with
rounded apices, often with a slight median constriction, averaging
14.7x 3.5 pm on Cannabis stems but ranging 16-22 x 3-4 pm in cul-
ture (Fig 5.24).Appressoria club-shaped, medium brown, edge ir-
regular to almost crenate, 11-16.5 x 6-9.5 pm, rarely becoming com-
plex.
Over a dozen additional synonyms of C. coccodes are
cited by von Arx (1957).The fungus is a powerful pathogen
of solanaceaous crops (tomatoes, potatoes, eggplants), as well

Figure 5.25: Acervuli with setae formed by Colletotrichum

dematium (LM x40, courtesy Bud Uecker).

DIFFERENTIAL DIAGNOSIS
C. coccodes and C. dernatit~rnare best differentiated by
their conidia (Fig 5.25). C . coccodes conidia are fusiform,
straight, and have rounded ends. C. demati~~rn conidia are
Figure 5.24: Colletotrichum conidia, x1500. A. C. falcate, curved, and have pointed ends. Both species produce
coccodes; B. C. dematium (McPartland redrawn from abundant sclerotia and setae, and their appressoria appear
Sutton 1980). similar (McPartland & Hosoya 1998).
122 Hemp Dis:eases and Pests

Anthracnose can be confused with blights, leaf spots, an upright branch which can reach 150 pm in height. Phialides
stem cankers, wilt diseases, and plant diebacks. Anthracnose hyaline, 19-35 pm long, 1.5 pm wide. Conidia arise singly but
may also mimic damping-off in seedlings. congregate in small droplets at tips of phialides, oval to ellipsoid in
shape, hyaline, aseptate, 2.5-8 x 1.5-3.2 pm (Fig 5.26).Microsclerot~d
arise by lateral budding of a single hypha into long c h a m of cells,
DISEASE CYCLE RE EPIDEMIOLOGY becoming dark brown to black, irregularly spherical to elongated,
Anthracnose fungi overwinter as sclerotia in plant de- 15-100 pm in length (Fig 5.26).
bris or soil. In the spring, sclerotia sporulate and conidia Reports of V. dahliae come from southern Russia
splash onto seedlings. Conidia form appressoria which di- (Vassilieff 1933, Gitman 1968b), Italy (Noviello 1957), the
rectly penetrate epidermal tissue or enter via stomates and Czech Republic (Ondrej 1991), the Netherlands (Kok et al.
wounds. Seed transmission does not occur. C. coccodes pre- 1994), and Germany (Patschke et al. 1997). V daliliae attacks
fers a cooler optimum temperature than C , den~atil~in. many cultivated, weedy and wild plants, in temperate zones
Expect anthracnose epidemics during cool damp and the tropics.
weather, especially in heavy soils. Hoffman (1958, 1959)
noted high losses in "bog" soils. Disease escalates in plants 2. Vevticillium albo-atvum Reinke & Berthier, Die Zersetzung
under stress from drought or frost damage. Anthracnose can der Kautoffel durcll Pike, p. 75, 1879.
rage in monocropped glasshouses, especially in hydroponic Description: Conidiophores abundant, mostly hyaline but with
systems (Smith el al. 1988). Conidia spread via splashing a darkened base (especially 111 zli~:o),with two to four ph~alides
water and wind-drilren rain. Hoffman (1958,1959)and Cook arising at regular intervals along an upright branch up to 150 pni
(1981) described heaviest infections after plants had tall. Phialides hyaline, variable, 20-30 (up to 50) pm in length and
flowered, with males succumbing before females. 1.5-3.0 pm wide. Conidia arise singly ellipsoid to short-cylindrical,
Concurrent infection by the nematode Heterodera schactii or hyaline, usually single-celled but occasionally one septum, 3.5-10.5
x 2.5-3.5 pm. No microsclerotia formed.
the fungus Rhizoctonia solaizi increases plant susceptibility
V alho-atriln~has attacked hemp in China (Tai 1979)and
to anthracnose (Smith et al. 1988).
the Ukraine (Gzebenyuk 1984). V. albo-atrilrrr prefers cooler
CULTURAL & MECHANICAL CONTROL (see Chapter 9) climates in northern Eurasia and North America, and attacks
many dicots. It causes more severe symptoms than V. iiahliae.
Sanitation is the cornerstone-methods 1 (eliminating
residues), 10 (careful pruning and rouging), 2a (deep
ploughing), 2b&c (sterilizing or pasteurizing soil), and 3 DIFFERENTIAL DIAGNOSIS
(eliminating weeds), in that order. Also observe methods V. dahliae produces slightly smaller conidia than V. albo-
7a&b (avoid drought or waterlogging), 8 (avoid heavy soils), atriini, but the two species are difficult to distinguish. Only
and 6 (avoid solanaceous crops if C. coccodes prevails). Do V. dahliae produces microsclerotia. No reports of Verticilliun~
not wet stems while irrigating. Stay out of Cnn~zabispatches on C a n n a b i s mention the presence or absence of
when plants are damp, since conidia spread by contact. microsclerotia. Thus either fungus could have been collected.
Clarke (pers. cornmun. 1995)noted Uniko-B was susceptible
to C . iieiizatiuni in China, whereas the local Chinese landrace
\vas resistant to the fungus.

BIOCONTROL & CHEMICAL CONTROL

According to Samuels (1996), Trichodernin harzianzlni
(Trichodexg)has controlled Colletofricllum disease (described
under Rhizoctonia sore shin disease). Yepsen (1976) sug-
gested a prophylactic spray of lime sulphur on plants dur-
ing susceptible weather. Kaushal & Paul (1989) inhibited a
related species, Colletotriclzl~nzt r ~ ~ n c a t u iwith
n , an extract of
Cnnnnbzs.

VERTICILLIUM WILT
Two organisms probably cause this disease in Cannabis.
Both species attack many crops, and both pathogens live
i\.orldwide.
Figure 5.26: Verticillium dahliae, x400. A. Condiophore
SYMPTOMS
with conidia aggregated in droplets; 6. Microsclerotium;
Leaves first turn yellow along margins and between
(McPartland).
veins, then turn grey-brown. Lower leaves show symptoms
first. Slightly wilted plants often recover at night or after
Prior to wilting, symptoms of Verticillium wilt may be
irrigation. These recoveries are transient as wilt becomes per-
confused with nutritional disorders. Verticillium wilt resem-
manent. Dissection of diseased stems reveals a brownish dis-
bles Fusarium wilt except the xylem discolouration is dif-
colouration of xylem tissue. If the fungus invades only a few
ferent. The wilt mimics symptoms caused by nematodes and
xylem bundles, only parts of the plant may wilt.
root-boring insects (root maggots, white grubs), or drought.
CAUSAL ORGANISMS & TAXONOMY
DISEASE CYCLE & EPIDEMIOLOGY
1.Verticillizlm dahline Klebahn, Mycol. Cei~tualb.,3:66, 1913.
= iirrticillu~n~ulbo-atruin rar. niediunz Wollenweber 1929; =Vertici/lium Both species overwinter in soil and invade roots of seed-
traclzeipliillln~Curzi 1925. lings in the spring. Once in roots Verticillil~n~
species spread
Description: Conidiophores abundant, completely hyaline, via xylem; they eventually clog the pipes and cause wilts.
with three or four rvliorled phialides arising at regular nodes along Verticillium wilt increases in moist soils rich in clay. Root-
 Chapter 5: Fungal Diseases 123

knot nematodes predispose plants to Vertlcllhl~ininfection (McPartland 1995a). Rataj's error reappears in Barna et al.
I/ nlbo-atr lrrn grows best at 23 5"C, the optimum for V dahl- (1982),Gutberlet & Karus (1995), and Bosca & Karus (1997).
me 1s 21°C
CAUSAL ORGANISMS & TAXONOMY
CULTURAL & MECHANICAL CONTROL (see Chapter 9) 1. A e c i d i u m c a n n a b i s Szembel, Conlnletztarii Institutl
Clean the soil-use methods 1 (but no burying!), 2b Astmchanensis ad defensiorlern plantarum, i, 5-6, p. 59, 1927.
(sterilize the soil), and 3 (weeding). Method 2a is not too Description: Aecia arise in pale leaf spots on undersides of
helpful since microsclerotia can live 75 cm deep in soil. lower leaves, "globose" [cupulate?],orange coloured, gregarious,
measuring 360-400 pm deep and 300-340 pm in diameter. Aecial
Method 2c (soil solarization) works well against V. dahliae,
peridium cells rectangular, often rhomboidal, 32-40 x 18-24 Fm,
but not heat-resistant V nlbo-atrl~n~ (Elmore et al. 1997).Flood- exterior walls 6-8 pm thick. Aecidiospores round to ellipsoidal,
ing infested soil for 14 days kills Verticillil~nlmicrosclerotia subhyaline, Xrerrucose, single-celled, 24-28 x 20-24 pm, with an
(Lucas 1975). epispore 1.5-2.0 pm thick.
Avoid planting in lieavy, poorly draining soils. Accord- Szembel collected A. cannabis from Cannabis r~~deralis
ing to Elmer & Ferrandino (1994),Verticillium wilt decreases growing near the Caspian Sea. The Cannabis was concurrently
when nitrogen is supplied as an ammonia [e.g., (NH&S04] infected by Septorin cannabis.
rather than a nitrate [e.g.,Ca(NO&].
Crop rotation with nonhost monocotyledonous plants 2. Uredo kriegeriana H . & P. Sydow, Osterreiciliscile botaniscile
must be long (four years for V. albo-atruin, twice that for V. Zeitschrift 52(5):185,1902.
d~lhline).Monocot crops must be weed-free, since almost all Description: Uredia sorl borne on undersides of leaves in
dicot weeds serve as alternate hosts. Root-knot nematodes irregular yellow to ocher spots, either sparsely or gregariously
(Meloirlogyze species) should be eliminated from the soil. distributed, initially subepidermal ("h~ddenin enclosing per~dium")
Method 5 is a future option-hops breeders have developed then erumpent and pincush~onshaped, covered by yellow spores.
~ ~ a r i e t resistant
~es to V albo-atrl~in.Kok et al. (1994) found Uredospres subglobose to ellipsoidal, finely echinulate, orange on
the inside, single-celled, 21-27 x 15-22 pm, bearing many
partial resistance to V. dahliae in fibre cultivar 'Kompolti germination pores.
Hibrid TC.'
Sydow & Sydow (1924) amended the description,
"Uredinia appear in loose aggregations within leaf spots, the
BIOCONTROL & CHEMICAL CONTROL
covering peridium consists of roundish to obtuse-angled
V e r f l c l l l i l ~wilt
i ~ ~ constitutes a real menace to world ag-
thin-walled cells, hyaline and membranous, 1 l m thick, 12-
riculture, because no chemicals can control the disease (Smith
17 yrn long by 10-13 pm wide. The uredinospores contain a
ct 01. 1988). Many soil bacteria and fungi antagonize
hyaline epispore 1.5 pm thick, and the germination pores
Vertic~ili~~nr-increasetheir populations by mixing green al-
are difficult to see." Father and son Sydow collected U .
falfa meal into soil. Windels (1997) mixed sudangrass into
kriegeriana from hemp near Schandau, Germany.
soil as a green manure to control V ilalzliae in potatoes. Grewal
(1989) reduced Verticilliliiil growth by mixing compost with
3. U r o m y c e s i n c o t r s p i c u t ~ s Otth, Mittlzeilungen d e r
d r ~ e dCnllrlabis leaves. natufo~sclzmdenGesellschaft in Bern. 1:69,1868.
A Trichoderrnn product (Bio-Funguss) reportedly con- Description: Leaf spots on undersides of leaves, very incon-
trols Verficilli~lii7wilt (described under Rhizoctonia sore shine spicuous, telia growing in punctiform tufts, contain~ngfew
disease), as does the FZB24 strain of Bacilll~ssubtilis (Rhizo- teliospores.Teliosporesborne on short hyaline pedicels; spores dark
Pluso, described under damping off). Talaronzyces flavus is brown-black, ellipsoid, venuciform, smglecelled, with a small hyaline
irregularly available (see below), and V e r t i c i l l i t ~ mnigresceirs apex, 32 x 18 pm.
is being developed. V. nigrescens is a nonpathogenic species Otth collected U . inconspic~~us from hemp, potatoes, and
that protects plants from infection by pathogenic Verticillil~rrl several genera of weedy plant species. his host range is
species (Howard et al. 1994). unusual for Uroi?zycesspecies. Saccardo (S.F. 14287) expressed
his doubts, "Quid sit haec species pantogena non liquet, certe
Tnlnromyces f l a v u s dubia res." Fisher (1904) thought Otth misidentified a
BIOLOGY: Kendrick (1985) reported a 76% reduction in hyphomycete.
eggplant wilt from V. dahliae by inoculating soil with spores
of the soil-inhabiting fungus T.flavus. The biocontrol fungus DISEASE CYCLE & EPIDEMIOLOGY
produces hydrogen peroxide which kills V . daliliae All three fungi have been described from a single spore
microsclerotia. state. Interestingly, each represents a different stage of the
APPLICATION: Supplied as spores in granules, which rust life cycle (aecia, uredia, and telia, respectively). They
are mixed into soil or potting medium. T.flaul~sis tolerant of could be different spore states of the same organism.
heat; heating soil (solarization) makes V.~dahliaemuch more
susceptible to the biocontrol DIFFERENTIAL DIAGNOSIS
True rusts should not be confused with "white rusts,"
which are related to downy mildews. Manandhar (pers.
commun. 1986) saw a white rust caused by an Albirgo spe-
RUST cies in Nepal.
Rust fungi cause many of our worst crop diseases. They
CONTROL
often exhibit complicated life cycles spanning several plant
Cultural control method 10 (careful pruning-see Chap-
hosts, and produce up to five distinctive spore states. A full
ter 9) may be sufficient. Also follow method 7c (avoid excess
rust life cycle includes spermagonia, aecia, uredia, telia, and
humidity, avoid overhead watering and dew condensation
basidia (Fig 3.3).
in glasshouses). No biocontrol is known. Sulphur helps if
Of the four rust fungi reported from Ca~znabis,one arises
one carefully dusts the undersides of affected leaves. Misra &
from a typographical error: Rataj (1957) misspelled Melano- Dixit (1979) used ethanol extracts of Cannabis to kill Ustilago
sporacnni?abis (the cause of red boot) as M e l a m p s o r a caniza- tritici (Pers.) Rostr. and Ustilago lzordei (Pers.)Lager., two smut
bis-Melanospora is an ascomycete, Melanlpsora is a rust fungi that are somewhat related to rusts.
124 Hemp Diss!ases and Pests

CONTROL
BLACK DOT This disease does not arise in healthy Cannabis--observe
Also called "small shot" o r "black spot," this disease cultural methods 7a a n d 8 (described i n Chapter 9). Cure
affects leaves and stems. Leaf disease occurs i n India (Nair harvested material in a dark (no UV light), dehumidified
& P o n n a p p a 1974, P o n n a p p a 1977) a n d M a r y l a n d chamber. Biocontrol is unknown. Treat badly infected plants
(McPartland, unpublished). Stem disease has been reported with copper-based fungicides.
in the Ukraine (Gzebenyuk 1984) a n d the Czech Republic
(Ondrej 1991). In Illinois, the fungus acts as a saprophyte,
colonizing male flowers after release of pollen (McPartland,
u n p u b l i s h e d ) . T h e c a u s a l f u n g u s a l s o rots h a r v e s t e d BASIDIO ROT
marijuana in storage. It grows worldwide o n a wide range Older Cannabis literature describes four Corticizim spe-
of plants, animals, a n d processed foodstuffs. cies o n hemp-they are all old names, now considered syno-
nyms. Two Corticil~mspecies are true pathogens-C. solani
SYMPTOMS (= Rhizoctonia solani) and C. roljsii (= Sclerotil~mrolfsii). They
Black dot disease is characterized b y small dark pus- are discussed under Rhizoctonia sore shin a n d southern
tules of fungal growth. Pustules reach a finite size (less than blight, respectively. T h e remaining t w o species act a s
2 mm) and become covered with black conidia. Pustules may saprophytes o r weak pathogens.
appear o n healthy green tissue o r in the midst of a small
chlorotic ring. O n leaves they concentrate near midveins or CAUSAL ORGANISMS & TAXONOMY
vascular tissue. O n stems they scatter randomly. 1.Athelia arachnoidea (Berkeley) Jiilich, Willdenowin Beih. 7:23,
1972; =Septobasidium nrachnoideum (Berkeley) Bresadola 1916,
CAUSAL ORGANISM & TAXONOMY =Hypochnus arachnoideus (Berkeley) Bresadola 1903, ~Corticium
Epicoccum nigrurn Link, Magazin Ges. Nuturf.Freunde, Berlin 7:32, arachnoideum Berkeley 1844.
1816; =Epicoccum pururascens Ehrenberg ex Schlechtendahl1824. anamorph: Fibularhizoctonia carotae (Rader) Adams & Kropp
Description: Sporodochia 100-2000 pm diametre. 1996; ~Rhizoctoniacarotae Rader 1948.
Conidiophores densely compacted, straight or flexuous, occasion- Description: Basidiocarp effuse, thin, bluish white when fresh,
ally branched, colourless and smooth but turning pale brown and surface arachnoid, 2-6 cm long and 1-3 cm wide, 100-200 pm thick
verrucose at the tip. Conidia formed singularly, monoblastically, in cross section. Subhymenial hyphae loosely interwoven, without
clamp connections, 3-5 km in diameter; basal hyphae somewhat
globose to pyriform in shape, golden brown to dark brown, with a wider, with thickened walls and occasional clamp connections. Ba-
warted surface obscuring muriform septa which divide conidia into sidia clavate, grouped like candelabra, normally producing two
many cells (up to 15), conidia 15-25 pm diametre (Figs 3.2 & 5.27). basidiospores borne upon pointed sterigmata, 20-30 x 5-7 pm.
Basidiospores hyaline, smooth, ellipsoid to pyriform, usually 6-10
x 4-6 um. Mvcelium is similar to that of Rhizoctonia,but with clamv
connections: Sclerotia globose, composed of dark brown
chymatous cells, 0.2-1.0 mm diam (larger in culture, 1-5 mm diam).
Lentz (1977) cited A. arachnoidea o n Cannabis. This fun-
g u s is common in North America a n d Scandinavia. It grows
o n leaf humus, parasitizes lichens, and causes a cold-stor-
age disease of carrots (Daticzis carota).

2. Athelia epiphylla Persoon, Mycol. Europ. 1:84,1822.

=Athelia epiphylla Persoon:Fries 1822; =Hypochnus cetrifugus
(Leveille)Tulasne 1861,~Corticiurncentrifugurn (Leveille)Bresadola
1903; =Butlerelfa eustacei Weresub & Illman 1980.
anamorph: Fibularhizoctonia centrifuga (Leveille) Adams &
Kropp 1996;=Rhizoctonia centrifuga Leveille 1843.
Description: Basidiocarp effuse, very thin, white to buff,
surface arachnoid to byssoid, 2-6 cm long and 1-3 cm wide, 75-150
pm thick in cross section. Subhymenial hyphae loosely interwoven,
without clamp connections, 2-3 ym in diameter, basal hyphae up
to 6 pm in diameter rarely with clamp connections. Basidia clavate,
grouped like candelabra, normally producing four basidiospores
Figure 5.27: Conidium of Epicoccum nigrum germinating borne upon pointed sterigmata, 10-20 pm long. Basidiospores hya-
with many germ tubes, next to a deflated Cannabis line, smooth, ellipsoid to pyriform, usually 4-8 x 2.54 pm. Myc-
cystolith leaf hair (SEM x1060, McPartland). elium and sclerotia are identical those of Athelia arachnoidea.
Endo (1931) recovered A. epiphvlla
. . " from Cannabis in
Japan. The fungus decays forest leaves, rots wood, and causes
DIFFERENTIAL DIAGNOSIS fisheye decay of stored apples. It is common in Eurasia, less
This disease can b e confused with early brown blight, s o in North America.
Curvularia leaf spot, corky leaf spot, o r black mildew. A. epiphylla a n d A. arachnoidea h a v e been shuffled
through many different genera. The t w o taxa may represent
DISEASE CYCLE & EPIDEMIOLOGY the same organism (Adams & Kropp 1996). Saccardo (S.F.
The fungus overwinters in crop debris. Conidia germi- 6:654) considered them identical 100 years ago.
nate many germ tubes (Fig 5.27) a n d quickly form colonies.
Optimal growth occurs at 23-28OC. Conidium production in- DIFFERENTIAL DIAGNOSIS & CONTROL
creases under UV light. The fungus produces a strong red- Symptoms of basidio rot may resemble southern blight
dish-brown pigment (mostly beta carotene) in host tissue. and Rhizoctonia sore shin. Follow cultural controls described
This pigment is visible in light-coloured substrates (maize for Rhizoctonia sore shin disease. N o biocontrol is known
kernels, apples). Metabolites of E. nigrum may cause hepatic a n d chemicals are ineffective.
and renal disorders (Rippon 1988).
 Chapter 5: Fungal Diseases 125

SYMPTOMS
RED BOOT Yellow leaves appear in late June to August, followed
This disease is a secondary problem and only arises on by wilting, leaf necrosis, and plant death. Roots develop de-
plants previously damaged by hemp canker. Some hemp pressed and discoloured lesions as the fungus destroys cor-
cultivators nevertheless consider red boot more dangerous tical tissue and invades xylem. Sifting soil around plants with
than hemp canker itself. Red boot has been reported in a 1 mm mesh sieve reveals small brown sclerotia. A tan myc-
Germany (Behrens 1891,Kirchner 1906, Flachs 1936) and the elial mat may form around the base of dead plants in warm,
Ukraine (Gzebenyuk 1984). damp conditions.

SYMPTOMS CAUSAL ORGANISM & TAXONOMY

Red boot appears on plants in late July or August (Beh- Phymatotrichopsis omnivora (Duggar) Hennebert, Persoonia
rens 1891).The causal fungus rapidly encases stems in thick, 7:199, 1973; =Pllynzatotric/l~~m ornnivoruni Duggar 1916; =Ozon~~iln
red mats of mould. Fibre harvested from infested stems is ornniz~orumShear 1907.
soft, frail, stained a red colour, and valueless. Description:Mycelium with septate hyphae lacking clamp con-
nections, forming thick-walled cruciform aerial setae and aggregat-
CAUSAL ORGANISM & TAXONOMY ing into subterranean cord-like funicles. Conidiophores borne di-
Melanosporn cannabis Behrens, Zeitscllrff Pfianzenkmnkheiten rectly on hyphae, hyaline, simple or branched, &ate to globose,
1213, 1891. hyaline, 20-28 x 15-20 pm. Conidia holoblastic, ovate to globose,
Description: Perithecia globose, orange red, averaging 210- smooth, thin walled, hyaline, with a broad base exhibiting a
230 pm diametre, ostiole atop a 60-90 pm perithecial neck. Asci detachment scar, 6-8 x 5-6 Fm. Sclerotia borne on hyphae, ovate to
described as swollen, no dimensions listed. Paraphyses not found. globose, at first yellow but turning reddish brown, 1-2 mm diameter,
.4scospores black, ell~ptical,22-26 x 15-17 pm. Conidiophores more often aggregated in clusters reaching 10 mm diameter.
or less erect, verticillately branched, multicellular, flask shaped, The teleomorph of D omnivo1.n is unknown. Unsubstantiated
narrowing at tips. Conidia arise at the apices of phialides, unicellu- reports link P. onnniuora to Trechisponz brinkrnailnii (Bresadola)Rogers
lal; forming in clusters or chains, red in colour, 4.4 x 3.0 pm. Beh- &Jacksonand Hyiinu~nornniz~orlnnShear.
rens noted that conidiophores bind together to form coremia in
artificial culture. DISEASE CYCLE & EPIDEMIOLOGY
The anamorph of iM. cannabis lias not been named. In a mono- The fungus is a soil organism. It grows in rope-like
grdph of Melai~ospouaspecies, Douguet (1955)considered M . canna- funicles, 20 to 60 cm underground. P. on~nivorainvades roots
bls slmilar to M ,zobelii and M . Jin~icola.Douguet did not examine of seedlings in the spring. In damp conditions the fungus
the type specimen, however, and expressed regret over Behrens's
failure to measure asci. emerges from the ground and spreads via conidia. In dry
conditions the fungus produces sclerotia in soil. These dura-
ble structures survive for years.
DIFFERENTIAL DIAGNOSIS
Red boot, with its thick, red mat of fungal hyphae and
red conidia, is rather unique and not easily confused with DIFFERENTIAL DIAGNOSIS
other hemp diseases. Initial symptoms (wilting) resemble those caused by
nematodes and soil insects (e.g., root maggots, white root
grubs). Texas root rot can also be confused with Fusarium
DISEASE CYCLE & EPIDEMIOLOGY
The fungus overwinters as ascospores in crop debris or wilt, Verticillium wilt, hemp canker, and southern blight.
Conidia of P. oinni~loraresemble those of B o t y t i s citlerea. The
soil. Epidemics occur in low, wet fields during damp grow-
cruciform setae and funicles are unique to P.otnniilora.
Ing seasons. Weedy fields reportedly promote the disease.
Conidia are splashed by rain to sites of secondary infections.
CULTURAL & MECHANICAL CONTROL (see Chapter 9)
M. cal~tinbiscannot infect Cnnnnbis in the absence of infec-
(Behrens 1891). M . cniirinbis Method 2a (deep tillage) exposes funicles to desiccation.
tion by Sclerotiizin sclerotiori~n~
Method 6 (crop rotation with monocots) must be of suffi-
also parasitizes mycelia and sclerotia of S. sclerotiorl~nz.
cient duration to outlast long-lived sclerotia. Also follow
methods 1 (sanitation), 3 (weeding), 7a (avoid drought), and
CONTROL
8 (optimize soil structure and nutrition).
Follon~cultural controls described in Chapter 9, espe-
cially methods 1 (sanitation), 2a (deep ploughing), and 10
BIOCONTROL & CHEMICAL CONTROL
(careful pruning). Methods 3 (destroying weeds) and 7c
Mixing organic matter into soil encourages natural
(a~roidexcess humidity) are also helpful. Eliminate hemp
biocontrol organisms. Use composted animal manure or a
canker (S. sclcrotiorl~m)and you eliminate red boot. See the
green manure of legumes or grasses. The biocontrol fungus
section on hemp canker. No biocontrols or fungicides have
Gliocladii~lnroseunz controls P. onznivorn (described under grey
been tested.
mould). No fungicide is effective. Move from Texas.

TEXAS ROOT ROT OPHIOBOLUS STEM CANKER

Texas root rot arises in the southwestern USA and north-
Four species of Opilioboll~sreportedly infest Cnrlnnbis.
ern Mexico. The causal fungus is limited to clay soils with
Gzebenyuk (1984) cited two species, 0. porphyrogoni~s and
little organic matter but lots of calcium carbonate, a high
0.vl~lgnris.These are old names of the species Leptospora rlL-
pH, and high temperatures. In the blackland prairies of
bella, discussed later. The remaining Ophiobol~lsspecies are de-
central Texas, the causal fungus attacks over 2000 species of
scribed below.
wild and cultivated dicots. Texas root rot attacks hemp in
Symptoms develop as weather warms. Lower stem sur-
Texas, Oklahoma, and Arizona (Chester 1941), causing 30-
faces turn brown-black. Plants senesce prematurely and die.
60% mortality rates (Taubenhaus & Killough 1923).Killough
After plants die, fruiting bodies of the causal fungi arise in
(1920) described an epidemic killing 95% of a crop during
blackened areas of stems.
the flowering stage.
126 Hemp Diseases and Pests

CAUSAL ORGANISMS & TAXONOMY Also follow methods 1 (remove or burn crop debris), and 7c
1. Oplziobolus cantlabintis G. Passerini, Rendiconti della R. (avoid excess humidity). Organic manuring and rotating
Accnderi~iadei Li~lcei4:62, 1888. with clover (method 6) encourages Ophiobolns-antagonistic
Description: Pseudothecia scattered, globose or conical, black, soil organisms. No fungicides work well.
immersed with conical apex and ostiole barely erumpent. Asci cy-
lindrical, short-stalked, eight-spored, 85 x 5 pm. Paraphyses cylin-
drical. Ascospores filiform, aseptate, hyaline, 65-85 x 1.0-1.25 ym.
Passerini found 0.cnnnabintlson hemp near Parma, Italy CHAETOMIUM DISEASE
His description of aseptnte ascospores would preclude this Chaetomillnt species are common soil organisms. They
fungus from the genus Opltiobol~~s. Many species have in- aggressively decompose cellulose, including hemp and
conspicuous septa, however, and he may have missed them. cotton fibres. During WWII the USA Army lost more tents,
tarps, rope, and clothing to Clzaetontiunl than to enemy forces.
2. Ophiobolt~sanguillidus (Cooke in Cooke & Ellis) Sacca- Most Clraetonlillnz species act as saprophytes, but reports of
rdo [as at~gt~~llides] Sylloge Fzinyorun~2:341, 1883; =Raplziodosporn Ckaetomil~mspecies acting as plant pathogens appear in the
rzngt~~llida
(Cooke 111 Cooke & Ellis) Cooke &Ellis 1878-9.
literature.
Description: Pseudothecia gregarious, globose to ampulliform,
black, immersed but quickly erumpent, papillate, 300-500 km
diameter. A s c ~cyliiidrical, short-stalked, eight-spored, 80-150 x 9- SYMPTOMS
12 Fn1. Paraphyses cylindrical, sliglitly longer than asci. Ascospores Ghani & Basit (1976) reported a root disease of Paki-
scolecosporous, yello~vish,ten to 14 septa, straight or curved, with stani marijuana caused by clzaetonlitlm,but did not identify
basal cells attenuated and apical cells swollen to an ovoid shape, the species nor describe symptoms. Chandra (1974) de-
80-120 x 2.5-4.0 pni (Fig 3.2). scribed Chaetomil~ntsl~ccine~lin causing a leaf disease of In-
0, anguillidlls commonly parasitizes members of the dian marijuana. Symptoms begin as light brown spots. Spots
Compositae, including Bidens, Anzbrosia, and Aster species. enlarge but remain limited by leaf veins. Spots gradually
Preston & Dosdall (1955) collected 0. angllillidi~sfrom feral darken, dry, and drop out, leaving irregular holes in leaves.
hemp near Judson, Minnesota. Chandra's citation is the first isolation of C. succineuln out-
side of North America.
DIFFERENTIAL DIAGNOSIS
0. n n g ~ ~ i l l i dhas
~ ~ smuch larger ascospores than 0. CAUSAL ORGANISM & TAXONOMY
cnnnnbinus. Ophiobolus stem canker may be confused with Chaetomium succineum Ames, Mycologia 41445,1949.
Leptosphaeria blight, brown stem canker, Phomopsis stem Description: Cltaetornitlir~perithecia are covered by extrava-
canker, and striatura ulcerosa (a bacterial disease). gant ornamental hairs (Fig 5.28). These hairs probably senreas de-
terrents against insect predation. Taxonomists use them to differen-
DISEASE CYCLE & EPIDEMIOLOGY tiate species. Perithecia are superficial, black, carbonaceous,
and 0. caizizabin~~s
0. ni~gl~illidi~s overwinter as mycelia ostiolated and globose to barrel-shaped. Asci of all species reported
or teleomorpl~sin crop debris, infecting plants in early spring. on Cannabis are clavate and eight-spored. Ascospores are olive brown
to brown when mature.
The disease worsens in fields retaining stubble from previ- l a e h r K Y i C 2 - -2 - L- - L - - ' ' 7 ~
-

DIFFERENTIAL DIAGNOSIS
0.nngl~illiclllshas much larger ascospores than 0 . CAUSAL ORGANISM & TAXONOMY
cailiiabii~l~s.
Ophiobolus stem canker may be confused with Chaetomium succineum Ames, Mycoiogia 41:445,1949.
Leptosphaeria blight, brown stem canker, Phomopsis stem Description: Cl~aeton~itlnl perithecia are covered by extrava-
canker, and striatura ulcerosa (a bacterial disease). gant ornamental hairs (Fig 5.28). These hairs probably serve as de-
terrents against insect predation. Taxonomists use them to differen-
DISEASE CYCLE & EPIDEMIOLOGY tiate species. Perithecia are superficial, black, carbonaceous,
0, nilg~~illidus
and 0. cnnnabitl~~s overwinter as mycelia ostiolated and globose to barrel-shaped. Asci of all species reported
or teleomorphs in crop debris, infecting plants in early spring. on Cannabis are clavate and eight-spored. Ascospores are olive brown
to brown when mature.
The disease worsens in fields retaining stubble from previ-
. . Slae@y,,,drfJ.A"fi@.YiC2: ----:-,. "--.-I---- - '-"'-
-"---:- -'
DIFFERENTIAL DIAGNOSIS
0. a n g ~ i i l l i d ~has
~ s much larger ascospores than 0. CAUSAL ORGANISM & TAXONOMY
cnnnnbinus. Ophiobolus stem canker may be confused with Chaetotnium succineutn Ames, Mycologin 41:445,1949.
Leptosphaeria blight, brown stem canker, Phomopsis stem Description: Cimetolnitllt~perithecia are covered by extrava-
canker, and striatura ulcerosa (a bacterial disease). gant ornamental hairs (Fig 5.28). These hairs probably serve as de-
terrents against insect predation. Taxonomists use them to differen-
DISEASE CYCLE & EPIDEMIOLOGY tiate species. Perithecia are superficial, black, carbonaceous,
0. nr~guillldtisand 0 . caiutabin~~s
overwinter as mycelia ostiolatrd and globose to barrel-shaped. Asci of all species reported
or teleomorphs in crop debris, infecting plants in early spring. on Cannabis are clavate and eight-spored. Ascospores are olive brou.11
to brown when mature.
The disease worsens in fields retaining stubble from previ- slae@hrA"fi.rC : L- L -- - - ' 1 1

DIFFERENTIAL DIAGNOSIS
0.nngl~illidllshas much larger ascospores than 0. CAUSAL ORGANISM & TAXONOMY
c~1ilnabn1lls.
Ophiobolus stem canker may be confused with Ckaetomiutn succineum Ames, Mycoiogia 41:445,1949.
Leptosphaeria blight, brown stem canker, Phomopsis stem Description: Chaetonziu~izperithecia are covered by extrava-
canker, and striatura ulcerosa (a bacterial disease). gant ornamental hairs (Fig 5.28). These hairs probably senJeas de-
terrents against insect predation. Taxonomists use them to differen-
DISEASE CYCLE & EPIDEMIOLOGY tiate species. Perithecia are superficial, black, carbonaceous,
0. nlzgnilliiil~sand 0. cannabinus overwinter as mycelia ostiolated and globose to barrel-shaped. Asci of all species reported
or teleomorphs in crop debris, infecting plants in early spring. on Cannabis are clavate and eight-spored. Ascospores are olive brown
to brown when mature.
The disease worsens in fields retaining stubble from previ- Slae@hdrf&A"figYiC2:
----:-- L --.-"--- -----I-'-' - '''~
DIFFERENTIAL DIAGNOSIS
0. nngl~illidltshas much larger ascospores than 0. CAUSAL ORGANISM & TAXONOMY
callr~ablnlls.Ophiobolus stem canker may be confused with Chaetomiutn succineum Ames, Mycologia 41:445,1949.
Leptosphaeria blight, brown stem canker, Phomopsis stem Description: Clmetonl~u~n
per~thec~a
are covered by extrava-
---I -.-A -L..:.L ..I /. 1 I ' 1 1'
 Chapter 5: Fungal Diseases 127

SYMPTOMS
Stem cankers begin light-coloured and slightly de-
pressed, with a distinct margin. Leaves near stem cankers
may wilt and become chlorotic. Stems then darken and be-
come peppered with small black pycnidia. Symptoms may
worsen after harvest (Gitman & Malikova 1933).By Novem-
ber, thin black "zone lines" form within stalks (Fig 5.29).
Perithecia extend tiny spike-like beaks to the stem surface.

CAUSAL ORGANISM & TAXONOMY

Phomopsis arctii (Saccardo)Traverso, Fl. Ital. Cnypt. 2:226,1906.
= P h a arctii Saccardo 1882; ?=Phomopsiscannabina Curzi 1927.
teleomorph: Diaporthe arctii (Lasch) Nitschke, Pyrenomycetes
Germanici 1:268, 1867; =Diaporthe tulasnei Nitschke f. cannabis
Saccardo 1897.
Description: Pycnidia subglobose to lens-shaped, immersed
then erumpent, ostiolated (20 pm diameter), upper peridium black
and stromatic, lower peridium parenchymous and pale sooty-brown,
200-450x 100-190 pm. Conidiophores conical to cylindrical, less than
10 pm in length. Conidiogenous cells phialidic, cylindrical to
obclavate. a-conidia fusifoim, straight orslightly curved, hyaline,
Figure 5.28: Perithecium of Chaetomium succineum with usually one septum and biguttulate, 7-10 x 2-3 pm (up to 11.5x 4.2
extravagant ornamental hairs (LM x150, McPartland). pm apud Gitman & Boytchenko). P-conidia unicellular, filiform-ha-
mate; 18-25 x 1 pm. Perithecial pseudostroma widely effuse over
stem surface, appearing blackened, carbonaceous; ventral margin
Chaetomium murorum Corda 1837, Chaetomium trilaterale of pseudostroma delimited by a narrow dark-celled
Chivers 1912 (=Chaetornilim aliretim Chivers 1912), and prosenchymatous "zone line." Perithecia immersed in stroma, glo-
Chaetomium piluliferum Daniels 1961.Characteristics of all bose to slightly flattened, solitary or gregarious, 160-320 pm in di-
six Chaetornitirn species are compared in Table 5.2. ametre with a conical ostiolated beak 280-480 pm tall. Asci
unitunicate, clavate,with an apical refractive ring, eight-spored,40-
DIFFERENTIAL DIAGNOSIS 60 x 6-10 pm. Paraphyses elongate, multiseptate,branching at base,
C. murortim, C. elaturn and C. pillilifertim resemble each deliquescing at maturity. Ascospores hyaline, biseriate, guttulate,
one septum whenmature, slightly constricted at the septum,fusoid-
other. The former is distinguished by ascospores bearing a ellipsoidal, straight or slightly curved, 10-15 x 2.5-4.0 pm.
longitudinal furrow. The latter is distinguished by the pres-
ence of its anamorph, Botyotrichtim pill~lifert~m Saccardo &
DIFFERENTIAL DIAGNOSIS
March. C. elattim produces hairs with dichotomous branch-
Phomopsis cannabina and Diaporthe tlilasnei f. cannabis are
ing. C. sliccineum is characterized by its loose cluster of am-
synonyms of D. arctii (McPartland 1995e).Researchers have
ber-coloured hairs. C. globoslim stands apart by its "permed
confused the fungus causing Phomopsis stem canker with a
hair" appearance.
different Phomopsis causing white leaf spot (Sohi & Nayar
Leaf spots described by Chandra (1974) could be
1971, Ghani & Basit 1976, McPartland 1983a, Gupta RC 1985).
confused with leaf diseases caused by Phoma, Ascochyta,
White leaf spot afflicts living leaves, and has not been iso-
Septoria, Colletotrichlirn,and Cercospora fungi, or Psetidomonas
lated from stems. See the following section.
and Xanthomonas bacteria. Microscopy tells them apart.

DISEASE CYCLE & EPIDEMIOLOGY

Chaetomilim species overwinter as perithecia in soil or
crop stubble. p he; are excellent saprophytes and thrive on
many organic materials. Ascospores are water-dispersed.

CONTROL
Control of Chaetomilirn is a simple matter of maintain-
ing vigourous plants. Stress-free plants will not succumb to
these fungi. Dead plants are another matter; anything in con-
tact with soil is subject to attack. See Chapter 8 regarding
retting and rotting of hemp fibres. The fungus Dicyma
pulvinata has been used experimentally as a biocontrol
against Chaetomitirn species (Jeffries &Young, 1994).

PHOMOPSIS STEM CANKER

This disease arises on senescent plants in late autumn.
It has been described on hemp in Italy (Curzi 1927, DeCorato
1997), the former USSR (Gitman & Malikova 1933, Gitman
& Boytchenko 1934), and Illinois (Miller et al. 1960). Her-
barium specimens (e.g., IMI no. 128315) show the disease Figure 5.29: surface of hemp stem blackened by
also strikes drug varieties in India (McPartland 1983b). In Phomo~sisstem canker, with patch of surface shaved
1995, Clarke collected a specimen on hemp stems in China, (arrow) to expose pockets of perithecia, and stem split to
coinfected with Colletotricht~mdematitim. reveal zone lines (McPartland).
128 Hemp Dis:eases and Pests

DISEASE CYCLE & EPIDEMIOLOGY globose to elliptical, ostiolate, peridium textura angularis-globosa,120-
P. arctii overwinters as pycnidia or perithecia in plant 220 x 120-300 pm. Conidiophores cylindrical and slightly tapering
debris. The anamorph is found more frequently than the towards the apex, simple or branched, 8-15 x 1-2 pm. Conidiog-
enous cells phialidic, cylindrical to obclavate. a-conidia hyaline,
teleomorph. Both conidia and ascospores are spread by wa- unicellular, fusiform to elliptical, biguttulate, usually 7-8 x 2.5 pm.
ter and wind-driven rain. Although the fungus is consid- j3-conidia hyaline, unicellular, filiform, mostly curved, 16-22 x 1.0
ered a weak pathogen, it infests many Compositae, pm (Fig 3.2). Cultures deposited at ATCC (#52587) and CBS
Umbelliferae, and Urticaceae in Europe and North America. (#180.91).

CONTROL Several cultural methods are useful (see Chap- DIFFERENTIAL DIAGNOSIS
ter 9)-methods 1 (sanitation), 2a (deep ploughing), 3 (weed- White leaf spot can be confused with yellow leaf spot
ing), 7 (avoid drought), and 5 (genetic resistance). Gitman & and brown leaf spot. Yellow leaf spot produces darker, more
Malikova (1933) noted that damp storage conditions accel- angular lesions peppered with randomly-arranged pycnidia.
erate the fungal destruction of harvested stems. Brown spot lesions are smaller, darker, and break apart leav-
Gitman & Malikova (1933) listed several resistant fibre ing holes in leaves.
varieties. DeCorato (1997) tested four fibre varieties: 'Yellow Ghani & Basit (1976) collected P. ganjae in Pakistan and
stem' (from Hungary) was the most susceptible, followed initially called it a Phoma species. They subsequently made
by 'Shan-ma,' 'Chain-chgo' (from China), and 'Foglia a second mistake by calling the fungus Phomopsis cannabina.
pinnatofida' (from Italy), but these differences were not P. cannabina is a synonym of P. arctii and causes Phomopsis
statistically significant. stemcanker (described in the previous section). P ganjae and
The biocontrol agent Streptomyces griseoviridis controls P. arctii are distinguished by differences in morphology
Phomopsis stem rot in other crops. It is described under (smaller pycnidia and conidia), modifications in
damping off fungi. Bordeammixtureinhibits Diaporthe/Phomap- pathogenicity (attacking young vs. senescent plants; leaves
sis species on other crops. vs. stems) and the inability of P. ganjae to form a teleomorph
in culture or on the host (McPartland 1983b).

WHITE LEAF SPOT DISEASE CYCLE & EPIDEMIOLOGY

P. ganjae overwinters in crop debris. Conidia exude from
White leaf spot has appeared in India (Sohi & Nayar pycnidia in slimy cirrhi, and spread by water dispersal. a-
1971, Gupta RC 1985),Pakistan (Ghani & Basit 1976),Illinois conidia germinate to form appressoria. Appressoria directly
(McPartland 1983b, 1984), and Kansas (McPartland 1995a). penetrate leaf epidermis. P-conidia of P ganjae sprout short
The fungus attacks fibre and drug varieties. germ tubes but do not form appressoria; their function re-
mains uncertain (McPartland 198313).Uecker (pers. commun.
SYMPTOMS 1994) studied the genetics of P. ganjae; he applied molecular
Symptoms begin as pinpoint white leaf spots on young techniques reported by Rehner & Uecker (1994). P. ganjae
plants in late spring. Spots enlarge, remain irregularly circu- was related to Phomopsis isolate no. 649 reported in Rehner
lar, and become slightly raised or thickened. They may
& Uecker (1994).
darken to a beige colour. Black pycnidia arise in concentric
rings within spots (Plate 68). Leaf spots coalesce together,
leaf tissue between spots becomes chlorotic and necrotic, and CONTROL
leaves drop off (Fig 5.30).White leaf spot rarely infests flow- Follow methods utilized for control of yellow leaf spot.
ering tops; THC and CBD inhibit the fungus (McPartland Male plants are more susceptible to P. ganjae than females
1984).White leaf spot's ability to completely defoliate plants (McPartland 1984)-so consider rouging males. Many Pho-
attracted biocontrol researchers (Ghani & Basit 1976). mopsis species are seedborne (M. Kulik, pers. cornrnun. 1988);
do not use seeds from infected females.
CAUSAL ORGANISM & TAXONOMY
Phomopsis ganjae McPartland, Mycotaxon 18:527-530,1983.
Description: Pycnidia immersed in stroma then erumpent, PEPPER SPOT
Pepper spot appears on Russian hemp (Gutner 1933,
Gitman & Boytchenko 1934)and Nepali plants (Bush Doctor
1987a).The report by Nair & Ponnappa (1974) from India is
a misidentification (McPartland 1995a).
Gutner (1933) described symptoms as round yellow-
brown leaf spots 2 4 mm in diameter, with pseudothecia
appearing as tiny black dots within the spots. According to
Gitman & Boytchenko (1934), pseudothecia form on both
sides of leaves.

CAUSAL ORGANISM & TAXONOMY

Leptosphaerulina trifolii (Rostrup) Petrak, Sydowia 13:76,1959.
=Sphaerufina trqofii Rostrup 1899; =Pleosphaeuufinacannabina
Gutner 1933.
Description: Mycelium in culture becomes grey and slightly
floccose, pseudothecia arise in black crusts, produced in concentric
rings of satellite colonies growing from the central inoculation point.
Figure 5.30: Healthy seedling (C), flanked by plants Pseudothecia globose, brown, erumpent at maturity, ostiolate,
inoculated with Phomopsis ganjae on left, and nonparaphysate, thin walled, 120-250 pm diameter.Asci bitunicate,
Macrophomomina phaseolina on right (McPartland). ovate, eight-spored, 50-90 x 40-60 pm. Ascospores hyaline
 Chapter 5: Fungal Diseases 129

(becoming light brown at maturity), oval to ellipsoid,smooth, with 2.'Curvularia lunata (Wakker) Boedijn, Bull. Jard. Bot. Buitenz.
three or four transverse septa and zero to two longitudinal septa, 111 13:127, 1933; =Acrothecium iunatum Wakker in Wakker & Went
25-50 x 10-20 pm (Fig 5.31). 1898.
L. trifolii displays a widevariation of spore and pseudothecium teleomorph: Cochiioboius iunatus [as lunata] Nelson & Haasis
morphology, resulting in a large synonymy. To that synonymy, 1964;=Pseudocochiiobolusiunatus (Nelson& Haasis) Tsuda et ai. 1977.
McPartland (1995e)added Pieosphaerulina cannabina. Description:Conidiophores up to 650 pm long. Conidia three-
septate, third cell from the base is curved and larger and darker
brown than other cells, fourth cell (end cell) nearly hyaline, surface
smooth to verruculose, 13-32 x 6-15 pm (Fig3.2).Pseudothecia soli-
tary or gregarious, black, ellipsoidal to globose with a tall beak,
ostiolate, up to 700 pm in height including a 210-560 prn long beak,
up to 530 pm in diameter.Ascibitunicate, cylindricalto clavate, with
a short stipe, one to eight spored, 160-300 x 10-20 pm. Ascospores
filiform, 6-15 septate, hyaline, arranged either straight, or coiling in
a helix within asci, 130-270 x 3.g6.5 wm.

DIFFERENTIAL DIAGNOSIS
C. ltinata conidia are smaller, less septate, and more
curved than C. cymbopogonis conidia. They also lack the
prominent hilum present in C. ymbopogonis conidia.

DISEASE CYCLE & EPIDEMIOLOGY

Both species overwinter in plant debris near the soil line.
Infectionbegins in warm, wet spring conditions. C. cymbopo-
gonis causes blights on dicots, monocots, and gymnosperms
Figure 5.31 : Asci and ascospores of Leptosphaerulina around the world. Its homothallic pseudothecia have only
trifolii from Russian hemp (LM x500, McPartland). been seen in culture. C. lt~natais cosmopolitan, and normally
causes blights on monocots. On rare occasions, C. llinata in-
fects humans. In one celebrated case, Schwartz (1987) won-
DIFFERENTIAL DIAGNOSIS
dered if the source of a patient's C. ltinata sinusitis was con-
Pepper spot may be confused with brown leaf spot or
taminated marijuana. Brummund et al. (1987) replied, "no C.
early stages of yellow leaf spot. Another Leptosphaertilina
ltinata has been cited in any Cannabis research we have seen."
species, L. chartartim, rarely occurs on Cannabis and is de-
scribed under Pithomyces chartanim.
CONTROL
Disease can be controlled with cultural methods 1(sani-
DISEASE CYCLE & EPIDEMIOLOGY
tation), 4 (escape cropping), and 7c (avoiding excess humid-
L. trifolii attacks a wide range of plants. It overwinters
ity). Researchers have controlled C. llinata disease in other
as pseudothecia embedded in dead leaves. In the spring, as-
crops by spraying extracts of Cannabis leaves (Pandey 1982,
cospores germinate and directly penetrate leaf epidermis.
Upadhyaya & Gupta 1990). Purified THC and CBD inhibit
CONTROL C. ltinata (Dahiya & Jain 1977).
Avoid overcrowding-crop losses increase in dense
stands (Smithet al. 1988).Sanitation is the key to control since
no effective fungicides are known. Cultural methods 1, 2a,
and 10 are most important.
PHYLLOPLANE FUNGI
Phylloplane fungi live in nooks and crannies of leaves,
but cause no disease. They feed on cellular leakage oozing
from plant epidermis. They may also feed on aphid honey-
CURVULARIA BLIGHT dew, pollen grains, and other airborne contaminants. Phyl-
Litzenberger et al. (1963) described a "Ct~rvtilariaspe- loplane fungi exist as epiphytes (living above the leaf epi-
cies" causing leaf spots on Cambodian Cannabis. Babu et al. dermis) or endophytes (living in spaces below the epidermis).
(1977) recovered Ct~rvtilarial ~ i n a t afrom Indian seeds. Some epiphytes, such as sooty moulds, indirectly harm
McPartland & Cubeta (1997)isolated Ctirvtilaria cymbopogonis plants by blocking sunlight and reducing photosynthesis.
from seeds in Nepal. No symptoms were described in any Other epiphytes protect plants by suppressing or destroying
of these reports. pathogenic fungi. These helpful epiphytes thus attract
attention as possible biocontrol organisms (Fokkema & Van
CAUSAL ORGANISMS & TAXONOMY den Heuvel1986).
1. Curvularia cymbopogonis (C.W.Dodge) Groves & Skolko, No one has systematically investigated Cannabis phyl-
Canadian I. Research 23:96,1945; =Heiminothosporiumcymbogonis [as loplane fungi. One protective epiphyte, Aureobasidium
cymbogoni] C.W. Dodge 1942. pullulans (deBary) Arnaud, appears in the hemp literature
teleomorph:Cochiioboius cymbopogonis Hall & Sivanesan, Trans. (Lentz 1977, Ondrej 1991).Many dematiaceous "pathogens"
BY.mycol. Soc. 59:315,1972. cited by Ponnappa (1977) and Gzebenyuk (1984), such as
Description: Conidiophores simple, septate,brown, up to 300 Alternaria alternata, Cladosporitim herbartim, Epicocclim nignlm,
pm long. Conidia acropleuroge~ious, smooth, straight or curved,
clavate to ellipsoidal, four (sometimes three) septa, averaging 40- and Stemphylitim botyostim, can also exist as nonpathogenic
50 x 12-15 pm, middle cells dark brown, base cell and end cell paler, phylloplane fungi (Fokkema & Van den Heuvel 1986). As
base cell obconical with a protuberant hilum. Pseudothecia scattered the old saying goes, "one plant's protective phylloplane
or aggregated in concentric zones on agar, black, globose with a fungus is another plant's latent pathogen" (paraphrased from
long cylindrical beak, up to 575 pm diameter. Asci bitunicate, Palm 1999).
cylindrical with a short stipe, eight-spored, 210-275 x 1523 ym. Control of phylloplane fungi is neither necessary nor
Ascospores filiform, 8-14 septate, hyaline, 195420 x 3.5-4.5 pm. advisable. Fungicides kill epiphytes and cause a "rebound"
130 Hemp Diseases and Pests

of pathogenic organisms. As in human medicine, this is trients'are immobile in soil, which causes a nutrient-deple-
termed "iatrogenic" disease. Heavy aphid or whitefly tion zone to form around roots. Mycorrhizal fungi grow be-
infestations may cause an overgrowth of sooty mould, which yond the root zone and draw nutrients back to the plant.
is best eliminated by controlling the insects. These fungi form an extensive network-Tisdall & Oades
measured about 150 feet of mycelium per cm2 of soil.
Phosphate ions are the most immobile soil nutrients.
Menge (1983)found that mycorrhizae-infected plants absorb
MYCORRHIZAE 60 times more P than uninfected plants. Uptake of zinc and
About 120 years ago, scientists discovered that some copper dramatically increases, and absorption of K, Ca, Fe,
fungi invaded plant roots without causing diseases. We now Mg, Mn and S also improves. Only the absorption of nitro-
recognize these strange mould-plant relationships as sym- gen (the most mobile soil nutrient) remains unchanged in
biotic, not parasitic. Termed "mycorrhizae" (Latin for "fun- the presence of mycorrhizae. Nitrogen-fixingAzotobacter spe-
gus-roots"), these associations occur in almost all plants and cies, however, synergistically increase plant growth when
are very important. Mycorrhizae nevertheless have escaped inoculated with mycorrhizal fungi (Linderman et al. 1991).
wide attention, because infected roots look normal and the The mycorrhizal mantle protects plants from some root-
fungi themselves grow with difficulty in artificial culture. feeding insects, nematodes, and many fungal pathogens,
includingFtlsarium oxysponlm, Ft~sariurnsolani, Macrophomina
CAUSAL ORGANISMS phaseolina, Pythium ultirnttm, Rhizoctonia solani, and Sclerotium
Mycologists describe two classes of mycorrhizae. iolfsii. ~ ~ c o r r h i z also
a e aid plants by reducing drought
~ctom~corrhizae associate with tree species, especially gym- stress, and even produce plant growth hormones.
nosperms. Endomycorrhizae associate with trees and her-
baceous plants. ~ndom~corrhizae are usually Zygomycetes. Application for crops
~ndomicorrhizaeproduce swellings (vesicles) or minute Mycorrhizal fungi serve as "biotic fertilizers," substi-
branches (arbuscules) within plant cells. Botanists call them tuting for nutrient supplements. Optimizing the growth of
VA mycorrhizae. mycorrhizal fungi requires a balance of proper pH, mois-
In 1925 Arzberger photographed mycorrhizae in Can- ture, light intensity, soil fertility, percentage of organic mat-
nabis roots. He died shortly thereafter, and his findings were ter, and soil flora and fauna-not a project for the neophyte
never reported. We recently rediscovered ~ r z b e r ~ e rnotes
's grower. Hayman (1982) described two pointers for begin-
and glass plate negatives in the USDA archives (Fig 5.32). In ners: adding organic fertilizers to soil improves mycorrhizal
1961 Mosse produced an artificial VA mycorrhizal relation- growth, whereas adding petrochemical fertilizers (e.g., am-
ship in Cannabis by inoculating roots with "an Endogone spe- monium nitrate) decreases mycorrhizal growth.
cies." McPartland & Cubeta (1997) documented naturally- Pesticides raise questions of practical importance. Some
occurring VA mycorrhizae in feral hemp. They identified the growers report stunting of plants after soaking soil with
fungus as a Glomus species, probably Glomus mosseae (Nic. fungicides. They attribute this to chemical toxicity. Review
& Gerd.) Gerd. & Trappe (Plate 69). of the symptoms, however, suggests phosphorus
.,
deficiency-the funnicides destroyed the mvcorrhizae
(reducing uptake of phosphorus).
Menge (1983) tested pesticides on several Glomus
species, including G. mosseae. He found six pesticides lethal
to mycorrhizae: methyl bromide, metam-sodium,
chloropicrin, formaldehyde, PCNB, and thiram. Benomyl
was only lethal as a soil drench-if sprayed on foliage it
caused little mycorrhizal destruction. Maneb was interme-
diate in lethality. Captan, terrazole, copper sulphate, and
nematocidal fumigants caused little damage and may have
improved mycorrhizal growth by eradicating mycorrhizal
parasites and predators.
Why isn't crop stunting a universal phenomenon
following heat sterilization of soil? Menge (1983)noted that
VA mycorrhizae survive deep in organic debris, insulated
from damage. In addition, earthworms, insects, and small
mammals carry mycorrhizae back into sterilized areas.
Almost all soils contain natural populations of
mycorrhizal fungi. Exceptions include soils laying fallow for
two or more years, and soils supporting continuous crops
of non-mycorrhizal plants such as Cruciferae (broccoli,
cabbage, cauliflower, mustard greens, turnips, etc.) and
Chenopodiaceae (quinoa, lamb's quarters, pigweed). Plastic
Figure 5.32. "Endotrophic mycorrhiza on roots of sacks of sterile potting soil, peat moss, builder's sand, and
Cannabis sativa" (photo by E.G. Arzberger, ca. 1925). perlite also lack mvcorrhizae.
Can we inoculate sterile soil with mycorrhizal fungi?
EPIDEMIOLOGY Some cultivators throw in a handful of "starteru-soil and
Merlin & Rams Das (1954) discovered that metabolites root fragments from a previously S U C C ~ S S ~Crop.
U ~ Wilson et

produced by Cannabis roots enhance the growth of mycor- (1988) compared this "sourdough starter" with Pure
rhizal fungi. In return, mycorrhizal fungi improve Cannabis mycorrhizae spores: after four months, test plants
growth by increasing the surface area of the root network (Andf'opogon gerardii) grown in sterile soil were stunted. Av-
and making several soil nutrients more available. Some nu- erage dry weight was only 0.02 g. Plants grown in sterile
 Chapter 5: Fungal Diseases 131

soil inoculated with pure spores weighed a n average of 5.40 mostly long, cylindrical, flexous, simple, septate, hyaline, 5-65 x 1-
g, or 270 times greater than plants g r o w n i n sterile soil. Plants 1.5 pm. Conidia borne on short sterigmata along the lengths of
inoculated with the sourdough method averaged 3.71 g each, conidiophores, lens-shaped, golden-brown with a hyaline band
around the perimeter, 8-12 pm diam.
185 times greater than plants i n sterile soil.
Persoon's b a s i o n y m a p p e a r s t o h a v e priority. A.
M e n g e (1983) harvested p u r e s p o r e s f r o m roots of
"mother plants." H e ground u p t h e roots a n d recovered the pkaeosperrn~irngrows worldwide o n sedges a n d reeds (Carex,
spores b y wet-sieving o r centrifugation. Subsequent disin- Glyceria, Phragmites species). Chandra (1974) described A.
fection of spores with streptomycin a n d s o d i u m hypochlo- pkaeosperrnllm erupting from C. sativa leaf spots in India. Spots
rite (bleach) assures a p u r e inoculum. The mycorrhizal fun- t u r n light-brown, gradually necrose, a n d finally collapse,
g u s Glomlls intraradices is n o w commercially available a s a leaving irregular holes i n the leaf.
biocontrol against Pythium a n d Fzlsaritlrn species (described
in the section o n Fusarium stem canker). Menge estimates Botyosphaeria obtusa (Schweinitz) Shoemaker, Canadian 1.
the cost for mycorrhizal inoculation equals current costs for Botany 42:1298,1964; =Physalospora obtusa (Schweinitz) Cooke 1892.
anamorph 1: Sphaeropsis malorum Peck, nomen nudum;
phosphorus application.
=Sphaeria cannabis Schweinitz 1832.
In hydroponic systems, a constant flow of soluble nu-
anamorph 2: unnamed.
trients around roots eliminates t h e nutrient depletion zone. Description: Pseudothecia embedded in stems, stromatic, dark
Nevertheless, some hydroponics operators inoculate their brown-black, solitary or gregarious in botryose aggregations up to
systems with mycorrhizae t o optimize nutrient uptake. Ojala 3 mm wide; individual locules globose, 150-300 pm diameter. Asci
& Jarrell (1980) reviewed techniques for establishing myc- bitunicate, clavate, eight-spored, 90-120 x 17-23 pm. Paraphyses
orrhizae in hydroponic systems. filiform. Ascospores broadly fusoid, hyaline, unicellular or
sometimes with one septum, 25-33 x 7-12 pm. Anamorph 1pycnidia
are immersed then erumpent, stromatic, black, on Cannabis
appearing unilocular and globose, averaging 235 pm diameter. Co-
MISCELLANEOUS LEAF, nidiogenous cells hyaline, simple, cylindrical, holoblastic, discrete,
determinate, 8-14 pm in length, 3-4 pm in width. Macroconidiaborne
STEM AND ROOT DISEASES in mucilage, elliptical or broadly clavate, base often truncate and
The 26 pathogens presented i n this section rarely cause bordered by a scar, thick-walled, verruculose to almost smooth-
disease. They don't even h a v e common names. They have walled, at first hyaline and unicellular with a large central guttule;
only been mentioned once o r twice i n t h e literature. Many rapidly becoming brown, at length becoming two-celled and
biguttulate, 19-21 x 10-11 pm (Fig 5.33). Anamorph 2 (the
lack herbaria-preserved voucher specimens, a n d their cor-
microconidial state) is rarely encountered; for a description see
rect identification is questionable. They are briefly described McPartland (199513).
and discussed: Historically, Schweinitz's 1832 publication of Sphaeria
cannabis is t h e first description of a Cannabis pathogen.
Arthrinium phaeospermum (Corda) M.B. Ellis, C.M.I. Myco- McPartland (1995b) examined Schweinitz's specimen and
logical Papw No. 103, ~3,1965;=Papulavia sphuerosperma (Persoon:Fries)
synonymized it w i t h the a n a m o r p h of B. obtzisa. Stevens
Hohnel1916.
Description: Hyphae hyaline to pale brown, smooth or (1933) provided a n extensive synonymy for this common
verruculose, septate, 1-6 pm thick. Conidiophore mother cells short, pathogen.
lageniform, smooth or verruculose, 5-10 x 3-5 pm. Conidiophores
Coniothyrium cannabinum Curzi, Atti lstit. got. Univer. Pavia,
ser. III(3):206,1927.
Description: Pycnidia scattered along stems, immersed then
erumpent, spherical or flattened from above, ostiole somewhat
sunken, sooty brown-black, peridium consisting of small parenchy-
mous cells finely woven together, 90-120 x 60-90 p.Conidiophores
scarcely apparent. Conidia thick-walled, olive brown, almost spheri-
cal to oval, with one large central guttule, 4-5 x 2.5-3.5 pm (Bestagno-
Biga et al. 1958 described conidia 5-5.5 x 2.5km).
Curzi's description of "scarcely apparent" conidiophores
suggests the determinant character of Microsphaeropsis phialides. But
the thick-walled conidia suggest a C o n i o t h y r i t ~ mspecies.
Unfortunately, Curzi's type specimen is missing (Curator Dr.ssa
Terzo, pers. commun. 1987).
C. cannabinum infests h e m p i n Italy ( C u r z i 1927,
Bestagno-Biga et al. 1958) a n d Russia (Gitman & Boytchenko
1934). G z e b e n y u k (1984) c i t e d t w o o t h e r s p e c i e s ,
Coniothyrium tenue Diedicke a n d Coniothyrium olivaceum
Bonorden, which a r e probably misheterminations
(McPartland 1995a). Fuller & N o r m a n (1944) reported a n
unspeciated Coniothyrium attacking field-retted hemp stalks
i n Iowa. N o disease symptoms are described i n any of these
reports. Coniothyritlrn species overwinter i n crop debris. Co-
niothyriurn diseases of other crops (notably Coniothyrillm
filckelii o n roses) are controlled b y avoiding injury t o stem
surfaces.

Figure 5.33: Sphaeria cannabis (=Botryosphaeriaobtusa) Cylindrosporium cannabina Ibrahimov, Akademii Nauk
macroconidia. A. LM optical section showing large internal Azerbaidzhanskoi SSR. Izvestiya 466-67,1955.
guttule, ( ~ 2 3 0 0 )B.; SEM showing surface details ( ~ 1 7 0 0 ) ; Description: Acervuli epiphyllous, gregarious, barely conspicu-
McPartland. ous, 55-96 pm wide. Conidiophores small, hyaline, cylindrical-
132 Hemp Diseases and Pests

acuminate, crowded together. Conidia unicellular, filiform, hyaline, H y m e n o s c y p h u s herbarum (Persoon:Fries)Dennis, Persoonia
curved, rarely straight, 18-54 x 1.2-2.0 pm. 3:77,1964; =Helotium herbarum (Persoon:Fries)Fries 1849.
Ibrahimov discovered this species causing leaf spots in Description: Apothecia common, gregarious, arising on a very
Azerbajian. Leaf spots are yellow-brown or cinnamon short stalk, pale yellow to ochraceous, minutely downy, flat to
slightly convex, disc up to 3 mm in diameter. Asci cylindrical to
coloured, 0.1-6 mm in diameter. Ibrahimov's illustration of
clavate, stalked, apical pore turns blue with iodine, eight-spored,
cupulate acervuli resembles Briosi & Cavara's illustration of up to 90 x 8 pm. Paraphyses hyaline, filiform, 90 x 2 pm. Ascospores
Septoria neocannabina. Ibrahimov's specimen could be a Sep- two-celled, hyaline, often biseriate, fusiform to cylindrical, 13-17 x
toria, Septogloetim, or Phloeospora species. 2.5-3.0 pm.
McPartland (1995a)examined the herbarium specimen Saccardo found H,herbartim fruiting on Cannabis stems
of "Cylindrosporium species" cited by Miller et al. (1960) and near Padova, Italy. He distributed exsiccati specimens as
Farr et al. (1989).The specimen proved to be a mix of Septoria Mycotheca italica No. 119 (BPI!). H. herbartim infests many
neocannabina and Psetidocercospora cannabina. Ghani & Basit herbaceous hosts, especially Urtica species. The fungus lives
(1975) described a Cylindrosporitlm species forming a black- worldwide in temperate climates. It sporulates on dead stems
ish fluffy mycelium on undersides of leaves in Pakistan. They in October, spreading ascospores by wind and water. The
described conidia as filiform, two to four septate, and bent fungus's r61e as a pathogen is questionable.Most mycologists
at the apex. consider H. herbartim a saprophyte, although some
Hymenoscypht~sspecies (e.g., H. ericae) act as mycorrhizae.
D i d y m i u m c l a v u s (Albertini & Schweinitz) Rabenhorst,
Deutsckland Kyptogamen Flora 1:280,1844. Lasiodiplodia theobromae (Pat.) Griffon & Maublanc, Bull.
Description: Plasmodium grey or colourless.Sporangia stalked trimest. Soc. mycol. France 25:57, 1909; =Botryodiplodia theobromae
(sometimes appearing sessile), discoid, greyish white with darker Patouillard 1892.
stalk, 0.5-1 mm diameter, up to 1mm tall. Peridium (cap) membra- teleomorph: Botyosphaeria rhodina (Berkeley & Curtis in Curtis
nous, adaxial surface nearly covered with light lime crystals, under apud Cooke) von Arx 1970; =Physalospora rhodina Berkeley &Curtis
surface without lime, brown. Stalk tapers upward, longitudinally in Curtis apud Cooke 1899.
striate, dark brown to black, paler near top. Capillitium (threads) Description: Pycnidia stromatic, simple or compound (uni-
delicate, hyaline to pale purple, sparsely branched. Spores black en or multilocular), often aggregated, carbonaceous black, ostiolate,
masse, pale violaceous brown under the microscope, nearly smooth, between 300 and 5000 pm in diameter. Conidiogenous cells hya-
6-8 pm diameter. line, simple, cylindrical, sometimes septate, rarely branched, holob-
Gzebenyuk (1984) cited D. clavus covering 4.6% of hemp lastic, 5-15 x 3 pm. Paraphyses hyaline, cylindrical, septate, up to
stems near Kiev. D. clavtls occurs across Eurasia and North 50 pm long. Conidia at first ellipsoid, hyaline and thin-walled, then
America. Slime moulds may crawl up plant stems in wet thick walled, later developing a median septum, dark brown pig-
weather, but cause little damage. mentation, longitudinal striations, and a truncate base, 20-30 x 10-
15 pm (Fig 3.2). Teleomorph perithecia stromatic, uni- or multilocu-
lar, gregarious. Asci bitunicate, clavate, eight-spored, 90-120 x 15-
Jahniella bohemica Petrak, Annales Mycologici 19:123,1921.
28 pm. Ascospores one-celled,hyaline, ellipsoidal, 24-42 x 7-18 pm.
Description: Pycnidia unilocular, brown to black, immersed
then erumpent, flattened subglobose, ostiolated, (310-) 600 (-850) McPartland (1995b)found L. theobromae infesting stems
pm diam., up to 400 pm tall, thick walled (50-60 pm thick), of feral hemp in Illinois. The hemp was also parasitized by
scleroplectenchymatic,peridium textura angularis. Ostiolum central, Ftisaril~mstilpht~rezim(see Plate 58). L. theobromae prefers
papillate, up to 110 pm tall. Conidiogenous cells holoblastic, dis- climates between 40" north and 40" south of the equator, but
crete, determinate, ampulliform to short cylindrical, 4-14 x 2-5 pm. is distributed worldwide. It decays harvested mangos,
Conidia hyaline, filiform, straight or curved, base truncate, smooth- bananas, avocados, melons, cocoa pods, citrus, and cotton
walled, finely guttulate, indistinctly (2-)3-4(-5) septate, (18.5-) 45
bolls (Holliday 1980).
(-55) x 1.0-2.5 pm.
Saccardo & Roumeguere (1883) misdetermined J. bohe-
Leptosphaeria acuta (Hoffman:Fries) P. Karsten, Mycologia
mica as Leptosphaeria actGa, on hemp stems near Leige, Bel-
Fennica 298,1873; =Sphaeria acuta E. F. Hoffman 1787,=Cryptosphaeria
gium. McPartland (1995e)inspected their specimen and rec- acuta (Hoffman:Fries)Greville Fl. Edin. p. 360,1824; ?=Leptosphaeria
ognized the mistake. For a full account see McPartland & acuta f. cannabis Roumeguere 1887.
Common (2000).J. bohemica has also been collected from fig- anamorph: Phoma piskorzii (Petrak) Boerema & Loerakker,
wort (Scrophl~larianodosa) and garden loosestrife (Lysimachia Persoonia 11:315,1981; =Diploplenodomuspiskorzii Petrak 1923, non:
vulgaris). Phoma acuta auct., nomen ambiguum.

Figure 5.34: Leptosphaeria acuta. A. Pseudothecia on stem, LM x40; B. Sectioned pseudothecium showing asci and
minute ascospores, LM x105; C. Ascospores, LM x410 (all courtesy Bud Uecker).
 Chapter 5: Fungal Diseases 133

Description: Pseudothecia conlcal, surface smooth, glistening Nair & P o n n a p p a (1974, P o n n a p p a 1977) isolated the
black, immersed then erumpent, ostiolate, peridium a thick-walled a n a m o r p h from marijuana leaves i n India. N o symptoms
textura globt~losanear ostiole, thereafter a radiating textuva prisrnatica, w e r e described. The f u n g u s attacks m a n y plants. It produces
300450 pm wide, beak 100-200 tall, topped by a 3 0 4 0 pm diam- a n oligopeptide, sporidesmin, which causes eczema in mam-
eter ostiole (Fig 5.34).Asci cylindrical, eight-spored, base constricted
mals (Rippon 1988).
to a short pedicle, 90-140 x 7-11 pm (Fig 5.34). Paraphyses filiform,
hyaline, septate. Ascospores pale yellow, guttulate, fusiform, usu-
ally seven-septate and constricted at central septum, 35-45 x 4-7 Leptospora m b e l l a (Persoon:Fries) Rabenhorst, Herb. ilu, nlycul
pm. The anamorph is described under P. piskovzii in the section on Ed. 11, No. 532,1857; =Ophiobolus poylzyrogonus (Tode)Saccardo 1883;
Brown leaf spot and stem canker. =Ophiobolus oulgnris (Saccardo) Saccardo 1881.
L. aciita is usually restricted to Urtica species (Shoemaker Description: Pseudothecia appear on blackened or purple-red
1964, Sivanesan 1984, C r a n e & Shearer 1991). Saccardo & stems, scattered or gregarious, immersed then more-or-less
erumpent, globose to conical, often laterally compressed, black, gla-
Roumeguere (1883) reported L. acuta o n h e m p stems a n d
brous, papillate; contain periphyses, paraphyses, and asci; 200-300
distributed a n exsiccatus, Reliquiae Libertianae n o . 56 (191). , ,
pm diameter. Asci bitunicate, borne upon short stalks, cylindrical,
Four years later Roumeguere eiected t h e taxon Leptosphaeria eight-spored, 140-160 x 4.5-6.0 pm (Holm says asci grow to 225 pm
aclita f. cannabis a s a n exsiccatus, Filngi selecti exsiccati n o . long). Ascospores slightly spiralled within the ascus, filiform, yel-
4172. We e x a m i n e d s e v e r a l exsiccati specimens-some low, with many obscure septa and guttules, nearly as long as asci
contain i m m a t u r e , unidentifiable ascomata (at BPI-see and 1 pm wide.
McPartland 1995e); other specimens contain misidentified H o l m (1957) listed t e n s y n o n y m s for this c o m m o n
Jniznlclla bohenlica ( a t F H a n d BR-see M c P a r t l a n d & fungus. Gzebenyuk (1984) reported L. rzibella twice o n Soviet
Cornmon 1999). Taxonomically, C r a n e & Shearer (1991) a n d h e m p s t e m s (as t w o synonyms, 0 . porphyrogom~s a n d 0.
McPartland (1995a) recognized the taxon L. acuta (Hoffm.:Fr.) vulgaris). The life cycle of L. riibella is similar t o Ophiobollis
Karsten, whereas Boerema & G a m s (1995) preferred t h e species described u n d e r Ophiobolus s t e m canker.
taxon L. aclitn (Fuckel) Karsten.
M i c r o d i p l o d i a a b r o m o v i i Nelen, Nov. Sist. nlz. Rast. 14:106,1977.
L e y t o s p h a e r i a cannabirza Ferraris & Massa, Annales Mycologici =Microdiplodia cannabina Abramov 1938 or 1939.
10:286, 1912. Description: Pycnidia scattered, thin-walled, papillate, brown, cells
Description: Pseudothecia epiphyllous, globose, black, around the ostiole become black, 130-180 pm diameter. Conidla
erumpent at maturity, ostiolate, 130-140 pm diameter, peridium ("stylospores") olive to pale brown, bicellular, w ~ t hrounded endi,
"membranous" [thin-walled?].Asci clavate, straight or curved, apex 8-11 x 4.0-4.5 pm.
rounded, base constricted to a short pedicle, 45-50 x 7-10 pm. Abramov found this fungus o n stems of male plants in
Pal-aphyses not described. Ascospores honey-coloured, fusiform, Russia. Dobrozrakova et al. (1956) described it again in Russia.
three-septate, slightly constricted at septa, 19-20 x 5 pm. Nelen reduced Abramov's species t o a nonlen n o i l ~ ~ n The z.
Ferraris & Massa f o u n d pseudothecia of L, cannabitza in genus also has taxonomic problems, see Sutton (1977).
irregularly-shaped leaf s p o t s of wilting plants near Alba,
Italy. Leaf spots w e r e dull w h i t e w i t h a n ochre margin, 3-5 M i c r o p e l t o p s i s c a n n a b i s McPartland, Mycological Research
inrn in diameter. The f u n g u s also infests Russian h e m p (Git- 101:854,1997.
m a n & Boytchenko 1934, G i t m a n 1935, Dobrozrakova e f al. Description: Ascomata catathecioid, 45-130 pm diam., flattened
1956).Ferraris & Massa considered L. cannabina a "probable" ampulliform, 25-46 pm high, dark brown to black, ostiolate, margin
teleomorph of Septoria cannabis. T h e y d i d n o t confirm this in entire; upper layer composed of radially-arranged quadrangular
culture. P e r h a p s t h e y f o u n d pycnidia of S . cannabis near L. cells, peridium textt~vaprisnzatica; basal layer of similar construction
c ~ ? n n a b i l lS,
~ ~cailnabis
. is a ubiquitous h e m p pathogen; prox- to upper layer but paler. Ostiole central, raised, composed of small
cells and bearing a crown of divergent setae, whlch are arranged in
lniity docs not coizstitutc a relationship. an inverted cone over the ostlole. Setae straight, subulate, th~ck-
walled, nonseptate, smooth, dark brown, 12-22 pm long. A s c ~
L e p t o s p h a e r i a zuoroninii Docea & Negru i n Negru, Docea bitunicate, ovoid to obclavate, four- to eight-spored, 2140 x 4-9 pm.
& Szasz, Novosti Sisteinntik~Niishikh Rnstenij 9:168, 1972. Ascospores hyaline, guttulate, ellipsoid, with a sii~glemed~ansep-
Description: Pseudothecia flask-shaped, brown-black, tum, nonsetulate, 11-12 x 2.5-3.0 pm.
carbonaceous, Immersed then erumpent, 150-180 pm diameter. Asci Roumeguere's syntype of Orbila luteola (see t w o species
cylindrical to clavate, eight-spored, 58-100 x 18-20 pm. Paraphyses b e l o w ) c o n t a i n e d thyriothecioid f r u i t i n g b o d i e s w h ~ c h
fillform, hyaline, with branching apices. Ascospores fusiform, proved t o b e a n e w species.
slightly curved, four to six septa, slightly constricted at septa,
guttulate, granular cytoplasm, yellow, 23-29 x 3.5-5.0 pm.
M y r o t h e c i u m r o v i d i ~ mTode:Fries, Systema Mycologicurn 3:217,
Docea & Negru (1972) described L. zuoroninii from seeds
1828.
of Cannabis satiaa near Cluj, Romania. Description: Sporodochia sessile or slightly stalked, cushion
shaped, light coloured to dark, diameter variable, 16-750 pm.
Leptosphaerulina c h a r t a r u m Rowc, Tmns. Br. Mycol. Soc. 86:319- Conidiophores hyaline, cylindrical, septate, with multiple branches
323,1986. bearing phialides in whorls. Conidia ellipsoid to elongate, subhyaline
anamorph: Pitllornyces cizartavunl (Berkeley & Curtis apud to olivaceous, aseptate, guttulate, 5-10 x 1.5-3.0 Fm.
Berkeley) M. B . Ellis 1960. Although M. roridum attacks cotton, coffee, a n d m a n y
Description: Mycelium dark olive brown, septate, smooth to other crops a r o u n d the world, it has only been reported o n
verruculose. Pseudothecla globose, dark olive brown, immersed or Cannabis in India (Nair & P o n n a p p a 1974, Ponnappa 1977)
superficial, with a large ostlole, five to seven asci per pseudothecium,
a n d Pakistan (Ghani & Basit 1976). Gzebenyuk (1984) cited a
nonparapliysate. Asci bitunicate, short ovate but elongating with
maturation, eight-spored, 100-150 x 60-100 pm. Ascospores hyaline related a n d easily mistaken f u n g u s o n Ukrainian h e m p ,
to light brown, broadly ellipsoidal, smooth, usually three transverse M y r o t h e c i u m verrilcaria (Albertini & Schweinitz) Ditmar.
septa and one longitudinal septum, slightly constricted at septa, M. roridum causes "corky leaf spot." Symptoms begin
23-27 x 7-12 pm. Conidiophores straight or slightly curved, cylin- a s small brown leaf spots in July. Spots enlarge a n d sometimes
drical, hyaline to subhyaliile, holoblastic, 2.5-10 x 3-3.5 km. Conidia coalesce. Masses of slimy conidia t u r n spots black. U p o n
borne singularly atop conidiophores, light to dark brown, broadly drying, they become corky a n d tough. H e a v y infection leads
ellipsoidal, verruculose to echinulate, usually three transverse septa to defoliation a n d death. M. roridunz overwinters in the soil.
and one longitudinal septum, 18-29 x 10-17 pm.
T h e f u n g u s is a w e a k p a t h o g e n a n d rnvades plants vra
134 Hemp Diseases and Pests

wounds. Optimum temperature for growth is 28-32°C; corky Phyllachora cannabidis P. Hennings, Hedwigia 48:8,1908.
leaf spot flourishes i n subtropical climates a n d w a r m glass- Description: Perithecia lie tightly bunched together, develop-
houses ing in a blackened plano-pulvinate pseudostroma under stem epi-
dermis, perithecial wall membranaceous, brown, 16 pm thick,
ostioles erumpent, 170 pm wide x 120 pm deep. Asci clavate with
Orbila luteola (Roumeguere) ~ c ~ a r t l a nMycological
d, Research
rotund apices, eight-spored, 45-55 x 10-12 pm. Ascospores in one
101:854,1997; =Calloria luteola Roumeguere 1881.
Description: Apothecia superficial, sessile, translucent yellow- or t w o rows, arranged at oblique angles to ascus axis, hyaline to
orange, margin spherical to ellipsoidal, up to 0.6 mm in diameter greyish blue, oval to almost fusiform, nonseptate, 12-17 x 5-6 pm. ,

and 100 ym in thickness. Excipulum consists of hyaline thin-walled Hennings discovered P. cannabidis o n stems near Sao
textura globulosa. Asci cylindrical, eight-spored, 26.0 x 4.5 pm. Paulo, Brazil. Pkyllachora perithecia merge together atop host
Faraphyses hyaline, filiform, slightly enlarged at the apex. epidermis to form a shield-like clypeus. Clypei are shiny
Ascospores hyaline, single-celled, fusiform, some indistinctly a n d black a n d called "tar spots." Theissen & Sydow (1915)
guttulate, 6.5 x 1.5 pm. n o t e d P. cannabidis's p e r i d i u m a n d e x c l u d e d it f r o m
Only Dr. Roumeguere h a s found this fungus o n h e m p Pkyllachora. Since then, however, peridia have been recog-
stems, near Villernur, France. But h e found enough of it t o nized i n Phyllachora species (Parberry 1967).
distribute nearly 100 specimens i n his Ftingi Gallici exsiccati. Phyllachora species are obligate parasites (feeding o n
living cells) a n d cause little host damage. Tar spots rarely
Penicillium c h y s o g e n u m Thom, Bull. USDA Bur. Animal Ind. cause economic losses. Parberry (1967) stated these fungi
118:58,1910; =Penicillium notatum Westling 1911.
have a w i d e host range a n d he proposed that many of the
Description: Conidial heads paint-brush-like, bearing conidia
in well-defined columns up to 200 ym long. Conidiophores smooth, 1100 Pkyllachora taxa described o n different hosts a r e
hyaline, up to 350 pm long by 3.0-3.5 pm wide. Branches (penicilli) synonyms.
usually biverticillate, measuring 15-25 pm by 3.0-3.5 pm. Metulae
borne atop penicilli in clusters of two to five, measuring 10-12 pm Rhabdospora cannabina Fautrey, Bull. Soc. mycol. France 15:156,
long by 2-3 ym wide. Phialides borne atop metulae in compact clus- 1899.
ters of four to six, flask-shaped, mostly 8-10 by 2.0-2.5 pm. Conidia Description: Pycnidia appear on blackened stems, intensely
elliptical to almost globose, smooth, pale yellow to pale green, mostly aggregate, numerous, immersed then erumpent, surface woven and
3.0-4.0 x 2.8-3.5 pm (Figs 3.2 & 8.2). thinly reticulate, grey to black, papillate, with a round ostiole.
Babu et al. (1977) described P. chysogentim "burning" Conidia variously curved, guttulate, 4 0 4 8 x 1.752.0 pm.
leaves of plants i n India. Small branches w e r e also affected, Fautrey isolated this fungus near Semur, France. His de-
causing a wilt. P. c k y s o g e n t ~ malso rots Cannabis seeds a n d scription of R. cannabina conidia resembles t h e conidia of
causes a marijuana storage disease (Fig 8.2). The fungus Septoria cannabis o r Jakniella bohemica (McPartland 1995d).
grows worldwide b u t prefers a n o p t i m u m temperature of Unfortunately, Fautrey's type specimen is missing.
23°C. P. c k y s o g e n ~ i mis a friend a s well a s a foe--Dr. Fleming G z e b e n y u k (1984) r e p o r t e d t w o o t h e r Rhabdospora
discovered "penicillin" w h e n this fungus floated into his lab. species o n h e m p stems: R h a b d o s p o r a origani (Brunand)
Saccardo 1884 a n d Rhabdospora hypochoeridis Allescher
Periconia byssoides Persoon, Synopsis Methodica Fungorum p. 686, 1897. Both species produce conidia smaller than Fautrey
1801; =Periconin pycnospora Fresenius 1850. described for R . cannabina.
Description: Conidiophores are long cylinders topped by swol-
len heads, erect, straight or slightly flexous, two to three septate, Rosellinia necatrix Prillieux, Bull. Sac. mycol. France 20:34,1904
dark brown at base and paling to subhyaline at the apex, 12-23 pm anamorph: Dematophora necatrix Hartig 1883.
diameter at base tapering to 9-18 pm below the head, then forming ~escri~tion:>erithecia densely aggregated globose, brown-black,
a septum with an apical head swelling to 11-28 pm diameter, entire ostiolate, 1000-2000 pm diameter. Asci unitunicate, cylindrical, long-
length 200-1400 pm (up to 2 mm). Conidiogenous cells hyaline, el- stalked, eight-spored, 250-380 x 8-12 pm. ~scosporesfusiform,
lipsoidal to spherical, monoblastic to polyblastic. Conidia cantenate, straight or curved, single-celled,brown, 30-50 x 5-8 pm, with a lon-
forming chains, spherical, brown, verrucose, 10-15 pm diameter. gitudinal slit. Paraphyses filiform. Conidiophores bound together
Gitman & Malikova (1933) described P. byssoides black-
e n i n g h e m p stems near Moscow. G i t m a n & Boytchenko
(1934) noted the conidiophores merged together into corernia.
The fungus infests stems in the Czech Republic (Ondrej 1991).
Fuller & Norman (1944) f o u n d a Periconia species o n h e m p
in Iowa.
Gzebenyuk (1984) reported Periconia c o o k e i Mason &
Ellis o n h e m p stems near Kiev. I? cookei differs little from P.
byssoides-it lacks a s e p t u m a t the apical cell a n d produces
slightly larger conidia (13-16 k m diameter).

Pestalotiopsis species
Description: Acervuli epidermal, often epiphyllous, circular
to oval, up to 200 ym in diameter. Conidiophores cylindrical, sep-
tate, occasionally branched, hyaline, up to 10 pm in length. Conid-
iogenous cells holoblastic, annellidic, hyaline, cylindrical. Conidia
fusiform, five-celled, smooth walled, averaging 24-26 x 5.6 pm; ba-
sal cell hyaline, with a simple hyaline appendage (pedicel) averag-
ing 6.6 pm in length; three median cells umber to olivacous brown,
thick-walled; apical cell hyaline, conical, with three or less commonly
two simple hyaline appendages (setulae)averaging 17.1pm in length
(Figs 3.2 & 5.35).
McPartland & Cubeta (1997) reported this fungus caus-
ing leaf a n d stem s m u d g e o n Cannabis near Pokhara, Nepal. Figure 5.35: Conidium of Pestalotiopsis species (LM
Paulsen (1971) isolated a Pestalotia species from h e m p i n Iowa. x1920, McPartland).
 Chapter 5: Fungal Diseases 135

111 upright synnemata, 40-400 pm thick and 1-5 mm tall, often di-
monly arises on dead herbaceous stems, especially on nettles
chotomously branched towards the apex. Conigiogenous cells poly-
blastic, integrated and terminal or discrete and sympodial. Conidia
(Urtica dioica). The fungus occurs worldwide, mostly in tem-
light brown, ellipsoid, one-celled, smooth, 3-4.5 x 2-2.5 pm. Hy- perate regions.
phae brown, septate, formlng pear-shaped swellings at ends of cells
next to septa. Trichodema viride Persoon, Ron~er'sNeues Mag. But. 1:94,1794.
This fungus reportedly causes "white root rot" of hemp =Pyrenium lignurzlrn Tode 1790.
in southern China (Anonymous 1974) and Japan (Kyokai Description: Colonies grow rapidly (covering a 9 cm petri plate in
1965, Kishi 1988). Above-ground symptoms include stunt- three or four days), at maturity a dark green or dark bluish green,
ing, wilting, and early plant death. Below-ground signs in- emitting a distinctive "coconut" odour. Conidiophores septate,
clude white spheres of mycelium forming on root surfaces branching pyramidally from a central upright stalk, with short
branches occurring near the apex and longer branches occurring
and a feather-like mycelium growing within roots. R. necatrix
below. Conidiogenous cells straight or bent, slender flask-shaped
lives in temperate and semitropical areas worldwide. It is a phialides, arranged in irregular whorls of two or three (rarely four),
hardy soil fungus, difficult to eradicate once established. Tri- size variable but mostly 8-14 x 2.4-3.0 ym. Conidia green en iinzsse,
cizoderinn knrzianlrni infests and kills R. ~iecatrix. globose, warted surface, 3.5-4.5 pm in diameter (Fig 5.10).
Prior to Rifai's revision of the genus, all green-spored
Torula Iierbarum (Persoon:Fries) Link, Mngazin, Gesellsci~aft species of Trichoderirza were cited as T.viride. This aggregate
h~fltuuforscizendevFreuiuie zu Berlin 3:21, 1809; =Monilia lzerbarum species flourishes anywhere from tropical forests to arctic
Persoon 1801; =Turulil nzonilis Persoon 1795 spud Hughes 1958. tundra, including swamps, dunes, and deserts. Domsch et
Description: Colonies olivaceous when young, dark brown to
black when old, velvety. Conidiophores septate, pale olive, 2-6 ym
al. (1980) reported it from volcanic craters and children's
\vide, enlarging at conidiogenous cells to 7-9 ym in diameter. Co- sandpits. McPartland (unpublished data 1993) found it
nidia cylindrical, resembling a chain of attached spheres, mostly sporulating on soil and rockwool in Amsterdam.
four to five septa (range two to ten), constricted at septa, straight or T. viride colonizes the root surfaces of many plants, but
cumed, pale olive to brown, venuculose to echuxulate, 2G70 x 5-9 pm. disease is rarely reported. It has been isolated from hemp
This fungus colonizes hemp leaves as a secondary para- stems in the Czech Republic (Ondrej 1991).Fuller &Norman
site and causes a post-harvest storage mould of marijuana. (1944)described a Trichodernla overrunning field-retted hemp.
Gzebenyuk (1984) reported T. herbartlnz colonizing hemp Dried leaves of Cannabis suppress T. uiride growth
stems in the Ukraine. Behrens (1902) counted the fungus (Grewal1989).The fungus produces toxic metabolites against
among hemp retting organisms in Germany. McPartland fellow fungi; Weindling first demonstrated this antagonistic
(unpublished data 1995) identified T. Izerbarzlnz overgrowing activity in 1932. T. uiride is now used for commercial
helnp stalks previously parasitized by Colletotriclnlnl denla- biocontrol of many pathogenic fungi (see the section on
tz~l~iz,
collected in China by Rob Clarke. T. l~erbar~~iiz com- Rhizoctonia sore shin disease).
 "Be very careful not to suffer weeds of any sort to ripen their seeds on or anywhere within gunshot of your mines,
or mints,for making money, which your manure-heaps and compost-beds may be styled,
almost without a metaphor. " -Helen Nearing

Chapter 6: Other Cannabis Pests & Pathogens

In this chapter we describe nematodes, viruses, bacteria, 1. SOUTHERN ROOT KNOT NEMATODE
parasitic plants, protozoa, non-insect arthropods, and ver- Meloidogyne incognita (Kofoid & White 1919)Chitwood 1949.
tebrates. Some of these organisms can cause significant dam- Description:Meloidogyne species are difficult to tell apart. Only
age, but most are uncommon problems. the head and anal (perineal) shields can be differentiated; every-
thing else is nearly identical (seeTable 6.1).Eggs are ellipsoidal, col-
ourless, 40 x 80 pm in size. Larvae emerging from eggs are long and
slender, averaging 15 x 400 pm. As larvae feed, a marked sexual
NEMATODE DISEASES dimorphism develops.Males enlarge into long, robust cylinders with
About 500 species of nematodes are plant parasites short round tails, averaging 35 x 1300 pm (Fig 3.4). Females turn a
(Cook & Qualset 1996),but only a few infest Cannabis. Most pearly-whitecolour and swell into pear shapes (averaging750 x 1300
pm), they may swell to the point of protruding from galls. Females
nematodes attack roots, causing subterranean damage. lay eggs in a jellylike sac that is yellow-brown in colour and bulges
Nematodes indirectly damage plants by destroying from their genitalia.
beneficial mycorrhizal fungi (Hayman 1982). Not all Perineal patterns of adult females (wrinkles in the cuticle
nematodes are bad news-some are beneficial and prey on around the anus) are relatively easy to prepare for observation un-
insect pests (see the chapter on biocontrol). der light microscopy. Eisenback (1985)presented line drawing, pho-
tographs, and charts of perineal patterns. Here in Table 6.1 we de-
scribe two characters of perineal patterns-the dorsal arch and striae.
Eisenback also presented drawings,photos, and charts of heads and
ROOT-KNOT NEMATODES stylets of females, males, and juveniles.
Root knot nematodes attack thousands of plant species Crop susceptibility tests can identify Meloidogyne spe-
around the world. They cause greatest damage in warm cies. The "North Carolina Differential Host Test" uses six crop
regions where summers are long and winters are short and plants to differentiate M. incognita, M . hapla, and M . javanica
mild. In the USA, root knot nematodes get nasty south of (see Sasser & Carter 1985).Recently, polymerase chain reac-
the Mason-Dixon line (39" latitude), especially in sandy soils tion (PCR) techniques have been used to differentiate these
along coastal plains and river deltas. species (Guirao et al. 1995).
Most "Meloidogyne species" in the Cannabis literature
SIGNS & SYMPTOMS refer to M. incognita, from Brazil (Richter 1911), Tennessee
Above-ground symptoms are nonspecific-stunting, a (Milleret al. 1960),India Uohnston 1964),and the former USSR
chlorosis that resembles nitrogen deficiency, and midday (Goody et al. 1965). M . incognita is the most widely distrib-
wilting with nightly recovery. Farmers may misinterpret uted Meloidogyne species worldwide; it prefers a warm cli-
these symptoms as mineral deficiencies or drought, myste- mate. In the USA, it remains south of the Mason-Dixon line
riously appearing despite adequate nutrients and moisture. (39"latitude), except for scattered appearances in warm glass-
These symptoms do not arise uniformly across fields, but in houses. M . incognita attacks hundreds of hosts. It produces
patches of scattered infestation. Greatest symptoms arise in larger root knots than other Meloidogyne species.
plants near the centre of these patches, blending to healthy
plants at the periphery.
Below-ground symptoms are more distinctive. Root
knot nematodes embed themselves in roots and cause roots
to form giant cells or syncytia. Syncytia swell into root galls.
Galls may coalesce together to form "root knotsu--
conspicuous, hypertrophied roots with lumpy surfaces. Root
galls also stimulate the formation of adventitious rootlets,
creating a "bushy" appearance. The excessive branching of
adventitious rootlets may lead to false impressions, since
diseased roots look more developed than healthy roots.
Nematodes can be seen by peering closely at galls with a
magnifying lens, or breaking galls open (Fig 6.1 & Plate 70).
Fungal root rot may arise in galls, especially late in the season.
Figure 6.1: Progressive growth stages of immature
Meloidogyne incognita larvae, from freshly hatched (on left)
CAUSAL ORGANISMS & TAXONOMY
to nearly-mature "sausage stage" (LM courtesy G.W. Bird).
Between 1855 and 1949, root knot nematodes were
known by different names: Ang~till~ila marioni Cornu 1879,
Heterodera radicicola Miiller 1884, H. javanica Treub 1885, An- 2. NORTHERN ROOT KNOT NEMATODE
gl~illulaarenaria Neal 1889, and Oxytlrtis incognita Kofoid & Meloidogyne hapla Chitwood 1949.
White 1919.In 1949 B. G. Chitwood moved thesenematodes Description:M. hapla larvae and adults are slightly smaller than
to the genus Meloidogyne and divided them into five spe- those of M. incognita, but they are difficult to tell apart. The head
cies-three of which attack hemp, described below. Sepa- and anal (perineal)shields differ, see Table 6.1.
rating Chitwood's species can be difficult; many research- Inoculation tests prove Cannabis is susceptible to M. hapla
ers simply identify them to genus, as "Meloidogyne species." (Norton 1966; de Meijer 1993). M . hapla eggs tolerate cold
137
135 Hemp Diseases and Pests

Table 6.1 : Some differential characteristics of Meloidogyne species.

SPECIES DORSAL
ARCH OF STRIAEOF PERINEAL STYLETCONE OF HEADCAP OF MALES
PERINEAL PATTERN PATTERN FEMALES

M. incognita squarish, high coarse, smooth to large, anterior half flat to concave
wavy to zigzaggy cylindrical, curved

M. javanica rounded, interrupted coarse, smooth to anterior half convex, rounded, broad
by lateral ridges slightly wavy tapered, less curved

M. hapla rounded, low fine, smooth to small, narrow and convex, rounded, narrow
slightly wavy delicate, slightly curved

better than M . i~zcognitn;the northern root knot nematode Table 6.2: Infestation severity index for root-knot nematodes,
lives in eLrery state except Alaska (in Hawai'i it only occurs estimated by 2 methods (described in text).
at higher elevations). It also lives in Canada, northern Eu-
rope, and southern Australia. M . lmpla generally causes less
damage than M. incognzta. M . lzapla galls remain small and
spherical; compound galls rarely develop. The species is a
serious pest of tomatoes and potatoes. M . hapla can be con-
fused with Meloidogyne cltitzuoodi Golden, a species dis- Very low
ROOTEXAMINATION
(% OF ROOT AREA
AFFECTED BY GALLS)

galls rare
SOILASSAY
(# OF NEMATODES
PER PINT SOIL)

0-20
1
tributed in the western USA. Light 1-10% 21-100
Moderate 11-25% 101-300
3. JAVA ROOT KNOT NEMATODE
iMeloidogyne jnualzica (Treub. 1885) Chitwood 1949. Heavy 26-50% 301-1000
Description: M. larni~icalarvae and adults are nearly identical Critical >50% > I 000
to those of iL1. incognita (see description above). The head and anal
(perineal) shields differ, see Table 6.1.
Decker (1972)reported M . javanica infesting hemp in the found worldwide but cause greatest damage in temperate
southern USSR. M. jniinnica has no cold tolerance; it lives in regions. Above-ground symptoms mimic symptoms of root-
semitropical regions (in the USA it has been reported in the knot nematodes. Underground symptoms consist of
Carolinas, the Gulf states, and southern California). distorted, knobbed, and bushy roots. Affected roots have
cysts embedded in them (Plate 71). Cysts are the remains of
ROOT KNOT DISEASE CYCLE female nematodes, filled with eggs. Roots damaged by cyst
Root knot nematodes overwinter as eggs. First moult nematodes become predisposed to other soil pathogens, no-
occurs in eggs, so second-stage larvae hatch and penetrate tably Fi~sari~irn
and Rhizoctonia fungi.
roots of susceptible plants. Once in roots, the larvae undergo
three more moults and develop a sexual dimorphism. Males 1. SUGAR BEET CYST NEMATODE
become migratory, whereas females stay sedentary and Heterodeva schaclztii Schmidt 1871
remain in roots. Mating may occur, but Meloiilogyne females Description: Cysts are spherical to lemon-shaped, coloured
can produce eggs parthenogenetically. Females usually white-yellow-brown (depending on their age), and average 500-
produce 300-500 eggs (rarely over 2000) in egg sacs. Sacs 800 pm inlength (Plate 71).Eggs within cysts are minute. First moult
occurs in eggs, second-stage larvae hatch out. Orrce feeding beg~ns,
often protrude from root surfaces (Plate 70). Females die and cyst nematodes develop a sexual dimorphism. Males reach 1600 pm
egg sacs decay, releasing eggs into the soil. Under optimal in length; females enlarge to the size and shape of cysts.
temperatures (25-30°C) the life cycle may turn in 20 days. H.schachtii attacks a wide range of hosts, principally
Nematode root wounds provide portholes for many plants in the Chenopodiaceae (e.g., sugar beet) and
root-pathogenic fungi, including Rlzizoctonia solaili, Pytlrizin? Cruciferae (e.g., cabbage). It lives in temperate zones; in
rlltinzl~ii~,
F11snriz~li1
O X ~ S ~ Oand
Y L Verticillinln
L~~, species. Studies North America it is found from southern California and
with tobacco show that weakly-pathogenic fungi (C~irulllaria, Florida north to Ontario and Alberta. H. schnclztii has only
Botrytis, Peizicillil~~i~,
and Aspergillus species) also invade roots been reported on Cannabis in Europe and southwest Asia
\.in nematode ~vounds(Lucas 1975). (Kirchner 1906, Goody et nl. 1965).
To estimate nematode infestation levels, two methods
can be used-root examination a n d soil assay. Root 2. HOPS CYST NEMATODE
cxaiiii~zntioilis done at harvest. Estimate the percentage of Heterodeva humuli Felipjev 1934
root area galled by nematodes. Use the index outlined in Description: H. lztlmt~licysts, like those of H . sclznci~ti~,
are
Table 6.2. The second method, soil assay, is also done at lemon-shaped and yellow to brown coloured, but smaller (300-600
har\.est. Collect samples from soil 15-20 cm deep, mix the pm long). H.izz~rnulilarvae are smaller than those of H. schaciltii
samples and ship one pint of soil to your agricultural extension (males reaching 1 mm long, females averaging 410 pm).
agent. Samples must remain cool (<27"C)and moist. For soil H. h u l n l ~ l inormally infests hops, but Filipjev &
samples, apply the index outlined in Table 6.2. Stekhoven (1941), Winslow (1954), Decker (1972), Spaar et
al. (1990), and Gutberlet & Karus (1995) report the species
CYST NEMATODES infesting hemp. Contrarily, Kir'yanova & Krall (1971)
Two species of cyst nematodes attack Cannabis. They are claimed H. lu~nznlidid not infest hemp.
 Chapter 6: Other Cannabis Pests & Pathogens 139

Figure 6.2: Disease cycle of Heterodera schachtii (McPartland redrawn from Agrios 1997). A. Eggs overwinter in cysts;
B. Eggs develop into 1st-stage larvae; C. Then moult into 2nd-stage larvae while still within eggs; D. Second-stage larvae
hatch in the spring; E. 2nd-stage larvae invade rootlets; F-H. After several moults, females become more rotund than
males; J. Females eventually rupture roots, their posteriors exposed to the soil; K. Males exit roots to inseminate the
exposed females; L. During the growing season, females extrude as many as 500 eggs in a gelatinous mass, and the
cycle repeats; A. In autumn, females do not extrude eggs, they die and their cuticles harden into egg-filled cysts.

CYST DISEASE CYCLE

Eggs overwinter in a cyst (the hardened, dead shell of
their mother). For the rest of the life cycle, see Fig 6.2. At an
optimal temperature of 25"C, the life cycle of H. schaclztii takes
less than a month. H. lz~~nzulireproduces slower, only one or
two generations arise per year.

STEM NEMATODE
This is one of those unusual nematodes that does not
llre in roots. It feeds on above-ground parenchymatous
tissue. Symptoms begin as inconspicuous thickenings of
branches and leaf petioles. Sometimes even the middle veins
of leaves become swollen. Stems subsequently become
twisted and distorted, with shortened internodes producing
malformed, stunted plants (Fig 6.3). Infected stems feel
spongy. Plants often wilt due to severed xylem vessels.
Lightly-infected plants may send up new shoots from below
infested areas (Mezzetti 1951).

Ditylenclzus diysaci (Kuhn) Filipjev 1936

dlpsnciKuhn 1857;=Tylenchus deaastatvis (Kuhn)
=Aily~i~llli/lna
Oerley 1880
Description: Males and females retain their filiform shape into
maturity Stylets are short and thin and difficult to see under a mi-
croscope; the stylet cone is about half of the total stylet length, stylet
knobs are small and rounded. Females may reach 1.6 mm in length,
11-1thonly a few eggs in the uterus at one time, and a long slender
tail. Eggs are oval, 60 s 20 ym.

STEM NEMATODE DISEASE CYCLE

The life cycle of stem nematodes begins like the life cy- Figure 6.3: Symptoms caused by the stem nennatode,
cle of other nematodes. Eggs overwinter in the soil, and first Ditylenchus dipsaci (from Mezzetti 1951 ).
140 Hemp Diseases and Pests

moult occurs within eggs, which hatch into second-stage NEEDLE NEMATODE
larvae. Thereafter, stem nematodes travel a different path- This nematode is an ectoparasite-most of its body re-
they migrate out of soil and infest above-ground plant parts. mains outside the root while its head penetrates root tips.
They enter stems through stomates, lenticels, or wounds. Underground symptoms include thickened and distorted
Invasion of plants is favoured by cool, moist conditions. ro0.t tips, with poor development of the root system as a
Mating with males is necessary for reproduction. The life whole. The disease is sometimes called "curly root tip." Root
cycle quickens to 19 days in wet weather and optimal tem- tips become curled or hooked, and turn a necrotic brown
peratures (15-20°C). D. clipsaci is one of the few nematodes colour. Above ground, infected plants are stunted and
that spreads by seed, but this has only been reported in bean develop chlorotic leaves. Heavily infested plants may wilt
and onion (Bridge 1996). and die during hot dry summers.
D. dipsac1 is found in North America, southern Africa,
Australia, and temperate areas of Eurasia. Canriabis disease Paralongidorzis maximtrs (Biitschli) Siddiqi 1964
has only been described in Europe (Kirchner 1906, Ferfaris =Longidort~sntaxirrilis (Butschli) Thorne & Swanger 1936,
1913 & 1935, Mezzetti 1951, Goidanich 1955, Rataj 1957, =Dolylainzus niarimus Biitschll 1874
Ceapoiu 1958, Goody et al. 1965, Dempsey 1975). The Description: P,nlaxiritus is a large nematode. Eve11 the eggs
nematode attacks several hundred plant species, including are big-measuring 260 x 65 pm. Females average almost a
onion, grains, legumes, and many weeds. Taxonomists centimetre (9840 pm) in length and 92 pm wide. The body tapers
separate D.dipsaci into different subspecies ("races"); slightly toward both ends. The head is offset from the body by a
deep constriction. Males are slightly shorter (9630 ym) and rarely
Ciintinbis is infected by the "flax-hemp race" (Kir'yanova &
encountered.
Krall 1971).The hemp flax race also infects flax (Linlinz spe-
cies), rye (Secalecereale), and may indeed be identical to the "rye
NEEDLE NEMATODE DISEASE CYCLE
race" (Nickle 1991).
Females produce few eggs. Larvae moult through four
stages before reaching adulthood. Reproduction (usually
ROOT LESION NEMATODE parthenogenetic) occurs in late summer and autumn. P. nzax-
Symptoms arise as spots on small feeder roots, first ap- iinlls has been reported on Cannabis in western Europe
pearing watersoaked or cloudy yellow. The spots turn dark (Goody et al. 1965). The nematode attacks a wide range of
brown and the root tissue collapses. Collapsed lesions enlarge dicots, monocots, conifers, and possibly ferns. It transmits
and coalesce as secondary organisms (bacteria and fungi) the alfalfa mosaic virus.
invade tissues. The entire root system may become stubby
and discoloured. Above-ground symptoms resemble those
of drought stress or mineral deficiencies. Plants gradually OTHER NEMATODES
lose vigour, wilt easily, turn yellow, and become stunted.
Dippenaar et al. (1996) found other nematodes in roots
of South African hemp, including spiral nematodes
Pratylenchtrs penetrans (Cobb 1917)Chitwood & Oteifa 1952
(Heliocotylenchus and Sctrtelloneti~aspecies) and reniform
(Cobb 1917)Filipjev & Schuurmans-Stekhoven1941
=I? penehn~~s
nematodes (Rotylenchulus species). These nematodes
Description: P. penetrilizs adults are 400-800 ym long and 20-
25 pm 111 diametre. They have blunt, broad heads and rounded tails, attacked roots of the cultivar 'Secuini' more than roots of
~ v l i ~ cmake
h the nematodes lookshort.The stylet is short (1619pm) wit11 other hemp cultivars ('Fedora-19,' 'Futura-77,' 'Felina-34,'
lvell-developed basal knobs. Eggs are elongate, 60-70 x 20-25 pm. 'Kompolti').
Scheifele et al. (1997)assessed nematode populations in
LESION NEMATODE DISEASE CYCLE Canadian soils, before and after a hemp crop (using cultivars
Eggs overwinter in temperate soil. In warmer regions, 'Unico B' and 'Kompolti'). The hemp crop suppressed
eggs and lnri~aeoverwinter in diseased roots. First moult oc- soyabean cyst nematodes (Heterodera glycitzes Ichinohe), pin
curs in eggs. All subsequent stages (second-stage juveniles nematodes (Paratylenclzlis species), and stunt nematodes
through to adults) remain migratory endoparasites, enter- (Tylenckorkynclzusspecies), but increased the soil populations
ing and leaving roots at will. As they move through roots, of spiral nematodes (Heliocotylencll~isspecies) and root knot
root lesion nematodes secrete a root-necrosing enzyme. When nematodes (Meloidogyiieimpla).
root necrosis becomes severe, the nematodes exit in search
of new roots. Adult females lay eggs singly or in small groups DIFFERENTIAL DIAGNOSIS OF NEMATODES
in roots or soil. Because of their migratory nature, total egg Root gall nematodes may be confused with cyst nema-
production per female is unknown. The life cycle is relatively todes. The cuticle of female Meloidogyne species remains
slo~v,taking 45-65 days. white and soft, whereas the white skin of Heteroiiera females
P. petietrans predominates in temperate potato-growing turns yellow then brown, hard, and crusty.
regions, especially in areas with sandy soil. The species is Chlorosis and wilting from root nematodes mimic
susceptible to desiccation, and becomes dormant during symptoms caused by root fungi, some soil insects (e.g.,root
droughts. P. peiletrails predispose roots to Verticilliilirl and maggots, white root grubs), nutrient deficiencies, or drought.
Rlzlzoctotlia fungi. According to Ferraris (1935),symptoms caused by D. dipsaci
P. penetrans has been reported on h e m p in the can be confused with those caused by Pse1idoperoi1ospori7
Netherlands (Kok et nl. 1994) and South Africa (Dippenaar etA
cailtlabinn, a downy mildew.
111. 1996). P.pelzetralzs attacks many crops, weeds, and wild
plants. Compared to other crops, Cannabis is a highly CULTURAL & MECHANICAL CONTROL
susceptible host (Kok & Coenen 1996).Dippenaar et al. (1996) (numbers refer to Chapter 9)
compared different cultivars, and reported more P. peiietrans Method 1 (sanitation) controls many nematodes, espe-
infesting 'Kompolti' roots than roots of 'Fedora-19,' 'Futura- cially D.dipsaci. Method 2a (deep ploughing) kills many
77,' 'Felina-34,' and 'Secuini.' Potato growers test soil samples nematodes (but P. nzaxinzl~srequires ploughing down to 40
prlor to planting; fields with >I00 P. penetrai~sper 100 ml soil cm). Method 2b (steam sterilization) can be implemented in
sustain significant crop damage (Howard et al. 1994). glasshouses. Soil solarization (method 2c) controls P pm-
 Chapter 6: Other Cannabis Pests & Pathogens 141

etrnrzs, D. dipsaci, H. schaclztii, and M. hnpln, but not M. incog- Burkholderia (Pseudomonas) cepacia
nztn (Elmore et al. 1997).Flooding soil (method 2d) also works. BIOLOG~: A soil bacterium that colonizes the root zone
Weeding (method 3) eliminates alternative hosts. of plants and protects roots from some nematodes (lesion,
Growing vigorous plants is the best control against sting, lance, spiral nematodes). It is described under Damp-
nematodes. Healthy plants can regenerate new roots. Avoid ing off disease.
water stress (method 7a). Keep soil nutrients in balance. Add
calcium against D, dipsaci, potassium against Meloidogytze Myrothecium vewucaria
species. Avoid planting in sandy soils that nematodes like BIOLOGY: This fungal saprophyte lives in a wide range
best. Adding compost and manure encourages the growth of soils. Strains selected for biocontrol do best around 20°C.
of organisms antagonistic to nematodes. APPLICATION: Available as an emulsifiable suspension
Rotating with monocot crops will stanre root knot nema- (DiTera E S ) or granules (DiTera G@),for the control of many
todes and cyst nematodes, but may take a while (two years nematodes, including Meloidogyr~e,Heterodern, Pratyleticlzl~s,
for root knot, seven for cyst). Finding nonhost plants for D. and Xiphinenla species. The fungus may be incorporated into
dipsaci is difficult, and nearly impossible for P. maximus. soil as a preplant mix or injected into the root zone of
Quarantine prevents the movement of nematodes into established plants.
new areas. Larvae left on their own only migrate 20 cm in a NOTES: Gzebenyuk (1984) cited M. verrl~cariaattacking
lifetime. But eggs and cysts can travel many miles on tractor Ukrainian hemp, but he may have confused it with
tyres, floating in streams, even blowin' in the wind. Myrotllecit~tnroridl~nz,the cause of corky leaf spot.
Quarantines must be rigorously enforced to be effective-
exclude plants, soil, even dirty shoes and spades from Arthrobotrys and Dactylaria species
uninfested areas. Breeding for crop resistance works has BIOLOGY: These two genera of soil fungi strangle nema-
barely begun in Caizilabis. De Meijer (1993,1995) studied the todes in hyphal nooses or trap them on sticky hyphal knobs.
resistance of hemp cultivars to M. hnpln. The most resistant Commercial products containing Arthrobotrys roblista (Royal
were Hungarian cultivars 'Kompolti Sargaszar6' and 3000, used against Ditylenchus species) and Artllrobotrys
'Kompolti Hibrid TC.' The most susceptible were superha (Royal 350@,used against Meloidog!/ne species) are
French cultivar 'Futura 77' and Ukrainian cultivar 'USO-13' irregularly available. Unfortunately, these fungi grow slower
('YUSO-13'). than most biocontrol fungi, and they are very sensitive to
desiccation.
BIOCONTROL (see Chapter 10)
Plants belonging to 57 families possess nematicidal Pasteuria (Bacillus) penetrans
properties (Bridge 1996). Cntznnbis plants can suppress Mel- BIOLOGY:A soil bacterium that infects numerous nema-
oldogyne species (Kok et al. 1994, Mateeva 1995) and Heterod- todes including Meloidogyne, Herterodera, Ditylei~clzl~s,
era rostoclziensis (Kir'yanova & Krall 1971). Akhtar & Alam Pnralotzgidort~s,and Prntylenchus species. Endospores of P.
(1991)intercropped mustard greens (Brnssica juncen) to sup- penetrnns infect immature nematodes in soil; endoparasitic
press M. incogrzitn. Marigold (Tagetes species, described be- nematodes such as Meloidogyne species die before they can
low) suppresses nematodes by acting as a "decoy crop." reproduce. The commercial availability of P. penetrnns is
Decoy crops cause nematode eggs to germinate, but the hampered by its expensive production costs (Cook et al.1996).
nematodes cannot complete their life cycle on the decoy
plants, so the nematodes die out. Palti (1981) decoyed M. Verticillium chlamydosporium
incogi~itaand 1l4. javaizica with marigold (Tagetes pntula, T. BIOLOGY: A soil fungus and facultative parasite of cyst
i i l i i l ~ ~ t ~sesame
z), (Sesutrz~~liz
orientale), castor bean (Ricitlusconz- nematodes and nematode eggs. Infected cyst nematodes turn
~iiziizis),and Clrrysaiztlzrnzir~i~ species. Prat!/lerich~is penetrans brown and shrivel. I/: chlanzydospori~~rntakes two weeks to
can be decoyed with marigold (Tngetes species) and blanket kill its host. Cadavers become surrounded by a halo of
flowers (Gaillardia species). chlamydospores in optimal conditions. The fungus has also
Bacteria and fungi that kill nematodes are on the been isolated from plant roots, snail eggs, and other soil
biocontrol horizon. According to Meister (1998), coating fungi-so it is not dependent on nematodes for survival.
seeds with Bzlrkholderia cepncin (described under damping
off) protects seeds from lesion, sting, lance, and spiral Hirsutella rhossiliensis
nematodes. Gliocladi~~nz rose~~ln infests cyst nematodes BIOLOGY: A soil fungus whose sticky spores adhere to
(described under grey mould). Seven other bacteria and fungi nematodes as they move through soil. Heterodern schaclztii
are described below. For more, see Carris & Glawe (1989). juveniles are killed within three days of infection at 20°C
(Tedford et nl. 1995). Dead nematodes sprout new spores to
Tayetes species repeat the H. rhossiliensis life cycle. The fungus has a broad
BIOLOGY: Marigolds are annual herbs that ooze host range, including Heterodera, Meloidogyne, Ditylenchlis,
nematode-repellent metabolites into soil. Some nematodes Prntyletzcht~s,and Xiphinenln species. Unfortunately, the
are attracted to marigolds and bore into the roots, but are fungus also kills biocontrol nematodes-Steinerizenzn and
unable to feed or reproduce (Howard et nl. 1994). The most Heterorhabditis species.
effective species are African marigold (Tngeteserecta), French
rnarlgold-(~a~etrs patula), and south ~ h e r i c a nmarigold Netnatophtlzora gynophila
(Tagrtes nnnuta). Crops interplanted with marlgold suffer BIOLOGY: This oomycete is an obligate parasite of
fewer nematodes, as do subsequent crops growing in the nematodes in the genus Heterodera. It does well in the damp
same location. But marigold is not a cure-all-some cultivars soil preferred by nematodes.
are susceptible to Meloidogyize infestation (Miller et al. 1960).
Marigold works best against Prntyletzclz~~s, Rotyle~~chl~s, and CHEMICAL CONTROL (see Chapter 11)
~/lr~zchorlzynchusspecies. It works poorly against Ditylenchl~s, Urea is a traditional cure for infested fields, mixed into
Heteioilrra, and most ectoparasitic~nematodes( ~ o w a r et d al. soil after the growing season. By spring, the urea has broken
1993) down and most nematodes are dead. Clandosans combines
112 Hemp Dise ases and Pests

urea with chitin from ground-up crab shells. Crab shell chitin
encourages the growth of nematicidal soil organisms when AMV
mixed into soil at a rate of 2-4 kg per 10 m2. Adding cow
dung and poultry manure to soil provides some control of
~Zil.lricognlta (Bridge 1996).
ern, normally used as an insecticide, has nematocidal
properties. Akhtar & Alam (1991) mixed neem oil and cas-
tor oil into soil to repel nematodes. Olkowski et al. 11991) ~,
used neem oil against M. incognita. Neem cake also works,
mixed into soil at rates as low as 100-250 kg ha-1 (Bridge
1996).The residual effect of soil treatments lasts for several
months. Aqueous extracts of azadirachin, the active ingre-
dient in neem, prevent M. incognita infection in a variety of
crops when used as a seed treatment and root dip ordu due
& Blackwell 1993).
Aqueous extracts of Cannabis leaves caused a high
mortality rate in M. incognita (Vijayalakshmi et al. 1979),
Heterodera cajani (Mojumder et al. 1989), M. javanica (Bajpai
& Sharma 1992),Hoplolainzlis i~zdicz~s, Rotylenchz~l~isreniforiizis,
and ~ ~ ~ ~ e ~ l c i z o brassicae
r ~ ~ r ~ c (Haseeb
l ~ l l s et al. 1978).Soil mixed
\\-it11 3"J0J.v/\.v dried Cannabis leaves suppressed M. incogrzita
(Gos~vam& i Vijayalakshmi 1986) a n d Aplzelenchoides
co~iipo~ticoln (Grewal 1989). In human medicine, "worms"
have been treated with juice extracted from crushed hemp
seed (Parkinson 1640, Culpeper 1814).This practice dates to
Imperial Rome (Pliny 1950 reprint).
Extracts of the pawpaw free repel nematodes (Meister
Figure 6.4: A menagerie of plant and animal viruses.
1998).Yepsen (1976) mulched soil with latex-bearing plants AMV =Alfalfa mosaic virus; TMV = Tobacco mosaic virus;
(goldenrod, milkweed, dandelion, Eiiphorbia species) to repel CMV = Cucumber mosaic virus; NPV = Nuclear polyhe-
nematodes. drosis virus (insects); Cox = Coxsackie virus;
HIV = Human immunodeficiency virus; Cold = human
Unfortunately, the egg-protecting cysts of cyst
nematodes renders them impervious to nearly all coronavirus, Ebola = Ebola (McPartland).
nematocides. Soil fumigation (injecting soil with volatile
liquids) works best against cyst nematodes but is very netically-altered plant viruses (Garrett 1994). RNA viruses
expensive and dangerous. ~ u m i g a n t sare usually injected mutate a million times faster than DNA viruses (Strauss &
into field soil to depths of 22-30 cm with row cl~isels. Strauss 1988).All RNA viruses, including plant and animal
viruses, probably descended from a single ancestor virus.
A separate concern is the contamination of plants by
VIRAL DISEASES human viruses. An outbreak of Hepatitis A in Washington
The virus world was discovered by Dmitri Iwanowsky in was linked to Mexican marijuana fertilized with faeces con-
1892. The he discovered infected plants, not animals. taminated by the Hepatitis A virus (Alexander 198%).
It caused "mosaic" symptoms in tobacco. Plant viruses are Viruses rarely kill Cannabis. They can, however, cause
named by symptoms they cause and plants they infect. serious symptoms and reduce yields. Once acquired, viruses
Iwanowsky's virus is now called the tobacco mosaic virus are nearly impossible to eradicate-they invade all plant
(TMV).For a while, viruses were named with Latin binomials parts, including seeds and pollen, enabling viruses to replicate
(Holmes 1939). Thus TMV was named "Marnior tabaci," in across host generations. Insects spread many viruses as they
the Phyllum Phytophagi of the Kingdom Vira. Since buzz from plant to plant. According to Ceapoiu (1958), the
biologists debate if viruses are truly living creatures (see worst vectors are the bhang aphid (Phorodon cannabis),green
Chapter 3), Holmes's binomial system has been abandoned. peach aphid (Myzlls persicae), greenhouse whitefly
About 1200 17irusesare able to infect plants (Cook & (Trialelidodesoaporarior~~nz),and onion thrips (Tllrips tabaci).
Qualset 1996).Amazingly, only five viruses regularly infect Hartowicz et al. (1971) tested 22 viruses for their ability
Caiz~iabis.Perhaps this is due to the presence of THC, which to infect Caruzabis. Paulsen (1971) reported four of the 22
inactivates viruses (Blevins & Dumic 1980, Lancz et a1.1990, viruses caused serious mosaic symptoms and dwarfing:
Lancz et al. 1991). Cannabis also contains terpenoids tobacco ringspot (TRSV),tomato ringspot (TomRSV),tobacco
(limonene, a-pinene) and flavonoids (apigenin, luteolin, streak (TSV) and cucumber mosaic (CMV). Two of the 22
quercetin) with antiviral activity (Che 1991). viruses caused mosaic symptoms without dwarfing:
Cannabis viruses are very simple-they contain positive- eunoymous ringspot (ERSV) and alfalfa mosaic (AMV).Elm
sense, single strand (ss) RNA covered by a protein coat (Fig mosaic (EMV) caused mild necrotic flecking. Back
6.3). In contrast, many human viruses contain double- inoculations to indicator plants proved TMV and foxtail
stranded (ds) DNA with a protein coat and an outer lipid mosaic (FMV)cause symptomless infections in Cannabis. The
membrane. Plant viruses currently pose no threat to humans, remaining viruses did not infect Cannabis: barley stripe mo-
but there is some concern of plant viruses exchanging RNA saic (BSMV), brome mosaic (BMV), maize dwarf mosaic
material with similar human viruses. The human viruses strain A (MDMV-A), panicum mosaic (PMV), wheat streak
containing ssRNA are dreadful-Dengue, Coxsackie, Polio mosaic (WSMV), soilborne wheat mosaic (SBWMV), bean
(all positive-sense), Rabies, Hanta, Lassa, Machupo, pod mottle (BPMV),potato virus X (PVX),strawberry latent
Marlburg, Ebola, and HIV (negative sense). Concern about ringspot (SLRV),tobacco etch (TEV),tobacco rattle (TRV-C),
mutant viruses has increased due to the engineering of ge- soyabean mosaic (SMV) and squash mosaic (SqMV).
 Chapter 6: Other Cannabis Pests & Pathogens 143

More recently, Kegler & Spaar (1997) tested three fibre DISEASE CYCLE & EPIDEMIOLOGY
cultivars ('USO-11,' 'YUSO-14,' 'YUSO-31') for their resist- Roder (1941) and Goidanich (1955) cited hemp aphids
ance to eight \riruses: AMV, CMV, PVX, TomRSV, potato vi- (Phorodon can~inbis)as the chief vectors of HSV. Seeds of dis-
rus Y (PVY),broad bean wilt fabavirus (BBWV),arabis mo- eased plants give rise to diseased progeny. Plants developed
saic ~ ~ i r (ArMV),
us and raspberry ringspot nepovirus (RRV). symptoms within six days of inoculation (Roder 1941).Male
The results of their inoculation experiments are presented plants suffer higher rates of infection than females.
in Table 6.3. Symptoms of AMV, CMV, and ArMV are de-
scribed below. TornRSV caused light-green checkmarks in ALFALFA MOSAIC VIRUS (AMV)
one cu1tir.x ('YUSO-31') and did not infect other cultivars. Schmelzer (1962)infected hemp with a strain of lucerne
PVX caused a light- and dark-green mosaic between leaf mosaic virus from Vibllrn~lnioplillls. Lucerne mosaic virus is
veins in one cultivar ('YUSO-31') and did not infect other a synonym for the alfalfa mosaic virus (AMV). Schmidt &
cultivars. PYV caused a light- and dark-green mosaic, as well Karl (1970) described naturally-occurring AMV on hemp in
as upturning of leaf tips. BBWV caused a yellow-green Germany. They identified the virus via graft and sap
mosaic at the base of young leaves. RRV caused a light-green transmission with test plants.
mosaic and curling of leaf petioles.
SYMPTOMS & EPIDEMIOLOGY
AMV symptoms have been described as a grey leaf
Table 6.3: Susceptibility of 3 fiber cultivars to 8 viruses.l mosaic (Schmelzer 1962), chlorotic stripes along leaves
(Schmidt & Karl 1970), or light green flecking with chlorotic
VIRUS NUMBER
OF PLANTS WITH SYMPTOMS (%) veins and slight puckering of young leaves (Kegler & Spaar
uso-11 YUSO-14 YUSO-31 1997). Spaar et al. (1990) illustrated symptoms of AMV in a
colour painting. AMV has a wide host range and is found in
AMV 0 57.1 11.1
nearly all temperate regions. Viral egression is primarily via
-CMV
- 18.2 66.7 50.0 aphids. Seed transmission, root grafts, and dodder (Cusclifn
BBWV 37.5 0 16.7 species) may also spread AMV. Virus morphology (the "vi-
ArMV 10.0 0 7.6 ral genome") consists ssRNA encapsulated in four particles-
all four particles are needed for infection-the first is spheri-
RRV 25.0
- 0 9.1
cal (18 nm diameter), the other three are bacilliform-all 18
PVX 0 0 14.5 nm wide but different lengths (58 nm, 48 nm, 36 nm).
PVY 11.1 33.3 18.2 1
1
I
TornRSV 0 0 42.9 1
I
CUCUMBER MOSAIC VIRUS (CMV)
Schmidt & Karl (1970) cited CMV causing leaf mottling
1. Results of inoculation experiments by Kegler & Spaar (1997). in German hemp. Kegler & Spaar (1997) described svmptoms
as light green check-marks diffusely covering leaf &faces,
especially young leaves. Spaar e t nl. (1990) illustrated
A grower in Seattle reported transmitting a virus from
symptoms of CMV in a colour painting. Paulsen (1971)found
tobacco to marijuana (Rosenthal & McPartland 1998). He
that both CMV and CMV-Xstrains can infect hemp, CMV is
used the same scissors to trim tobacco and take Cannabis
vectored by the bhang aphid (Schmidt & Karl 1970), green
cuttings for clones. Symptoms included dwarfing, early se-
peach aphid (Myzlls persicae), and black bean aphid (Aphls
nescence, and reduced yields. The virus was probably TRSV,
fabne) in a nonpersistent manner, CMV also spreads in seeds
TolnRSV, or TSV.
and pollen of infected plants (Lucas 1975).
he virus infects at least 470 plant species and is
HEMP STREAK VIRUS (HSV) distributed worldwide, especially in temperate regions. CMV
Roder (1941) originally described hemp streak virus is a multicomponent virus, consisting of ssRNA packaged
(HSV) in Germany. HSV has caused serious losses in Italy in three isometric polyhedral particles, each 30 nm in
(Ferri 1963).In the Czech Republic, Rataj (1957)said the virus diametre. CMV replication is favoured by high temperatures,
was limited by the range of its aphid vector. Farther north in long day length, high light intensity, and excess nitrogen
Russia, Gitman (1968b) considered HSV a rare disease. HSV (Lucas 1975). Breeders have developed tolerant varieties of
is probably the "curly leaf virus" that Clarke (pers. commun. many susceptible crops (cucurbits, tobacco, spinach).
1993)found in Hungary. The viral genome and morphology
of HSV remains unknown. ARABIS MOSAIC VIRUS (ArMV)
Schmidt & Karl (1969)described ArMV causing yellow-
SIGNS & SYMPTOMS
green leaf spots and stripes on "weakly growing hemp" near
Foliar symptoms begin as a pale green chlorosis. Chlo-
Potzdam, Germany. Kegler & Spaar (1997) described
rotic areas develop into a series of interveinal yellow streaks
symptoms as yellow-green check-marks or mosaic on young
or chevron-stripes. Sometimes brown necrotic flecks appear,
leaves. Spaar et al. (1990) illustrated symptoms of ArMV in
each fleck surrounded by a pale green halo. Flecks appear
hemp. ArMV causes "nettlehead" disease in hops. The virus
along the margins and tips of older leaves and often coa-
also infects many vegetables. It occurs in Europe, North
lesce. Streak symptoms predominate in moist weather,
America and New Zealand. ArMV is packaged in three
whereas fleck symptoms appear during dry weather.
isometric particles each 28 nm in diameter, much like CMV.
Eventually, leaf margins become wrinkled, leaf tips roll
ArMV is transmitted via infected seeds and soil nematodes
upward, and leaflets curl into spirals. Whole plants assume
(Xiphitiema species).
a "wavy wilt" appearance (Plate 72). Diseased plants are
small and the yieidof fibre and seed is greatly reduced. R6der
(1941) and Ferri (1963) provided excellent black-and-white HEMP MOSAIC VIRUS (HMV)
photographs of disease symptoms. Spaar et al. (1990) Not much is known about HMV. It may be a
illustrated symptoms in a colour painting. Cucumovirus such as CMV, or a Nepovirus such as ArMV,
144 Hemp Diseases and Pests

TRSV, and TomRSV. Ceapoiu (1958) described symptoms in Thus, virus-free material can be obtained by cutting off a tiny
Romania, beginning as punctate, chloroticlesions which turn piece of shoot tip and regenerating it in tissue culture. This
necrotic, coalesce and, finally, the entire leaf wilts. Blattny et technique has not been attempted in Cannabis (Mandolin0
al. (1950) described the virus causing leaf enation (torsion) & Ranalli 1999). Virus-resistant plants are a possibility-
in Czech hemp. Ghani & Basit (1975) reported a mosaic vi- agronomists are testing fibre c u ~ t i ~ afor
r s resistance to virises
rus in Pakistani plants causing leaf curl, bunchy top, and (Kegler & Spaar 1997). Other crops have been genetically
"smalling of leaves." Traversi (1949) inoculated hemp with engineered for resistance to AMV, CMV, and other viruses
the Argentine sunflower virus, and it caused ~ ~ ~ - 1 i k e s ~(Goodman
m ~ - et al. 1987).
toms. Traversi's virus was transmitted by sap, seeds, the If virus infestation is suspected, minimize the
greenhouse whitefly, green peach aphid, and onion thrips. mechanical spread of viruses by dipping hand tools in a
HMV is vectored by Pkorodon cannabis (Ceapoiu 1958). dilute solution of skim milk (100 g skim milk powder per 1
Schmidt & Karl (1970) described a "hemp mottle virus" water). Milk inactivates many plant viruses (Howard et al.
similar to HMV; the virus was vectored by the aphid Phorodon 1994).Eliminate virus vectors, especially aphids, dodder, and
cannabis. nematodes. Cover seedlings with mesh netting to protect
them from flying insects.Many weeds harbour symptomless
DIFFERENTIAL DIAGNOSIS infections, so eradicate them (method 3). Some researchers
Viral diseases may be confused with symptoms of bac- claim viruses can be eradicated from infected seeds using
terial blight, early brown leaf spot, or yellow leaf spot. The thermotherapy, see method 11.
"wavy wilt" appearance of HSV seems unique, but ArMV
and HMV may cause similar symptoms. HSV, ArMV, HMV, BIOLOGICAL & CHEMICAL CONTROL
and AMV cause mosaics, making exact identification diffi- No biocontrol or antiviral chemicals have been effec-
cult. Using an electron microscope is effective but expensive. tive. Protect plants from virus vectors with insecticides.
Cross-inoculation studies with other plants are time-inten- Control ArMV vectors with nematocides.
sive. Serological tests for HSV and HMV are not available.
Christie et al. (1995) used simple stains to detect viral
inclusion bodies in plants. Inclusion bodies are intracellular
structures consisting of aggregated virus particles and coat
BACTERIAL DISEASES
proteins. Inclusions induced by a specific virus produce a Dozens of bacteria have been reported from Cannabis over
characteristic appearance, even in different hosts. Christie the years. Most of these species are retters of harvested hemp.
stained inclusions with Azure A, or a combination of Others bacteria rot harvested female flowers. Retters and
Calcomine Orange 2RS and Lux01 Brilliant Green BL. After rotters of harvested products are discussed in Chapter 8.
staining, viral inclusion bodies are easily seen with a light Kosslak & Bohlool (1983) isolated two species of
microscope. diazotrophic bacteria from the rhizospheres of marijuana
plants: Azospirillum brasilense Tarrand, Kreig & Dobereiner
CULTURAL & MECHANICAL CONTROL (see Chapter 9) and Azospirillum lipoferum (Beijernick) Tarrand, Kreig &
Destroy obviously infected plants and do not clone them Dobereiner. These mutualistic bacteria live on the surfaces
or use their seeds. Viruses migrate into developing seeds, despite of roots, where they fix nitrogen for their host.
assurances to the contrary (Rosenthal High Times 278:95). Subtracting the aforementioned saprophytes and
Virologists discovered that apical shoots of many plants mutualists, we only found four species of true pathogens
grow slightly faster than viruses can migrate cell-to-cell. (with one species split into four "pathovarieties"), described
below.

BACTERIAL BLIGHT
Also known as bacterial leaf spot, this disease may be
limited to Europe. Bacterial blight has been described in Italy,
Germany, Hungary, Bulgaria, Romania, Yugoslavia, and the
former USSR. It affects plants of all ages, from seedlings to
full-flowering females.

SIGNS & SYMPTOMS

Symptoms begin as small watersoaked leaf spots. Spots
enlarge along leaf veins (rarely crossing them) and turn
brown or grey. Dead tissue breaks apart causing leaf perfo-
rations (Fig 6.5).An Italian strain of the bacterium ("variety
italica") produces "ulcerous stripingo-small protuberances
arising in rows between leaf veins, turning into dark, necrotic
stripes (Sutic & Dowson 1959).Stem lesions may also form.
Plants become stunted and deformed.

CAUSAL ORGANISM & TAXONOMY

Pseudomonas syringae pv. cannabina (Sutic& Dowson 1959)
Young, Dye & Wilkie 1978, New Zealand lournal of Agricultural
Research 21:153-177.
=Pseudornonascannabina Sutic & Dowson 1959; =Pseudornonas
cannabina Sutic & Dowson var. italica Dowson 1959
Figure 6.5: Symptoms of bacterial blight caused by Description: Cultures white, convex, circular,some strains pro-
Pseudomonas syringae p ~cannabina
. (from Sutic & duce green fluorescent pigment on King's medium B agar, other
Dowson 1959). strains exude a nonfluorescent brown pigment (Smith et a/. 1988).
 Chapter 6: Other Cantiabis Pests & Pathogens 145

Aerobic gram (-) rods, size usually 1.5 x 0.3 pm, straight or slightly nese ancestry seemed more susceptible to striatura ulcerosa
curved with rounded ends, exhibiting one to four polar flagella (Fig than hemp of strictly Russian heritage (Hillig, pers. commun.
3.1). Non-proteolytic, starch weakly hydrolysed, acid (but no gas) 1998). P.s. pv. inori commonly blights mulberry, attacking
produced from xylose, dextrose, galactose, mannose, sucrose, raffi- leaves and stems. On trees, the disease resembles fire blight,
nose, and glycerol. No sheaths or spores are produced. Bacteria of-
oozing white or yellow masses of bacteria. It spreads by
ten form in chains.
wind-driven rain and overwinters in crop debris. P. s. pv.
DIFFERENTIAL DIAGNOSIS tnori was the first bacterium isolated and identified from dis-
Klement & Kaszonyi (1960)and Klement & Lovrekovich eased hemp (Peglion 1897).
(1960)noted the close relationship between P.s. px7. cnnizabina
and P.s. pv. nzori. Bacterial blight can be confused with early
symptoms of Wisconsin leaf spot and Xanthomonas leaf spot, XANTHOMONAS BLIGHT
or even early wildfire. It may also be confused with some Xanthomonas blight has been reported in Japan and
viral or fungal diseases (notably brown leaf spot), or insect Korea (Okabe 1949, Okabe & Goto 1965, Kyokai 1965). In
damage. Stem symptoms can be confused with striatura ul- Romania, the disease appears as a wilt (Sandru 1977) or as
cerosa (see next). small, brown leaf spots (Severin 1978). The bacterium
produces a mucilaginous, extracellular polysaccharide
DISEASE CYCLE & EPIDEMIOLOGY known as xnllthan gum, which plugs xylem and disrupts fluid
Goidanich & Ferri (1959) reported no seedborne infec- flow.
tion. Subsequent researchers refute this (Noble & Richard-
son 1968),and consider seedborne infection a primary mode CAUSAL ORGANISM & TAXONOMY
of dissemination. The bacterium overwinters in plant de- Xanthomonas carnpestris pv. cannabis Severin 1978, Arci1i.r:
bris and infects seedlings in the spring. During the growing ftir Pizytopnthology cind Pflanzenschutz 14:7-15.
season, rain droplets pick up bacteria oozing from lesions, =Xnnt/zornonascannabis (Watanabe) Okabe & Goto 1965,
=Bacterium cnnnabis (Watanabe) Okabe 1949, =Racilltis cnrlnabls
and splash them around. Wataliabe 1947-8; =Xant/zomonascannabis Mukoo npt~dKishi 1988.
Cross-inoculation studies with P.s. pv. cnniiabiizn by Sutic Approx~mately12 years before Severilx's publication, Okabe &
& Dowson (1939) demonstrated lethal susceptibility in Goto described Xnntkomonas cnnrlabis (Watanabe) Okabe & Goto 111
kidney bean (Phaseolz~salllgaris), and slight symptoms in Japan. Okabe's taxon was overlooked by Severin, by Bergey's 7th
common vetch (Vicia sntivn),grass pea (Lnthyrl~ssatizills), pea and 8th editions, as well as by Young, Dye & Wllkie (1978).
(Pis~lrnsativzrnz) and hyacinth bean (Doliclzos Inblab). Plants Description: Cultures usually yellow, smooth and viscld.
related to Cailt~abis,such as hops (Ht~inl~ll~s I I L ~ Z L ~nettle
ZLS), Aerobic gram (-) straight rods, with single polar flagella, usually
(Urtica~iioicn),and mulberry (Morusnlba), were not susceptible. 0.7-1.8 x 0.4-0.7 km. No denitrification or nitrate reduction.
Proteolytic, ~ionlipolytic;acid production from arablnose, glucose,
mannose, galactose, trehalose, cellobiose.
STRIATURA ULCEROSA
Kirchner (1906) first described this disease in Italy and DIFFERENTIAL DIAGNOSIS
Germany. Gitrnan (1968a) called the disease "bruzone" in Differentiating Pse~~doinonas from Xanthonzonns is diffi-
Russia. Ferri (1957a & b) and GoidBnich & Ferri (1959) cult. Xantlzonzot~asspecies produce a yellow pigment in cul-
published important monographs on striatura ulcerosa. ture and only grow one flagellum per cell. Xanthomonas leaf
spot is easily confused with bacterial blight (caused by Psell-
SIGNS & SYMPTOMS domoi~nssyriilgne pv. caiz~znbiiia).Bacterial leaf diseases can
Striatura ulcerosa arises in adult plants, beginning as also be confused with some fungal problems such as brown
waxy, dark grey, o l d lesions which elongate along stems leaf spot or yellow leaf spot.
(Plate73). These lesions rarely encircle more than half the
stem circumference, but may run 10 cm along its length. DISEASE CYCLE & EPIDEMIOLOGY
Small pustules (1-2 x 2-3 mm) arise within lesions, filled X,carnpestris infests many crucifers and other plants.
with a yellow mucilaginous ooze of bacteria. These pustules Okabe (1949) said the species easily infects hemp and flax.
rupture and shred the epidermis. Secondary pustules arise Severin (1978) found the hemp pathogen could also infect
around the shredded remains of earlier pustules. hops, mulberry, pinto bean, soyabean, cucumber, tobacco,
and Pelargoiiil~mplants. Disease worsens in summers with
CAUSAL ORGANISM & TAXONOMY high temperatures and high humidity (Okabe 1949).X. cainp-
Psezido~nonassyringae pv. mori (Boyer & Lambert 1893) estris overwinters in crop refuse and infected seed (Smith et
Young, Dye & Wilkie 1978, Neu: Zealnnd /oclrnai of Agriczlltl~rnl al. 1988).During the growing season it spreads by splashing
Rescarcl~21:153-177. water, wind-driven rain, insects, and field workers.
=Psel~dolrlonasnloui (Boyer & Lambert 1893) Stevens 1913,
=Bacillus n l o ~ (Boyer
i & Lambert 1893) Holland 1920;
=Pseudoniotlnsctibonli~ni(Macchiati 1892)Krasil'nikov 1949, =Rncillus
ctibonint~lisMacchiati 1892 non Stelnhaus 1942. WILDFIRE and WISCONSIN LEAF SPOT
Description: Pathovars of P sy~ingaeare morphologically indistin- These two diseases are difficult to distinguish from Bac-
bwisliable from each other (see description of I! s. pv, cannabiru above). terial Blight, from Xanthomonas blight, and from each other.
Symptoms begin on lower leaves and spread rapidly in wet
DIFFERENTIAL DIAGNOSIS weather. Watersoaked spots turn into small necrotic lesions
Klement & Lovrekovich (1960) noted a close relation- surrounded by chlorotic halos. Lesions may coalesce into
ship between striatura ulcerosa and bacterial blight. The two irregular necrotic areas. Alternately, leaf lesions may grow
causal organisms are now considered pathovars of the same into "angular leaf spots" without halos, limited by leaf veins.
species, P.syriilgnr.Striatura ulcerosa resembles early symp- Affected leaves become twisted and distorted.
toms of Cladosporil~n~,Phonzopsis, or Ophioboll~sstem cankers.
1. WILDFIRE
DISEASE CYCLE & EPIDEMIOLOGY Pseudomonas syringae pv. tabaci (Wolf & Foster 1917)
In greenhouse studies, plants with a percentage of Chi-
146 Hemp Dise'ases and Pests

Young, Dye & Wilkie 1978, New Zealnnd lournnl of Ayricultz~ml tlinzefaciens infects plants, the bacterium injects Ti plasmids
Research 21:153--177. into plant cells, and the plasmids become incorporated into
=pseudor~~onas tabnci (Wolf & Foster 1917)Stevens 1925; =Pseu- the host plant genome. The plasmid genes cause plant cells
~ion~oilnsar~gtilntn(Fronime & Murray 1919)Stevens 1925. to proliferate into cancer-like galls and produce nutrients
Description: Cultures white, slightly raised, with translucent for the bacteria.
edges al-rd opaque centres, producing a green fluorescent pigment.
Patho\,arsof sy~ingneare morphologically indistinguishablefrom Scientists can bioengineer foreign genes into Ti
each other (see description of s. p\<canrlnbina above). plasmids, genes coding for insect and disease resistance. In
Some authors (Lucas 1975, Smith et al. 1988) consider P. the laboratory, A. tzlnlefaciens then splices these desirable
nrzglllnta a nontoxin-producing strain of P. s. pv. tabaci, so we genes into plant protoplasts, which regenerate into resistant
place P.angulnta in synonomy here. Toxin-producing strains plants (Goodman et al. 1987).
destroy chlorophyll, causing chlorotic halos. Nontoxin
strains cause angular leaf spots without halos. BACTERlAL WILT
P.s. pv. tabncz normally infects solanaceous crops Gitman (1968a) cited this disease and causal organism
(tobacco, tomato, potato), but can infest Cnnizabis (Johnson in Russian hemp. Ghani & Basit (1975) described bacterial
1937).Johnson noted, however, that Ca~zizabisplants require wilt of Pakistani C n t ~ n a b i scaused by an unidentified
stressful water soaking " for successful infection. Succulent bacterium. Sands et al. (1987)described an Eriuinia causing a
watersoaked leaves become hypersensitive to infection. vascular wilt in Oxford, Mississippi, affecting at least 1% of
Wind-driven rain splash from infected plants creates plants in the plantation.
epidemics and spreads like wildfire. Symptoms begin as dull green leaf spots, followed by
sudden wilting and necrosis of leaves and stems. Some plants
2. WISCONSIN LEAF SPOT exhibit incipient wilting, others suffer complete wilting and
P s e ~ r d o r n o n a ss y r i n g a e pv. tnellea (Johnson1923)Young et al. die. Seedlings are especially susceptible. Bacteria ooze in a
1978, Ne7il Zetllnnd \ournnl of Agricult~lralResearck 21:153-177. highly viscid fluid from cut stems. The bacterium normally
=Pseudonzonas lr~ellaJohnson 1923, =Bacterittnl nzelleurt~Johnson 1923. grows in the vascular system of cucurbits (cucumber,
P. s. pv. nzrllra normally attacks tobacco in Wisconsin, pumpkin, squash, etc.).
Japan, and the former USSR. Gitman (196%) reported it on
hemp but did not describe symptoms. Lucas (1975) and CAUSAL ORGANISM
others consider the organism a strain of P. s. pv. tnbaci. E r w i n i a t r a c h e i p h i l a (Smith 1895) Bergey, Harrison, Breed,
Hammer & Huntoon 1923, Zentralblntt filr bakteriologie,
CROWN GALL pamsitenkunde, infecktionskmizk/leiten und hygiene, Abteilung 11,1:364-
373.
Lopatin (1936) and Gitman (1968a) described this =Bacterium tvacheipizilz~s(Smith 1895) Chester 1897, =Enoinia
disease in Russia. Lopatin placed Callnabis in the "extremely an~ylovoravar. tvacheipl~ila(Smith)Dye 1968.
susceptible" category. The causal organism attacks dicot Description: Cultures grow slowly in nutrient agar.
plants around the world; 643 species from 331 genera are Facultatively anaerobic, gram (-) straight rods, size 1.0-3.0x 0.5-1.0
recorded as hosts (Smith et 01. 1988). Monocots and pm, with four to eight peritrichous flagella. Nonproteolytic,
gymnosperms are rarely infected. nonlipolytic, acid (but no gas) produced from glucose, tructose,
galactose, methylglucoside, and sucrose. Optimum growth
SIGNS & SYMPTOMS temperature is 27-30°C.
Twelve to 15 days after infection, abnormal cancer-like
growths appear on plants. These beige-coloured, granular- DIFFERENTIAL DIAGNOSIS
surfaced, spherical galls form at the soil line (crown) or grow Bacterial wilt can be confused with drought, nematodes,
on roots underground. Galls rarely arise on above-ground soil insects, some nutritional problems, or wilts caused by
stems and rarely exceed 10 mm in diameter. Crown galls Fusarilln~,Verticillilrnl, and Sclerotil~nlfungi.
stunt plants but rarely kill them.
DISEASE CYCLE & EPIDEMIOLOGY
CAUSAL ORGANISM E. trncheiphiln is transmitted via striped cucumber bee-
A g s o b a c t e r i u m t ~ l m e f a c i e n s(Smith & Townsend 1907) Conn tles (Acalynlnla vittata). Bacteria overwinter in the gut of bee-
1942, / Rlzcteriology 44:353-360. tles and spread via insect excrement, entering through feed-
=Racte~zurntu~nefnciensSmith & Townsend 1907. ing wounds or leaf stomata. E. trackeiplzila lives world wide.
Description: Cultures white, convex, circular, glistening and The Erzvir~ia isolated in Mississippi grew best at tempera-
translucent. Aerobic gram (-) rod, size 1.0-3.0 x 0.4-0.8 pm, with tures above 28°C (Sands et al. 1987).
one to hr~eperitrichous flagella.Nonproteolytic, nonlipolytic, starch
not hydrolysed, acid (but no gas) produced from glucose, fructose,
arabmose, galactose, mannitol, and salicin. Some strains reduce
nitrate, others do not. Optimum growth temperature is 25-30°C.
PHYTOPLASMAS
These organisms resemble bacteria, but lack cell walls
DIFFERENTIAL DIAGNOSIS when viewed under an electron microscope (Fig 3.1).Because
A, tzlii~efnciensis split into three biovars; biovar 1prob- of their superficial resemblance to mycoplasmas, botanists
ably infects Calzizabis. Symptoms of crown gall are unique. provisionally called them mycoplasma-like organisms
(MLOs). Now they are called phytoplasmas.
DISEASE CYCLE & EPIDEMIOLOGY
A. tllnlefacietls may spread via seedborne infection, as SIGNS & SYMPTOMS
demonstrated in hops. A. tunzefaciens can live as a sapro- Typical phytoplasma symptoms include chlorosis,
phyte in soil, unlike most phytopathological bacteria. Bac- dwarfing, and phyllody, which is a distorted hypertrophy
teria in soil are attracted to root sap oozing from wounded of leaves or flowers, also called a rosette or witch's broom.
roots. The bacteria enter through wounds. Phatak et al. (1975) described these symptoms near Delhi,
A. i~inlefnriensharbours extrachromosomal plasmids India. Flowers on male plants proliferated into small
branches causing a bushy appearance. Electron microscopy
coded with "Ti" (tumour-inducing) genes. After A.
 Chapter 6: Other Cannabis Pests & Pathogens 147

revealed phytoplasmas in phloem tissue of infected plants. and may stunt seedlings. Harman et al. (1987) rid Brassicn
Sastra (1973) previously described these symptoms from seeds of Xanthonzonas can~pestrisby soaking seeds for 30 min-
northeastern India but attributed the disease to viruses. utes in NyolateG. This hospital disinfectant is packaged as
Ghani & Basit (1975) reported "bunchy top" on Pakistani two components. Mix ten parts of component A (2.7% so-
plants but blamed viruses. Bush Doctor (unpublished data dium chlorite) and three parts of component B (15.1%lactic
1983) found similar symptoms on feral hemp in Illinois. acid) in 90 parts water. After soaking, rinse seeds in water
and air-dry.
CAUSAL ORGANISM Drenching soil with Bordeaux mixture or fixed copper
Cannabis Mycoplasma-like Organism, Phatak et al., will control wildfire and Xatltllornonas disease but may cause
Zeit. 83:281, 1975.
131~ytop17t/~. phytotoxicity in Cannabis. Copper and streptomycin have
Description: Pleomorphic cells observed only in phloem ele- been used as foliar sprays, but the copper sprays are largely
ments of infected plants, membrane-bound, roughly circular and ineffective. Treating phytoplasma-infected plants with tet-
less than 0.5 pni In cross-section; cell membrane is electron dense as racycline causes temporary remission of disease but does not
visualized by electron microscopy. cure plants (Lucas 1975).
Extracts of Cannabis suppress many bacteria (Krejci 1950,
DIFFERENTIAL DIAGNOSIS Ferenczy 1956, Ferenczy et nl. 1958, Kabelik et al. 1960, Gal et
Symptoms can be confused with those caused by vi- al. 1969, Velicky & Genest 1972, Velicky & Latta 1974,
ruses, especially HSV. Some genetic disorders also produce Klingeren & Ham 1976, Fournier et al. 1978, Braut-Boucher
phytoplasma-like symptoms. et al. 1985, Vijai et al. 1993, Krebs & Jaggi 1999).In the Ckraine,
aqueous extracts of feral hemp were sprayed on crops to
DISEASE CYCLE & EPIDEMIOLOGY protect them from bacteria (Zelepukha 1960, Zelephukha et
Leafhoppers commonly vector phytoplasmas but al. 1963). The extracts worked best against gram (+)
Phatak et al. (1975) could not indict any specific insect. Coryizebacteriz~nzspecies, worked okay against gram (-)
Phytoplasmas also spread by seed transmission. Xanthon~onnsspecies, and were least effective against gram
(+) Bacilll~sandgram (-) Pseudomonas, Erwinia, ~arterinnz;and
CULTURAL & MECHANICAL CONTROL (see Chapter 9) Agrobacteril~irlspecies (Bel'tyukova 1962). Aqueous extracts
Exclude bacteria and phytoplasmas by employing also inhibited soft rot of potatoes, caused by gram (-) Ereoinia
sanitary measures (method 1).Avoid planting seeds from ca~otovoraand other bacteria (Vijai et al. 1993). An aqueous
diseased plants; if seeds must be used, disinfest them with extract of hemp (50 g dried flowers soaked in 1 1 water) in-
hot water (method 11).Long-term storage of seed (for at least hibited E. carotozlora more than the essential oils extracted
18 months) may kill Psel~doirronasin seeds (Lucas 1975). from cinnamon oil (Cintlanzonzliinzeylanicl~nz),thyme (Tlzynuls
Losses from Pseltdornonns increase in hemp planted on al~lgaris),and peppermint (Mentlza piperita) (Krebs&Jag@1999).
unworked soil (Ghillini 1951), so observe methods 1
(sanitation), 3 (weeding) and 2a (deep ploughing). Deep
ploughing in the autumn is effective against some bacteria
(e.g.,P, s. pv. tabaci).Method 2c (soil solarization) works well WEED PLANTS
against A. tz~inefacieizs,but control of Psel~donzonasis Several plants compete with Cannabis, despite the oft-
unpredictable (Elmore et al. 1997).Follow methods 7c (avoid repeated claim by Dewey (1914) that "hemp smothers all
excess humidity) and 8 (optimize soil nutrition). weeds." Giant ragweed (Ambrosia trifida) and bamboo
Protect plants from insect vectors. To prevent crown (Bambusa, Dendrocalamus, and Pllyllostaclzys species) can
gall, avoid wounding plants near the soil line. Bury carrot outgrow Canizabis and shade it from sunlight.
slices in soil to monitor for A. tliinefnciens; in infected soil the Canizabis subdues slower-growing plants after canopy
carrots will develop galls. Genetic engineering has produced closure-when leaves of adjacent Cannabis plants mesh
tobacco and tomato plants with resistance to A.tl~inefnciens. together and shade the soil. Weed suppression works best
Sandru (1977) evaluated different cultivars for their when the canopy closes early, when plants are approximately
susceptibility to X. cnnzpestris. 'Fibramulta' and 'Kompolti' 50 cm tall. This depends on proper seeding density. At lower
were the most susceptible, the most resistant cultivars were seeding densities (<40 kg ha-1or ~ 3 Ibs/acre)
6 the canopy of
'USO-6' ('YUSO-6'), 'USO-13' and 'Afghan hemp.' fibre hemp closes late (not until plants are 100-150 cm tall),
allowing weeds to proliferate (Bocsa & Karus 1997).
BIOLOGICAL CONTROL (see Chapter 10) Short-statured seed cultivars (e.g., 'FIN-314') may never
Agrobacterittm radiobacter close their canopy, so they become susceptible to weed
BIOLOGY: A nonpathogenic soil bacterium that pressure. Grasses can choke the growth of young seedlings
suppresses A. tl~iizefaciei~s (Nogall@,Galltrol-A&, Norbac if hemp is sown late. Any weed may cause problems in thin
83C'E).A.tilii~efacleilsand A. radiobacter are nearly identical stands, where gaps exist in the canopy.
except for a lack of Ti plaslnids in the latter. Some weeds damage Cannabis by harbouring pests or
APPLICATION: Supplied as agar plates or aqueous sus- spreading diseases. For instance, wild mustard (Brassica
pensions, to be mixed into nonchlorinated water. Plates can kaber) is frequently infected with a fungus, Sclerotinln
be stored for 120 days in a cool (8-10°C), dark place. Apply sclerotionltn. The weed disease spread to hemp crops in
as a pre-planting dip, or spray on seeds and seedling roots, Manitoba, Canada (J. Moes, unpublished data 1996).
or apply as a post-planting soil drench. Vines damage hemp by climbing stems, binding plants
together, and sometimes girdling branches. Dewey (1913)and
CHEMICAL CONTROL (see Chapter 11) Dempsey (1975)singled out several bindweeds-Calystegia
For chemical treatment of infested Caiznabis seed, Kotev sepium, Co~zvolvulzisaruensis, and Polygonttm convolvu-
& Georgieva (1969) recommended seed soaks in formalin lus, which are noxious cousins of morning glories. Dewey
(1300 dilution). Maude (1996)soaked infested seeds in strep- (1914)noted that bindweed seeds are the same size as hemp
tomycin for two hours, using a 500 yg ml-1 solution against seeds, and difficult to separate by screening. Seed contami-
Xniztlloi~~oizascninpestris and 1000 pm ml-1 against Pseli- nation commonly occurs in Chinese hemp seed (Clarke, pers.
do~norlnssyriirgae. Unfortunately, streptomycin is phytotoxic commun. 1999).Bindweeds are deep-rooted, difficult to pull
7 48 Hemp Diseases and Pests

up, and can reproduce from roots remaining in soil. In 1988 The ability of hemp to serve as a "smother crop" is
the USDA imported a caterpillar, T y t a lucttrosa, which usually attributed to physical characteristics, such as its
defoliates wild morning glory and (supposedly) nothing else. dense canopy and root system. But Cannabis also produces
Catiizabis Itself can become a difficult weed, despite as- allelochemicals-water extracts of Cannabis inhibit the
surances by Hackleman & Domingo (1943): "Since the crop growth of other plants (Stupnicka-Rodzynkiewicz 1970,
is an annual and is harvested before seed is formed, there is Vimal & Shukla 1970, Srivastava & Das 1974, Pandey &
not much danger of its becoming troublesome." These re- Mishra 1982).Haney & Bazzaz (1970) speculated that terpe-
searchers did not look out their laboratory windows! A field noids produced by Cannabis may suppress the growth of
survey conducted in the same Illinois county located nearly surrounding vegetation. Pulegone, 1,8-cineole,and limonene
900 stands of feral hemp (Haney & Kutscheid 1975). Feral suppress plants (Asplund 1968), and these terpenoids are
hemp grew in central Illinois as early as 1852 (Brendel1887). produced by Cannabis (Turner et al. 1980). Latta & Faton
Schweinitz (1836) found feral hemp growing near Bethle- (1975)suggested that cannabinoids may also play a role; they
hem, Pennsylvania in the early 1800s. The feral hemp de- measured increased THC production in Caizilabis compet-
scribed in these early reports was probably of European an- ing with weeds.
cestry, what Dewey (1902) called 'Smyrna' hemp. In the late Could allelochemicals in Cannabis inhibit the growth
1800s, European landraces were replaced by Chinese hemp, of subsequent crops grown in a crop rotation with hemp?
from seeds obtained by American missionaries (Dewey 1902). Probably not. Barley is another smother crop, it produces an
Chlnese hemp became the breeding stock for 20th century arsenal of allelochemicals, yet it serves in a variety of
American hemp (Dewey 1927).Descendants of Dewey's Chi- cropping systems and rotation schemes (Overland 1966).
nese hybrids now grow wild across central North America. Allelochemicals may overwinter better in no-till situ-
Indeed, Chinese hemp has grown wild in China for almost ations, where crop residues remain in place on the field.
2000 years (Li 1974). Haney & Kutscheid (1975) estimated Muminovic (1991) found that a mulch of Cannabis straw
that feral hemp colrered 5-10 million acres (2-4 Mha) in the inhibited the growth of Agropyron repens, a noxious weed.
USA. At that time, the acreage was expanding. Since then, Allelochemicals may intensify in crops being attacked by
l a ~ venforcement has spent millions to eradicate feral hemp insects; the insects feed on leaves and concentrate the
(Bush Doctor 1986a). allelochemicals in their frass (Silander et ai. 1983).

WEED CONTROL
Do not allow weeds to flower and set seed-"One year
seeds, eleven years weeds." If weeds do seed, the weed seeds
PARASITIC PLANTS
must be separated from hemp seeds prior to sowing. This is Two plants, dodder and broomrape, are genuine plant
accomplished by careful screening. Some weed seeds in soil parasites. have no and On other
can be killed using soil solarization (method 2c in Chapter plants for survival. Dodder and broomrape sink specialized
9). Solarization works best against annual weeds with low (haustoria) into the host's and phloem to
heat tolerance (Elmore et 01. (1997). withdraw fluids and nutrients.
Weeds can be controlled by timely tillage of soil. Dodge
(1898) recommended tilling soil twice-when hemp plants DODDER
are 5 cm tall and again when plants are 10-20 cm tall. Some Five species of this alien-looking plant have been
weed seeds require a brief exposure to light to break pulled off hemp around the world. Dodders are related to
dormancy. Thus, German research suggests that tilling at morning glories. But while morning glories spiral up fence
n~gl~ reduces
t weed germination. USDA researchers posts, dodders have no chlorophyll and must twine around
experimented with night tillage, driving tractors while living plants. In addition to causing mechanical damage,
wearing military-issue night goggles. Night tillage reduced dodders vector viruses as they grow from plant to plant.
the germination of small-seeded broadleaf species (ragweed,
lambsquarters, pigweed, black nightshade), but did not SIGNS & SYMPTOMS
reduce annual grasses and large-seeded weeds. Dodders arise as conspicuous tangles of glabrous yel-
Quackgrass, Agropyron repens, is a creeping perennial low filaments (Plate 74), bearing vernacular names such as
problem of late-planted hemp in New England, the north "gold thread," "hair weed," "devil's ringlet," and "love vine"
central states, and the prairie provinces. It spreads by (Lucas 1975). Robust specimens girdle branches and pull
underground rhizomes. Standard tillage equipment down hosts.
(rototillers, rotovators) merely cut a n d distribute the
rhizomes. Alternatively, use Danish s-tines, which pull 1. Cuscuta campestris Yunker 1932
rhizomes to the soil surface where they dry out and die. This is the common field dodder in North America. It
lives on marijuana in Oregon and Pennsylvania, and
on feral hemp in Illinois (Bush Doctor, pers. commun.
1987). C. campestris has been introduced into Europe
ALLELOPATHIC PLANTS (Ceapoiu 1958).Like most dodders, C.catnpestris infests
Certain plants inhibit Cannabis by secreting allelochemi- a wide range of plants, mostly herbaceous dicots such
cals, ~vhichare toxic. Allelochemicals ooze from roots and as clover (Trifolii/rn)and alfalfa (Medicago).
emanate from leaves as a gas or liquid. Leaf trichomes act as 2. Cuscuta europea L. 1732
repositories for these "natural herbicides." Heavy rainfall col- This species lives in Europe (Kirchner 1906, Transhel et
lapses the trichomes, releases their contents, and washes the al. 1933, Ferraris 1935, Flachs 1936, Ciferri 1941,
contents to the ground. Allelochemicals inhibit seed germi- Dobrozrakova ef 01.1956,Barloy & Pelhate 1962, Serzane
nation and plant growth. Good (1953) noted that Cannabis 1962, Vakhrusheva 1979, Gutberlet & Karus 1995). It
grows poorly near Spinacin oleracea, Secale cereale, Cicia sativa frequently infests hops (Humlili~s)and nettles (Urtica).
and Lepldilint sntiiui~n.Peculiarly, some plants are "anti- 3. Cuscuta pentagons Engelman
alle1opaths"-extracts of Geranii~nzdissectt~rrlactually im- This may be a subspecies of C. canzpestris. It is native to
prove germination of hemp seeds (Muminovic 1990). North America but now lives everywhere (Parker &
 Chapter 6: Other Cannabis Pests & Pathogens 149

Riches 1993). C. peiztagolza attacked hemp in Serbia germinate in late spring after soil is w~ell-warmed.
Seedlings
(Stojanovic 1959). grow a rudimentary root and send up a heliotropic (sun-
4. Cuscutn e p i l i n u ~ nWeihe 1824 seeking) tendril, which gropes for a host. After the tendril
This species has been reported on Cannabis in Italy (Fer- makes contact with a host, the dodder root atrophies and
raris 1935) and Kenya (Nattrass 1941). It normally at- the parasite escapes from the soil (Fig 6.6). Dodders are an-
(Parker & Riches 1993).
tacks flax, Linllnz llsitatissinz~~rn nual plants.
5 . Cilscutn suaveolens Seringe 1840
Dewey (1914)reported "a Chilean dodder" (he called it DODDER CONTROL
Clisc~itaracen~os>)tangling hemp in California. This Observe the following techniques described in Chap-
species normally infests Trijoli~~nz and Medicago species. ter 9: method 1 (burning, especially if Cl~scutahas shed seed),
It originated in Chile but now occurs across the Ameri- 2a (deep ploughing), 2b (steam-sterilize soil), 4 (plant late
cas and Europe. after the spring flush of Czlsctlta germination), 7c (avoid moist,
Description: Dodder stems are threadlike, twine up host stems, 1- shaded ground near streams or rivers), 9 (cut and pull, no
2 m m \vide, orange-yellow in coloul; sometimes tinged with red or fragments should remain on stems or ground). Ceapoiu
purple, sometimes almost white. Tendrils arise at frequent inter-
vals along the stems, tw1n111garound host stems. Leaves reduced to (1958) said hemp seed from infected crops should not be
mmute scales, arising opposite tendrils. Separating Cuscl~taspecies planted. If planting such seed is necessary, carefully screen
by t h e ~ flowers
r is not easy. They all produce small (14 mm), short- all alfalfa-sized dodder seed. Rotate with crops resistant to
11ved flowers. The species cited on hemp produce two styles per dodder-bean (Phaseolus aulgaris), squash (Cucurbita pepo),
flo~~rer; some stigmas halve knobs, some d o not. Even experts make tomato (Lycopersicori lycopersicllnz), and cotton (Gossypi~1171
mistakes in identihcation, as exampled by Rataj (1957),who cited
C~isclitntinnjo~;a 11onexlstent species. Seeds are reddish-brown, 1-2
/zirsutilin) are resistant to C. cnm~estris.
mm long. For more informati011see Yunker (1932)or Parker & Riches The Ch~nesereleased a dodder-speclflc b~ocontrolfun-
(1993). gus, Colletotrichumgloeosporiodes f.sp. cusctltae (Lubao 1 )
New formulations and hypervirulentstrains of the fungus,
DODDER DISEASE CYCLE Lubao 1 S22' and Lubao 2', are now in use (Quimby & Birdsall
Each plant produces thousands of seeds from late sum- 1995). Russian researchers have controlled dodder with an
mer until frost. Seeds survive in soil for up to ten years. Seeds Alterliarin species (Parker & Riches 1993).

Figure 6.6: Disease cycle of dodder, Cuscuta campestris (McPartland & Clarke redrawn from Agrios 1997).

As dodder

Dodder stems twine up host

plant and sink haustoria into
the host for sustenance

Flowers mature
into clusters of

Dodder over-
winters mixed djacent healthy plants

Dodder seeds
overwinter in soil Dodder seed germinates and
climbs up host plant
150 Hemp Disea:;es and Pests

BROOMRAPE 0.ranzosa originated around the Mediterranean, but

Host plants become s t u n t e d , sickly, and d i e now ranges from England to South Africa, and across Asia
prematurely. Broomrapes starve their hosts, provide (Kreutz 1995). 0. raniosa was introduced into Kentucky
portholes for root rot fungi, and vector viruses. Dewey (1914) around 1880 from hemp imported from China (Garman
called broomrape "the only really serious enemy to hemp." 1903). It still persists in Kentucky, California, and Texas
Broomrape infestation seriously threatened the hemp indus- (Musselman 1994).
try in Kentucky (Dewey 1902, Garman 1903, Musselman
1994).According to Dodge (1898), "the Germans call this ter- 2. PERSOON'S BROOMRAPE
rible tireed the Harlfniorder." Orobanche aegyytiaca Persoon 1806
Broomrapes do their damage underground (Fig 6.7). Description: 0. aryyptlacn closely resembles 0, mntosa, but
Only briefly do they appear above ground, sending up shoots tends to be taller (20-45 cm) with larger leaves (5-12 mm). The co-
lvhich quickly flower and seed (Plate 75).Strangely, Fournier rolla is longel; usually 20-35 mm; anther filaments densely hairy.
& Paris (1983) detected THC and CBD in flowers and seeds
Parker (1986) reported this species on Cannabis. 0.
of broomrape parasitizing French hemp aegyptiacn lives in the eastern Mediterranean region,
extending south into Africa and east to India. It attacks the
1. BRANCHED BROOMRAPE same plants as 0, ranzosa. Future research may prove it to be
Ovobanche valnosa L. 1753 a subspecies of 0. ranzosa with hairy anthers.
=P/lrliparnrarrlosa (L.) C.A. bleyer
Description: Broomrape shoots arise near the base of hemp stems. 3. TOBACCO BROOMRAPE
They resemble short shoots of pale, branching asparagus. Shoots Orobanche cernua Loefling 1758
ar,erage 10-20 cm in height (range 8 4 5 cm), and bear brownish- =Orobanchecilirlana Wallroth
yello~;scalelike leaves 3-8 mm long (Plate 75). They produce five Description: Shoots robust, rarely branched, 20-30 cm tall, leaves
to seven pale blue-purple flowers, with a tubular corolla supported 5-10 mm long, bearing numerous flowers. Deep blue-purple flowers
by both a bract and bracteole, less than 20 mm long. Anther fila- do not have bracteoles, corolla 12-20 mm long, anthers arise well
ments not densely hairy, arising from base of corolla. Flowers ma- up the corolla at least 4 mm distal from base.
t ~ l r einto resinous capsules filled with 600-800 tiny, oblong seeds This species parasitizes Cannabis in Romania (Ceapoiu
(PJCII only 260 pm long), see Fig 6.7. 1958) and India (Marudarajan 1950). Parker & Riches (1993)
Kreutz (1995) called this species "hemp broomrape," used hemp as a trap crop to induce 0. cerizua germination.
and noted many experts place it in the genus Phelipaea or The species is a serious pest of tobacco, sunflower (Heliantlzl~s
Pilellpnizclze. Dempsey (1975) considered 0. ralizosa a major annuus), a n d Artemisin species. It lives around the
pest. Rataj (1957) described 0. ranzosa "raging" on hemp in Mediterranean region, east to the Arabian Sea, India, and
the Czech Republic. Barloy & Pelhate (1962) considered a Australia.
combination of 0. ranzosn knd F~lsari~l~n solarzi the greatest
hazard to hemp in southern France. BROOMRAPE DISEASE CYCLE
Several 0, ranzosa subspecies together attack over 60 0. raniosa seeds can remain dormant in soil for up to
plants (Lucas 1975).0. rnnlosa subspecies ranzosa parasitizes 13 years (Garman 1903).Root exudates from hosts stimulate
hemp, tobacco (Nicotinna fnbacunz), tomato (Lycopersicon seed germination. Germinating seeds produce a "radicle"
l~/copt~rs~c~~m), and Salnmil~species. The ranzosa subspecies is 3-4 mm long which drills haustoria into Cannabis roots, pen-
further divided into races; Paskovic (1941) said the Italian etrating xylem and phloem. Ten or more parasites may be
race virulently infested all hemp cultivars, whereas the found attached to a single host. Broomrape spends most of
R u s s ~ a nrace only caused problems in Russian hemp its life cycle underground and unseen. Shoots emerge from
cultivars, not Ukrainian or Italian cultioars. the soil in summer and quickly flower and go to seed. Each
parasite can produce up to 500,000 seeds (Barloy & Pelhate
1962).Mature seed capsules open and disperse the dust-like
propagules to wind and surface water. The sticky seeds also
adhere to Cannabis seeds; Berenji & Martinov (1997)reported
an Orobariclie species infesting seed hemp in ~ u ~ o s l a i i a

CULTURAL & MECHANICAL CONTROL (see Chapter 9)

Observe methods 1 (sanitation) and 2b&c (sterilize or
pasteurize the soil). Method 11 (cleaning seed) requires
nzetic~rlol~s
attention because broomrape seeds easily adhere
to hemp flowers (Dewey 1914, Dempsey 1975). Garman
(1903) killed broomrape seeds by immersing contaminated
hemp seeds in a 60°C water bath for ten minutes; this
treatment did not harm hemp seeds.
Hand-pulling broomrape (method 10)prevents broom-
rape from setting seeds, but crop damage has already taken
place. Broomrape shoots arise all summer and the shoots go
to seed within two weeks, so hand-pulling must be repeated
frequently. Be careful not to pull up the hemp with the
broomrape, especially in light, sandy soil. Method 8 (nutrient
supplementation) offers hope: Berger (1969) said potassium
supplements promote resistance; Abu-Irmaileh (1981)
suppressed broomrape with manure, ammonium nitrate
Figure 6.7: Orobanche ramosa.
(NH4N03), and ammonium sulphate [(NH4)2SO-1].
A. Roots of hemp plants parasitized by 0. ramosa, which is
Method 6 (crop rotation) takes years to be effective.
sending up shoots; 5.Fully-floweringshoot (from Lesik 1958).
Breeding for resistance (method 5) is difficult. Chinese vari-
 Chapter 6: Other Can1iabis Pests & Pathogens 151

eties are very susceptible, according to Bocsa & Karus (1997). day; you will find them under the cardboard, congregated
Barloy & Pelhate (1962)reported a loss of resistance in French for easy picking. Frank (1988)rid pillbugs and sowbugs with
crops. Senchenko & Kolyadko (1973) tested 26 cultivars for a rotenone soil drench.
resistance and ranked them accordingly. Resistant cultivars
include the Italian cultivar 'Carmagnola' (Paskovic 1941), MILLIPEDES, SYMPHYLANS & SPIDERS
the Hungarian cultivar 'Kompolti,' Russian cultivars 'JUS- According to Yepsen (1976),millipedes cause problems
1 ' and 'JUS-87,' and the monoecious cultivar 'Juznoja during dry spells in the summer. Bush Doctor (unpublished
Odnodomnoja' (Dempsey 1975). Most of these cultivars are data 1986) found dark brown millipedes, probably Oxidus
no longer available (see de Meijer 1995). gracilis (Koch), chewing roots of feral hemp in Illinois.
Garden symphylans,
. * .
Sctltigerella immaczllata (New-
BIOLOGICAL & CHEMICAL CONTROL port), are slender, pearly-white, centipede-like arthropods,
Garman (1903)suggested controlling broomrape with 8 mm long. ~ymphylanshave 12 pairs of legs; centipedes
Plzytolnyza oroba~zchiaeKaltenbach. This fly lives every- have 15. Symphylans feed on plant roots and kill young seed-
bvl~ere0. rnnzosn arises (Lekic 1974).Releasing as few as 500- lings. They are troublesome in glasshouses and hop yards in
600 flies per ha provides significant biocontrol (Kapralov the-pacific Northwest. They survive for years in damp loca-
1974).Broomrape infested with I? orobalzclliaemaggots should tions such as decaying logs and deep humus.
be pulled up and laid to the side allowing the fly to mature Spiders are close relatives to mites. They normally feed
and reproduce (Parker & Riches 1993). Unfortunately, the on other arthropods. Bantra (1976)reported finding Argyodes
maggots are hyperparasitized by chalcids. Two fungi may and Cyclosa species on Indian Cantlabis. We d o not kill
urovide biocontrol of broomrape, Fusarium lateritiuln and spiders ... most do more good than harm.
Fzisarilifn o x y s p o r u m f.sp. o r t h o c e r a s (Bozoukov & Control millipedes by applying peat compost (Yepsen
Kouzmano\,a 1994).The latter fungus achieved commercial 1976). Symphylans,
. -
on the other hand, are attracted to com-
use in the former USSR (Quimby & Birdsall 1995). post (move compost piles niuay from gardens). Hops grow-
Trap crops reduce broomrape seed banks in the soil. ers in the Willamette valley reduce symphylans with soil
Traps for 0.mnlosa include bean (Phaseolusolilgaris) sorghum solarization and repeated rototilling (Jackson,pers. commun.
(Sorglz~inzbicaloiir) (Parker & Riches 1993), or white mustard 1997). Eliminate millipedes with nicotine soil drenches and
(B~.nssicnalba) (Lucas 1975). For 0. cernun use cowpea (Vigna repel symphylans by drenching soil with garlic or tobacco
titzg~iiculntlz)or sorghum (Parker & Riches 1993). Tobacco tea (Yepsen 1976).
farmers in ~ u l g a r i aallow geese to graze in infected fields.
Two geese per ha eliminate broomrape (Kircev in Lucas SLUGS & SNAILS
1975):~umiiatingsoil kills seeds, but this draconian measure Slugs can be vexing in humid corners of the globe. Selgnij
is dangerous and expensive. (1982) described slugs attacking y o u n g sprouts in
Watsonville, California. Bush Doctor (pers. commun. 1989)
found grey garden slugs destroying seedlings in
ANIMALS OTHER THAN Pennsylvania, Deroceras reticulatum (Miller) or Deroceras
INSECTS AND MITES agreste (L.).Adults are grey-white or flesh-coloured with grey
markings, growing up to 45 mm long. They can cause serious
damage until plants are 1 m tall (Plate 76).
PROTOZOA Most North American slug pests were introduced from
PJ1yto~zonnsis a newly-discovered genus of protozoans. Europe. The European giant garden slug (Lintax maximus
These species principally infect lactiferous ducts, but also L.) is a frequent offender in vegetable gardens. Other
invade xylem and phloem vessels. Thus far no Plzyton~onas aggressive slugs include Agriolinzax and Arion species. The
species have been discovered in hemp, even though Cannabis brown garden snail, Helix aspersa, is a problem in California.
produces vestigial lactifers (Zander 1928, Furr & Mahlberg Slugs cause two peaks of maximum damage-in April-
1981, Mesquita & Santos Dias 1984, Lawi-Berger et al. 1984). May and September-November. One or two generations
Plzytoiilonns species attack closely-related plants in the arise per year. They overwinter as eggs. Slugs prefer wet
Llrticaceae (Dollet 1984).Insects vectoring these protozoans, weather and crops planted adjacent to meadows, pastures,
such as Nyslzis and Stenoceplzallis species, are common or w o o d s . They usually feed at night. They are
Cnili~abispests. Pl~yton~oizas species are found between hermaphroditic-each slug is equipped with both male and
l a t i t ~ ~ d e50"
s north and 50" south, girdling the globe in female reproductive organs. They can mate with themselves
distribution. if no other slugs are around.
Please note: until recently, several oomycetes were
misidentified as fungi (e.g., Pytlrillnz and Psel~doperoi~ospora MECHANICAL & BIOCONTROL OF SLUGS
species). Because of this historical tradition, oomycetes are Protect small gardens from slugs with a circular strip of
still studied by mycologists and described in Chapter 5. copper (the best), zinc, or steel. The metal interacts chemi-
cally with slug slime, and repels them, if the metal is kept
SOWBUGS & PILLBUGS clean. Many commercial brands are only 5 cm broad, yet Ellis
Members of the arthropod (but not insect) order Isopoda & Bradley (1992) suggested using a band 7-10 cm wide.
are also called called woodlice. Frank (1988) described sow- Chervil (Alltlirisclis cerrfolitinz) serves as a trap crop for
bugs killing Cnnizabis sprouts. These crustaceans have slate- slugs (Ellis & Bradley 1992).Yepsen (1976)repelled slugs with
grey segmented, robust bodies. They are omnivorous and a border planting of common wormwood (Arfeniisin
normally feed at night on decaying vegetable matter. absintlzitlm). Many organic gardeners let ducks or chickens
Sowbugs and pillbugs are controlled with cultural meth- roam their gardens to chug slugs. Researchers are working
ods 1 (sanitation),2a (spring tillage before planting), 3 (weed- on Tetatzocera species, flies that parasitize various slugs.
ing), and 9 (hand removal).Frank & Rosenthal(1978)trapped
pillbugs and sowbugs by placing pieces of cardboard on soil. Phnsmarhabditis lzermaphrodita (=I? nzegid~s)
The pests gather in cool, moist areas during the heat of the BIOLOGY: Asoil nematode (Nemaslug') that infests slugs
152 Hemp Diseases and Pests

(mostly Deroceras and Arion species) and snails (Helixaspersa pheasant (Phasianus cblchicus) depend on feral hemp seed
and miny others). It does best in moist but not waterlogged in the American midwest (Hartowicz & Eaton 1971). The
soil that is loamy or sandy, not clay, and soil temperatures dependence of game birds on feral hemp has led wildlife
between 5-20°C (15°C is optimal). agencies to oppose police eradication efforts (Vance 1971).
APPEARANCE Sr DEVELOPMENT The nematodes are ap- Birds caused substantial losses in Tasmania (Lisson &
proximately 1 mm long. They actively move through soil in Mendham 1995).Sorauer (1958)cited many seed-eating Eu-
search of slugs. within three to five days of infection, slugs ropean birds including the hemp linnet (Carduelis can-
get sick and stop feeding; they die ten days later. nabina), magpie (Pica pica), starling (Sturnus vtrlgaris), com-
APPLICATION: Supplied as infective juveniles in pack- mon purple grackle (Quiscalus quiscula), tree sparrow
ets. They can only be stored for one or two days in a cool (Passer rnontarzus), English sparrow (Passer domesticas),
(2"C), dark place. I? hrrniapllrodita must be poured or sprayed nuthatch (Sitta europaea), lesser spotted woodpecker
into moist soil and not exposed to UV light (apply in evening), (Dendrocopus minor), and turtledove (Streptopelia turtur).
and soil must be kept moist for at least two weeks after In Nepal, Mattheissen (1978) described Himalayan
application. The usual recommended dose is 300,000
A A
goldfinches (Carduelis spinoides) feeding on Cannnbls seeds.
nematodes per m2, every six weeks. P. hernzaphrodita works
best on young slugs and snails, so apply when juvenile slugs BIRD CONTROL
are present. French hemp breeders treat seeds with anthraquinone,
a bird repellent (Karus 1997). Netting protects seedlings in
Rulnina decolla ta small seed plots. Rataj (1957)lamented, "Birds cause the most
BIOLOGY: A decollate snail that preys on snail eggs and serious damage to hemp seed. In spite of their harmfulness
immature garden snails. It does best in moderate humidity we lack protective measures against this danger. It is usu-
and warm temperatures. ally necessary to guard seed-plots and drive the birds away."
APPEARANCE & DEVELOPMENT: X . decollata resembles Chinese farmers keep a constant daytime watch, hang
elongate versions of its prey, growing to 25-50 m m in length. brightly coloured cloth, or use firecrackers to scare away
It feeds at night. During the day it hides under rocks and birds (Clarke 1995).Lisson & Mendham (1995)deployed owl
well-drained rubble. The species reproduces slowly and may decoys and used "bioacoustics"-the broadcasting of bird
take one or two years to control large pest populations. distress calls over loudspeakers. Stretching iridescent rib-
APPLICATION: Supplied as dormant adults. Store up to bons across fields (which vibrate in wind and make annoy-
a week in a cool (8-1O0C),dark place. Release one to tensnails ing sounds), floating helium balloons with large owl eyes,
per m' in moist, shaded areas with adequate organic matter. ringing "bird bells," firing "bird cannon," deploying scare-
Occasionally they eat seedlings, so d o not release in a freshly crows that spray water when set off by motion detectors ...
seeded area. To protect native molluscs, the sale of snails is all work, for a while. Spraying birds with soapy water re-
restricted in some areas. pels birds but may kill them by removing protective oils.
Some birds, such as sea gulls, die after swallowing pieces of
CHEMICAL CONTROL OF SLUGS Alka Seltzers. Synthetic bird repellents and avicides are also
Setting a pan of stale beer in the garden, edges flush available.
with the ground, is a time-honoured slug trap. Slugs wallow
into the beer and drown. For slugs not falling for this old MAMMALS
saw, set out toxic bait-moisten bran withbeer then mix with Antibacterial terpenoids and THC in Cannabis may harm
metaldehyde, a molluscicide. Sluggo@and Escar-Goo kill slugs domesticated ruminants. Cattle in India (Bos indicus) &fused
xvith ferric phosphate (the form of iron utilized in nutritional to eat cattle feed mixed with ganja (Jain & Arora 1988).
supplements). Other commercial slug traps are available.You Driemeier (1997) described Brazilian cattle (Bos taurtls)
can handpick slugs off plants at night. Kill them with a spritz of dying after consuming bales of dried marijuana. Cardassis
saturated salt solution or Bordeaux mixture. (1951) reported the deaths of horses (Eqtius caballus) and
mules (E. caballus crossed with Eqzrus asinus) who grazed
BIRDS on Cannabis in Greece.
Any ornithologist worth her weight in binoculars Conversely, Siegel (1989) described cattle and horses
knobvs that birds devour Carinabis seeds. Darwin (1881)noted happily feeding on flowering tops in Hawai'i. Patton (1998)
that hemp seeds can germinate after passing through the reported cattle and horses eating hemp seed meal. Goats
digesti~etract of birds, if not cracked by beak or gizzard. (Cnpra species) devour Cannabis without ill effects (Clarke,
According to old folk tales, birds sing better songs after pers. commun. 1996).
ingesting a few hemp seeds. The American birdseed industry Dogs (Canisfamiliaris) have sickened and occasionally
used 2 million tons of Cnizilabis seed per year before the died from eating hashish and marijuana (Meriwheter 1969,
Marihuana Tax Act of 1937. After lobbying by the birdseed Clarke et al. 1971, Goldborld et al. 1979). Dogs also suffer
industry, sterilized seed was exempted from restriction. allergic reactions from breathing marijuana dust (Evans
Parkinson (1640)and Martyn (1792) reported, "Hemp seeds 1989).Ferrets (Mustela fiiro) become comatose and die after
are said to occasion hens to lay a greater quantity of eggs." ingesting marijuana (Smith 1988).Cats (Felis cattrs) have no
Lindley (1838)claimed, "Hemp seed ... has the very singular use for Callnabis, but may dig up transplanted seedlings,
property of changing the plumage of bullfinches and attracted to fish emulsion in soil mixes (Bush Doctor, pers.
goldfinches from red and yellow to black if they are fed on it commun. 1993).
for too long a time or in too large a quantity." Non-domesticated mammals cause more problems.
Early reports from Kentucky described the passenger Some are protected by wildlife laws and cannot be killed
pigeon (Ectopistes migratorius) feeding on hemp seeds except under rare circumstances. Wildlife pests include deer
(Allen 1908). Feral hempseed is the most important food of (Odocoileus species) in the USA (Frank & Rosenthal 1978,
mourning doves (Zenaidura macroura) in Iowa (McClure Alexander 1987, Lassen 1988, Siegel 1989),dik-dik (Madoqua
1913).Captive doves thrive for long periods of time on hemp species) in Kenya (Quinn, pers. commun. 1996), and
alone. Bobwhite quail (Colinus virginiantls) and ringtail unidentified monkeys in South America (Siegel 1989).
 Chapter 6: Other Cannabis Pests & Pathogens 153

Other mammals are considered vermin and not pro- tate with hair, urine, and faeces from deer predators (e.g.,
tected by wildlife laws. Rabbits (Sylvilagtls species) cause dogs, humans), or bars of scented soap. ~ a n at k deer breast
problems in the USA (Hartowicz & Eaton 1971, Alexander height (1 m above ground). Scents work best when hung
1987, Lassen 1988).During droughts in Hungary, rabbits strip before males mark their territory in early spring.
bark from the base of stalks in search of moisture (Clarke, Fencing deer requires a 2.5 m tall barricade, or a three-
pers. commun. 1996, Plate 77). Siege1 (1989) cited raccoons line electric fence. Groundhogs can be fenced with wide poul-
(Procyon lotor) and Rattus rattus attacking Cnnnnbis. Rats try netting, 75-90 cm above ground and 30-45 cm buried. A
strip bark from stalks to build nests. Field voles (Microtus ring of tin (i.e., cans with the tops and bottoms removed)
species) and hamsters (Cricetus cricetus) feed on sown hemp protects seedlings from most rodents. Discourage rabb~tsby
seeds in Europe (Sorauer 1955). Mice (Mus mtlscttlus) have removing refuge sites such as brush piles, stone and trash
broken into police vaults to feed on seeds in confiscated mari- heaps, and weed patches. Rabbits shy from bloodmeal. They
juana (Siege1 1989). In the USA, gophers (Citellus species), can be humanely trapped in cages for removal. Set empty
moles (Scalopus species), voles (Microtus species), and pop bottles in mole holes; bottles whistle in the wind and
groundhogs (i.e., woodchucks, Marmota monax), attack vibrate moles away (they work better than those silly plastlc
plants (Selgnij 1982, Alexander 1987, Lassen 1988). windmills).
Groundhogs can quickly destroy a young Cannabis stand. Planting spurge ( E u p h o ~ btnnrgi?inta)
i~~ will stupefy moles
They seem to feed on some plants and roll around in the (Yepsen 1976). Moles and rodents are repelled by castor oil
rest. (Frank & Rvsenthal 1978). The biocontrol bacterium
Salmonella enteritidis var. issatschenko kills rodents. Mice
MAMMAL CONTROL and rats can be eliminated with live traps, spring traps, or
Contact repellents such as Hinders and Deer Awaya poisons. Warfarin and cholecalciferol are much less
can be applied to stalks making them unpalatable to deer. dangerous than single-dose poisons (e.g., sodium
Repel deer with scents-ring the field perimeter with small fluoroacetate, red squill, and strychnine, all of which are
mesh bags filled with bloodmeal. For repellents to be effec- produced by plants). Single-dose rodenticides should be
tive, you must change scents every two or three weeks. Ro- avoided for the sake of wandering dogs.
 "The use ofpesticides is a n act ofdespevatioiz i n a dying agricultz~ve.
It's izot tlze ovei~o7clerilzginvader w e m u s t fear but the zoeakened coizditiolz of the vrctim." -William Albrecht

Chapter 7: Abiotic Diseases

Ab~otic(nonliving) causes of hemp disease and injury in- Plant analysis requires precise sampling protocols. Sam-
clude imbalances of soil nutrients (either deficiencies or ex- ples should be mature leaves, from the upper part of the
cesses), climatic insults (drought, frost, hail), air pollution, plant, free of disease, insect darnage, and dead tissue. Col-
soil toxins, pesticides, and genetic factors. These abiotic prob- lect a composite sample by selecting one or two leaflets from
lems also predispose plants to infectious diseases caused by
other organisms. Drought-stressed plants, for instance, be-
come more susceptible to pathogenic fungi (McPartland &
Table 7.1 : Proposed guidelines for nutrient levels in
Schoeneweiss 1984)
Cannabis leaf tissue.'
Symptoms of abiotic diseases may resemble disease
symptoms caused by living organisms. Conversely, symp- DEFICIENTSUFFICIENT TOXICITY
toms caused by disease organisms have been blamed on NUTRIENT LEVEL RANGE LEVEL
abiotic causes. Early obserl'ers of Dutch elm disease in Eu-
rope, for example, attributed elm death to poison gas released N, '10 <2.0 2.0-5.0
during WWI. Some hemp diseases have unknown causes, P, "/o <0.2 0.2-0.5
such as "grandine" disease. K, % <1.0 1.O-5.0
Mq, % -<o.
-- 1 0.1-1.0
NUTRITIONAL IMBALANCES Ca, % <O.1 0.1-1 .O --
S, % <O.1 0.1-0.3
Poor soil causes most abiotic problems. This chapter
Fe, ppm <50 50-250
treats soil fertility in three sections: 1) symptoms caused by
Zn, ppm 15-20 20-100 s400
nutritional imbalances, 2) correction of imbalanced field soils,
Mn, ppm 10-20 20-300 >300
and 3) correction of glasshouse/growroom soils. The latter
CU, ppm 3-5 5-20 >20
two sections come from different perspectives: the field soil
B, ppm <lo 10-1 00 >I00
section emphasizes "soil balancing" to prevent nutrition im-
Mo, ppm <O.1 0.1-0.5 >0.5
balances, whereas the glasshouse section emphasizes trent-
CI, % <0.2 0.2-2.0 >2.0

t
i i l e ~ of
t imbalances after symptoms arise in plants.
Si, % <0.2 0.2-2.0
The section on field soils was contributed by Bart Hall,
Na, % -<-
I .O 1.o-10
a soil scientist with nearly 30 years experience as an organic
agronomist. His company, Bluestem Associates, is based -CO, ppm <0.2 0.2-0.5 20.5
in Lawrence, Kansas, USA (tel: 785-865-0195), and provides V, P P ~ <0.2 0.2-0.5 s1
consulting for organic, biological, and conventional produc- Adapted from general crop guidelines proposed by Bennett
ers worldwide. (1993).

SYMPTOMS at least a dozen plants (Jones 1998). Collect samples from

Generally, deficiency symptoms of mobile nutrients (N, plants with symptoms and without symptoms, for cornpari-
P, K, Mg, B, Mo) begin in large leaves at the bottoms of plants. son in the laboratory. Samples for soil testing should be col-
Deficiency symptoms from less mobile nutrients (Mn, Zn, lected at the same time and location as leaf samples are taken.
Ca, S, Fe, Cu) usually begin in young leaves near the tops. Standard plant analysis is conducted in a laboratory It
The most common deficiencies arise from shortages of is accurate, but slow and expensive. Alternatively, leaf sap
macronutrients-N, P, K, Ca, Mg, and S. can be quickly tested in the field using leaf tissue analysis
Too much of a nutrient may cause toxicity. Symptoms (LTA). LTAfield kits contain either chemically-treated paper
of toxicity often mimic s y m p t o m s of deficiency. strips or plastic vials with indicator reagents. Battery-
Micronutrients cause most toxicities, especially Mn, Cu, B, powered ion meters for the determination of nitrate,
Mo, and C1. Excesses of anything, however, may stress plants. potassium, and calcium are also available. For information
Obviously, symptoms may be misleading, because on plant analysis and LTA, see Jones (1998).
different problems give rise to identical symptoms (Bennett
1993). Thus, observing symptoms is only the first step in NITROGEN
determining a diagnosis. Always check soil pH. Soil pH Lack of nitrogen (N) is the most common deficiency in
directly affects the availability of nutrients (see Chapter 2 Cnnrznbis (Frank 1988). Symptoms begin with chlorosis of
and Fig 2.1). The next step is soil testing and plant analysis. lower leaves (Plate 78). Yellowing starts down the midrib
Soil testing and plant analysis may uncover hidden and then expands. In extreme cases, whole plants turn pale
hunger (Jones1998). Hidden hunger is a nutrient deficiency yellow-green, and leaves grow sparse and small. Stems and
m,ithout symptoms. Hidden hunger affects the quantity and leaves may accumulate a red pigment called anthocyanin,
quality of crop yield, but no plant abnormalities are seen. but this symptom may also indicate a phosphorus deficiency.
Plant analysis of Caiznabis is problematic, because analysis Frank & Rosenthal (1978) claimed N deficiency sh~ftssex
guidelines ha1.e not been published for this plant. Bennett ratios towards male plants. Because of N's high mobility In
(1993)proposed general guidelines for most crop plants, and soil, N deficiency is common in loose, sandy soils.
these are presented in Table 7.1.
155
156 Hemp Diseases and Pests

Deficiency symptoms reverse within four days of ap- thesis, but Coffmann & Genter (1975) found a negative cor-
plying N fertilizer. Ammonium (NH4+)is best utilized by relation between soil Mg and plant THC.
young plants, whereas nitrate (NO3-)is utilized during most
of the plant's growth period (Bennett 1993). Manure, blood CALCIUM
meal, fish emulsion, and guano are excellent organic sources Berger (1969)says hemp requires large quantities of cal-
of N. See Table 7.3. Free-living, nitrogen-fixing bacteria have cium (Ca).Deficient leaves are distorted and withered, their
been sprayed on plants to serve as "biofertilizers" (Fokkema margins curl, and tips hook back. Apical buds may wither
& Van d e n Heuvel 1986). A z o s p i r i l l l ~ n i species are and die (Clarke 1981). Plants are stunted, stalks are brittle,
commercially available. Azospirillilin brasilense a n d roots are discoloured and excessively branched. Ca
A z o s p i ~ i l l l t n z lipoferzll?~have been isolated from the deficiency predisposes plants to stem and root pathogens
rhizosphere of Canizabis (Kosslak & Bohlool 1983). (Botrytis, F~lsariunz,and Rllizoctoiiia fungi; D ~ f y l e t ~ c h z i ~
Excess N causes lush, dark green growth, susceptible to nematodes). Deficiencies are most common in acidic, sandy
insects and diseases. The tensile strength of fibre decreases soils low in organic matter. Deficiencies may arise in growing
(Berger 1969), and plants become susceptible to stalk tips during periods of very high humidity, because Ca is
breakage (Scheifele 1998, see Plate 79). Roots of N-overdosed carried by transpiration-in 100% RH, stomates close down,
plants discolour and rot. I n extreinis, N-toxic plants turn a transpiration ceases, and tips grow deficient. Excess Ca
golden-copper colour, wilt, and die. Ammonium tends to causes wilting and symptoms of Mg, K, iron, and maganese
become toxic in soils with an acid pH; nitrate is a problem in deficiencies.
neutral to alkaline soils (Agrios 1997). Toxic levels of N can
be leached from soil by frequent, heavy irrigation. A com- SULPHUR
mon drug cultivar of Cannabis ('Skunk No. 1') produces lime Symptoms of sulphur (S) shortage resemble N defi-
green leaves often mistaken for N or iron deficiencies ciency-chlorosis of leaves-except S symptoms begin with
(Schoenmakers 1986, Kees 1988). Beware-'Skunk No. 1' is small leaves at the top of the plant. Stems are thin, brittle,
easily burned by N amendments (Alexander 1988). and etiolated. Drought worsens S deficiency, spontaneous
recovery often arises after rainfall washes S from the sky.
PHOSPHORUS Symptoms of excess S mimic those of sulphur dioxide air
Phosphorus (P) deficiency causes stunted growth, and pollution, described below. S overdose can be partially cor-
plants produce small, dark, bluish green leaves. Branches, rected with Ca amendments.
petioles, and main leaf veins develop reddish purple tints.
But purple tints from anthocyanin are normal in some hemp ZINC
varieties (Dewey 1913) and drug varieties (Frank 1988).The Ainong the micronutrients, zinc (Zn) causes the most
tips of lower leaves may turn brown and curl downward. P problems in Cannabis. Deficiency causes interveinal chlorosis
deficiency delays flowering, flowers become predisposed to of younger leaves. New leaves grow torqued and twisted,
fungal diseases, and seed yield is poor. P deficiency worsens and may drop off the plant prematurely. Flowers grow small
in heavy soil, in acidic soil, and cold, wet soil (Plate 80). and deformed, stems are short and brittle. Zn deficiency
Excess P reduces the availability of zinc, iron, magne- arises in alkaline soils low in organic matter. Adding lime or
sium, and calcium in new leaves. Watch for deficiency symp- excess P may precipitate Zn shortages. Excess Zn causes dark
toms of these elements, especially zinc. mottled leaves.

POTASSIUM MANGANESE
Potassium (K or "potash") deficiency symptoms change Manganese (Mn) shortage leads to interveinal chloro-
with age. Young leaves, according to Rataj (1957), exhibit a sis of small leaves (Coffman & Genter 1977). Extreme Mn
nonspecific chlorotic mottling. Older leaves develop a brown deficiency resembles Mg deficiency-leaf margins turn
"burn" of leaf tips and margins, with curling (Plate 81). nearly white, while veins remain green and boldly stand out
Schropp (1938) described dark-coloured foliage with brown against the pale background. Weigert & Fiirst (1939) and
margins. Small branches and petioles may turn red, stalks Wahlin (1944) described leaves developing necrotic spots
weaken and lodge easily. Flowers are small and sparse. K which expanded and destroyed the plants. Weigert & Furst
shortage becomes more pronounced in dry weather, cold (1939)and Wahlin (1944)described extremely stunted plants
weather, and under low light intensity. K leaches easily, so K (20-30 cm instead of 60 cm in height), and described bad
deficiency arises in sandy soil. Recovery is slow, often weeks Mn shortages in hemp grown on reclaimed marshland
after applying potash. (alkaline soils), especially during cold, wet periods.
Excessive K scorches plants and causes wilting. Excesses Excess Mn arises in acidic soils and in soils that have
may lead to deficiencies of calcium or magnesium. been steam-sterilized. Excess Mn causes mottled leaves with
orange-brown spots. This toxicity appeared in our plants,
MAGNESIUM which grew in compost naturally high in Mn; the glasshouse
Deficiency causes older leaves to develop interveinal air was dehumidified, causing extra transpiration, which
chlorosis-the margins and tissues between leaf veins turn pulled excess Mn into the lea\& Plant analysis revealed Mn
yellow, leaving only dark green veins (Frank & Rosenthal levels of 1500 ppm (Potter, pers. commun. 1999). Overdos-
1978). In extreme cases, leaves turn white (see Plate 61).Berger ing Mn locks up Zn and iron. Mn may play a role in
(1969) described grey-white spotting in lower leaves of cannabinoid synthesis (Latta & Eaton 1975).
stunted plants. Deficiencies worsen after heavy application
of ammonia or K to soil. Sandy, acidic soils commonly lack IRON
Mg, especially soils leached by heavy irrigation or rainfall. Young leaves deficient in iron (Fe) develop interveinal
Plants recover quickly after Mg supplementation. chlorosis with green veins, like symptoms of Mn deficiency.
Excess Mg creates small, dark green leaves with curled Leaves may turn completely white with necrotic margins
edges. Adding calcium acts as an antidote. Latta & Eaton (Storm 1987).Fe shortages arise in alkaline soils (Fig 2.1) and
(1975) considered Mg a co-enzyme required for THC syn- soils with excess P or heavy metals. Adding an acidifying
 Chapter 7: Abiotic Diseases 157

agent to the soil improves Fe availability. Plant synthesis of SELENIUM

THC requires optimal Fe levels (Kaneshima et al. 1973, Latta & Selenium (Se) is worth mentioning. Plants do not re-
Eaton 1975). Excess Fe causes leaf bronzing. quire it, but we do-our RDI is about 70 pg per day. Plants
absorb Se from the soil; soyabean seeds and cereals (wheat,
COPPER rye, rice) are good sources, ifgrown in Se-rich soil. In gen-
Copper (Cu) deficient plants have been called eral, soil in rainier areas have less Se (e.g.,the Pacific Korth-
"gurnmihanf" (Kirchner 1966).Xylem loses its rigidity causing west and the east coast of the USA). Unfortunately, hemp
stems to become rubbery, and plants bend over. Berger (1969), oil is a poor source of Se (Wirstshafter 1997).
on the other hand, described Cu-deficient stems as brittle and
easily broken. Young leaves wilt and necrose at tips and CORRECTING FIELD CROP NUTRITION:
margins. In extreme cases, roots become affected and whole
plants may wilt. Berger described Cu shortages in peat soils. SOIL, FERTILITY, AND GROWTH
Kircl~ner(1966) applied 50-100 kg ha-1 copper sulphate to soil
in autumn to correct deficiencies. by Bart Hall
Excess Cu results in Fe deficiency, especially in acid soils. Bluestem Associates
Toxicity may arise from repeated use of copper sulphate Lawrence, Kansas, USA
fungicides.
Healthy soil is the basis of all successful agriculture.
BORON It is not my intention to provide the reader with a complete
Boron (B) shortages cause grey specks on leaves, or chlo- discussion of soils and fertility, as this information is readily
rosis, with twisted growth. Terminal buds turn brown or grey available in numerous professional and introductory texts
and then die, lateral buds follow suit (Frank & Rosenthal1978). to be found in any decent university library (e.g., Donahue
Stalks swell at their bases, tend to crack, and then rot (Clarke 1973, Berger 1978). Rather, after an introductory overview
1981). B shortages arise in marshland soils (Weigert & Fiirst of soils and how they work, it is my intention to focus on
1939).B shortages also arise in growing tips during periods of soil management approaches oriented towards soil health
excessive humidity. Excess B causes discolouration and death and crop quality, particularly as they relate to hemp
of leaf tips, then leaf margins. production. Those familiar with soil science and its
principles may wish to skim or skip this section.
MOLYBDENUM
Deficiency of molybdenum (Mo-not Mb abbreviated by What is soil, anyway?
Frank iYr Rosenthal 1978) causes "whiptail diseaseu-young Soil is the highly variable result of complex interac-
leaves grow pale, twisted, and withered. Mo shortage tends to tions between parent material, climate, plant cover, and
arise in acid soils. Excess Mo is not common, it causes Fe and time. These interactions eventually result in the develop-
Cu deficiency ment of different layers to make a 'stack' characteristic of
each soil type. The stack is called the soil profile, and fami-
CHLORIDE lies of similar profiles define soil types, groups, and orders.
Chloride (Cl) shortage causes wilting and chlorosis or Of the ten soil orders in the world, only two are gen-
bronzing of leaves. This deficiency is rare and easily erally suited to large scale commercial hemp production
01,ercorrected. Excess C1 also causes bronzing! Overdosing to- on a sustainable basis, the Alfisols and the Mollisols. Hemp
bacco plants with C1 has an adverse effect on cigarette com- production is certainly possible on other soil types, but is
bustibility; C1 fertilizers are eschewed (Lucas 1975). proportionally more challenging.

Table 7.2: Summary of symptoms caused by nutrient deficiencies and overfertilization

Yellowing of young leaves 1 1 2 1

Yellowing of middle leaves 2 2 2 1
Yellowing of older leaves
-- 1 1 1 1 1
Yellowing between veins 1 1 1 1
Leaf pale green 1
Leaf
-- -dark green or redlpurple 1 --
Leaf
---- colour
- mottled 1 1
Leaf small, stunted 1 1 1
Whole leaf brown 2 1 2 1
Leaf wedge brown 1 1 1
Leaftipledges curl 1 2 2 1
Leaf wrinkled - 1
Stem soft, pliable 2
- - 2 - 1
Stem
-- - brittle, stiff -- 2 2 1 1 1
Root stunted 2 1 1
Whole plant wilted 1 1 1

Chart modified from table posted by N.P. Kaye (http:llnpkaye.home.ml.org).

1 = primary symptoms, 2 = secondary symptoms
158 Hemp Diseases and Pests

Alfisols are the second largest order, accounting for 13% limited in unfertilized systems, the solubility of those
of rvorld soils, and primarily found in a broad band centred nutrients inakes them readily available to soil microbes and
on 5OoNin Europe and Russia, the USA maize belt, and much growing plants.
of the Canadian wheat belt. Significant patches are also to be As it is incorporated by a soil microbe or growing plant,
found in the tropics, primarily India, sub-Saharan Africa, a nutrient mineral transfers from the chemical to the
eastern Brazil, and southern Australia. Alfisols can be iden- biological component of the soil. Because it is closely
tified in soil surveys when the longer taxonomic name for a interfaced with living systems, this biological component is
soil type includes the ending "...alf," as (for example), Typic the most dynamic of the three. The biological component
Hapludalf. consists of growing plants, their residues, stable and
Mollisols are the fourth largest order, accounting for 9% decaying organic matter, and a wide range of both soil
of world soils. These are some of the most productive soils microbes (fungi, bacteria, etc.) and soil macrobes [the big
in the world, and probably the most cultivated. Mollisols guys, like worms and grubs).
are predominant in central Asia (south of the Alfisols), Soil organic matter is a delightfully complex subject, but
southeastern Europe, the North American Heartland, and one largely beyond the scope of this chapter. Readers inter-
Argentina. Mollisols always end in " ...011," as in Typic ested in more information about soil organic matter should
Hapludoll. consult texts and journals in a good university agricultural
Soil can be thought of as having three major components library. Two of the real leaders in this field have been Maurice
(and I depart somewhat here from classic soil science Schnitzer (Canada, 1960s through 1980s) and MM Kononova
approaches)-physical, chemical, a n d biological. The (Russia, 1950s and 1960s). For a recent compilation, see
physical component of soil is the particulate mineral matter Jackson (1993).
comprising the bulk of soil mass: sand, silt, clay, gravel, and Soil organic matter is stored solar energy, the remains
so on. Sand, silt, and clay are each precisely defined terms of formerly living plants and animals, mostly plants. These
describing a particular size particle without reference to its remains are present in a variety of forms, ranging from fresh
composition. What soil scientists call "texture" is a shorthand plant tissue, through partially decomposed plant parts, to a
description of the relative proportions of each size class in dark substance bearing no remnants of the original materials
the soil unit. from which it was derived. As a general rule, the ensemble
Sandy soils are considered light-textured (because of these residues is referred to as-"organic matter," and the
they're open and don't hold much water), while clay soils dark, undifferentiated matter, as "humus." In addition to
are considered heavy-textured because they're dense and functioning as the substrate for soil microbial activity, organic
retain a lot of water. Loamy soils have about 40% each of matter I ~ a senough negative charges that it contributes
sand and silt, along with roughly 20% clay. Loams are significantly to overall CEC as well.
medi~~m-textured, and are generally excellent soils to farm. The microbial activity in healthy soil is remarkable, and
The most common Alfisols and Mollisols tend to be silty 1 kg of such soil will be home to 300 million algae, 1000
loams and silty-clay loams; generally easy to work (if not million fungi, 20,000 million actinomycetes, and 4 million
too wet) and highly productive soils. millionbacteria. Under favourable conditions (a freshlv in-
The chenzical component of soils is less precise, and is corporated green manure crop, for example) soil bacteria can
often included with the physical. Because particle names, double their population in less than half an hour.
however, describe only size and not chemical composition,
two soils of similar texture can behave very differently, How does it work for hemp?
depending on their native chemistry. Sand-sized particles With hemp, as with any crop, there is a limited amount
might be made of nearly pure silica, or calcitic limestone, or any grower can do to alter the soil. Changing the physical
even dolomitic (magnesium-rich) limestone. Each of those component is generally impossible, or too expensive to be
soils ~villhave a different response to the same fertilizer practically possible. The chemical fraction can be adjusted,
application because, for example, high magnesium levels sometimes easily, sometimes not. It is helpful to think in
may obstruct potassium uptake by the crop. The silica-based terms of soil correctives as being in a different category than
soil bvill be much more acid than the limestone based soil of cropfertilizers. Generally speaking, crop fertilizers can be used
exactly the same texture. much more efficiently when they are not also interacting with
Additionally, the smallest (clay-sized) particles tend to a soil that is chemically out-of-kilter. One such example
be dominated by a family of minerals with surplus negative would be the case of trying to provide calcium to hemp in a
electrical charges along their rather ragged edges. Positively very acid (low-calcium) soil. Most of the calcium will react
charged elements (cations) in the soil are understandably chemically with the soil in such a way that renders it unavail-
attracted to those negative sites and can be held there until able to the crop.
rernoved by locally-altered soil chemistry. These cations can Adjusting the biological fraction of the soil is relatively
be exchanged between two negative sites or between simple, though neither easy nor quick. For the most part,
negative sites and the water component of the soil, or 'soil more organic matter is better than less, and fewer chemicals
solution.' are generally better than more. Soil organic matter levels are
This chemical capacity of a soil to store and exchange substantially controlled by the rotation used, and within that
cations depends on the amount and specific types of clay, framework by the extent to which soil is open and/or culti-
and can be measured. The resulting measurement, called vated. Soils rich in organic matter are almost always an ex-
Cation Exchange Capacity (CEC), is the most important cellent substrate for a vibrant microbial community. The one
chemical feature of a soil after the composition of the parent exception is poorly drained soils, rich in organic matter be-
material. Heavier soils have a higher CEC and can store and cause there is too little oxygen to break it down.
exchange more cations than lighter soils, roughly two to three Crops can be broadly divided into three categories -
times more, which is one reason the Alfisols and Mollisols soil-builders, neutral crops, and soil-destroyers. Soil-build-
are so fertile and productive. ers are the sod crops: grass, clover, alfalfa (lucerne), and so
The soil solution (watery component) contains dissolved on. Because sod crops cover the soil more or less completely,
nutrients, and although the amounts of such nutrients are there is less exposure to heat and excess oxygen, neither of
 Chapter 7: Abiotic Diseases 159

rvhicl~is good for soil organic matter. Sod crops also produce production. As stated above, hemp is most productive in
prodigious quantities of roots, so on sum, they generate more the Alfisols and'Mollisols so common in the agricultural
organic matter than they allow to be destroyed. heartlands of Europe and America. These are usually some
Row crops like maize (corn), soyabean (soybean), and variant of silty loam (best) or silty clay loam (somewhat
vegetables are at the other end of the spectrum. Much of the harder to work). Local extension agents or other
soil is exposed to the sun for substantial segments of the professional agronomists in the field can be of considerable
growing season. Row crops are often cultivated (especially on assistance in determining the soil order, characteristics, and
biological and organic farms), exposing the soil to degradation suitability to hemp production of the various soils found
by excess oxygen. Furthermore, row crops tend to produce on your farm.
fairly minimal root mass. As a result, under row crops, more Fertilizing hemp for optimum production and quality
organic matter is destroyed than created, and soil health suffers will be substantially easier if the soils in which it is to be
as a consequence. grown are reasonably well balanced. The whole topic of
Drilled crops, such as grains and green manures, fall more "soil balancing" has been somewhat contro-versial for a
or less halfway between the soil-destroyers and the soil- generation or two, and still engenders heated discussion
builders. On balance they will probably be generally neutral in some circles. The concept was largely developed by
in their influence on overall organic matter levels. It is worth William Albrecht, a professor of soils at the University of
noting again that green manures are not particularly effective Missouri from the 1920s through the 1950s (Albrecht &
as soil builders. Their role in a healthy farm system, however, Walters 1975). In general, the proponents of soil balance
is vitally important since their incorporation furnishes soil hold that soils are most productive when exchangeable
microbes with the abundant fresh organic matter they require calcium, magnesium, and potassium levels in the soil
as their primary source of energy. correspond to pre-deterinitzedoptirn~lnllevels ofbase saturation.
Because hemp is a row crop it could be considered a soil- Opponents contend that none of it makes any difference.
destroyer, were it not for one important exception. Much of Base saturation refers to "base" minerals (as in acids
the masslye above-ground production of organic matter and bases), especially calcium, magnesium, and potassium.
remains in the field as a result of the retting process. If that In virtually all cases (fruit being a notable exception) these
organic matter is incorporated into the soil after the harvest soil saturation optima are 70-80% calcium, 12-15% mag-
(instead of being left on the surface to oxidize) hemp's soil- nesium, and 3-5% potassium. The optimum 70-80% for
destroying effects can be substantially mitigated. Hemp also calcium means that calcium occupies 70-80% of the total
has a more extensive root system than other crops considered possible cation exchange sites in-the soil. While calcium
as soil-destroyers. The cautious approach is nevertheless to saturation of 68% is nothing to worry about, calcium in
consider hemp as a soil-destroyer in spite of these mitigating the mid 50s will lead to problems with crop quality. Mag-
factors, and to manage the rotation accordingly. nesium at 17% is similarly not a problem, but above 25%
For a farming system to be sustainable, the crop rotation Mg the soil will indeed be much tighter and compact than
must have at least as many years of soil-building crops as of at lower levels.
soil destroying crops. The number of years dedicated to neutral Particularly with Alfisols and Mollisols, which (after
crops has no significant bearing on sustainability, provided the all) are the soils with which the theory was developed, I
grower remembers that open, uncovered soil is the ultimate have worked with many field situations in which paying
soil-destroying 'crop.' Some illustrative examples may serve attention to soil balance has made a real difference. With
to illustrate this dynamic. out wishing to universalize those observations, the
Albrecht principles of soil balance probably lend
Maize - Soyabean (the most common 'rotation' in the USA themselves rather well to hemp simply because hemp's
Corn Belt) is strongly soil destroying. Maize is a row crop, and optimum zone of production corresponds so closely to that
the soil is left open over winter. On conventional farms the in which the theory was developed.
soya are most usually drilled, making them a neutral crop, but Out-of-balance soils, according to the Albrecht model,
the soil is left open over winter again. usually have one of five problems, or some combination
thereof -low calcium, low magnesium, high magnesium,
Maize - Soyabean - Oats - Clover is substantially better low potassium, and high potassium. Many of the soil
(and substantially more rare). Here, oats (a neutral crop) are correctives that follow are not acceptable for organic
used as a nurse for the soil-building clover. On a conventional certification. Growers for whom certification is important
farm with drilled beans, this is getting close to a sustainable should verify acceptability of materials with their agency.
rotation. On an organic farm, however, with its row-cropped Materials generally (but not always) acceptable in
beans and frequent mechanical cultivation of both the maize organic certification are followed by a 5 symbol. Amounts
and the beans, this would not be a soil-building rotation. are appropriate for silt loams and silty clay loams, and are
listed in kg ha.' (kilograms per hectare). Increase the
Maize - Soyabean - Oats - Alfalfa -Alfalfa -Alfalfa is amounts listed by 10% to arrive at a nearly-exact lb/ac
strongly soil-building, in comparison to both of the other crop equivalent. Reduce the amounts by 40% for sandy soils,
rotations. and increase them by 50% for clays and clay loams.
Substitution of hemp for maize in any aforementioned ex-
amples would be a significant improvement, although it is im- CALCIUM
possible to recommend a hemp-soyabean rotation as be- Ca-saturation 65-75% - Major corrections are unnec-
ing any more agronomically sound than maize-soyabean. essary. Apply 1000 kg ha-' hydrated lime (calcium hydrox-
ide) just before planting hemp and also ahead of any leg-
Correcting soil "balance": umes in the rotation. Monitor every five years.
the step before crop fertilization Ca-saturation 55-65% - Apply 1500 kg ha.' finely
ground calcitic limestones (not magnesium-containing
Within that framework of general soil concerns, it becomes dolomitic lime) plus 1000 kg ha-' hydrated lime. Monitor
possible to examine some specifics as they pertain to hemp after two years.
160 Hemp Diseases and Pests

Ca-saturation 45-55'10 - Apply 3000 kg ha-' finely magnesium correction, and is to be preferred over SPMs
ground calcitic limestones (i~otmagnesium-containing unless the Ca-saturation situation clearly militates in favour
dolomitic lime) plus 1500 kg ha.' hydrated lime. Monitor after of SPM.
two years. In most cases, coriecting for low calcium will
simultaneously correct for low pH (acid soil). A common
MAGNESIUM error in conventional agriculture is to see low soil pH as the
ME-saturation >30% - This is a serious problem soil, problem, rather than as a symptom of low calcium. With
but not an uncommon one in certain areas, such as the USA such an approach, calcium additions to the.soil have tended
upper Midwest. The challenge is high pH, coupled with to be incidental to limestone applications focused on
resulting low calcium. Use the same criteria as for calcium correcting pH, rather than governed by the calcium/
(above), except replace all limestone with fertilizer-grade magnesium regime in the soil.
gypsum4 pellets, and increase the application rate by 50%. This is unfortunate, since calcium as a nlltrienl plays an
Fertilizer-grade gypsums is more expensive than field grades, important role in crop health and quality. Of particular
but because it is much more finely ground (and pelleted), interest to hemp producers is calcium's action in
the calcium is vastly more available to react with the chemical strengthening cell structure. This occurs both at the level of
fraction of the soil. The amounts of hydrated lime are low plant framework (fibre in this case) and at the level of cell
enough that they will not significantly affect pH. Monitor chemistry. Adequate levels of available calcium significantly
after two years. improve the strength of the pectin that is such an important
ME-saturation 20-30% - Use the same criteria as for component of plant cell walls.
calcium (above), except replace all limestone with fertilizer- Strengthening pectin with calcium is a chemical reaction
grade gypsums pellets, and increase the application rate by independent of whether or not the pectin is part of a living
200,;.Monitor after two years. plant or not. Anyone who has used soluble calcium to set
Me-saturation 10-20% - No action specific to the low-methoxy pectins used in making sugarless jams can
magnesium is necessary. With soils having >15% Mg- attest to this fact. In living plants with abundant available
saturation it may be desirable when adjusting calcium calcium, pectin is much more robust. Amongst other things,
saturation to replace 1000 kg ha.' of limestone with fertilizer- it makes cell walls more resistant to the polygalacturonase
grade gypsum? When Mg-saturation is below 15% and enzyme used by most fungal germ tubes to break down plant
limestone is needed to adjust calcium saturation, dolomitic defences when attempting to exploit an infection court. In
limestones5 are acceptable when their magnesium contents plain English that means that there is a greater chance that
are below about 5%. germinating fungal spores will dry out and die before they
Me-saturation <lo% - Significant magnesium shortage. get into the plant and start to grow. In practical terms, the
Use the same criteria as for calcium (above), except replace plant is more resistant to fungal diseases.
all limestone with dolomitic limestone" If dolomitic If a soil has been "balanced" for calcium, soluble calcium
limestone is unavailable in the region, or if Ca-saturation is added as a crop fertilizer will not get tied up trying to adjust
high enough that no limestone of any sort is needed, it will a disequilibrium in the soil, but instead remains available
be necessary to furnish potash in the form of sulphate of for uptake by the plant and subsequent incorporation within
potash-magnesia*in order to supply magnesium. pectin or other elements of plant structure.
Total soil balance requires not only a balance of "base"
POTASSIUM cations (calcium, magnesium, and potassium), but also a
K-saturation >10% - This is often a serious problem balance of anions, particularly phosphates and sulphates:
soil, probably suffering from alkali difficulties. High sodium
may also be a problem, and the p H is often over 8. Such PHOSPHORUS
situations are not at all common in Alfisols and Mollisols. Another common w a y in which soils remain
The most practical solution is an attempt to flood some potash underproductive is a shortage of phosphorus. Phosphorus
out of the soil with heavy applications of fertilizer-grade takes many forms in the soil and is highly sensitive to
gypsum' and modest amounts of hydrated lime. Adequate chemical conditions. As a result, testing for soil phosphorus
rainfall (or irrigation) is necessary for this approach to work. can be a somewhat confusing exercise, given that there are
K-saturation 5-109'0 - Good soil for fruit, but hemp fibre half a dozen possible lab tests that could be used. The trick
quality and disease resistance are likely to be compromised is in finding a test that will give a usable and halfway decent
unless calcium levels are in the 70s or higher. Adjusting for representation of what is actually going on in the soil.
calcium and magnesium will usually decrease potassium For soils anywhere on the acid side of the ledger, the
saturation levels, particularly under the rainfall regimes com- most common test is the Bray, which itself is really two tests,
rnon in prime hemp production areas. one for available phosphate and the other for reserve phos-
K-saturation 3-5% - Ideal levels. No soil correction for phorus. These tests employ two different concentrations of
potassium is necessary though this will not eliminate the the same reagent and are known as Bray P, (weak) and Bray
need to furnish appropriate amounts of potash as a crop fer- P, (strong). Other common phosphorus tests are the Olsen,
tilizer. organ, and Mehlich tests. It is important to know what
K-saturation <39b Correct simultaneously for Ca and
- test has been used, and to remember that phosphorus num-
Mg if these are out of balance. For potassium, use a 50-50 bers for the same soil will vary tremendously according to
mix of potassium chloride (muriate, 0-0-60) and potassium which test is used. Because the Bray P, test is so common in
sulphate"0-0-50), applying as a corrective 100 kg ha I of this USA Maize Belt soils, it will be the focus of these discus-
mix at 3016K-saturation and increasing proportionally by 100 sions.
kg ha for each percent K-saturation below 3%. If Mg-satu- Most soil scientists consider that P, levels above about
ration is below 10% and Ca-saturation levels preclude use of 10 ppm are more than adequate for successful production
limestonei replace each 50 kg ha-' potassium sulphate' with of most crops. While a good case can be made for this posi-
100 kg ha- sulphate of potasll-magnesia5 (SPM, 0-0-22-11Mg- tion, its primary weakness is that it focuses exclusively on
22s). Dolomitic limestones is the least costly source of crop yield, ignoring not only crop quality, but the needs of a
 Chapter 7: Abiotic Diseases 161

healthy soil microbial environment as well. Soil microbes volved aren't too great, this approach can be an extraordi-
need one part of usable phosphorus for every part of narily effective method for kick-starting a worn out soil.
nitrogen, and one part of nitrogen for every 20 to 30 parts of
carbon. Efficiency of carbon cycling, therefore, is ultimately SULPHUR
dependent on phosphorus. For this reason I usually This often forgotten nutrient has two key roles in the
encourage growers to consider corrective action if healthy hemp production system. Most importantly for all
phosphorus P, test levels are much below 50 ppm. If levels crops in the rotation, sulphur is essential for enthusiastic
are below 30 ppm, 1 would fairly strongly urge correction of microbial activity. Most microbes need one part of sulphur
wme sort for each ten parts of nitrogen and phosphorus. What is
For 5011swith pH below about 6 4, perlodi: appl~cat~ons commonly described as the carbon:nitrogen ratio (C:N) is
of soft-rock phosphate' at a rate of 1000 kg ha (every three really the carbon:nitrogen:phosphorus:sulphur ratio
to f ~ v e1 ears, depending on the rotation) are usually (C:N:P:S) and should be approximately 250:10:10:1 for
adequate. Rock phosphates are not only the least expensive optimal microbial health.
source of pliosphate on a kilo-for-kilo basis, but supply about The second important role for sulphur is its significant
2100 calcium and a range of micronutrients (particularly influence on protein qilality. This will affect seed yields, and
molybdenum) as well. Above p H 6.4, however, soil in particular the feeding value of hempseed oilcake for
availability of the phosphate in rock phosphates is limited, livestock. With attentive crop fertilization, sufficient sulphur
and abo\.e pH 6.9, virtually nil. can be provided on a regular basis by adding gypsum6 or
Now is a good time to highlight a common source of one of the potassium sulphatess into the crop fertilizer blend.
confusion in regard to phosphate fertilizers in general, and Correction of soil sulphur levels is rarely needed, and is
the rock phosphates in particular. In order to a l l o ~ l usually ineffective in climates sufficiently humid for
comparison of phosphate materials, many agronomists have profitable hemp production.
agreed on a common basis for expressing phosphate content
of any fertilizer material. The term "available phosphate" MICRONUTRIENTS
has a very specific meaning, much narrower than that used Apart from rare occasions, it is not particularly wise to
in more everyday agronomic speech, referring to the amount attempt micronutrient correction of any soil. Most
of phosphorus than can be extracted using a 2% solution of micronutrients are very expensive, and several are toxic if
citric acid, expressed as P,0; (= P x 2.29).Citric acid solution applied to excess. Even distribution is essential to avoid
is used because, as a weak acid, it roughly approximates localized toxicity, and this requires precisely calibrated
nrhat plant scientists think might be going on as the plant sprayers or spreaders. Since the micronutrient requirements
root interacts with soil phosphate. of each crop in the rotation are different,it makes the most sense to
Total phosphate in a material may or may not be actually supply modest amounts of micronutrientswith the crop ferhlizer.
available to the plant, and ironically, the same caution applies
to the standardized "available phosphate." Most highly Fertilizing the crop
soluble phosphate fertilizers react with the chemical fraction Hemp is a nutrient hog, requiring e\.en more nutrients
of the soil, up to 85% of the "a\railable" phosphate becoming for productive growth than an equivalent crop of makze.
unavailable-to the plant. In acid soils the phosphates react While so-called nutrient extraction measurements are of lim-
~ v i t hiron and aluminium to form extremely unavailable ited utility in evaluating a fertility programme, they can be
compounds. In alkaline soils these same phosphates react somewhat indicative. Whole plant nutrient uptake for a 20
instead ~vithcalcium, forming compounds quite similar to tonne ha-1 hemp crop will be on the order of 180 kg ha-1
those found in rock phosphates. Suggested application rates nitrogen, 50 kg ha-1 phosphate (P,O,), 185 kg ha-1 potash
for conventional phosphate fertilizer material account for (K,O), 120 kg ha-l calcium, 20 kg ha-1 magnesium, and 20 kg
this problem and consequently exceed actual crop needs by ha-1 sulphur (Berger 1969).
several fold. ~oitunately,the retting process is such that the nutrient-
Natural phospl~ateavailability peaks in modestly acid rich leaves generally remain in the field after harvest. If the
soils (pH 6.1-6.7), with abundant organic matter and an leaves are quickly incorporated into the soil (and their
enthusiastic microbial community. If a soil is not yet in that nutrients captured by a wlnter cover crop such as autumn
condit~on,there is an excellent corrective approach available, rye), most of the nutrients contained in the leaves can be
tl~uughit is not particularly cheap or easy. Natural rock phos- rktained in the biological fraction of the soil system. Nutrient
phates?can be mixed with manure and composted to render removal by a 6 tonne ha-I crop of retted stems is substantially
nearly all of the total phosphate available for crop growth. lower, being approximateli50 kg ha-I nitrogen, 15 kg ha''
The organic acids in manure solubilise the calcium pl~os- phosphate, 100 kg ha-1 potash, 45 kg ha-1 calcium, 10 kg ha-I
phates in the natural rock, while the phosphates stabilize magnesium, and 10 kg ha-1 sulphur (Berger 1969). These
and protect much of the manure nitrogen against loss dur- nutrients must ultimately be returned to the soil, lest the
ing composting and application. system be unsustainable over the long term.
Chemically what's going on is that most of the nitrogen Although it is impossible (and rather foolhardy!) to
in manures is in the form of ammonium carbonate, which is provide precise recipes for fertilization of a hemp crop, i t is
unstable. Adding calcium phosphate in the presence of or- often helpful to provide an exaniple as a starting point, which
ganic acids produces calcium carbonate (limestone) and can be modified as soil, rotation, climate, and crop specifics
arnmoniurn phosphate, which is stable, yet available to dictate. Lsine " the above nutrient extraction data as a base,
plants. As a general rule it works well to add about 25 kg of this example assumes that the hemp crop follows a healthy
rock phosphate per m o f manure (50 lb/yd3), composting clover crop providing 70 kg ha-1 nitrogen. Furthermore,
to a medium finish by turning two or three times over two estimated nitrogen release from the breakdown of an
months before field application. Ten to 15 tonnes per hec- hypothetical 3% organic matter content will provide another
tare (1-6 tons/ac) is a good application rate. It is almost cer- 25 kg ha-1 (Menghini pers. commun. 1999, where the ENR is
-I ' ,
tainly not worth doing this if the manure must be bought, estimated at 9 kg ha nltrogen for each percent organic matter
but f; manure is free for the hauling and the distancesin- in the soil, via the Walkley-Black test).
162 Hemp Diseases and Pests

Allow~ingfor the clover carryover and the natural helpful.Although analysis guidelines can be broadly inferred
estimated nitrogen release, a productive hemp crop will nev- from other crops, the accuracy of such tests is highly sensitive
ertheless require an additional 85 kg ha-l nitrogen. This could to the timing of sampling, and particularly to the specific
be provided from about 10 tonnes ha-1 cattle manures, or parts of the plant sampled. We simply do not know whether
about half as much poultry manures. Other sources of nitro- it is the third leaf cluster below the top, or the seventh, or
gen acceptable for organic certification are not economical some other part of the plant entirely that will prove most
for hemp production. The most common dry materials sup- helpful in assessing the nutritional state of hemp. Nutrient
plying nitrogen (bloodmeals, feather meals, alfalfa meals, and levels in plants also shift widely as the plant matures, and
so on) are pricey enough that the cost of their nitrogen tissue analysis results clearly demonstrating deficiency in
approaches £3 kg-' (Euro 5 kg-lor US$2 per lb). Fish solubles early July might indicate a very healthy crop six weeks later.
are more expensive, commonly costing £18 kg-' nitrogen. By Until these parameters have been defined, tissue analysis is
comparison, conventional nitrogen and nitrogen taken out probably more of an expensive curiosity piece than it is a
of the air by a healthy crop of alfalfa costs the grower only useful management tool.
around £0.40 kg-' nitrogen. Before deciding that nutrient deficiency is indeed the
In addition to 85 kg ha-1 nitrogen, the hemp grower cause of symptoms in the field, it is advisable to rule out
should plan to provide 25 kg ha-1 phosphate, assuming the common biotic diseases (see Chapters 4-6), as well as abiotic
soil is well-balanced soil with Bray P, at 15 ppm (amount of damage. Apart from accidental (or intentional) herbicide
P,O, determined as 25[0.70-0.035 (Bray P, in ppm)](stem damage, the most common abiotic distress in hemp will
yield goal in tonnes ha-'). The grower should add 65 kg ha-1 probably be ozone damage, discussed later in this chapter.
potash, assuming a soil with exchangeable potash at 100 ppm Experienced field crop growers should find hemp a
(amount of K 2 0determined as 25[1.166-0.0073 (exchange- relatively easy crop to manage. When sufficiently supplied
able K in ppm)] (stem yield goal in tonnes ha-1).The grower with nutrients, hemp provides a full, dense canopy very
should add at least 45 kg ha-1 calcium, 10 kg ha^' each of effective in controlling weeds during that year of the rotation.
magnesium and sulphur, and an array of micronutrients. Where production is allowed, hemp is an excellent substitute
This can be provided by 250 kg ha-1 (banded at plant- for maize, while being rather less damaging to the soil. If
ing) of a 10-10-10 type fertilizer with 2 Ca-2 Mg-4s-0.5 B- hemp fields are planted to winter cover (such as rye) soon
0.3Cu-0.5Mn-0.4Zn. One tonne of 10-10-10 agronomically after retted steins are removed from the field, hemp
ideal for hemp production can be formulated from the fol- production can strengthen the rotation while increasing
lowing recipe. sustainability of the enterprise both in terms of soil health
and diversified sources of revenue.
360 kg ammonium sulphate Bart Hall is a soil scientist with nearly 30 years
200 kg mono-ammonium phosphate experience as an organic agronomist. His company, Bluestem
110 kg sulphate of potash-magnesia Associates, is based in Lawrence, Kansas, USA (tel: 785-865-
90 kg fertilizer grade gypsum 0195), and provides consulting for organic, biological, and
60 kg potassium chloride (muriate) conventional producers worldwide.
55 kg potassium sulphate
50 kg sodium-calcium borate
15 kg magnesium oxide FERTILIZING GLASSHOUSE
14 kg manganese sulphate AND GROWROOM SOILS
14 kg zinc sulpl~ate
12 kg copper sulphate As Frank & Rosenthal(1978) stated, "Most indoor grow-
ers prefer to buy their soil, while some prefer to dig it." Field
Fertilizer can be formulated more cheaply than from this soils are too heavy for plants grown in containers. Most
recipe, b u t the a m m o n i u m sources of nitrogen are bagged commercial soils are also too heavy. Field soils and
particularly beneficial to hemp and the three different sources bagged commercial soils benefit from the addition of soil
of potash provide a phased release early in the season when conditioners. Soil conditioners balance water retention and
potash demands are low. drainage, and promote root g r o w t h . Popular soil
When the crop is 30 cm tall, sidedress at cultivation with conditioners (in decreasing cost) include granular rockwool,
either of two blends: a) 250 kg ha-1 ammonium sulphate (21- perlite, vermiculite, peat, composted manure, and sand.
0-0) plus either 90 kg ha-1potassium sulphate (agronomically Three examples of soil mixtures:
ideal, but a bit expensive) or 75 kg ha-I muriate of potash, ov
b) 175 kg 11a-l ammonium nitrate (34-0-0) plus 75 kg ha-1 5 parts soil/2 parts perlite/l part composted manure
muriate of potash. Do not mix nitrate and sulphate materi- 5 parts soil/l part sand/l part peat/half part blood meal/
als, as they will usually clump. half part wood ash
Properly fertilized under normal field production con- 5 parts soil/l part vermiculite/l part sand/l part peat/quarter
ditions, hemp will only rarely suffer nutrient deficiencies. part 12-12-12chemical fertilizer
The most probable deficiency to occur in this example is
potash deficiency, induced by an excess of magnesium in Manure and peat are rich sources of organic material.
certain soils. If the grower has previously paid attention to Organic material improves the moisture-holding capacity
questions of soil "balance," this is less likely to be a problem. of sandy soils. Organic material improves the drainage of
Flushing the soil with relatively high levels of soluble cal- clay soils, and make root penetration and tillage easier.
clum - for example 1500 kg ha-1 hydrated lime - will re- Particles of organic material have negatively charged
duce the phenomenon in future. Shifting from one side-dress- surfaces, which keep K, Mg, Ca, and other positively-charged
ing to two (at 25 cm and 50 cm), splitting the nitrogen be- cations from leaching out of soil. Organic material brings
tween the two but doubling the potash by applying the origi- soil to life by providing nutrients for saprophytic microor-
nal amount in each of the two applications, will also help, ganisms. Soil should consist of 4-10% organic material. Or-
especially if done in conjunction with calcium flushing. ganic material must be added to soil every year, since soil
In my experience, plant tissue analysis is probably not microorganisms break it down.
 Chapter 7: Abiotic Diseases 163

To decide on nutrient supplementation, first evaluate organically-grown plants contained 79'10 more THC than
plant symptoms. Next, verify symptoms by testing with pH their sample of 18 plants grown inorganically. Organic crops
meters, chemical soil tests, and leaf tissue analysis. Some cause less laryngitis in smokers than chemically-fertilized
growers forego verification and simply supplement with crops (Clarke, unpublished research 1996). Chemically-fer-
equal amounts of the "big three" nutrients-N, P, and K. tilized, hydroponically-grown Catztznbis contains higher mo-
The results are often suboptimal. lybdenum concentrations than field crops (Watling 1998).
Different crops vary in their demands upon the soil, and Not all organic fertilizers are safe. Frank & Rosenthal
vary in thelr needs for supplementation. Fibre production (1978) recommend spraying leaves with organic fish
requires high soil N, at least 150 kg ha-l. According to emulsion. But Farnsworth & Cordell (1976) warned against
Hendrischke et 01. (1998),supplementing N over this baseline spraying marijuana plants with fish emulsion-virtually all
does not increase yield significantly but restores soil N. At liquid foliar fertilizers, including fish emulsion, contain
harvest, wrliole hemp plants contain about 150 kg N ha-l (= nitrates. Leaves of Cannabis convert nitrates into carcinogenic
16.6 g N m-2). Field retting releases 40% of N back to the N-nitrosamines.
soil, but the soil is still left with a significant N deficit, and Macronutrient deficiencies should be corrected with a
nitrate released by retting is susceptible to further loss from mix of rapidly-absorbed fertilizers and slow-release
leaching (Hendrischke et 111. 1998). Fibre production makes fertilizers. See Table 7.3. For N deficiencies, side-dress plants
equally high demands for K, then in descending order: Ca, with a mix of bird guano (rapid) and composted manure
P, Mg, and micronutrients (especially Si). (slow).Supplement P with a mix of wood ash (fast) and bone
Seed crops, compared to fibre crops, require the same meal (slow). Supplement K with wood ash (fast) and
amount of soil N, less K, and more P (see Table 2.2 in Chap- greensand (very slow).Supplement Mg with dolomitic (high-
ter 2). Drug crops are probably similar to seed crops in their Mg) limestone. If liming is contraindicated, use magnesium
nutrient requirements. Coffman & Genter (1977)found posi- sulphate (Epsom salt) or sul-po-mag (Il%Mg,22%K, 23°10S).
tiare correlations between soil N, soil P, and plant THC. Haney Ca can also be supplemented by liming. If you need Ca with-
& Kutsheid (1973)also correlated soil N with plant THC but out changing pH, apply calcium sulphate (gypsum-21%Ca,
found a ~zegnti;>ecorrelation between soil P and plant THC. 16910s).Sulphur deficiency is rare thanks to acid rain; if
T h ~ disagrees
s with most experts' advice to add-P to soil as needed, apply the aforementioned sul-po-mag or gypsum.
Cnrltlabis begins flowering (Frank & Rosenthal 1978, Frank Micronutrient supplementation is tricky. They are eas-
1988).Coffman & Genter (1975) found a negative correlation ily over-supplemented, causing toxicity. Composted manure
between soil Mg and plant THC. This is surprising, since contains adequate micro-nutrients. Maintaining organic
\lg, along with Mn and Fe, probably plays a role in matter and proper soil pH is usually sufficient for most
cannabinoid synthesis (Latta & Eaton 1975). micronutrients.
In Holland, we supplement with organic fertilizers The pH of soil affects the availability of soil nutrients
rather than chemical fertilizers. Most organic fertilizers, such (see Fig 2.1).In many humid and semihumid areas, soil acid-
as composted manures, improve soil texture and nurture soil ity increases to the point that many nutrients become una-
microorganisms. Chemical fertilizers harm soil texture and vailable (Wolf 1999).Acid soils need to be limed. Limestone
kill soil organisms. Marshmann el nl. (1976) analysed Jamai- (calcium carbonate, CaCO,) is a cheap way to neutralize
can plants cultivated with organic fertilizers, and compared acidic soils. Wolf (1999) provides charts that estimate the
them to plants cultivated inorganically. Their sample of 19 amount of limestone required per acre, depending on soil
pH, soil class (clay loams, silt
loams, sand loams, etc.), and
Table 7.3: Some fertilizers, their N-P-K ratio, and nutrient aI v a i ~ a b i l i t ~ . ~
percentage of organic material in
the soil.
FERTILIZER PERCENTAGE (BY WEIGHT) AVAILABILITY
N P205 K20 TO PLANT
POLLUTANTS,
Natural:
manure (dairy cow) 1 0.75 1 medium TOXINS &
activated sludge 5 3 0.1 medium
----

worm castings 3.5 1 1 mediumlrapid

PESTICIDES
blood meal 15 3 0.75 mediumlrapid Cniznnbis does not toler-
----- -
bird guano 12 8 3 rapid ate excess salt (NaCI), or saline/
bat guano 6 9 1 brackish water, despite the fact
------ --
rapid
fish emulsion 8 3 3- slow/medium that C1 is an essential nutrient
------ - and Na may be beneficial in trace
cottonseed m e x 8 3 3 slow/medium
amounts. Dewey (1902) applied
bone meal (raw) 4 21 0.2 medium
small amounts of KaC1 to soil
-
seaweed 4 21 0.2 medium
and reported an increase in cel-
wood ash 0 2 7 rapidlmedium
lulose production in hemp
-----
phosphate rock 0 30 0 slow
plants. He emphasized caution,
greensa;;;lP 0 1.5 5 slow
--- because "salt is likely to prove
very injurious on light soils."
Chemical: anhydrous ammonia 82 0 0 rapid
Hancer (1992)claimed that "poi-
urea 44 0 0 rapid
sonous salty breezes" killed
amrno-phos(Grade A) 11 15 0 rapid seedlings growing near the Ha-
potassium sulphate 13 0 39 rapid waiian shoreline.
---
triple superphosphate 0 46 0 rapid Many air pollutants are
"Rose Food" 12 12 12 rapid injurious to plants. Injury
, , from
a h pollution peaks d u r i n g
Data collected from various references.
daylight hours, and worsens in
164 Hemp Disealses and Pests

warm, humid conditions. "Acid rain" doubles the damage contaminated with cadmium and copper. The crop efficiently
by acidifying soil and creating various nutrient deficiencies. extracts the toxic metals from soil and accumulates the metals
According to Agrios (19971, the lowest rain pH reported (pH in seeds. The crop is harvested, and the metals are recovered
1.7 in Los Angeles) is more acidic than vinegar (pH 3.0). by leaching seeds with hydrochloric acid (Kozlowski 1995).
Sulphur dioxide gas (SO,) causes interveinal leaf In 1999 a hemp crop will be cultivated near Chernobyl, to
chlorosis in Cannabis (Goidanich 1959). It is toxic at extract radioactive caesium-137and strontium-90 from the soil.
concentrations as low as 0.3 ppm. SO, combines with water Some of the most dangerous toxins are agricultural
to form toxic droplets of acid rain. pesticides. Many insecticides, miticides, and fungicides are
Hydrogen fluoride gas (HF) causes interveinal chlorosis phytotoxic. Pesticides are mixed with solvents, emulsifiers,
and necrosis of leaf tips and margins. Alternatively, leaves diluents, and carriers, which may also cause plant injury.
may develop a white speckling (Goidanich 1959). Young Common symptoms include petiole malformation, leaf burn,
growth is the most susceptible, especially when wet. HF chlorosis, stunting, and bud necrosis.
vapors are produced by various industries and factories Herbicides deserve special mention. Farmers apply tons
processing ore or oil. of herbicides to their fields, and tons of herbicides move off
Nitrogen dioxide gas (NO,) causes symptoms similar target by vapour drift and water runoff. Hemp is very
to those of SO, toxicity, at levels as low as 2-3 ppm. NO, sensitive to herbicide drift (Bocsa & Karus 1997). High
combines w i t h b , in bright sunlight to form ozone. concentrations of herbicides are sprayed on suburban lawns,
Ozone (0,)is the most destructive air pollutant to plants fields, roadsides, railroads, and power line rights-of-way.
(Agrios 1997).Ozone initially causes chlorotic spots on leaves, Herbicides are purposely sprayed on Cannabis for two
primarily on upper surfaces. The spots enlarge and turn reasons. Henzp ci~ltivntorsoccasionally apply herbicides as
brown, leaves defoliate prematurely, and plants become defoliants to make stalks easier to harvest (Goloborod'ko
se~.erelystunted. 1986). Treflanais popular for this purpose (Nazirov &
Ozone binds with incompletely combusted gasoline to Tukhtaeva 1981). Growth-retarding chemicals are also used
produce peroxyacyl nitrates (PANS,a.k.a. smog). PANSmay (Keijzer et al. 1990).Laiv enforcement officials spray herbicides
combine with atmospheric water to create nitric acid rain. on marijuana to render crops unusable. Horowitz (1977)
Sharma & Mann (1984)studied Cannabis indica growing next tested nearly 50 herbicides against Cannabzs and listed 20
to a smoggy Himalayan highway. Roadside plants suffered chemicals which kill or severely injure feral hemp. Most
chlorosis and necrosis. Plants produced smaller leaves with herbicides are absorbed by roots, limiting their effectiveness
shorter petioles than plants growing in the hills. to young plants. A few herbicides work directly on foliage-
Microscopically, polluted plants produced fewer stomates glyphosate (Roundup"), amitrole (Weedazolm),
but inore trichomes per leaf area. Because of increased phenmedipham (Betanal), 2,4-D, and paraquat.
trichome density, S11arma & Mann concluded that auto From 1975 to 1978 the USA government used paraquat
pollution might increase THC concentration. Roadside plants to eradicate Mexican marijuana crops. Paraquat causes plants
collected in the USA were mostly female (Haney & Bazzaz to completely necrose within two days of application (see
1970).This observation agrees with Heslop-Harrison (1957), photograph in Science 119:863). But crops harvested
who reported that carbon monoxide (CO) may cause a shift in~i?zediatelyafter exposure look normal. As a result, normal-
of sex expression in plants, from male to female. looking Mexican marijuana tainted with paraquat appeared
Carbon dioxide ( C 0 7 ) is produced by automobiles, in the USA. In 1983 the Drug Enforcement Agency (DEA)
industries, and burning raihforests. Global levels of CO, are began spraying glyphosate on illicit crops in Georgia and
rising. This may promote photosynthesis, carbon Kentucky. Widespread outrage forced the DEA to stop
accumulation, and plant growth. But it dilutes the plant's N spraying. Recently, the DEA has sprayed feral hemp with
concentration, which forces insects and other N-seeking the herbicide triclopyr.
herbivores to eat more plant tissue (Roberts 1988).Increased
CO" also exacerbates weed problems.
Cnnilabis absorbs many heavy metals from the soil, with-
out ill effects. These toxins deserve mention for their effects CLIMATIC HAZARDS
on us. Ingesting plants laced with lead (a contaminant found As summarized by Hackleman & Domingo (1943),
In chemical fertilizers) causes anaemia and brain damage. hemp is susceptible to lnoistl~restress-too little ortoo much.
Ingesting mercury, another cumulative toxin, also causes Drought can be corrected by irrigating accessible plants.
neurologic deficits. Mercury is released from combustibles Avoid using tap water filtered through a water softening
such as coal and volcanoes. Siege1 et 01. (1988) measured 440 system-it contains too much sodium. Flooding causes the
ng mercury per gram of marijuana grown on Hawai'i's vol- same symptoms as drought-wilting. Wilting may be tem-
canic soil. They noted that mercury is absorbed ten times porary, pending quick correction of the problem. But it of-
more efficiently by the lungs than by the gut, and calculated ten becomes permanent. Overwatering is common. Canna-
that smoking 100 g of volcanic marijuana per week would bis is very sensitive to water soaked, wet soil conditions
lead to mercury poisoning. (Scheifele 1998). Plants can sometimes be revived with
Jurkowska et al. (1990) found high levels of lithium in Oxygen Plus", a hydrogen peroxide product. Flooding is best
hemp (1.04mg kg-'), higher than any other crop plant tested, prevented by intelligent land use and careful field drainage.
including barley, maize, mustard, oats, radish, rape, sorrel, Frost is poorly tolerated by mature plants; temperatures
spinach, sunflower, and wheat. Catlnabis crops grown near below -2 to -3OC are harmful (Hanson 1990).Frosted flowers
Australian uranium mines become enriched with uranium blacken and develop a harsh taste (Frank & Rosenthal1978,
and thorium, whereas crops grown in gold producing re- Selgnij 1982, see Plate 82). 'Gouda's Glorie' and 'Early Bird'
gions contain a significantly enhanced gold signature were two Dutch cultivars resistant to cold and early frost
(Watling 1998). (Kees 1988). Seedlings are less susceptible to frost than ma-
Hemp's ability to hyperaccumulate metals may prove ture plants; they tolerate temperatures down to -7'C (Urban
useful for decontaminating toxic waste sites. In Silesia, 1979, Hanson 1990). Perhaps cold tolerance is related to the
Caiznnbis is deliberately cultivated on wasteland fact that hemp seeds generate more heat upon germination
 Chapter 7: Abiotic Diseases 165

than other crops plants (Darsie et nl. 1914). Seedlings that monoxide shifts sexual expression towards females. Frank
sur\.ive late frosts often have chlorotic leaves. (1988)said root-bound seedlings mature into predominantly
Cnnnnbis tolerates hail and high winds better than other male plants.
large-leaved plants. Nevertheless, high winds can be a Cannabis is normally dioecious-plants produce a 50:50
problem during stages of rapid stalk elongation, before stalk ratio of male (staminate) flowers and female (pistillate) flow-
strength is fully developed. Plants may lodge at the base, in ers. Hemp breeders have developed monoecious plants with
which case they right themselves fairly readily; or the stalk both female and male flowers. These are often called "her-
may kink somewhere above the base-in this case stalks will maphrodites," incorrectly. As Borodina & Migal (1987)
often curve upright again, "gooseneck" fashion (Moes, pers. pointed out, monoecious plants with unisexual flowers (male
cornmun. 1999). and female) are intersexual, not hermaphroditic. Truly
Hail can severely damage young hemp fields (Plate 83). hermaphroditic (bisexual) flowers in Cannabis are abnormal,
Hail injury may ruin fibres for textile use, but the fibres can usually sterile, and rarely seen. Miller (1970) explained
still be harvested for cellulose (paper). Immature seed crops monoecism as the expression of heterozygous genes on X-
suffer little damage from hail, because the injured plants chromosomes or autosomes in XX plants.
branch out and produce more seed (Bocsa & Karus 1997). The deleterious effects of inbreeding were first described
Plants damaged right before harvest, however, may not have by Fleischmann (1934),who reported a 50% reduction in seed
time to rebranch and recover (Moes, pers. commun. 1999). yield. Crescini (1956) illustrated many incestuous
Split branches and broken stalks of valuable plants can be phenomena, such as strange pinnate phyllotaxy and
hand-repaired with splints and tape. ramification of hemp stems. Bocsa (1958)blamed inbreeding
Lightning sometimes strikes hemp fields, leaving for losses of seed and fibre yield, short plant stature (only
circular patches of dead plants. Do not mistake these patches 68% the height of normal hemp), shortened lifespan
for diseases or deer damage. (vegetative growth nine weeks shorter than normal plants),
Poor soil aeration (low soil oxygen) causes lower leaves production of sterile seeds, and increased susceptibility to
to turn grey-green, then wilt and ?all from plants (Frank disease. Recently, Lai (1985) confirmed these mutagenic
1988). Cnniznbis grows in a wide range of relative humidity, effects.
but extremely dry air causes leaf tips to turn brown (Frank The Hungarian cultivar 'Kompolti Sargaszaru' is a
1988). yellow-stemmed mutant (de Meijer 1995). Yellow stem is
Short photoperiods (daylight less than 11 hours) cause caused by a monogenic recessive mutation (Sitnik 1981).The
Cni~izallisto flower prematurely, before full flower and seed gene involved has a pleiotropic effect on plant yields,
yields are attained Night light as low as 0 03 Lumen keeps decreasing biomass, fibre and seed production.
plants from flow~ering(Frank & Rosenthal 1978) Cannabis Mutations may also cause plant fasciation, a twisting
requires a much higl~erlight intensity to keep growing, of stems and leaves (Crescini 1956).Borodina & Migal (1987)
however. Plants become spindly and chlorotic in light illustrated flower fasciation and other flower teratologies (Fig
intensity less than 350 Lumen (see Chapter 2). Too much 7.1). Lyster Dewey collected fasciated plants in Virginia, of
light causes "sun scald." This may happen when indoor the 'Kymington' variety (specimens deposited in the USDA
seedlings are transplanted outside. Undersides of fast- herbarium). The cause of fasciation might be pathogenic
growing leaves are vulnerable if wind turns them over to rather than genetic-Corynebacterit~rn fascians (Tilford)
exposure in brilliant sunlight. Scalded tissues become dry Dowson, a small bacterium, causes fasciation in other crops.
and brown, and resemble "hopperburn" caused by potato For more information concerning the genetics and
leafhoppers. breeding of Cannabis, see Chapter 1 ("host" side of the crop
damage triangle), and Chapter 9 (method 5).
GENETICS
Cntiiinhis is normally diploid, its 27.1 = 20 chromosomes
(18 + XX or XY). Polyploid strains can be artificially pro-
duced. Generally, XX plants are female and XY plants are
male. But sex determination in Carztzabis is not as rigidly ge-
netic as it is in humans. It can be modified by the environ-
ment. Mohan Ram & Sett (1982) found the balance of ethyl-
ene and gibberellins in plants determined sex expression.
They used chemicals (silver nitrate, cobalt chloride, etc.) to
induce male or female flowering. Severe defoliation, cold,
and photoperiods of less than six to eight hours also pro-
duce sex reversal (Clarke 1981). Laskowska (1961) reported
that pollen stored for over two weeks produced more fe- Figure 7.1 : Flower teratologies caused by genetic
male offspring. Haney & Bazzaz (1970) suggested carbon mutations (from Borodina & Migal 1987).
 "Hemp looselzs the soil and lrlnkes it irlove rnello.iu."
-Lyster Hoxie Dewey

Chapter 8: Post-harvest Problems

Carl~labls1s a plant of Inany uses. Nearly all above-ground Retting is accomplished by microbial, mecl~anical,or
parts can be utilized-stalks provide fibre (from phloem cells) chemical means. Microbial water retting is the most com-
and hurds (from xylem cells), achenes ("seeds") provide ed- mon method, described later. Steam retting was practised in
ible seeds and polychotomous seed oil, fenzalef7ozuers pro- the former USSR; it is expensive, takes several hours, and
vide marijuana (wit11 seeds) and sinsemilla (without seeds), produces stiff, unpliable fibre. The new "steam explosion"
and resir1 glands extracted from female flowers provide hash- method is much quicker than old steam retting, but the fibre
ish jcharas) and cannabinoids. All of these products, how- is shredded and unsuitable for spinning. Chemical retting is
e\ er, are subject to decay and destruction by fungi, insects, also called enzymatic retting or alkaline digestion. Chemical
and other organisms. retting produces a high quality fibre, but is t-ery expensive.
Microbial retting is a technical term for rotting. As mi-
croorganisms digest resins and pectins in hemp stems, they
HEMP FIBRE give off a bad odour. Microbial retting can be accomplisl~ed
The history of hemp cultivation fills many pages (Able two ways-either dew retting or pond retting.
1980, Canapasemi 1988, Herer 1991).Agricultural fibres are Dew retted stalks are cut close to the ground and spread
currently making a comeback. Hemp fibres (bast cells) are on fields in thin, uniform layers to catch dew. Robinson (1946)
the hemp plant's phloem or sap-conducting cells. Primary noted dew retting is difficult in dry parts of the USA and
fibre cells are only 10 pm (1/15OU)wide, but up to 3" (7500 Canada west of the 100th meridian (where the great plains
pm) long. Zylinski (1958) described "secondary" fibres, begin). Microorganisms normally dew-ret hemp in four to
which are shorter, stiffer, and weaker than primary fibres. six weeks (Dodge 1890). This happens faster in warmer
Secondary fibres are useful for paper production but conditions and slower in cooler climates. Farmers in northern
undesirable for most textile production. latitudes (e.g., Wisconsin, Ontario, and Siberia) have
IJrimary and secondary fibres overlap in bundles con- practised "snow retting'l-stalks are spread on fields and
tainlng ten to 40 cells per bundle Fibre bundles often run remain under snow until springtime. The disadvantage of
-
the entlre length of plants, from roots to tops About 25
A

dew retting is its dependence on ideal weather.

bundles lle arhund siems, embedded in a ring of phloem Pond retting has been practised in Europe and China.
1-7arenchyma.This ring is sandwiched between stem Pond-retted fibre is superior to dew retted fibre-the retting
;pidermis ("bark") and the cambium. The cambium layer is more consistent and the fibre is not discoloured (dew retted
merges with thick-walled woody xylem cells. This woody fibre, in contrast, is stained grey). Henry Clay trled
laver thins into pith, which surrounds a hollow centred stalk introducing pond retting to ~ e n t u c k i b u t considered
(Fig 8.1). ~ibre-bundles,cambium, and woody xylem are pond water poisonous and an unnecessary hazard (Dodge
glued together by plant resins and pectins. Loosening fibres 1890,1898). French peasants retted hemp in running streams
from the bark and wood is called retting. and even in the Lo~reRlr er, " although publ~cclplnlon IS
generally against river retting because it renders the waters
foul" (Dodge 1890).
II cambium W O O ~ Y X Y ~ ~ ~ I e s rafted in water 1.5 m deep, free of
~ e r n ~ b u n d l are
aquatic growth, and low in calcium and chlorides. The Ini-
phloem \ pith
I tial tem~eratureshould be above 15°C. Within a week the
water turns acldic (from production of butyric acid) and fer-
mentation bubbles appear around bundles. After two weeks
stems lose their chlo>ophyll and turn white
Retting in suarln water speeds the process. Ponds in
China are passively heated by sunlight to 23-30°C (Clarke
1995). Stalks are pond retted for one to three days and then
spread on fields to continue retting a few days more.
Hungarians use geothermally-heated water, contained In
concrete basins. In Poland, warm water retting takes place
in concrete or steel tanks for 24 hours, then fres11 water is
added and brought to an elevated temperature for two or
three days. Approximately 50 metric tonnes of water are ~ ~ s e d
per metric tonne of fibre (Kozlowski 1995).Air compressors
aerate the water, and tanks are periodically inoculated trith
pure cultures of retting organisms.

HEMP RETTING ORGANISMS

Researchers have isolated many retting organisms. Be-
hrens (1902, 1903) and Fuller & Norman (1944, 1945, 1946)
surveyed the dew retting inicroflora in Europe and Korth
Figure 8.1: Cross section of a hemp stalk (Mepartland). America, respectively. Behrens isolated R ~ ~ Z L Jstolon~f~~l.,
~LLS
167
168 Hemp Disslases and Pests

C l n d o s p o r i ~ i l~zie~r b a r ~ ~ ~and
i i , several Mllcor species from dew Recently a number of basidiomycete fungi have been evalu-
retted hemp. R. stolorzifer was more active in warm weather, ated as biopulping organisms. De Jong et al. (1992) degraded
while M ~ r c o rhieiizalis predominated in snow retting. Other hemp lignin with the fungus Plzntzeroclzaete clzrysospori~inz.
Mzicor species involved in retting included Mzicor plz~nzbezis, Less effective fungi included Tranzetes slersicolo~~r,
Tranzetes
M I L C Onzlicedo,
Y and Mucor corticola. On the contrary, in Iowa, villosa, and Bjerkarldera adusto.
Fuller & Norman never isolated M z ~ c o rspecies from retted
hemp. They only found Mticor in artificial laboratory condi- CONTROL OF FIBRE DECAYING ORGANISMS
tions. In fields they isolated 15 genera of fungi, predomi- Retting must be monitored carefully to avoid rotting.
nantly species of Alternaria, Hornzodendr~~nz (=CladosporilLnz), Finished fibre must be dried quickly to stop the retting proc-
Fusarizinz, Plzoiiza, and Cephalospori~iin. ess. Hemp fibres should be stored in cool, dry conditions.
Other fungi have been isolated from dew retted hemp Damp conditions permit dormant fungi to reactivate and
by other researchers, including several Acreinoniz~nzspecies, renew their retting and rotting. Properly stored hemp does
Acrenloiziella atra, Azlreobnsidit~irzpullt~lans,Clzaetostylt~iizfuesitiii not need pesticides. Romanian growers sprayed processed
(=Ascopizorn pulcllra, Bulbotlzanznidiunz elegans), Diplodiella fibres with pentachlorophenol (PCP) to prevent fungal in-
ranzeiztacen, Epicoccnnr nigrunz (=E.pl~rpl~rasce~zs), Go~iatobotys festations, but this was discontinued (Bhcsa & Karus 1997).
silrzples, Plzialopliora species, Pericotzia species, Staclzybotrys
lobzilata, Stnclzybotrys nlter?iaizs, T l i a ~ n n i d i z ~ nelegans, i OCCUPATIONAL HAZARDS
Tor~lla
lzcrL~~1ulcr1z, and many Mortierelln and Ulocladilinz species. Fields workers may develop allergies to hemp. Because
Several of these fungi are cellulose-destroying organisms and of its prolific pollen production, Cannabis is an important
therefore undesirable for retting (Gzebenyuk 1984). cause of hayfever (Durham 1935, Wodehouse 1945, Freeman
Bacteria are less important than fungi in dew retting. 1983).During the late summer and autumn, pollen from wild
Behrens (1903) only identified Bacilli~sasterosport~s (= B. hemp constituted 17% of all pollen in the air over Nebraska
pol?/~izyra).Fuller & Yorman (1944) described bacteria but (Maloney & Broadkey 1940). Significant amounts of Canna-
did not identify the organisms. Caminita et al. (1947) once bis pollen appear over Ann Arbor, Michigan (Solomon 1976)
reported Aerobncter cloacae (=Enterobacter cloacae) from retted and Delhi, India (Malik et nl. 1990).The allergic agent in hemp
hemp but failed to specify whether it was from dew or pond pollen is a water-soluble agent (ANbarro & Fontela 1996),
retted hemp. possibly a lectin (Tumosa 1984),but pure THC and CBD also
In pond retted hemp, scientists cite about 20 different elicit allergic reactions (Liskow et al. 1971, Watsonet al. 1983).
bacteria. All of these names, however, are synonyms of just Individuals allergic to hemp pollen often cross-react with
five species: Clostridiutn acetobl~tylicl~nz, Clostridillm butyriclim,
hops (Lindemayr & Jager 1980).
Clostri~jilinlp e c t i n o v o r ~ t m ,C l o s t r i d i t ~ r np e c t i n o v o r i ~ mvar.
Factory workers handling retted hemp often suffer from
pnr;lztnl, and Clostridii~nifelsine~im. The first two organismsbyssinosis. This serious disease, also termed the "Monday
commonlq rot various substrates. In contrast, ~ . f e l s i n i ~ has im
syndrome," is common to all textile workers exposed to fibre
only been isolated from the mud of Italian retting ponds. dust. After a weekend away from work, renewed exposure
causes chest tightness, shortness of breath, wheezing, and
FIBRE DECAYING ORGANISMS coughing. These symptoms usually dissipate as the work
Retting must be stopped the moment pectins and resins week progresses, only to reappear the following Monday.
are gone. Otherwise, some microorganisms will begin rot- In Italian h e m p workers, the s y n d r o m e w a s called
ting the fibre, ruining it. Since hemp fibre is pure cellulose, "cannabosis." The British physician Schilling (1956)equated
its attraction is limited to organisms " with cellulolvtic en- cannabosis to byssinosis, a syndrome shared by flax, cotton,
zymes. Fungi are notorious destroyers of cellulose fibre jute, and sisal workers. Hemp processors, however, suffer
(Marsh & Bollenbacher 1949).Some fungi cause double trou- the worst (Zuskin et al. 1976). Thomas et al. (1988) report
ble-they cause disease in living plants, then rot fibre after 15% of hemp workers experienced byssinosis ("pousse" in
plants die. These fungi include Botrytis cinerea, Alternaria al- Ireland), whereas only 2.8% of sisal workers were afflicted.
trrnnta, Triclzotheciiinz roseiinz, Epicoccllnz pzlrpllrascens, and A causal agent in fibre dust has not been identified.
Cla~iospori~lnz lzerbnr~inz.Fuller & Norman (1944, 1945) con- Dimitrov et al. (1990) found a high concentration of fungal
sidered Piclzotlzeci~ilnrosetiln particularly destructive. Behren spores in hemp mill air (mostly Cladosporil~in,Altertzarza,
(1902)named C l a d o s p o r i ~ ~/z;rbarz~iiz
n~ as-the most destructive Aspergilllls, Penicilliiiirz, and Fztsarilinz species). Castellan et
and dangerous ceilulose-degrading fungus, although al. (1984) blamed byssinosis on an endotoxin-producing
Thaysen & Bunker (1927) disagreed with his opinion. gram negative bacteria. Carninita et 01. (1947) isolated gram
Other retters that may become rotters include Aspergil- (-) Enterobacter cloacae from hemp dust. They blamed E. cloacae
111s iziger, A s p e r g i l l ~ ~gsl a u c ~ i s ,Acremotziella a t r a , a n d for "hemp fever," a byssinosis-like syndrome with fever,
Staclzybotrys
. . ntrn. Oudemans (1920)listed several rare hemp- chills, nausea, and vomiting. Nicholls et al. (1991)found that
rotting fungi: Dendvylplzillln nlacrosporzim, ~ i ~ l o d i e f l a hemp fibres contain ten times more bacteria than cotton, flax,
or iute fibres. Nicholls isolated E, cloacae and four other
pizlirelinz, Spororl?zia catlnabina, Sor~iariawiesneri, Perisporillm species of Enterobacteriaceae, three species of Psezidonzonas,
dilabens, Perisporiuiiz fiinicz~latuin, and Perisporii~nz klltzzei. and assorted Bacilllls, Coryt~ebacteril~m, Staphylococcz~s,and
(Some of these species requlre taxonomic re-evaluation.) Acinetobacter species-a very pathogenic bunch.
Some fungi rot hemp textiles, such as Staclzybotrys Many researchers fear byssinosis leads to chronic ob-
lohiilati~and A l t e r ~ z a ~ zalternnta
a (Agostin] 1927). Not many structive lung disease. Bouhuys & Zuskin (1976)stated, "We
insects attack finished hemp fibre. carpet beetles cause mi- believe that deterioration of lung function among hemp
nor problems. In Italy, larvae of the Mediterranean flour moth workers begins before the age of 45 and it continues even if
(Eplzrstin kiilziziella) can infest stored fibre (Ferri 1959a). exposure to dust ceases." In one surveyed Spanish town,
the average life span for hemp workers was only 39.6 years.
BIOPULPING ORGANISMS Nonhemp farm workers in the same town lived an average
Removal of lignin from pulp is the most toxic process in of 67.6 years (Schilling 1956).
paper pulping. Most paper companies use toxic chemicals.
 Chapter 8: Post.-harvest Problems 169

microflora of stored seeds. Both researchers isolated Alter-

SEED izarin nlternnta, Rhizopl~sstolon<fern,Cladosporil~mherbarum, and
Plants consist mainly of carbohydrates. Compared to several Fl~saril~in species. Stepanova (1975)also investigated
animals, plants contain much less protein and oil. Herbivores stored seed, but he only identified fungi to genus: Alternaria,
must therefore consume large volumes of plant material to Cladosporil~m,Aspergillus, Penicillium, and Fl~saril~m. Babu et
gain sufficient protein and oil. Seeds contains higher levels al. (1977) identified 16 fungi and a Streptonzyces species
of protein and oil than the rest of the plant (see Table 3.2 in growing on the sl~rfacesof seeds. More importantly, they
Chapter 3), so seeds becomes attractive-to us and to pests. uncovered ten organisms from internal parts of seeds:
Cannabis seed is technically an a c h e n e a small, dry nut. Penicillium chrysogeizl~m,Penicillinrrl frequentans, Penicillium
Carinnbis plants are prolific seeders. Haney & Kutsheid (1975) clzerinesirzi~~nz,
Peizicillium lavitl~nz,Peizicilli~~mfell~~tan~~~v~,
Peni-
counted nearly 7000 seeds per plant in scrubby feral hemp. cilliuin cl~rlicliii,Aspergilllls iziger, Aspergillus sl~lplzlrre~is,
Herer (1991) claimed over half the weight of a well-polli- Cladosporil~mherbar~~til, and Ceplzalosporilrrn clirtipes.
nated female turns to seed. Field-grown crops yield an av- Harvey (1925) discovered that hemp seeds served as ex-
erage of 400 g seeds per plant (Watson, pers. commun. 1998), cellent "bait" to attract water moulds. He steam-sterilized
or 0.5-1.0 metric t ha-1 (Pate 199910).Males (staminate plants) seeds and suspended them in pond water. Since Harvey's
are prolific pollen producers; an average male sheds 40 g discovery, many moulds new to science have been described
pollen, or over 500 million pollen grains (Miller 1970). from "baited" hemp, including a dozen Aclllya species,
Special landraces for seed oil have been selected in Eu- A1lon1yces arb~~scl~la, Isoaclzlya anisospora, Pyflriz~nzspecies
rope, West Asia, Chile (De Meijer 1999), and China (Clarke, (Ogbonna & Pugh 1982), Plzytoplzthora, Pytlziunz, and other
pers. commun. 1997). A new cultivar, 'FIN-314,' has been Allomyces species (Fatemi 1974), Pytlzinnz perigynosuin
bred for seed production in northern climates (Callaway & (Sparrow 1936), Mortierelln rnpllaili var. canizabis (=M.
Laakkonen 1996). Canadian agonomists are currently oantiegheinivar. catznnbis) (Zycha et al. 19691, Geolegizin iizjlata,
breeding new seed cultivars. The drug cultivar 'Skunk No. and Geolegnin septisporangia (Lentz 1977). Olpidil~nllusl~rians
1' is a prolific seed and oil producer (Latimer 1996). Many was isolated from similarly-treated hemp pollen (Sparrow
fibre varieties produce seed consisting of 25-35% oil by 1960). None of these fungi cause decay in properly stored
weight (Dempsey 1975, Deferne & Pate 1996, Pate 1999b).A seeds.
Russian cultivar, 'Olerifera,' reportedly contains 40% oil Many insects feed on immature seeds in the field, such
(Small et 01. 1975). Hemp yields average 455 1 oil per ha (48 as hemp aphids (Phorodon cannabis), hemp flea beetles
gallons/acre), compared to 560 1 ha-1 from sunflower, but (Psylliodes ntfennntn), hemp borers (Grnpholitn delineaizn),and
yields of hemp oil up to 800 1 ha-1 have been reported. European corn borers (Osfrinia nlibilalis).In Kansas, Eaton et
Hemp seed oil has been used for industrial applications al. (1972) also found Clzaetoizenzn flea beetles, treehoppers
such as lightzng, lubrication, and a base for soaps and (Membracidae),leafhoppers (Cicadellidae), cucumber beetles
detergents. The fatty acids in hemp oil are "quick drying," (Dinbrofica species), and stinkbugs (Neznrn species) feeding
~ r h i c hmakes hemp oil useful for paints, varnishes, and on seeds. Seed-destroying mites include Acarzrs sir0
printing inks. Hemp oil can also be burned as fuel; it has (=Tyroglyplphl~s fariizae), Glycyphagl~sdestrlictor (Boczek et al.
combustion and viscosity ratings similar to # 2 heating oil. 1960), Tyrophng~~s noxius (Dombrovskaya 1940), Tyrophagl~s
Hemp seed is very nutritious. In China, whole hemp longior (Chmielewski 1984), Tyrophagi~sputrescentiae
seed is commonly eaten, roasted and raw. Li (1974)reported (Chmielewski & Filipek 1968),Acl~lopscannnbicola (Hartowicz
a traditional Chinese belief regarding the consumption of et al. 1971), and Epitetmnychus species (Rataj 1957).
hemp seeds, "...if eaten over a long term ... it makes one Some insects are only associated with seeds in storage,
communicate with spirits and lightens one's body." not in the field. Arnaud (1974) listed the Indian meal moth
Per 100 g, Carlilnbis seeds contain 421 calories, 21.7-27.1 (Plodia inteup~~nctella), foreign grain beetle (Ahasverl~sadvetzn),
g protein, 27.1-34.7 g carbohydrate (including 18.8-20.3 g rusty grain beetle (Cryptolestesferr~~gineus), flat grain beetle
fibre) and 23.6-30.4 g fat, plus many vitamins and minerals (Cryptolesfes pl~sillus),saw-toothed grain beetle (Oyzaeplzilus
(Duke 1985). Wirtshafter (1997) offered slightly different st~rinnmeilsis),milichiid fly (Desinonzetopa species), fruit fly
nurnbers: 503 calories, 22.5% protein, 35.8% carbohydrate, (Drosoplziln bl~sckii),fungus gnat (Bmdysia species), and
and 30°/0 fat. Wirshafter also analysed vitamins, minerals, scavenger flies (Scatopsefi~scipesand Desrnonzetopa species).
amino acids, and fatty acids in hemp seeds. Glover (1869) cited Laenzophloe~~s nzodesf~ls,a cousin of the
The primary hemp seed protein, edestin, is easily di- flat grain beetle. Strong (1921) and Smith & Olson (1982)
gested and contains all eight essential amino acids (Pate found the confused flour beetle, Triboli~~nz confi~sl~nz,
1999b). Hemp seed oil contains a rich array of fatty acids- bumbling in marijuana seeds. Schmidt (1929) reported
80% polyunsaturated, 100;" monosaturated, and 10% Syrphid larvae in hemp seeds.
saturated (Pate 1999b).Some polyunsaturated fatty acids are Rataj (1957) characterized birds as the most damaging
essential nutrients, such as omega-3 fatty acids (e.g.,linolenic pests of outdoor seed crops. Mice are the greatest destroyers
acid) and omega-6 fatty acids (e.g., linoleic acid, gamma- of stored hemp seeds. See Chapter h for a discussion of birds
linolenic acid, stearidonic acid). The percentage of omega-6 and mammals in Cannabis crops.
and omega-3 fatty acids in hemp oil are present in a 3:l ratio,
considered optimum for human nutrition (Pate 199913). CONTROL OF SEED DETERIORATION
Hemp seed, like finished fibre, should be stored in cool,
SEED DETERIORATION dry conditions (Toole et al. 1960). Seed maintained near 1°C
Many seeds are lost to other organisms. As neatly retains highest germination rates. Scheifele (1998) recom-
summed up in an olde English verse: mended drying seed to 4 2 % moisture before storage, and
heat used for drying seed should not exceed 30°C.
"One for the rat, one for the crow, Seeds harbouring bacteria and fungi can be disinfested
one to rot, and one to grow." with thermotherapy (see method 11 in Chapter 9). Seed not
destined for food use can be treated with pesticides (see "Pes-
Fungi may ruin entire lots of poorly-stored hemp seeds. ticide Application" near the end of Chapter 11).Review arti-
Pietkiewicz (1958) and Ferri (1961b) investigated the
170 Hemp Diseases and Pests

cles b y Ferri (1961), Robinson (1943a), and Koehler (1946) (Thurston 1900). To treat Caiitzabis overdoses, l e m o n juice
cite the extensive literature o n this subject. has b e e n recommended b y homeopaths (Hamilton 1852) and
Ironically, researchers h a v e used Cannabis leaves or Can- Ayurvedic practitioners (Shanavaskhan et al. 1997).
ilnbis extracts t o protect other crops. Z e l e p u k h a (1960) T h e t a x o n o m y o f medicinal Canizabis is s o m e w h a t
described a preparation f r o m feral h e m p that protects potato confused, as described i n Chapter 1. True Cannabis indlca w a s
and t o m a t o seeds f r o m bacterial diseases. Pandey (1982) utilized b y Ayurvedic practitioners i n India; ancient healers
protected millet seeds f r o m over 25 species o f f u n g i w i t h carried this plant t o southeast Asia and Africa over 1000 years
extracts o f Indian Cannabis. Riley (1892) and MacIndoo & ago (Clarke 1999). In t h e 1800s, Indian servants brought C.
Stevers (1924) scattered Cannabis leaves a m o n g bags and iiidica t o British Jamaica, where it spread t o Mexico and South
heaps o f grain t o protect against grain weevils. A m e r i c a . Cannabis afgharlica, i n c o n t r a s t , c o m e s f r o m
Afghanistan and Pakistan. Tbvo m o d e r n cultivars, 'Afghan1
#I' and ' H i n d u Hush,' are stable progenies o f C . afghaizica,
bred for t e n years i n California ( D e Meijer 1999). Stable,
EUPHORIANTS & inbred varieties o f C , indica include 'Durban,' f r o m South
Africa, and 'Haze,' a multihybrid w i t h ancestors f r o m Co-
MEDICAMENTS lumbia, Mexico, Thailand, and southern India ( D e Meijer
Several investigators believe Hoino sapiens w a s originally 1999). ' S k u n k #I' is a cross o f C. indica ( f r o m Mexico and
attracted t o Cannabis for its euphoriant qualities, not for its Columbia) and %. afglianica. ' N o r t h e r n Lights' is one-fourth
fibre or seed. Reininger (1946) e v e n speculated, " U s e o f C. ivdica ( f r o mThailand) and three-fourths C. afghanica ( D e
Canizabis for fibre occurred rather late." T h e Scythians w e r e Meijer 1999).
likely aware o f its intoxicating properties. Indian u s e o f Stockberger (1915),w h o probably grew C. indica, reported
Clznnlzbis as a d r u g m a y date back 12,000 years (Morningstar a yield o f 400-500 lbs o f dried, unseeded flowering tops per
1985). M o r n i n g s t a r h y p o t h e s i z e d t h a t t h e m e d i t a t i v e acre (448-560 k g ha-1). U n d e r intensive indoor cultivation,
practices o f H i n d u culture helped m a k e Cannabis socially m o d e m indica-afshanica hybrids yield 250-500 g flowering tops
acceptable. In contrast, t h e plant's euphoric e f f e c t s w e r e per m2,or 2,500-5,000 k g ha-l(D.Watson, pers. comrnun. 1998).
incompatible w i t h Chinese philosophy (Li 1974). T h u s , the Dried flowers yield 10-15% T H C , or 250-750 k g T H C ha-1
Chinese utilized t h e plant for fibre, f o o d , and medicine, b u t Field-cultivated Cannabis yields a b o u t 1.3 litre o f
not as an intoxicant. essential oil per metric tonne fresh weight,or about 10 1 ha '
U s e o f Caizilabis as a medicine extends into prehistory. (Mediavilla & Steinemann 1997). The yield o f essential oil
In Cai~nabinoidsas Therapelltic Agents, M e c h o u l a m (1986) increases at l o w seeding rates (optimally 5 k g ha-'= 15 plants
listed 20 medicinal uses o f Cannabis e m p l o y e d b y traditional per mz), and increases w h e n pollination is prevented (18 1
societies. See Table 8.1, reprinted f r o m Mechoulam's preface ha.' i n sinsemilla crops, versus 8 1 ha-1 i n pollinated crops).
( w i t h Raphael's permission). In pollinated crops, yield peaks w h e n about 50% o f seeds
In Western medicine, Cannabis enjoyed a 100-year heyday, reach maturity (Meier & Mediavilla 1998).
beginning w i t h t w o Irish physicians stationed i n India
(O'Shaughnessy 1839, D o n o v a n 1845), and ending w i t h the MEDICINAL PLANT ANATOMY
U S A Marihuana Tax A c t o f 1937. T h e inaugural issue o f t h e Trichomes (leaf hairs) cover nearly all above-ground
I O I L Y I I ~oJtlze
J Ainericaiz Medical Association praised t h e valuable parts o f Cannabis. There are t w o t y p e s o f trichomes, glandll-
properties o f Canizabis indica ( B r o w n 1883). Sir W i l l i a m Osler, lor and notlglandillar.
recognized as t h e finest physician o f his time, recommended ~ l a n d u l a rtrichomes are t h e site o f cannabinoid
Cniliznbls indica as t h e " m o s t s a t i s f a c t o r y r e m e d y " f o r synthesis. Glandular trichomes c o m e i n three sizes
migraine (Osler 1918). By 1937, t h e year it w a s restricted, at ( H a m m o n d & Mahlberg 1977).All are divisible into a n upper
least 28 pharmaceutical preparations i n t h e A m e r i c a n secretory section ("head") subtended b y a n auxiliary section
Pharmacopeia contained Cannabis iizdica (Sasman 1938). ("stalk"). T h e smallest and simplest glandular trichomes,
termed bt~lbousglands, usually consist o f two-celled heads
o n two-celled stalks. T h e i r overall height is 25-30 p m ,
Table 8.1: Medicinal use of Cannabis in folklore. including a 20 p m diameter head. Bulbous glands arise o n
all aerial parts o f the plant except hypocotyls and cotyledons.
Analgesic* Antiparasitic* T h e second t y p e , capiiate-sessile glands, produce heads
Anaesthetic Antirheumatic* t w o or three times larger t h a n bulbous glands. These heads
Antiasthmatic* Alleviation of memory loss consist o f eight t o 16 secretory cells arranged i n a flat disc,
Antibiotic* Appetite promoter* covered b y a t o u g h b u t distensible sheath. Accumulation o f
Anticonvulsive* Facilitation of childbirth secretionsbeneaih t h e sheath swell it into a spherical head
Antidepressive Hypnotic* u p t o 60 p m indiameter. Capitate-sessile glands are not truly
Antidiarrhoeal* Reduction of fatigue sessile b u t possess a short stalk o n e cell i n height and t w o t o
Antimigraine Sedative* four cells thick. T h e y arise o n leaves and flowers.
T h e third t y p e , capitate-stalked glands, produce heads
"Modern research substantiates the medicinal uses similar t o those o f capitate-sessile glands ( b u t larger-up t o
marked by an asterisk. 120 p m diametre), w h i c h are subtended b y large multicellular
stalks 100-200 p m tall (Fig 3 . 2 ) . S o m e researchers s a y
Alternative medical systems utilizing Callnabis include capitate-stalked and capitate-sessile glands are identical, just
Ayur\,eda ( D a s h 1989a, Kapoor 1990), Tibetan m e d i c i n e d i f f e r e n t d e v e l o p m e n t a l stages o f t h e s a m e structure
(Molvray 1988, Dash 1989b), Chinese medicine (Reid 1987, (Dayanandan & K a u f m a n 1978, Pate 1994). Differencesexist:
Bensky et al. 1993), and homeopathic medicine ( A l l e n 1875, capitate-stalked glands d o n o t f o r m until plants flower,
Sukul et n1. 1986). A m o n g 19th-century American herbalists, whereas capitate-sessile glands (and bulbous glands) f o r m
the Eclectics praised Cannabis (King 1854, Scudder 1875),b u t m o n t h s earlier. capitate-stalked glands arise only o n bracts
the Physiomedicalists c o n d e m n e d t h e herb as a neural toxin and t i n y s u b t e n d i n g l e a f l e t s o f f l o w e r s ( H a m m o n d &
 Chapter 8: Post-harvest Problems 171

Figure 8.2: Storage fungi

isolated from marijuana.
Left to right: Rhizopus
stolo.nifer, Mucor hiemalis,
Penicillium chtysogenum, P
italicum, Aspergillus flavus, A.
niger; A. fumigatus. Top row:
sporophores cross-sectioned
to reveal internal structures
(~400).Bottom row: natural
habit of above fungi (x25),
McPartland.

Mahlberg counted up to 842 glands on a single bract!). Lastly, They studied confiscated Mexican marijuana, at the request
stalked glands contain higher levels of THC and CBD than of DEA agents whose offices were overrun by these insects.
sessile glands. The predominant pest, Tribolillnz confitsztnz (confused flour
Nonglandular trichomes have been studied from a fo- beetle), only fed on seeds (not marijuana). Two other beetles
rensic viewpoint, since they survive combustion and serve cited in thestudy, Adisternzia zuatsoni and Microgramn~ear'ya,
as a marker of marijuana ashes. Nonglandular trichomes are were fungus feeders (the marijuana was mouldy). It seems
unicellular with silicified walls a n d warty surfaces. the researchers found n o insects that actually fed on
Nonglandular trichomes come in huo sizes. Shorter ones (70- marijuana proper-no Cannabis equivalent to ~nsiodernza
125 pm long) contain cystoliths of calcium carbonate and serricorne, the tobacco cigarette beetle. Arnard (1974)
are generally restricted to adaxial (upper) leaf surfaces (Fig investigated "infested evidence" in San Francisco. He
3.6).Long trichomes (250--370 l m in length) resemble long discovered scavenger flies (Scatopsefuscipes, Drosophila bitsckii,
curved glass needles. They arise on ad- and abaxial leaf Desinometopa species, and Bmdysia species),two fungus-eating
surfaces, flowers, petioles, and stems. beetles (Microgramnze arga and Typhaea stercorea), and a mite,
Maclzrocheles nzztscudomesticae. Piao (1990)examined Cantzabis
STORAGE DISEASES A N D PESTS in Chinese herbal medicine stores and found three mites,
Marijuana crops are unusual considering they may not Glycyphnglts destrztctor, Tydens koclzi, and a Tarsonenzlts species.
reach consumers for years after harvest. This interlude pro- Group 2 bacteria include Clostr~diumspecies which rot
\.ides wide berth for other "consumers" to cause decay and Cannabis in anaerobic conditions, such as damp marijuana
destruction. These "consumers" can be categorized into two stored in an airtight container (Bush Doctor, unpublished
groups. Group 1 consists of field organisms which infest data 1977).Many aerobic bacteria grow on damp marijuana.
living plants and carry over as storage problems. Group 2 Some of these bacteria harm humans-Klebsiella plzelliizoniae,
consists of storage organisms which are saprophytes and Eizterobacter cloacae, group D Streptococclrs (Ungerleider et al.
only invade dead plants after harvest. 1982) and Sal~notlellanznenchen (Taylor et al. 1982). Kurup et
Group 1 fungi include Botryfis cirzeren, Sclerotinia sclero- al. (1983) isolated several actinomycetes from marijuana
tioru~n,and Alternuria alternata. In living plants, these fungi cigarettes-Tlzernzoactinotizyces candidus, Tkermoactinomyces
cause grey mould, hemp canker, and brown blight, respec- vulgaris, and Micropolyspora faetzi. These actinomycetes cause
tively. Less damaging Group 1 fungi include Cladosporiltnl allergenic reactions and "Farmer's lung" disease. Ramirez
lzerbarunz, Epicoccilm izigrltnz, Stachybotrys lobulatn, Stem- (1990) reported four policemen contracting pulmonary
pllyliunz botryos~inl,and several Fusariltnz and Mncor species. histoplasmosis (from Histoplasnza mpsltlatllnz) after pulling
Group 1 bacteria contribute to the curing of harvested up marijuana in Puerto Rico.
Callnabis. Curing represents the breakdown of chlorophyll, Group 2 storage fungi are the primary cause of storage
polyphenols, and plant starches, which eliminates the harsh, losses-Aspergilllis, Penicilliltm, Rhizoplts and Mltcor species
green, "homegrown" taste of fresh marijuana. Plant enzymes (Fig 8.2). "Storage moulds" predominate in storage because
may also be involved; one possible enzyme may be o-dipheol they have evolved as such. They thrive under low oxygen
oxidase (=polypheno1oxidase), the enzyme involved in the levels, limited moisture, and intense competition for
curing of tea leaves. substrate. For instance, Grewal (1989) found that dried
Group 1 insects, such as aphids and spider mites, rarely Cannabis leaves suppressed the growth of most mesophilic
damage marijuana in storage. Like many other fastidious fungi, except the storage fungi Aspergillils and Peizicilliltnz.
organisms (e.g., viruses, nematodes, phytoplasmas, etc.), Dahiya & Jain (1977)tested the effects of cannabinoids on 18
these insects die with their host at harvest time. Exceptions species of fungi; THC and CBD inhibited the growth of all
include Grapiiolit~7deli~iear~a(hemp borers) and Helicoiierpa but two organisms, the storage fungi Aspergilllts niger and
niivigera (budworms). Pelzicilliurn clzrysogen~~m.Indeed, some storage moulds can
Group 2 insects were studied by Smith & Olson (1982). metabolize THC (Robertson et al. 1975, Binder 1976).
172 Hemp Diseases and Pests

Aspergillus rot fruits and vegetables, poison livestock that eat infected
The genus Aspergilltls includes over 500 species. These grain, and cause opportunistic infections in people. On the
ubiquitous fungi grow on anything from rocket fuel to as- positive side, Penicillitlm species are cultivated in huge vats
tronauts. The genus is millions of years old; while Homo sa- to provide us with antibiotics and vitamins. And Penin'llitrm
piens may come and go, Aspergilltls will remain. These meek roqtleforti puts the "blue" in its namesake cheese.
moulds, like their sister genus Penicillium, are found Kagen et al. (1983) and Kurup et al. (1983) isolated
worldwide. Aspergilltls species grow better in warmer Penicillitlm from marijuana cigarettes. Babu et al. (1977)
climates, Penicillitlm in cooler climates. identified Penicillitrm chrysogentlm attacking marijuana. P.
Westendorp (1854a) first described an Aspergilltls chrysogentlfn grows abundantly in nature. It provided
species mouldering Cannabis;he called it Aspergilltls conoidezrs. . Alexander Fleming with penicillin. Bush Doctor (1993a)
Aspergilltls niger has been isolated from mouldy marijuana isolated Penicillitlm italictlm from marijuana that was stored
(Kagen 1983) and mouldy hemp (Vakhrusheva 1979). with an orange peel at 3°C. Adding peels to marijuana
Margolis & Clorfene (1975) and DuToit (1980) described a imparts a "pleasant bouquet" (Frank & Rosenthal1978). In
mould that increased marijuana potency. Bush Doctor (1993a) this case, the peel imparted a nidus of infection. P. italicum,
found A. niger moulding marijuana and determined the the "blue citrus mould," is notorious for its ability to spread
mould did not increase potency. The mould does, however, by contact (i.e., "one bad apple spoils the whole bunch").
cause human disease. Schwartz (1987) scraped A. niger from
the sinuses of a marijuana smoker suffering severe headaches. Mucor
Kagen (1983) isolated Aspergilltls ftlmigattls from About 50 Mtlcor species are recognized worldwide. These
mouldy marijuana. A. filmigatus-moulded marijuana has fungi resemble yellowish-grey cotton candy. Mtlcor species
caused bronchopulmonary aspergillosis (Llamas et al. 1978). grow on almost any organic matter. They specialize in
Chusid et al. (1975) reported A.fi~migatuscausing near-fatal dung-Mtlcor spores survive passage through animal guts,
pneurnonitis in a 17-year-old.They noted the patient buried his which gives them first access to animal dung. A very
marijuana in the earth for "aging." No doubt the patient hoped existential life cycle.
for a potency-enhancing mould but A.firmigahls found him instead. Five Mtlcor species have been described on Cannabis.
Lastly, Kagen (1983) isolated Aspergilltls flavus from Mtlcor hiemalis, a common soil fungus, was originally
mouldy marijuana. A . flavtls produces nasty metabolites discovered on hemp (Wehmer 1903).It has been isolated from
called aflatoxins. Aflatoxins are acutely poisonous, with an leaves (Saccardo 1904, Oudemans 1920),stems (Behrens 1902,
LD5, of 0.25-0.55 mg kg-1 (nearly 2000 times more toxic than Wehmer 1903)and hemp fibres (Oudemans 1920).M. hiernalis
parathion, the most toxic insecticide). Aflatoxins are also prefers cooler climates. The Japanese use M . hiemalis for
carcinogenic (Moss 1996).Llewellyn & O'Rear (1977) found cleaning crud from pearls. The fungus metabolizes soyabean
aflatoxins in marijuana contaminated by A. flavtrs and curd into products like tempeh. M . hiemalis regrettably
Aspergilltls parasiticus. bioconcentrates (and cannot metabolize) paraquat from
tainted substrates (Domsch et al. 1980).
Penicillium The "Mucor species" recovered by Kagen et al. (1983)
There are half as many Penicillitlm species as Aspergilltls and Kurup et al. (1983) may have been M . hiemalis. Other
species in the world, but they are twice as difficult to tell possibilities include Mtlcor mucedo, a hemp retter cited by
apart. As previously mentioned, Penicillitlm species grow Lentz (1977),or three species that Gzebenyuk (1984)collected
worldwide but prefer cooler climates. Refrigerator conditions from Soviet hemp stems: Mtlcor corticola, Mtlcor genevensis,
may actually encourage Penicillitlm infestation. These fungi and Mtlcor pltrmbens.

Figure 8.3: A young Aspergillus conidiophore with many conidia (SEM x800, McPartland).
 Chapter 8: Post-harvest Problems 173

Rkizopus ficient leaves also release exudates. Expect more storage

This genus contains approximately 30 species. These mould problems in poorly-grown plants.
fungi live in soil, on ripe foodstuffs and, occasionally, on The secret to stopping bacteria and moulds is nloistiire
people (especially diabetics). Rlzizopus sporangiophores arise control. Flue-cured (or oven-dried) marijuana suffers less con-
in clusters that grow from rootlike rhizoids. These clusters tamination than air dried or sweat-cured crops. Sweat-cured
are connected by stolons, like strawberry plants. The pres- Cannabis (common in Colombia) maintains a "tradition" of
ence of rhizoids and stolons differentiates Rhizopi~sspecies Aspergillus contamination (Bush Doctor 1993a).Hashish must
from Mllcor species. be pressed carefully, with no air trapped inside. Poorly hand-
Gzebenyuk (1984)isolated Rhizopus stolonifer from hemp rubbed charas nearly always contains air pockets with
stems. This species is an important hemp-retting organism moisture, resulting in-fungal-and bacterial spoilage (Clarke
(Behrens 1902, 1903). Bush Doctor (1993a) infected damp 1998).
marijuana with a colony of R . stolonifer found growing on Oven drying inevitably leads to a harsh-smoking prod-
stale bread. The fungus is found worldwide. uct. So most people air-dry marijuana. Drying rooms should
be cool and dry,-preferably in llrlinterrilpted darkness (most
SIGNS & SYMPTOMS OF CONTAMINATION storage fungi require UV light to sporulate). Electrical fans
Marijuana consumers identify microbiological contami- improve air circulation. Drying marijuana in a 10% C0,-sup-
nants using several crude but seemingly effective screening plemented atmosphere will reduce mould and insects
techniques (Bush Doctor 1993~1,McPartland 1994, Conrad (Wilson & Chalutz 1991).
1997).The techniques mimic those used by tobacco growers Living Cannabis plants are about 80% water. According
(Lucas 1975). to Bush Doctor (1993a), perfectly dried marijuana contains
Visual screeiz: A golden colour means the herb was cured about 10-12% water or moisture content (MC). Material
In sunlight; green or purple material was cured in darkness below 10% becomes too brittle and disintegrates. Fungi
(Conrad 1997). Contaminated marijuana often darkens in cannot grow below 15% MC. Unfortunately, many growers
colour and becomes crumbly. Anaerobic bacteria turn mari- prefer to market their crop above 15%. Catmnbis, like corn
juana into brown slime. Fungi appear in mouldy material as flakes, is sold by weight, not volume. Tobacco farmers also
tufts and strands of fungal hyphae. Hyphae look white to allow their products to gain weight by resorbing moisture
light grey when marijuana is stored in complete darkness. before sale. They term this risky business "coming into order"
But if exposed to light, the tufts turn into velvet clumps of (Lucas 1975).Freezer storage will not protect material above
coloured spores. A slight tap sends these spores into billow- 15O/0 MC. Placing lemon peels in stored marijuana is
ing clouds. Clumps of Rhizopus and M i ~ c o rspores generally discouraged; the peels may raise the MC above 15% and
appear grey-black, Penicilli~~ni spores (conidia) are light blue- inoculate the marijuana with pathogens. Properly dried
green, and Aspergillirs conidia are dark green-black. marijuana also discourages insects and mites.
Improperly prepared hashish may also mould (Clarke
1998). Grey veins of mould become visible when hashish is BIOLOGICAL CONTROL
broken. Cl~erniak(1979) illustrated a piece of mouldy None of the organisms listed below has been tested on
hashish-see figure 8.17 in his book. harvested marijuana. They have great potential as
Fluorescence screen: Plant material freshly contami- biocontrols, b u t may taste bad o r pose a threat to
nated by aflatoxin will fluoresce a bright greenish-gold hue immunocompromised individuals.
under ultraviolet light (Bush Doctor 1993a). Wood's lamps
("black lights" emitting a spectrum of 365 nm) made Piclzia (Candida)guilliermondii (?=Debaryon~yceshansenii)
specifically for aflatoxin detection are sold in agriculture BIOLOGY: An epiphytic yeast that colonizes numerous
catalogues. plants and controls postharvest rot in fruit caused by Botrytis
Olfactory G teinperature screen: Rotting marijuana pro- cinerea, Alternaria alternata, Penicillii~nzdigitatum, Penicillinin
duces a spectrum of odours, from stale to musty to mouldy. italicllnz, and Rhizopl~sstolonifer.
P.italicl~inproduces a lavender or lilac odour. Lucas (1975) APPLICATION: Supplied as a powder or aqueous suspen-
reported A. f l n o l ~ s as smelling particularly offensive. sion. Plant material is sprayed or dipped in a yeast
Anaerobic bacteria stink like carrion. Marijuana undergoing suspension, then dried. Adding CaC12 (2% w / v ) to the
rapid decay may feel warm to touch. (At this stage the solution improves control (Wilson & Ehalutz 1991).
product is ready for composting.)
Smoke screen: Infested material produces a brown, black, Candida oleophila
or sooty smoke. Healthy marijuana produces milky white BIOLOGY: An epiphytic yeast (ASPIRE@)that protects
or light blue smoke (Conrad 1997). apples and citrus fruits from postharvest decay caused by
Screen screen: To expose insects, shake samples in a No. Botrytis and Penicillluli~species.
10 steel sieve. Not all bugs found in marijuana cause storage APPLICATION: Supplied as a wettable powder. Plant
damage. Some are simply "innocent bystanders" caught material is sprayed or dipped in a yeast suspension, then dried.
during harvesting. Crosby e f al. (1986) found fig-pollinating
iLJn5ps in confiscated marijuana. These rare insects helped Pseudomonas syringae
police determine the marijuana grew in Thailand or south- BIOLOGY: A bacterium (Bio-Save 11s) that protects ap-
ern Burma. Living insects in marijuana are more suspicious. ples and citrus fruits from postharvest decay caused by Bot-
Spreading samples on a sheet of white paper often reveals ~ y t i scinerea, Milcor species, and Penicilliunz species. It is a
live insects as they crawl away. A hand lens is often helpful nonpathogenic variety of a common plant bacterium. Strains
for the identification of small insects. ESC 10 and ESC 11 protect against postharvest decay; strain
742RS has a different purpose--protecting orchards from ice-
CULTURAL & MECHANICAL CONTROL crystallizing Pseudon~onasspecies.
Avoid damaging plants before they completely dry. APPLICATION: Supplied as spores in pellets or a
Wounded tissues release exudates upon which fungi estab- wettable powder. Plant material is sprayed or dipped in a
lish a foothold. Lucas (1975) said diseased and nutrient-de- suspension, then dried.
171 Hemp Diseases and Pests

CHEMICAL CONTROL marijuana cigarettes. People using medical marijuana should

Dipping Peilicilliuin-infested marijuana in a baking soda take extra precautions. McPartland (1984)suggested patients
solution will inhibit the acid-loving fungi, but the marijuana switch to synthetic THC. But Doblin & Kleiman (1991) re-
must be rapidly re-dried. Synthetic pesticides shol~ldnever be ported dissatisfaction with synthetic THC among patients
spri~!{e~i or1 rnauijunna. Spector (1985) reported youngsters and their physicians. Nearly half of all oncologists polled
spraying marijuana with formaldehyde. The treated weed by Doblin & Kleilnan recommended their patients switch to
caused anoxia and psychomotor retardation in smokers. illegal marijuana.
Fumigants (gases, not sprays or aerosols) may remain as Moody et 01. (1982) tested water pipes with Aspcrg~l-
gaseous residues in air pockets of marijuana flowers. 111s-contaminated marijuana and found only a 15';" reduc-
Cngerleider et 01. (1982) fumigated marijuana with ethylene tion in transmission of fungal spores. McPartland & Pruitt
oxide. They reported complete sterilization of fungi with no (1997)concluded that vaporizer devices are superior to wa-
loss of THC, but did not comment on gaseous residues. ter pipes for protecting smokers. Vaporizers heat marijuana
Harvested tobacco is fumigated with high pressure C02, to 180-190°C, a temperature range which vaporizes THC,
elther 4000 kPa for 30 minutes or 3000 kPa for 50 minutes. but is below the ignition point of combustible plant mate-
This "Carvex process" leaves no dangerous residues. rial. Gieringer (1996) determined that vaporizers deliver a
Ungerleider et al. also sterilized marijuana with high-dose higher cannabinoid-to-tar ratio than cigarettes or water
Cobalt 60 irradiation (15,000 to 20,000 Gray Units). This pipes. The best vaporizer delivered a 1:10ratio, whereas ciga-
iizetllod is i~otrecoirime,~dedfornovices. Seeds are killed by 20,000 rettes averaged 1:13, and water pipes averaged 1:27 (the
Gray Units, and the presence of radiolytic substances in worst pipe delivered 1:40).
irradiated marijuana has not been evaluated. Levitz & Diamond (1991)suggested baking marijuana
in home ovens at 150°C (300°F) for five minutes before
CONSUMER HAZARDS smoking. This temperature kills Aspergillus spores without
Immunosuppressed individuals and asthmatics should vaporizing THC. Unfortunately, this temperature is not suf-
ne\,er be exposed to moulds, especially Aspergillus (Figs 8.2 ficient to degrade Aspergilllls antigens, so sensitized patients
& 8.3).Kurup et nl. (1983) showed that spores of Aspergillils may still develop asthma (McPartland 1994).
and Mllcor species survive in smoke drawn from pyrolysed
 "Since insects for tlze m o s t part lack ct~iznilzgou intelligence, insect traps are often st~rprisinglysililple. "

-C. L. Metcalf

Chapter 9: Cultural & Mechanical Methods

of Controlling Diseases and Pests

Cultural and mechanical methods encompass a wide variety farm (McEno 1990). Several methods for sterilizing
of techniques that are used by both organic farmers and con- large quantities of nutrient solutions are reviewed by
\rentlonaifarmers. Cultural and mechanical methods can be Jarvis (1992). Precipitation of calcium is a problem
applied to any aspect of the crop damage trlangle described common to all methods. Jarvis also reviewed ultra-
In Chapter 1 Ciilt~~rnl methods are usually. preventatwe, and
-
violet irradiation, which is feasible for small
consist of ordinary farm practices that encourage healthy operations but it precipitates iron. Ultrafiltrat~onot
plant growth. These methods alter the landscape and make nutrient solutions through pores of about 10 pm at
it less favourable for pest reproduction and survival, such 300 kPa eliminates fungi but not viruses. Ozone
as tilling the soil, crop rotation, regulating moisture levels, bubbled through nutrient solution at a concentrat~on
and balanced use of fertilizers. Mrcizunicnl methods can be of 1.5 mg 1-1 for 30 minutes kills pathogens but may
preventative (e.g., steam sterilization of glasshouse soil) or be phytotoxic to Canrzabis. Sodium hypochlorite at
curati~re(e.g., vacuuming bugs off leaves or pruning away concentrations of 1-5 mg 1-1 kills many pathogens but
fungus-infected branches). Mechanical methods can be as not all. Sterilization of nutrient solutions with chlorine
simple as handpicking pests from plants or as complicated gas kills pathogens but causes phytotoxicity ~f concen-
as electronic insect traps. trations of free chlorine exceed 10 mg 1-1 (Jamis1992).
Some of the most useful cultural and mechanical
neth hods are l~stedbelow. Many of these methods are ancient. Table 9.1: Thermal inactivation of selected pathogens and
Many were abandoned with the advent of chemical pests. (from Jarvis 1992).
pesticides, but are enjoying a resurgence in popularity. David
West points out that some of these methods pertain to
ag~ot~oriiiccrops (e.g.,fibre and oil seed crops), whereas other
methods are better suited for horticult~irnlsituations (e.g.,
Botrytis cinerea 55 15
pl~arrnaceuticalCannnbls).
Sclerotinia sclerotiorum 50 5
For sources of sticky materials, traps, electric fencing,
Pythium ultimum 46 20-40
etc., obtain the annual Directory published by BlRC (Bio-
Fusarium oxysporum 60 30
Integral Resource Centre), P.O. Box 7414, Berkeley, California
Rhizoctonia solani 53 30
91707, telephone: (510) 523-2567.
most other fungi 60 30
most bacteria
1. Sanitation:
most viruses
a. Destroy crop residues after harvest. Shredding of
stalks and roots exposes overwintering pests and most nematodes 48-56 10-15
most insects and mites 60-70 30
pathogens to their natural enemies a n d the
most weed seeds 70-80 15
environment. Shredding hemp stalks requires special -

tillage equipment, however, because standard

2. Work the soil after harvest:
shredders cannot adequately cope with the fibre
a. Cultivation with tillage equipment exposes soil pests
(Moes, pers. commun. 1999).
and pathogens to their enemies and to the weather.
Covering crop residues with soil, using a
Properly done, tillage also reduces soil compaction,
n~ouldboardplough, will kill many pests and patho-
which reduces root rot. Improperly done, tillage leads
gens (see comments under 2a, below). Cart residues
to a loss of soil structure, organic matter, moisture,
off site for burial, burning, or composting.
and beneficial microorganisms. Repeated tilling may
Compostlng must be done properly-compost piles
accelerate soil erosion from wind and water (Cook et
may become trash heaps and turn into pest nurseries
al. 1996).
(Jarvis 1992). Remove anything that may shelter
Rotary tillers fluff surface soil nicely, but they
o\,ermrinteringpests, such as weeds and surface trash
are slow and not very energy efficient. Chisel ploughs
(dead leaves, brush heaps, boxes, equipment, etc.).
with 10.cm twisted chisels work the same depth as
b. Wash residues from glasshouse walls, cultivation
rotary tillers, but tend to get plugged with crop
equipment, and clothing (especially boots). Heat-
residues. Disk ploughs with standard-sized disks
sterilize hand tools, previously-used rockwool, and
penetrate soil 15 cm or more; unfortunately, disks may
polystyrene trays. Pests and pathogens are killed at
compact soil worse than mouldboard ploughs.
different temperatures, see Table 9.1. Rockwool
Mouldboard ploughs, considered oiitri by manv
should be as dry as possible before it is steam-steri-
people, suck up and turn over the top 20 cm of soil,
lized, since high temperatures are attained more
but mouldboards capable of reaching a metre down
quickly; slabs stacked 1.5 m high on pallets (with
have been used against deep-living soil pathogens
plastic covers removed) take two hours to reach
(Wolf 1999).Soil damage caused by annual ploughing
100°C in an autoclave (Runia 1986).
has prompted the use of "no-till" techniques. No-till
c. Hydroponic cultivators should disinfect recirculating
has soil benefits (especially on land with little organic
nutrient solutions. Plant diseases caused by water
material), but pests, pathogens, and weeds also
moulds can quickly shut down an entire hydroponic
benefit. 175
176 Hemp Dis:eases and Pests

b. Heat-sterilize the soil. Heating soil to 80°C for 30 laying plastic provides a combination of mechanical
minutes kills most soil organisms (Table 9.1). Steam and chemical control. Winter rape (Bmssicn iznp~ls)
heat is economical. Batches of soils, compost, and rock and certain broccoli cultivars ('Brigadier') produce
wool can be steamed in autoclaves. ~ u h c l a v e come
s high levels of glucosinolates. In the soil,
in various sizes, including walk-in models. For small glucosinolates decompose into isothiocyanates and
fields, steam can be injected into soil through hollow- nitriles, which kill fungal spores, weed seeds, and
tined soil injection devices, or through perforated some insects.
pipes buried permanently underground. d. Flooding the soil. This technique is like
Small batches of soils can be heat-sterilized in pasteurization-it kills plant-pathogenic fungi,
conventional ovens: Place moist soil in a shallow pan, nematodes and soil insects while maintaining a
cosrer the pan with aluminium foil, and insert a meat population of nonpathogenic microorganisms. Arti-
thermometre into the soil. Ellis & Bradley (1992) ficial inundation can be costly for large fields. But
suggested preheating the oven to 93°C (200°F), and some land, such as fields alongside rivers and lakes,
heating soil to a temperature between 60-82°C (140- may be naturally flooded or neighbouring waters can
180°F) for 30 minutes. If soil temperature exceeds be diverted easily (Bridge 1996).
82"C, remol-e the pan from the oven and let it cool
below 82'C before returning it to the oven. To use a
microwave oven, heat 1 kg of damp soil for 150
seconds, using a 625 watt microwave oven at full
power. Rockwool slabs can also be microwaved, but
microwaves may be poorly distributed in slabs, pre-
venting effective sterilization (Runia 1986).
Soil sterilization has drawbacks-it eliminates ben-
eficial organisms and creates a biological vacuum. An
accidental reintroduction of pathogens will cause
epidemics in sterilized soil. Avoid reintroduction by
washing all equipment touching soil-including
shovels a n d (s)hoes-in a solution of sodium
hypochlorite (bleach). But even with the best
precautions, pathogen reintroduction may occur via
seedborne infection.
Another drawback to sterilization is nutrient
toxicity. Manganese in soil heated above 80°C
becomes available to plants in toxic amounts,
particularly in acidic, clay soils (Jarvis 1992). Unless
this free manganese is leached from the soil, it may Figure 9.1: Row cultivation of hemp (from Lesik 1958)
remain toxic for 60 days or more and contribute to
iron deficiency. Nitrogen in steamed soil also
undergoes changes. Nitrifying bacteria are killed, so 3. Work the soil during the early growing season, between
ammonia may build up to phytotoxic levels (Jarvis seedling emergence and canopy formation. Soil tillage
1992). exposes soil pests and eliminates weed seedlings. Many pests
c. Pasteurization may be better than sterilization-it and pathogens are attracted to weeds first, then move to
kills pathogenic fungi while maintaining a rich Cannabis. Weeds also hold humidity which favours mould
inicroflora of beneficial organisms. Pasteurization is development. Keeping the soil clean eliminates insects that
accomplished by mixing steam with air at 60°C. cannot lay eggs on bare ground. Of course, bare ground may
Pasteurization is performed in the field by soil also accelerate soil erosion from wind and water (Cook et 01.
solnrlzntiolz, where the soil is covered by sheets of 1996). Rarely, pests liixz~riatein weed-free fields, because
plastic and heated under the sun for four six weeks certain weed-dependent biocontrol organisms are lacking
(Elmore et a1.1997).Intense summer sunlight can heat in these situations (Altieri et nl. 1977).
the top 5 cm of soil to 55'C, and to 37°C as deep as 45 For small plots, a standard hoe works well. Cultivate large
cm. Solarization works best in sunny climates, fields with tractor-drawn tines, spike-toothed harrows, or
obviously. The field must be carefully disked and its rotary brush weeders (Fig 9.1). Alternatively, cover rows of
surface smoothed, removing any rocks or clods that soil with rolls of black woven plastic, or plastic ground cover
might raise the plastic or puncture it. Treated soil must ("plastic mulch"), and grow plants through holes punched
be kept moist so that the heat will penetrate evenly in the plastic.
(consider laying drip irrigation lines before laying
plastic). Plastic sheets should be polyethylene, 4. Escape cropping-vary the time of planting and/or har-
transparent, and at least 76 cm wide. Proper thickness vesting. Early planting permits susceptible seedlings to
ranges from 1 mil (0.001 inch or 0.0025 m m thick) to harden before pests arrive. Late planting avoids cool
2 mils, depending on the wind. Plastic sheets must temperatures that predispose seedlings to damping off and
be laid tightly against the soil; row edges are anchored root rots. Late planting also eludes the egg-laying period of
by burying the edges in a shallow trench. some pests. Early harvest avoids the endless autumn rains
Solarization controls many soilborne fungal and that predispose plants to grey mould in the Pacific Northwest
bacterial pathogens, as well as some nematodes, weed and the Netherlands. Early harvest also reduces flea beetles
seeds, a n d insects. Beneficial fungi, such as and theft.
Trichodernzn and mycorrhizae, are very heat resistant.
Earthworms survive by escaping to lower soil depths. 5. Use resistant varieties. Most hemp breeders qualify and
Carefully disking a Brassica crop into the soil before quantify their cultivars by fibre yield and THC content, with
 Chapter 9: Cultural & Mechanical Methods of Controlling Diseases and Pests 177

little regard for pest resistance (Bocsa & Karus 1997). by not overcrowding them. Humidity always
Nevertheless, a few varieties have been selected for t h e ~ r increases after canopy closure, which is when leaves
resistant traits. In most cases, the mechanism of resistance is of adjacent plants touch each other and shade the soil.
physiological and poorly understood. But sometimes it is Canopy closure is unavoidable in fibre crops, but
sim~le--forinstance, slimmer buds hold less moisture which can be avoided in seed crops and seedling beds.
protects them from grey mould. Another mechanism of Properly-spaced plants keep the canopy open,
resistance is chemical-the terpenoids, ketones, and increase air circulation, and reduce humidity and leaf
cannabinoids svnthesized in cannibis trichomes. Selecting - wetness. Plant rows in the direction of prevailing
winds, or in an east-west or northeast-southwest
plants for trichomes poses a dilemma, however, because
these features also suppress biocontrol organisms, including orientation to promote solar drying. Air circulates
predaceous mites, ladybeetles, Chrysoperla canzea, Encarsia better on sloped hillsides or high points than on low
fo~nzosn, Geocorls pliizctlpes, and various Trichogratnnia and or flat ground. Avoid overhead irrigation during
Ornis species (Campbell & Duffey 1979). flowering.
~raditionall~, w e have manipulated the Cannabis genome Indoors, proper ventilation is critical so humidity
using sexual propagation. Sex introduces genetic variability, can escape. In glasshouses, rzarroiuing the ternperatilre
the cornerstone of breeding programmes. Breeders promote differential between night and day will lozuer the chance of
desired traits by methodicalli hand-pollinating choice dewformation. Keep glasshouses cool during the day
females with pollen from choice males. Cantlabis is wind- and heat them at night. Turn on heat (electric,
~ o l l i n a t e ds;, contamination of females with unselected propane, butane or natural gas, not kerosene or
pollen must be avoided. Lemeshev et al. (1995) recommend gasoline) before sundown. This slows the drop in
a distance of at least 2 km between fields to provide spatial temperature and prevents moisture condensation on
pollen isolation. If this is not possible, Clarke (1981)detailed plants. Heat all night and keep all vents open-
the use of pollen-collecting bags and protection of female although a "waste of heat," this is the only way to
flo~vers.Since most desirable traits (e.g., high yield) are drive moisture out of the night air. During the day
quantitative, more than two plants are needed for a breeding use air conditioning-cool air holds less water and
programme. Breeding techniques well-suited for Cannabis lowers the atmospheric humidity. Avoid dew at all
include the Ohio technique, Bredemann method, modified costs during flowering. Do not irrigate plants late in
Bredernann method (Bocsa & Karus 1997), and reciprocal the day or at night.
recurrent selection (Seven Turtles 1988). The latter strategy Humidity increases in low-ceiling structures such
maintains two select populations while crossing'them to as Dutch VenloTMglasshouses. Never allow plants to
produce hybrid offspring. fill more than 1/ 3 the volume inside a glasshouse or
Once a choice hybrid is selected, it can be preserved by growroom. Have a high roof and short plants.
asexual propagation. Clones from a single are geneti- Humidity increases in glasshouses using polythene
cally identical. Clones are propagated by cutting-rooting or blackouts (which restrict day length and induce flow-
by air-layering. Clarke (1981) and Frank (1988) described ering). Try using blackouts made of porous woven
both propagation methods in detail. Clones are maintained cloth, instead of plastic sheeting. Humidity also
under artific~allycreated long daylength. increases in glasshouses using thermal screens to
To induce mutations in a-search for useful traits, hemp reduce heat losses, a n d in glasshouses with
breeders hare blasted pollen and seeds with gamma (y) microscreens installed to exclude outdoor pests. Keep
radiation (Zottini et al. 1997). Genetic engineers have begun all screens clean to optimize airflow.
sequencing Cnniznbis DNA (Gillan et 01. 1995, Faeti et 01. 1996,
Jagadish et 01. 1996, Siniscalco Gigliano et al. 1997, Linacre & 8. Optimize soil structure and nutrition:
Thorpe 1998).In the future we will manipulate the Cannabis a. Balance nutrients carefully. Excess nitrogen
genome via recombinant DNA technology. Researchers are predisposes plants to bacteria, fungi, and leaf-eating
currently having trouble regenerating Cannabis callus tissue insects. Indiscriminate use of phosphorus to promote
into whole plants (Mandolino & Ranalli 1999). flowering is ill-advised for the same reason.
b. Improve soil structure (see Chapter 2 ) . Adding
6 . Crop rotation keeps pests moving, which is a lethal in- organic amendments to soil augments naturally
convenience. It eliminates many pests and pathogens that occurring biological control of pests and pathogens.
specifically attack Cannnbis. H e m p crops cultivated This tactic works particularly well against soilborne
continuously in monoculture suffer more pests than crops fungi and nematodes. Adding well-composted
grown in rotation (Bocsa & Karus 1997).Crop rotation is less material is safe and effective. Conversely, poorly
effective on general feeders, migratory pests, and pathogens composted materials may contain pests and
that survive for long periods in the soil. pathogens. Adding municipal sludge to soil has
increased the incidence of root-rot diseases (Windels
7. Maintain proper moisture levels that are optimal for plant 1997).Patented formulations of organic materials and
growth: minerals (e.g., S-H MixTM)have been sold for the
a. Drought kills plants outright and also predisposes suppression of soil pathogens. Some formulations
plants to diseases and pests. also contain urea and other pesticidal compounds
b. Overwatering also kills plants and predisposes them (e.g.,ClandosanTh')and are described under chemical
to diseases and pests. It is a common problem with control.
novice growers. Allow soil to dry between waterings,
and do not plant in poorly-draining soils. If poorly- 9. Remove insect pests by hand-eggs, larvae, pupae, and
draining soils are unavoidable, plant in raised rows. adults. Crush them or drop them into a bucket of soapy water.
c . Avoid excess atmospheric humidity Excess humidity (Be sure they are pests and not beneficial insects!) Chilly
is a common problem and permits many fungi and mornings are a good time to shake beetles and bugs off plants
some pests to flourish. and onto a ground cloth. Stem borers can also be removed
Outdoors, increase air circulation between plants by hand-carefully split galls lengthwise, remove borers,
178 Hemp Diseases and Pests

then bind the stem back together. Some weakly-flying in- different than living insects or standard illustrations.
sects can be removed with a hand-held vacuum cleaner. Gill & Sanderson (1998) assembled over 25
Whiteflies, for instance, will conspicuously hover in the air photographs of insects on sticky traps. You must sort
next to a plant that is shaken. Evolution did not prepare through the mix to separate pests from other insects.
whiteflies for vacuum cleaners. Suck them out of the air. Unfortunately, some beneficial insects may also be
A\-oid tearing foliage in the vacuum by covering the nozzle attracted to yellow, including Apkidoletes nphidinzyza,
lvith gauze. Vacuum cleaners are available in gas-powered Apkidius nzatricariae, Dacnusa siberica, Diglyplil~sisaea,
backpack models. Encarsiaforinosa, Eretinocerl~serernicus, and Metapkycus
helvoll~s.Cherim (1998) removed sticky traps before
10. Limbs with isolated fungal infections may be pruned releasing these biocontrols, and thereafter oily hung
from otherwise healthy plants. Many moulds establish a traps for two days per week for monitoring purposes.
toothold in senescent plant tissues, so pruning yellow leaves Pimentel et al. (1991) recommended tilting yellow
and injured branches is preventative. Coat all wounds and traps 45" off vertical to avoid trapping syrphids.
pruning cuts with a fungicidal tree sealer or biocontrol agent Alexander (1984a) warned that yellow clothing also
(e.g., Triclzoderriza harzianl~nzor Gliocladi~~in
rosel~nz).Whole attracts pests; unwitting growers carry them into
plants can be rouged from otherwise healthy fields or uninfested areas.
glassl~ouses.When rouging an infested plant, prevent the c. Attract thrips and root maggot flies with yellow
spread of airborne fungi or insects by quickly covering the sticky traps or light blue traps (reflecting a 440 nm
plant with a big plastic bag, and cinching the bag tightly wavelength). Blue traps are less attractive to benefi-
around the stalk before removing the plant. cial insects.
Recently, Cherim (1998) recommended using hot
11. Avoid seedborne infection. Do not save seeds from plants pink sticky traps to catch thrips. Prepare sticky traps
infested by viruses, bacteria, seedborne fungi, or mites. If as described above. Hang traps near tops of plants
seeds must be used, try disinfecting them with heat. Soaking to catch flying thrips. Catch flea beetles and tarnished
seeds in hot water (50°C) for 30 minutes eliminates most plant bugs with sukite sticky traps. Suspend traps near
pests and pathogens with minimal damage to seeds. But the top of the canopy to catch flea beetles; suspend
thermal damage may delay seed germination or stunt traps beneath the canopy to catch tarnished plant
seedlings. Minimize damage by plunging seeds into cold bugs. Unfortunately, some predators that eat white
water after treatment, then carefully dry them (Maude 1996). may also be attracted tb white sticky traps (e.g.,
Maude also disinfected seeds in microwave ovens (try Cryptolnenzus nzontrouzieri).
625 W for three minutes then plunge into cold water). For d. Attract nocturnal moths with white light (a mix of
chemical seed treatments, see Chapter 11. wavelengths) or ultraviolet light. If coupled with an
electric grid, a light trap can zap hundreds of moths
12. Mechanically trap or repel insects with light and colour: a night. Each female moth dies with dozens to hun-
a. Cover soil with reflective material laid between dreds of eggs. Many electrocutor traps are simply
plants to confuse flying insects and prevent them light bulbs surrounded by an electric grid. People
from depositing eggs. Aphid alatae (winged forms), find them vaguely entertaining.
female tl~rips,leafminer flies, and some leafhoppers Ligkt traps can nJfecf plant photoperiod so tkey nlust be
are particularly susceptible. Unroll aluminium foil used carefi~llyduringflowering. Different moth species
on the ground and perforate it with holes through tend to fly at different heights and speeds; they are
which seedlings can grow. Keep the foil in place by caught preferentially by different trap designs (Young
covering its edges with soil. Foil must extend 10 cm 1997). The attraction of light traps is limited to a
beyond the plant dripline. Reflective material loses radius of about 25 m; they are least effective at full
its effectiveness on taller plants. moon (Young 1997), and most effective between the
b. Some winged pests have a fatal attraction for yellow hours of 11 PM and 3 AM (Ellis & Bradley 1992).
objects, especially aphids, whiteflies, leafhoppers, e. Bait traps with synthetic insect pheromones, which
leafminer flies, and fungus gnats (Plate 6). Catch them are described in Chapter 11. Pheromones can be ap-
with "sticky traps," which are rectangular yellow plied to sticky traps or placed within rubber septums,
cards coated with mineral oil, vaseline, Tanglefoot8 nylon or wire mesh cones, wing traps, aerial water
or other sticky materials. The rectangles should be pans, or funnel-bucket traps. Funnel-bucket traps
25-50 cm wide and constructed of heavy cardboard, hold the most carcasses.
Masonite, or thin plywood. Rustoleum Yellow No.
659 reflects a wavelength of 550 nm and works the 13. Mechanical barriers protect plants from insects and other
best (Olkowski et 01. 1991).Hang rectangles vertically, pests. Yepsen aptly described mechanical barriers as "traps
with the centre of the rectangles level with the top of without the power of attraction." To stop crawling insects,
the crop. Support the squares on stakes or wire wrap stems with a sheath of aluminium foil and coat the foil
l~olders,and raise them as the crop grows. Yellow with Tanglefoot@(do not apply Tanglefoot directly on
sticky ribbons (tapes)are also commercially available. stems-the product is a mixture of castor oil, gum resins,
Sticky traps work best in areas with little wind, and wax-it stresses plants and may translocate to flowers).
otherwise they become covered by dust. Stir up Protect seedlings from flying pests and their egg-laying
winged insects by shaking plants periodically. For mothers. Outdoors, use tent netting, tightly woven cloth,
iilo~litilrii~g
purposes, distribute one trap per 200 m2 polyethylene screens (Green-Tek@),or floating row covers
crop area. For control purposes, distribute one trap (Harvest-Guard@,ReemayQ).Indoors, install micro-screen-
at least every 20 m2, to a maximum of one trap per 2 ing to seal all openings, including louvres (vents), open win-
m:, and replace the traps frequently. Monitoring for dows, and exhaust fan openings.
pests is not easy-sticky traps may reap a heap of To screen the smallest pests (thrips and mites), mesh
dead, distorted insects. Their twisted bodies look holes need to be about 192 ym in diameter; finer mesh
 Chapter 9: Cultural & Mechanical Methods of Controlling Diseases and Pests 179

impedes glasshouse ventilation (Gill & Sanderson 1998). flowers are found in four families: Umbelliferae choices in-
lmmediately repair any tears in the screening. Install tight- clude flowering fennel (Foenicl~llinzvulgare), Queen Anne's
fitting entrance doors. lace (Dal~cuscarotn), and wild parsnip (Pnstinaca sntiun).Rec-
A wire fence bars most mammals from crops. Mole traps ommended Compositae include tansy (Tanacetllm vulgare),
guard against underground marauders, whereas "bird bells" sunflower (Helialithus species), yarrow (Achilles species),
protect against avian attack. Australians have taken cosmos (Cosmos species), and coneflower (Echiriacea species).
mechanical barriers to the limit by stretching "vermin fences" Among the Leguminosae we suggest alfalfa (Medicagosntiun),
across their entire continent to control the spread of rabbits. sweet clover (Melilotl~sspecies), fava bean (Vicia fava), and
hairy vetch (Vicia species). These legumes also fix nitrogen
1 4 . Provide food, water, a n d habitat for biocontrol from the atmosphere and improve soil nutrition. Beneficial
organisms. These cultural and mechanical techniques Brassicaceae include sweet alyssum (Lob~~laria innrtitima),
enhance the effectiveness of biocontrols described in the next other alyssums (Al~riniumand Berteroa species), and mustards
chapter. Entice some predators (e.g., ladybeetles and (Brnssica species). Other flowers attractive to biocontrol
lacewings) to stay in crops and lay eggs by providing artificial organisms include buckwheat (Fagopyrum sagittatl~nz),
honeydew-mix honey and brewers yeast, apply to sticks milkweeds (Asclepias species), and cinquefoil (Potentilln
or wax paper, and place among plants. Commercial products species).
include Wheast", Formula 573, and Enviro-Feast@. Of course, these general recommendations must be
Provide vegetational diversity by intercropping or row tailored to your specific needs-some of the aforementioned
cropping other plants with Cnfirinbis.Mixed assemblages of flowers are also attractive to pests in some situations. Also,
plants disrupt pests and enhance biocontrols. Intercropping some of the flowers have cultural and environmental
with flowers provides nectar and sanctuary for biocontrols. requirements that may be contraindicated for Calznabis crops.
Biocontrols can only sup nectar from certain flowers, how- Lastly, be sure to include plants that flower early in the
ever, because their mouthparts differ from bees and butter- season, such as Alztennnria species, Setlecio species,
flies. Suitable flowers are small, relatively open (not tubular), CIirysogorli~inspecies, coltsfoot (Tt~ssilagofarfarn), and
and have relatively long blooming periods. The most suitable goldenseal (Hydrastis canadensis).
Plate 1. Leaf stippling caused by the two-spotted spider mite, Tetranychus urticae (Clarke).
Plate 2. Webbing and golf-ball-sized cluster of diapausing T: urticae mites (Clarke).
Plate 3. Different stages of the two-spotted spider mite, T: urticae (courtesy Koppert B.V.).
Plate 4. Predatory mite, Phytoseiulus persimilis, feeding on T: urticae mite (courtesy Koppert B.V.).
Plate 5. Hundreds of hemp russet mites, Aculops cannabicola, covering a leaf petiole (Hillig).
Plate 6. Winged migrants (alatae) of green peach aphid, Myzus persicae, caught on a yellow sticky trap (McPartland).
Plate 7. Wingless females (aptera) of black bean aphid, Aphis fabae, on axil of male flower (Clarke).
Plate 8. Aphid lions, larvae of Chrysoperla carnae, feeding on aphids (Clarke).
#ate 9. Eggs of green lacewinglaphid lion, Chrysoperla carnae (clarke).
Plate 10. Ladybeetle larva, Hippodamip,mnvergm, Sewding on aphids (Clarke).
M e 11. Ladybeetle adult, Wippodarnb mnlvetpens, feeding an ap'hhde (Clarkeb.
Piate 12. Slug-like larvae of Aphido.I@&waphidimyzafmding on aphids {Clarke).
M e 13. Whitefly adults ( T ~ ~ d & ~ m vaporariorumj
ih8 on underside of leaf (Clarice).
Plate 14. Encarsia ;Formosa, e pams&dd dwh-ittafl"ylaws@(Ktvppert B.V.1.
Plate 115. Predatory ladybeetle, De!pha@m pusif"EQI$,
next to pupa and llama of gm~nhsumMkefly, FiaJwmi&g
vaporariorum (Kupprt B.V.).
Plate 16. Big-eyed bug, Geocoris punctipes, feeding on larvae erf meetpotato whitefly, Bensbfa tabmi (USDA).
Plate 17. Greenhouse whiteflies, Trialeurodes vaporariorum, infested by biocontrol fungus Beauveria bassiana (USDA).
Plate 18. European corn borer (ECB), Ostrinia nubilalis (Vogl).
Plate 19. Female (L) and male (R) moths of ECB, Ostrinia nubilalis (McPartland).
Plate 20. Beneficial wasp, Trichogramma ostrinae, parasitizing ECB eggs (Chenus).
Plate 21. ECB larvae killed and discoloured by Bacillus thuringiensis (Bt)(Koppert B.V.).
Plate 22. Stem gall caused by hemp borer, Grapholita delineana (McPartland).
Plate 23. Moth of hemp borer, Grapholita delineana (McPartland).
Plate 24. Budworms in bud, young and old, Helicoverpa armigera (Clarke).
Plate 25. Adult moth of budworm, Helicoverpa armigera (McPartland).
Plate 26. Cutworm, Spodoptera litera, assuming characteristic "C" shape when a~srurbed(Clarke).
Plate 27. Thrips damage, caused by Thrips tabaci (Clarke).
Plate 28. Predatory mite, Neoseiulus (Amblyseius) cucumeris, attacking a thrips (Koppert B.V.).
Plate 29. Pirate bug, Orius insidiosus, impaling a thrips (Koppert B.V.).
Plate 30. Hemp flea beetle, Psylliodes attenuata (Clarke).
 . ,'

- .

SteEnern~maglaseri (Klein, USDA).

mat^ 32. Stem damage by grubs ~f turnbling flower ts[eetle, Mordelli~tenamicans (Clarke).
Shte 33.Southern gfwn stink bug, Nezara viridde, wi* rus$@and white wlo~wrrnarpk~ (McSartland).
Plate 34. Lygus bugs, related to tarnished plant bugs (Clarke$.
'Plate 36. teafm.iner tunneds hy Agjxomyza ~ep@ms: (McPartbnd).
F3aae 36. Pighlyphu@iiaw, a pa rasitold of I&miners (Koppert 6-V.).
Mato 37. Hungarian Ireafhopper,famjlly Cicadzefllidae!(Cl~arke).
Plate 38. Prdetory betle, Cryptobemu~rnontrouzi8ri, feeding on scale larvae (Koppert B.V.). .
Frrte 38.Termite damage in equatorial Africa (Clarke).
 .(Jal!aw aP)
zznl Aa16 e u! padolanua awoaaq saneal pue sJannoy 'h!p!wny q6!y ul ' s d o ~6 u ! ~ a ~ ojol jplnolu Aal9 ' 9 aaeld
~
'(160~)eaJaup s!Ai~ofl Aq pasnea 'plnow A a ~ 6Aq palsaju! pla!j d w a *pp ~ aaeld
- ( . ~ uaddoy)
q etuel s d ! ~ yuo
~ 6u!Aa~d'sal!ur s!dseodA~ 'aqw h o ~ e p a ~ * d~aaeld
p
,'(ayJel3) aeu!p!Jav A~!we~qns '~addoysse~6 paaej-luejs ue!~e6uny-( Aq pasnea a6euea aaeld
'(ay~el3)e a ! ~ j vu! sjsnaol Aq p a u n ~ ds d o ~
6 u ! ~ a ~ o l j aaeld
'(puelyedaw) sa!lj slaaju! ley$ sn6unj e 'ae3snur ~ J O ~ J ~ ~ O WwOoJ ~Usa~ods
j~ ayqM 40 oleH 'OP aaeld
I.
grey coiour (Vogl).
Pl~a'Bg47. Grey rncwld.af stalks. In high humidity, infested sta!lks k o m a envelop@din a grey fuzz (de Meijer).
C M e a.Hemp canker, caused by Sclerothia sc~rodorum,stalk covered with white rnydliiurn and black sclerotia
, (Sebifele).
Plate 49. Damping off of seedling (pire-emergent), note water-6oaked appmwanco of rmtlwt CMcPartIand).
-,< , @a.
Plate' hmptlng off (pcme~meqpnut)hit8 34mg5 after they bmargp M?.Ph8 mill QMcPartIand).
Plate 511.Yellow leaf spot, cauecd by .S@ptofi&n~ocannabinaon the left, and $@p&~rlawnnaM 017the right
(McPdand).
R e e 52. Yellow leaf spat (Syptoria cannabis) on fan leaves of flw&rng taps in Wepal CMcPartlad).
Plate 53. Root rot caused by a binucleate Rhizoctonia species (McPartland).
Plate 54. Yellow leaf spot (Septoria cannabis) and brown leaf spot (Ascochyta arcuata), mixed infection on the same
plant (McPartland).
Plate 55. Brown leaf spot, Phoma cannabis, infesting flowering tops, a herbarium specimen collected by Westendrop
in 1854 (McPartland).
Plate 56. Brown stem canker, Phoma exigua (McPartland).
Plate 57. Brown leaf spot, Phoma glomerata (McPartland).
Plate 58. Fusarium stem canker, with black dot-like perithecia of the sexual stage, Gibberella cyanogena (McPartland).
Plate 59. Fusarium root rot, Fusarium solani (McPartland).
Plate 60. Fusarium wilt, Fusarium oxysporum (McPartland).
Plate 61. Powdery mildew, Sphaerotheca macularis (McPartland).
 Plate 62. Olive leaf spot, Pseudocercospora cannabina, upper side of leaves (McPartland).
Plate 63. Southern blight, caused by Sclerotium rolfsii (Grassi).
Plate 64. Microscopic leaf pustules of black mildew, Schiffnerula cannabis, next t o a white leaf spot caused by
Phomopsis ganjae (McPartland).
; . Plate 65. Microscopic view of Fusicoccum marconii, a cause of twig blight (McPartland).
Plate 66. Pink rot, Trichothecium roseum (Clarke).
Plate 67. Anthracnose, Colletotrichum dematium, on stalk (Clarke).
Plate 68. White leaf spot, Phomopsis ganjae (McPartland).
Piate 69. Spores of a mycorrhizal fungus, Glomus mosseae (McPartland).
Plate 70. Root-knot nematodes, Meloidogyne incognita, embedded in roots, with egg sacs protruding (McPartland).
Plate 71. Cysts of the cyst nematode, Heterodera schachtii, assorted ages
Hate 72. Symptoms of the hemp streak virus (Clarke).
Plate 73. Striatura ulcerosa, caused by Pseudomonas syringae (Hillig).
Plate 74. Dodder, Cuscuta europea, a parasitic plant (Berenji).
Plate 75. Broomrape, Orobanche ramosa, a parasitic plant (Berenji).
 Plap 76. Slug, Limax mudmus Nql).
Plate 77, Stalk damage caused by rabbits, Sylviiagus species (Clarke).
+L PlaW 78. Nitrogen dleficiencyI yell@wilng
.of leaves (Clarke).
Plate 79. Nitrogen excess, causing stalk beakage (Schlelife).
-
Pie* %DaPhasphorus deRcliency small, dark Ilmndw ~ n petioles
d -
c m d by lo# ~ f f l ~ a(Clarke).
t u ~
. PI* 81. Potasojum defici-wyI b c w n edges and tips k ddw 1 - w IMea@bndj.
Plate 82. Frost damage of mature flowering tops (Clarke).
Plate 83. Hail damage (Vogl).
Plate 84. Spittle mass of the spittlebug, Philaenus spumarius (Clarke).
Plate 85. Koppert B.V. biocontrols for thrips and spider mites (Clarke).
Plate 86. Novartis Encarsia formosa biocontrol for whitefly (Clarke).
 "Bioco~ztsoluses living organisms to kill pests-turn yozlr garden into a bug-eat-btig wosld."
-Bush Doctor

Chapter 10: Biological Control

Ma12q c o n z p a ~ ~ i sell
e s bioco~ztsolosga~zisms.Unforttinately, entomologists have selected biocontrols with resistance to
nlany companies quickly go o u t of business. For a n up-to- pesticides, so the two controls can be combined (see Table 10.1).
date list ofconzpanies, please obtain tlze Annual Directory Several categories of biocontrols are useful-predators,
priblislzed b y B l R C (Bio-l~ztegrnlResot~sceCentre), P.O. Box parasitoids, microbial pesticides, companion plants, trap
crops, and autocidal controls. Some categories can be
7414, Berkeley, California 94707, telephone: (510) 524-2567. combined. Heinz & Nelson (1996) achieved much better
control of whiteflies by combining a parasitoid wasp (Encarsin
Biocontrol organisms must be reintroduced into our
foun~osa)with a predatory beetle (Delphastl~sptlsillus) than by
unnatural, annual monocropping systems. We employ two releasing either biocontrol alone. E. fornzosa does not
release strategies-inoculative release and inundative
parasitize D. pusill~is,and D. pusilllis does not prey on E.
release. See "Application rates for biocontrol" later in this
fornlosa adults. D, pz~sillt~s does eat some young E,forlnosa
chapter.
larvae within whiteflies, but as the parasitoids mature within
The best example of inoculative release happened over
their hosts, D. p~isillllsavoids them.
100 years ago. Albert Koebele released a handful of Vedalias
Knipling (1992)stated that biocontrol works as a ratio of
(Australian lady beetles) in a Californian orange grove, to
biocontrol organisms to pest organisms. The density of pests
control the marauding pest lcerya purckasi. The Vedalias
is of little consequence. This is a radical departure from tra-
thrived in Orange County and, in a few years, multiplied
ditionalists who claim a pest's density on plants determines
and nearly eradicated Iceuya ptirclzasi. This strategy is also
the success of biocontrol. For inundative biocontrol to be
called permanent introduction or classical biological
effective, Knipling estimated that the ratio of adult parasitoids
control-a long-term solution to pests. This strategy works
to adult pests must be at least 2:l. For predators, the optimal
best on pests with steady-state populations, not those with
predator-to-pest ratio is probably reversed, 1:2 or greater.
boom-and-bust cycles (17-year cicadas being an extreme
Some biocontrols arise naturally, others are purchased,
example). Permanent introduction can be applied to glass-
and some can be reared by growers (see Scopes & Pickford
houses if proper environmental conditions are careful main-
1985). The USDA has recently encouraged the use of
tained. But many biocontrol organisms cannot overwinter.
biocontrols over chemicals. The current approval time for
Vedalias cannot overwinter in northern California, so the
new biocontrols is two to ten months, while chemical
beetles must be reintroduced every spring. This is called
pesticides take 25-30 months for approval (Reuveni 1995).
seasonal inoculation, comparable to an annual "booster shot."
To achieve rapid pest control, a different tactic is used-
PREDATORS
inundative release. This tactic involves the repeated release
Pitting insect against insect is a most satisfying form of
of many, many biocontrol organisms. Inundative biocontrol
biocontrol. By definition, a predator must consume more than
uses beneficial organisms as "living pesticides." The strategy
one pest before r e a c l n g its adult stage. Predators have ei-
is relatilrely expensive, but appropriate for high-income
ther chewing mouthparts (e.g., lady beetles) or piercing-suck-
crops, such as medical marijuana. Inundative release works
ing mouthparts (lacewing larvae, assassin bugs). Piercing-
best for controlling localized areas of heavy pest infestation,
sucking predators suck the fluids out of pests rather than eat
especially in enclosed spaces.
them whole, so dead pests remain attached to plants, which
lnundative and inoculative biocontrol strategies benefit
may detract from the aesthetic value of ornamental plants.
from "habitat managementu-uniform temperatures and
Many predators arise naturally in outdoor crops: centi-
year-round plant cultivation-which makes glasshouses
pedes, spiders, predatory mites, lady beetles, ground beetles,
ideal for biocontrol. Biocontrol under glass began in 1926
rove beetles, tiger beetles, lacewings, bees, and wasps. Some
with the discovery of Encausia founiosa, the whitefly parasitoid
predators are general consumers. Praying mantids (Mantis
(Plate 86). But growers abandoned E.formosa for cheap DDT
religiosa) are well-known generalists (Fig 10.1).Generalists,
in the 1940s. The appearance of DDT-resistant spider mites
unfortunately, may eat other biocontrols and beneficial
around 1968 revived interest in biocontrol, using predatory
honeybees. Lacewings (e.g.,Cizrysoperla carnm) and ladybugs
mites (Phytoscililiis peusinlilis). Van Lenteren (1995)estimated
or ladybirds (more properly, ladybeetles) are less general and
that glasshouses covered 150,000 ha around the globe; the
more finicky. They rarely eat other beneficials. But given a
glasshouse area treated with P. persinzilis was greater than
7500 ha. E. forlnosa was reintroduced in 1970 and has been
released in over 2500 glasshouse ha. Cannabis cultivators in
the USA began using beneficial insects, such as ladybugs
and mantids, in the mid-1970s (D. Watson, pers. cornmun. 1995).
To \vork effectively, biocontrols should be introduced
before the appearance of pests, or early in a pest infestation.
Most biocontrols reproduce faster than pests, and soon
overtake and consume a pest infestation. But if pests get a
head start, crop damage occurs before the biocontrols can
catch up. Hence, preiletztative biocontrol works best. Heavy
populations of pests may need chemical controls before Figure 10.1: Praying mantid on a serrated leaf
biocontrols can work effectilrely.To meet these emergencies, (from Comstock 1904). 181
182 Hemp Diseases and Pests

choice between large and small prey, they'll eat the largest Fungal MPs are the exception. Fungi such as Verticillitlrn
ones first. A lacewing larva, even if surrounded by millions lecanii do not have to be ingested. They work on contact,
of baby aphids, would rather eat a plump caterpillar. This is infecting insects right through their skin. Unfortunately, most
one reason why selectii~epredators are the best, such as preda- fungi require high humidity, and some are infected by
tory mites and Aplzidoletes aphidinnyza. dsRNA viruses that decrease their effectiveness.
The most popular mail-order predators are mites and The use of MPs against other microorganisms is termed
ladybeetles. Popular mites include Pkytoseiulus persiniilis, antibiosis. Bacteria such as Bncill~lssubtilis, Pseudon~onasfluo-
Mesosezi~ll~s loizgipes, Neoseii~ll~s
californicz~s,and Hypoaspis rescetls, and Agrobacteriil~nradiobacter are commercially
nziles. Popular ladybeetles include Hippodainia conuergens, available. They produce antibiotics that suppress the growth
Cryptolnentus nloiztroi~zieri,Roifolia cardinalis, and Stethor~~s of other bacteria and fungi, and they ooze lytic enzymes
pic~pes.See Fig 4.4 for illustrations. which puncture the cell walls of pathogens. In the biocontrol
Some predators go dormant (diapause) when crops future we will select specific strains of these microorganisms
begin to flower in autumn. This is triggered by a short for their performance on Cannabis, so the host plant and MP
photoperiod and affected by temperature (see Fig 4.2). work as a unit against pathogens.
Scientists have overcome this trait in some species by selective MPs formulated for soil application survive best if applied
breeding. Nondiapausing breeds are becoming available. during a heavy rain (or through an irrigation system). MPs
Winged predators may ultimately fly away from crops can be applied to soil five different ways (see Table 11.7).
but, hopefully, they leave behind eggs which hatch into more MPs formulated for foliage application often require high
biocontrols. Entice them to stay and lay eggs by providing humidity for germination and survival. This is bad, because
water and artificial honeydew or nectar. high humidity encourages the growth of plant pathogens
such as the grey mould fungus (Botrytis cinerea). New
PARASITOIDS glasshouse research has shown that cycling two night's
Parasitoids, in contrast to predators, kill their prey from elevated humidity with two night's normal humidity is
within. Parasitoids only consume one individual host to reach adequate for reliable biocontrol without increasing mould
their adult stage. Adult parasitoids usually insert individual problems (van Lenteren 1995). Of course, MPs requiring
eggs into multiple hosts. The eggs hatch into larvae which high humidity work great in cloning chambers, which are
eat hosts alive, leaving vital organs for last. Parasitoid lar- always humid.
vae usually pupate in pest cadavers and emerge as adults, To spray MPs on foliage requires the use of compatible
off to lay more eggs. Examples of commercial parasitoids spray carriers and spray adjuvants. Spray carriers safe to
include Encarsin forl7zosn, Trichogral7linn wasps, and braconid most MPs include non-chlorinated water and spray oils.
flies (e.g., Cl~eloill~s
texaill~s).These wasps and flies d o not Categories of spray adjuvants include spreaders (wetting
bite, sting, or otherwise bother people or plants. Some agents), stickers (adhesives), extenders (UV protectants),
biocontrols serve as parasitoids and predators-larvae only buffers, and feeding attractants. MPs are more chemically-
consume one individual host, but moult into adults which sensitive than most pesticides, so they have their own special
feed on many pests. array of adjuvants. See the approved lists published in
Parasitoids are more efficient at finding prey than preda- Hunter-Fujita (1998),or read your MP label.
tors. Parasitoids aggressively hunt until prey are nearly eradi- Recently, MPs have been added to moth pheromone
cated. Many predators, on the other hand, prefer being sur- traps. The moths become coated with MP spores in the traps,
rounded by many prey. When the prey population abates a then are released to fly to fields where they deliver the MPs
bit, the predators migrate in search of happier hunting to larval populations (Hajek 1993).
grounds, and leave many pests behind. Parasitoids stick
around. Thus they are well suited for preventative control, EPIPHYTE ANTAGONISTS
but tend to work too slowly for large infestations. Epiphytic organisms are microscopic. They colonize
Parasitoids are more pest-specific than most predators. the surface of plants, living off cellular leakage oozing from
Some may be pest-crop-specific. As parasitoids emerge from plant epidermis. They do no harm to plants even though
pupae, they imprint the odour of a specific crop damaged they may be present in huge numbers. Wilson & Ehalutz
by a specific pest. Imprinted adults remain in that crop to (1991) estimated 1000 to 10,000,000 epiphytic bacteria live
search for prey, and will not fly off for prey in another crop. per cm2 of leaf surface.
In the future we may purchase "customized biocontrols" that Using epiphytes to suppress pathogens is a lot like
have been mass-reared from pests specifically raised on using microbial pesticides but the rnodlls operandi is different.
Cnizizabis, or raised on substrates sprayed with Cannabis Epiphytes do not infect pathogens or produce toxins, they
terpenoids. simply oiltgrosu pathogens. Many pathogens depend on
cellular leakage for spore germination. Epiphytes consume
MICROBIAL PESTICIDES the cellular leakage, leaving the pathogens starved for
If handling predators and parasitoids makes you nutrients and squeezed for space. Since epiphytes do not
squeamish, try biocoiztrol in a can. Containers of microbial produce toxins, they may be safer than microbial pesticides.
pesticides (MPs) contain millions-to-trillions of freeze-dried Some organisms act as both epiphytes and microbial pesti-
bacteria, viruses, protozoans, or nematodes. Using MPs is cides, such as Pickia gl~illiernzortdii(Wilson & Ehalutz 1991).
like using chemical control-mix with water (tlon-clilori~iated The use of epiphyte antagonists, although attractive, is
water) and spray onto foliage or pour into soil. In fact, many exacting. Epiphytes differ from plant to plant (Fokkema &
growers and governments treat microbials as pesticides Van den Heuvel1986). No one has investigated the epiphytes
rather than biocontrols. Some new MPs are genetically engi- of Cannabis. One plant's epiphyte is another plant's pathogen
neered organisms. (see the section on Phylloplane Fungi in Chapter 5). Cannabis
MPs rarely harm beneficial organisms. Nearly all MPs, may not support a rich diversity of epiphytes since it
such as Blzcillzis thl~ririgiensis(Bt, a bacterium) and NPV (a produces antifungal and antibacterial compounds, including
virus), must be ingested to kill pests. Thus, they do poorly THC and CBD (McPartland 1984).Non-glandular trichomes
against sucking insects (e.g.,aphids, whiteflies, leafhoppers). also restrict microbial growth (Fokkema &Van den Heuvel1986).
 Chapter 10: Biological Control 183

Table 10.1: Effects of some pesticides on some biocontrol organisms

PESTICIDE NAME APHIDOLETES ENCARSIA

CHRYSOPERLA NEMATODE PHYTOSE~ULUS TRICHO- VERTICILLIUM
AND CLASSIFICATION* APHlDlMYZA CARNEA FORMOSA SPECIES~ PERSlMlLlS GAMMA SPP. LECANII
LARVA / ADULT LARVA I ADULT PUPA / ADULT LARVA EGG 1 ADULT PUPA I ADULT SPORES

abamectin (I-M, B) 214 [-I 5 114 [-I 114 [3] 1 [I] 114 [2] -14 [-I
acephate (I, S) 214 [8-121 414 [6-81 414 [8-121 2 [I ] -14 [3-41 214 [24] 3
aldicarb (I-M-N, S) -14 [8-121 414 [8-121 4 [28] 414 [8-121 -
%=us thuringiensis (I, B) 111 [O] Ill [0] 111 [0] 1 [ol 111 [0] 1/1[0] 1
benomyl (F, S) 111 [0] 112 [-I 111 [0] 1 [o] 113 [2-31 4
Bordeaux mixture (F, B) 113 [-I
captan (F, S) 111 [0] 111 [0] 111 [0] '111 [0] 4
carbaryl (I, S) 314 [-I 314 [4] 314 [4] 1 [ol -14 [2] 414 [-I 4
chlorthalonil
--- (F, S) 111 [O] 111 [O] 111 [O] 1 [OI 111 [O] 4
c o p p e r compounds (F, B)- -11 [O] 212 [-I 111 [0] 1 [ol -14 [0] 4
cypermethrin + polybutene
(Thripstick + Cymbush) (I, B) - 112 [0] 113 [2-41 -
d
- z-am- e- t h r i n (I, B) 414 [8- 121 414 [8-121 414 [8-121 1 [O] 414 [8-121 414 [8-121 1
drazinon (I, S) 414 [6-81 414 [4] 214 [4-61 2 [-I 212 [ I ] 414 [2] 4
dichlotvos (I, S)-fumigant -14 [0.5] 414 [0.5] 414 [ I ] 2 [-I 114 [I] -/4[1] 1
dicofol (M, S) - 114 [-I 112 [ I ] 114 [I -21 1 [ol 314 [2] 313 [3] 1
dlenochlor (M, S) 112 [0] 414 [6-81 1 [ol 213 [2] 1
diflubenzuron (I, B) Ill [0] 413 [-I Ill [0] 1 [ol 111 [0] -11 [-I 1
fenbutatin oxide (M, S) 111 [O] 111 [0] 111 [0] 111 [0] lll[O] 1
fosetyl-aluminium (F, S) 31- [-I 1 [o] 111 [0] 4
horticultural (petrol) oil (I-M, B) 111 [O] 111 [0] 111 [0] 1 [o] -13 [0] 1
imidacloprid (I-M, B) 414 [0] 41- [4] 414 [>2] 1 [ol 114 [0] 314 [-I
iprodione (F, S) 111 [0] 111 [0] 111 [0] 1 [o] Ill [0] 111[0] 2
kinoprene (I, B) 11- [-I 211 [0.5] -11 [0]
malathGn (I, S) 312 [3-41 414 [-I 414 [8-121 1 [0] 212 [I-21 414 [8- 121 3
maneb (F, S) Ill [0] 111 [0] 111 [0] 1 [ol 111 [0] 4
metalaxyl (F, S) -11 [0] 112 [-I 1 [o] -13 [-I 1
methomy1 (1, S) 414 [8-121 414 [a-121 414 [6-lo] 4 [-I 414 [4] 414 [8-121 I
methoprene (I, B) 11- [0] 112 [0] 1 [ol 111 [0]
- neem
--- (azadirachtin I, B) 212 [O] 11- [<3] 113 [-I 1 [ol 111 [0]
n~cotinesulphate (I, B) 414 [4] 213 [-I 113 [0.5] -14 [ I ]
nlcotine (I, B)-fumigant -14 [O] 113 [0.5] 111 [0]
y l S)
p a r a t h i ~ n ~ e t h(I, 414 [8-121 414 [-I 414 [8-121 3 [-I 212 [0.5]
permethrin (I-N, B) 414 [6-81 414 [6-81 414 [8-121 2 [0] 414 [8-121 414 [8-121 1
pKzcarb - (I, S) - -114 [ I ] 212 [-I 113 [0.5] 212 [0.5] 114 [ I ] 1
propargite (M, S) 21- [0] 111 [0] 313 [I] 3 [-I 413 [0] 111[0] 4
pyrethrum + PBO (I, B) -14 [6-81 212 [I] 214 [I] 1 [ol 114 [I] -14 [-I -
pyridaben(1-M, S) 111 [0] 414 [-] -13 [-I
Soap (potassium salts
of fatty acids) (I, B) -14 [0] 414 [0] 214 [0] 4 [-I 214 [0]
rotenone (I, B) 214 [-I 414 [2] 314 [-I
sulphur (F, B)-spray 212 [-I 111 [0] 114 [>4] 3 [ol 111 [0] 114 [-I 4
sulphur (F, B)-dust 113 [3-41 3 [-I 112 [ I ]
sulphur (F, B)-fumigant -11 [-I -11 [0] -13 [0.5] 112 [ I ]
thiram (F, S) 211 [-I 211 [-I 113 [0.5] 1 [OI 211 [O] 213 [-I 4
triforine (F. S) 112 [-I 111 [0] Ill [0] 1 [OI 112 [O] -11 [-I 3
Jinciozo~in (F, S) -11 [OI 111 [OI 111 [OI -11 [OI -11 [o] I
zlneb (F. S) -11 [-] 111 [0] 111 [0] -11 [-I -11 [-I 4

*Classification: I=insecticide, M=miticide, N=nematocide, F=fungicide, B=biorational, S=synthetic.

tNematode species include Steinernema spp. and Heterorhabditis spp.
§Toxicity of pesticide to immature 1 mature biocontrol organisms. Numerical ranking: 1=harmless (<25% biocontrol organisms affected),
2=slightly harmful (25-50°h affected), 3=moderately harmful (51-75%affected), 4=very harmful (>75%affected). Bracketed [numerals]
estimate the number of weeks a pesticide remains harmful after application. A dash (-) indicates unknown data. Pesticide toxicity includes
mortality and decreased fertility, when the pesticide is applied at its proper recommended rate. All applications are sprays unless
otherwise noted. Compiled from information by Koppert (1998, 1999). Data presented here are only estimations and do not guarantee
safety if followed.
184 Hemp Diseases and Pests

COMPANION PLANTS & TRAP CROPS Many people reject this "simultaneous introduction" or
Plants cannot run away from their enemies. Forced to "pest-in-first" concept, but it works. Most glasshouse grow-
stand their ground and fight, plants have evolved incred- ers already have sufficient pest populations to maintain
ible defence mechanisms, including many repellent chemi- biocontrols. But for growers who don't, Koppert sells 500 ml
cals. The chemicals produced by companion plants are pow- bottles containing 40,000 spider mites (Spidex-CPRB), "to
erful enough to repel pests from the entire neighbourhood. provide a controlled infestation of the crop."
Two popular companion plants are marigolds (Tagetes spe- Using "banker plants" is another inoculative strategy
cies) and tansy (Tatiacet~iinaulgare). Many people intercrop that prevents die out (Van Lenteren 1995). Banker plants are
companion plants w i t h crop plants to repel pests. noncrop plants infested with noncrop pests, which serve as
Counterintuitively, Bush Doctor (pers. commun. 1984) sus- alternative food sources for biocontrols. For instance, if you
pected pest-repellent chemicals may detrimentally affect anticipate Myzus persicae in your Cannabis, release Apliidoletes
Cannabis. He thought the allelopathic effects of Tagetes on ap1zidinzyza. To prevent die-out, Koppert sells boxes of wheat
Cannabis may outweighed the beneficial effects of repelling plants infested with grain aphids (AphiBanks). The bank of
pests-a potential Master's Thesis research project. barley and grain aphids will sustain A. aphidlnzyza until M.
Others plants work by their attrnctiz1eness to pests, such persicae appears.
as the Iapanese beetle's affinity for Zinnia elegaiis. These trap The inundation approach releases biocontrols after pests
plants draw pests from neighbouring crops. Trap plants can have appeared. "Hot spots" of pests become release sites for
then be sprayed with pesticides or removed, taking the pests biocontrols. The biocontrols inundate the hot spots, then
with them. Many trap plants are only attractive to pests diffuse into the surrounding crop. Inundation provides an
during a specificplant growth stage. Planting new trap crops immediate but nonsustainable reduction in the pest
In "waves" every two weeks presents pests with a variety of population. Because most biocontrols only parasitize pests
plant stages to colonize. An effective trap crop will attract during part of their lifecycle, repeated inundations are
70-8530 of a pest population while only covering 1-1O0/0 of necessary.
the total crop area (Hokkanen 1991). Please note trap crops Application rates for inundative biocontrol depend on
require careful monitoring, otherwise they turn into pest many factors. First in importance is the number o f pests,
ilurserzes. Spray or remove trap crops before a new generation measured as the Infestation Severity Index (ISI) for each
of pests hatch out. pest. Other factors include crop biomass, crop location (field,
Decoy crops are nonhost crops planted to control nema- glasshouse, or under artificial lights), local environmental
todes. Decoy crops cause nematode eggs to germinate, but conditions (do temperature and humidity favour the
the nematodes cannot complete their life cycle on the decoy biocontrol or the pest?), and the longevity -of biocontrol
plants, so they die out. Palti (1981) killed Meloidogyrie organisms versus the longevity of pests.
iizcogizita and Meloidogyize javanica with marigold (Tagetes If biocontrol organisms are released infield crops, much
pntiila, Tagetes nziruita), sesame (Sesnlnlinz orientale), castor care must be taken in providing an immediate food source
bean (Riciiz~~s conzin~~iiis),
and Clzrysaizthen~~inz
species. for them; otherwise the biocontrols will exit crops in search
of shelter and food. This is especially true with winged
AUTOCIDAL CONTROL biocontrols. Dispersion is less of a problem in glasshouses
This technique is heavy-handed but effective. Male in- with screened vents and limited exits.
sects raised in captivity are sterilized with gamma radia- Many nonwinged biocontrols such as predatory mites
tion, then released to mate with normal females, producing have the opposite problem. They cannot disperse enoiigh. If
infertile eggs. If the ratio of sterile males to normal males is plants are touching it is much easier for biocontrols to walk
2:1, then 67% of females will mate with sterile insects, as- in search of prey. If plants are not touching it may be
suming sterile males are fully competitive with native males necessary to hand-disperse biocontrols onto each plant. Small
(Knipling 1992). cups or sacks attached to each plant can hold small doses of
The greatest disadvantage of autocidal control over biocontrols (assuming thefare not cannibalistic, like
other forms of biocontrol is the lack of progeny; sterile insects lacewing larvae), which crawl out onto each individual plant
must be recreated each pest generation. Another autocidal (see Plate 85).
technique utilizes studly male insects which are related to Biocontrols can be released by hand in small glasshouses.
pests. When Heliotizis s~ibji'esamates with the budworm It is easy and inexpensive to walk around a small garden
Helioth1s viuesceils, the offspring are sterile. These "mules" and shake beneficial insects out of their little bottles. But
are released to mate with native insects and produce infertile hand-releasing biocontrols in large glasshouses or field crops
eggs. becomes expensive. Furthermore, the even distribution of
biocontrols across large acreages is difficult to do by hand
APPLICATION RATES FOR BIOCONTROL (Mahr 1999).Thus, mechanized, calibrated delivery systems
As described earlier, there are two approaches to releas- have been devised. The mechanized delivery of some
ing biocontrols, inoculation and inundation. The inoczilation biocontrols, such as microbial bacteria and fungi, is simple-
nppromh releases biocontrols in the beginning of the season, they can be sprayed on crops using conventional pesticide
allowing them to establish a sustainable breeding popula- equipment. Mechanized delivery of insect predators and
tion. The inoculation approach works better in warm cli- parasites is more difficult. Mahr (1999) reviewed the
mates (e.g., glasshouses) that favour biocontrol reproduc- mechanized delivery of lacewing eggs and Triclzogmmina-
tion over a long season. Inundation works better in cooler, parasitized caterpillar eggs. The eggs can be glued with
short-seasoned situations. mucilage to bran flakes or vermiculite, and scattered with
Inoculation is frequently done proplzylactically, before hand-cranked applicators or compressed air blowers. Un-
pests become a problem. Several prophylactic releases must fortunately, the biocontrols end up on the ground instead of
be scheduled per season ("dribble release"), since biocontrols remaining on the foliage. This problem can be alleviated by
~ v i l ldie out if no pests are available. To prevent die out, mixing biocontrols in sticky liquid formulations and spraying
Hussey & Scopes (1985) deliberately released small them with guns fitted with large nozzles (the Bio-Sprayers).
populations of pests to assure the survival of biocontrols. Liquid carriers, however, also have problems-such as keep-
 Chapter 10: Biological Control 185

ing biocontrol organisms uniformly suspended in the liq- Bacillus stibtilis, bacteria suppress damping off fungi
uid, and keeping the organisms from suffocating in the fluid. Bacillus thiiringiensis (Bt-k, Bt-a, Bt-t, Bt-i, Bt-m), bacteria con-
For very large-scale operations, no form of ground applica- trol many caterpillars, some beetles, and a few flies
tion may be practical. In the former USSR, hemp g$wers Beauveria bassiana and B. globulifera, two related fungi con-
scattered Tricho~raininafrom an airplane (Marin 1979). trol many insects
Pl~~,toseiuluspershlilis has been mixed with corncob grits, Beauveria brongniartii (=B,tenella), fungi control beetle grubs
refrigerated, and released by conventional light aircraft onto Brassica alba, white mustard, serves as a trap crop against
spider-mite infested fields (Mahr 1999). broomrape
Burkholderia (Pseudomonas) cepacia, bacteria suppress
Pythiunz, Fusarit~mand Rhizoctonia fungi
SOME BIOCONTROL ORGANISMS Candida oleophila, yeasts control postharvest decay
Below is a list of biocontrol organisms mentioned in the Cannabis sativa has been used as a companion plant to repel
text. They are described at length under the pests they con- insects from neighbouring crops (Riley 1885, Berling 1932,
trol. Many biocontrols listed here have not yet been tested Stratii 1976,Pakhomov & Potushanskii 1977).Cnrlnabis plants
on Carztlabis. Please nzail us feedback for the next editioiz of our also suppress nematodes in soil (Kir'yanova & Krall 1971,
book. Carlilnbis can be tough on biocontrol organisms. The Kok et a/. 1994).
leaf hairs (trichomes) make it difficult for small predators to Chelonus instilaris, C. texantls, two related parasitic braconid
find and pursue prey. This also happens on leaves of toma- wasus control budworms
toes and cucumbers; entomologists have bred cucumbers Chysoperla (Chysopa) camea, green lacewings control aphids,
with fewer trichomes for improved biocontrol activity. whitefly nymphs, budworm eggs, thrips, and spider mites
Bredemann et al. (1956) began breeding hemp plants with Chysoperla comanche, Comanche lacewings control aphids and
fewer glandular trichomes, but their work was abandoned. other small insects
Furthermore, the anti-pest arsenal produced by Cannabis- Chysoperla (Chysopa) rufilabris, lacewings control aphids and
such as terpenoids, ketones, and cannabinoids-may also other small insects
repel biocontrol organisms. Some pests absorb plant chemi- Coccinella undecimpunctata, eleven-spotted ladybird beetles
cals and turn the chemicals against their biocoiztrol enenzies control aphids and other small insects
(Campbell & Duffey 1979, Clayton & Wolfe 1993). Colletotrichtrm gloeosporiodes f.sp. cuscutae, fungi control
dodder (parasitic plants)
Certain biocontrols require permits for shipment to
~ o n i o t l z y r i ~minitans,
m fungi control Sclerotinia sclerotiort~~rt
certain areas (especially to Hawai'i). Some biocontrols listed Cotesia marginiventris, braconid wasps control caterpillars
below are not yet commercially available, b u t their Cryptolaelitus montrouzieri, a ladybeetle known as the
a\-allability is changing rapidly. Rather than list currently mealybug destroyer
available biocontrols with an asterisk (and render this book Cuctrrbita foetidissima, wild buffalo gourds control cucuntber
instantly outdated), please obtain the Annual Directory beetles
published by BIRC (Bio-Intergral Resource Centre), P.O. Box Dacnusa sibirica, braconid wasps control leafminers
7414, Berkeley, California 94707, telephone: (510) 524-2567. Dactylaria species, predaceous soil fungi control nematodes
UDL addresses are ephemeral things, but four current sources Delphastus pusillus, beetles control whiteflies
of excellent information on the Internet are Koppert Delphinium exaltatlitn, larkspur plants attract Japanese beetles,
( w w ~ ~ . k o p p e r t . n l / e n g l i s h ) , Cornell University fatally
(www.nysaes.cornell.edu/ent/biocontrol), Midwest Deraeocovis brevis, predatory lnirid bugs kill aphids and thrips
Biocontrol News (entomology.wisc.edu/entomology/ Dicytiza plilvinata (=Hansfordia pulvinata), fungi control
~nbcn.html), and UC-Davis (www.ipm.ucdavis.edu/). Cercospora species and other fungi
Diglyphus isaea, chalcid wasps control leafminers
Adalia biputzctata, two-spotted ladybeetles control aphids Encarsia formosa, mighty little wasps control whiteflies
Agrobacterium radiobacter, nonpathogenic soil bacteria control Encarsia ltiteola, wasps colitrol the whitefly Bemisia nrgentiJolii
crown gall disease Entoinophtlzora exitialis, fungi control aphids
Agrotis segetutn Nuclear Polyhedrosis Virus is a NPV strain Entoinophaga g y l l i , fungi control grasshoppers
that kills A.se,getlini Entotnophthora praxibuli, fungi control grasshoppers
Aeolotlirips inteumedius, predatory thrips control Tkrips tabaci Entomoplzthora thripidum, E. pawispora, fungi control thrips
Aleocltara bilineata, staphylinid rove beetles whose larvae Erettnocerlcs eremicus, parasitic/predatory-wasps cont;ol
parasitize root maggots whiteflies
Alliilwz satizltlm, garlic, repels leaf beetles Eucelatoria b y a n i , tachnid flies control budworms
Ampelomyces quisq~ralis(=Cicinnoboltis cesatii), a fungus Euphorbia marginata, spurge plants repel moles
infecting powdery mildews Euseius hibisci, predatory mites control thrips
Aiuzgr~rsatomlls, ~nynaridwasps control leafhoppers Erynia neoaphidis (= Entoiizoplzthora nplzidis), fungi control
Anagyrus pseildococci, parasitoid wasps of mealybugs aphids
Anaphes iole, wasps that parasitize eggs of tarnished plant bugs E y n i a (Zooplzthora) radicans, fungi control leafhoppers and
Antlzoconls tzemorum, minute pirate bugs kill thrips aphids
Aphelitiils abdominalis, parasitic wasps control aphids Franklinothrips vespiformis, parasitic wasps control thrips
Aphidills colemnni, braconid wasps control aphids Fusarium Iateritiuin, fungi control broomrape
Aphidius matricariae, braconid wasps control aphids Fiisarium oxysportim (nonpathogenic), nonvirulent strain
Aplzidoletes aplzidimyza, maggots of syrphid flies control aphids controls hemp wilt fungi
Aphytis melintis, wasps control hard scales Ftlsarium oxysporum f.sp. orthoceras, fungi control broom
Alzllytas mannoratus, tachinid flies control budworms rape (parasitic plants)
Artct~zisiaabsintlziiim, common wormwood, repels flea beetles ~ a 1 e n d ; o m u s( ~ e t a s e i l i l u s occidentalis,
) predatory mites
and slugs control spider mites and some russet mites
Artl~robotrysspecies, predaceous soil fungi control nematodes Galendromus (Typhlodromus) pyri, predatory mites control
.4scl1ersonia aleyrodis, fungi control whiteflies and scales spider mites
Bacillus cereus, bacteria suppress damping off fungi Geocoris punctipes, predatory lygaeid bugs feed on white flies,
Bacilltls popilliae, bacteria control flea beetles aphids, and spider mites; a related species from India, G. tri-
colour, may also be useful.
186 Hemp Diseases and Pests

Gliocladium roseum (=Clonostachys rosea), fungi control other Pasteuria (Bacillus)penetrans, bacteria control nematodes
soil fungi Pelargonium hortorum, geranium, repels leaf beetles and
Gloinlrs intraradices, mycorrhizal fungi suppress Fusarium fungi leafhoppers
Gonatobotrys simplex, f~uigicontrol other fungi, such as Cer- Phasmarhabditis hermaplzrodita, nematodes control slugs
cospora species Phytoseiulus persirnilis, popular predatory mites control spider
Heliothis zea Nuclear Polyhedrosis Virus is a NPV strain that mites
kills H,zea and H. arrnigera Pichia guilliermondii [=Candid0 guilliermondii, Debaryomyces
Helioveya arrnigera Stunt Virus (HaSV) kills H. arrnigera and hansenii], yeasts control postharvest rot caused by Botrytis
other lepidopteran caterpillars cinerea, Alternaria alternata, Penicillium digitaturn, P italicurn,
Heterorlzabditis heliotlzidis (=H. bncteriophora), H. megidis, two and Rhizopus stolonifer
nematodes control white root grubs and many other soil insects Podisus maculiventris, predatory bugs control caterpillars and
Hippodamia convergens, convergent ladybeetles control aphids beetle grubs
Hirst~tellarhossiliensis, fungi control nematodes Pseudacteon species, parasitic flies control fire ants
Hirsutella thompsonii, fungi control spider mites and eryophid ~seudomonasfluorescens,bacteria suppress Py thiuvl, Fusariu~~z
mites and Rhizoctonia fungi
Hypoaspis aculeifer, mites prey on thrips Pseudomonas syringae, bacteria control postharvest decay
Hypoaspis miles, mites control fungus gnats Pyemotes tritici, mites control fire ants
Iplziseius (Amblyseius) degenerans, predatory mites control Pythium oligandrum, a biocontrol against the damping off
thrips pathogen Pytlzitrtlz ultimurn
Leptomastix dactylopii, parasitic wasps control Plnnococcus citri Rhizobius ventralis, black ladybeetles control mealybugs and
mealybugs scales
Lydella tlzoinpsonii, tachnid flies control European corn borer Rodolia cardinalis, Australian ladybeetles control mealybugs
Macrocentrus ancylivorus, braconid wasps control hemp borers and scales
and other caterpillars R t ~ m i n adecollata, predatory decollate snails control garden
Macroloplzifs caliginosu, mirid bugs that prey on whiteflies snails.
Melanopus saizglrinipes Entomopoxvirus, a virus that kills mi- Saccharopolyspora spinosa, bacteria control armyworms and
gratory grasshoppers other Noctuidae
Mesoseiulus (Phytoseiulus) longipes, predatory mites control Salmonella enteritidis var. issatscheilko bacteria control rodents
spider mites Scambus pterophori parasitize beetle grubs and caterpillars
Metarhizium anisopliae, fungi control spittlebugs, aphids, Steinerneha ( ~ e o a p ~ c t a ncarpocap&e,
a) S. feltiae, S.glaseri,
whiteflies, termites, ants, beetles, and other insects S, riobmvis, S. scapterisci, nematodes control caterpillars,
Metaphycus alberti, parasitic wasps control scale insects beetle grubs, root maggots, cricket nymphs, and other soil
Microcentus grandii, braconid wasps control European corn insects
borers Stethorus picipes, S, punctlrm, S. punctillum, ladybeetles
Microcronus psylliodis and M. punctlrlatus, two braconid wasps control spider mites
control flea beetles ~treptomycesgriseoviridis, actinomycetes control Riiizoctonia
Microplitis croceipes, wasps control budworms and Pythium fungi
Microtefljs flavus, parasitoids control brown scales Sycanus collaris, assassin bugs control caterpillars, beetles and
Myrotheciu~nvewucaria, fungi control nematodes many other pests. They take out an occasional honeybee and
Nematophthora pjnophila, fungi control nematodes sometimes bite a gardener's finger, but do far more good than
Nicandra physalodes, shoo-fly plant, repels whiteflies harm.
Neoplectana species see Steinernema Tagetes species, marigolds repel nematodes.
Neoseir~ll~s (Amblyseius) barkeri(=A. mackenziei), predatory Talaromycesflavus, fungi control Verticilliunz dalzliae
mites that prefer thrips but also eat spider mites Tanacetlrm vt~lgareL., tansy plants repel many insects, especially
Neoseiulus (Amblyseius) californicus, predatory mites control caterpillars and leaf beetles
spider mites and aphids Therodiplosis persicae, midges control spider mites
Neoseiulus (Amblyseius) cucumeris, predatory mites control Thripobius semileuteus, wasps control greenhouse thrips
thrips Tiphia popilliavora, T.vernalis, wasps control ~a~anese-beetles
Neoseiulus (Amblyseitrs) fallacis, predatory mites control Trichoderma viride, fungi control pathogenic fungi, especially
spider mites Rhizoctonia solani.
Neozygitesfloridana, fungi control spider mites Trichoderma harzianum controls Rhizoctonia solani, Sclerotillm
Nepeta cataria L., catnip plants, repel flea beetles ro2fSii and, to a lesser degree, Macrophomina phaseolina and
Nomuraea rileyi, fungi control Spodoptera litura and other Sclerotinia sclerotiorurn
nocturid caterpillars Trichoderma lignorum controls Fusariurn species
Nosema locustae, microscopic protozoans control grasshoppers Trichoderma ( Gliocladium ) virens, fungi control damping off
and crickets fungi Pythium ultimunz and Rhizoctonia solani, as well as
Nosema acridophagus, a related species, also kills grasshoppers Botrytis cinerea-the grey mould plague.
Nosema melolonthae controls certain beetle grubs (Melolontha Trichogramma maidis, parasitic wasps control European corn
species) borers
Nosema pyrausta controls European corn borers Trichogramma pretiosum & T , minutum, wasps control corn
Nuclear p o l y h e d r o s i s v i r u s (NPV) controls cutworms, borers, budworms, and other caterpillars
budworms, and many other Nocturids (sub-types include Trichopoda pennipes & T. giacomellii, parasitic wasps control
MbNPV, AcNPV, HzNPV, SeNPV, HcNPV, EtcNPV) stink bugs
Ocypus olens, devil's coachmen, staphylinid beetles control Trissolcus basalis. a tiny wasp that parasitizes eggs of stink bugs
garden snails Tropaeolum majus, nasturtiums are nasty to whiteflies.
Ooencyrtils subnzetallict~s,parasitoids of stink bugs Urtica dioica, nettles attract hemp flea beetles as a trap crop
Orius insidiosus, pirate bugs, control thrips and spider mites, Vairimorpha necatrix, a microsporidial protozoan that infects
as have 0. albidipennis, 0. laevigatus, 0. majusculus, 0. European corn borers and other caterpillars
tristicolour, and other related species Verticillium biguttatum, fungi control damping off fungi such
Paecilornyces farilzosus (=P. furnosoroseus), fungi control as Rhizoctonia solani
whiteflies
 Chapter 10: Biological Control 187

Verticillium c h l a m y d o s p o r i u m , soil fungi that infest soil

nematodes
Verticillium (Ceplzalosporium)Iecanii, fungi control aphids and
whiteflies and help control scales, mealybugs, thrips, beetles,
flies and eriophyid mites
Zinnia elegans, zinnias attract Japanese beetles as a trap crop

DARK SIDE
Biocontrol is not a panacea. All pest-control techniques
affect other organisms, and pose some degree of environ-
mental risk. Overuse of biocontrols, especially inundative
biocontrols, may reduce their effectiveness. Pests can develop
resistance to bihcontrol organisms if confronted with suffi-
cient selection pressure (Gould 1991). Biocontrols may ig-
nore pests and attack nontarget hosts, including other
biocontrol organisms. This side effect has actually led to the
extinction of nontarget organisms. H o w a r t h (1991)
documents dozens of extinctions of nontarget species caused
by rogue biocontrols, far more than the number of extinctions
caused by chemical controls. Off-target biocontrols may
become pests themselves. This is especially true among
insects and fungi released for the control of weedy plants
(McPartland &West 1999).
Biocontrols, like chemical controls, may disrupt local = harmless to biocontrols (<lo% mortality);
ecologies, allowing an outbreak of secondary pests. The Eu-
ropean ladybeetle ~occinellaseptempunctata, fir-instance, may
disrupt established biological control systems in the USA % = relatively safe (10-30% mortality);

(Schaefer et al. 1987%).Lastly, biocontrols may become public

health problems: the predatory snail Euglandi~za rosea,
released to control plant-eating snails, is now known to carry
I = relatively harmful (31-90% mortality);

-
a lung worm which infects humans (Howarth 1991).
Microbial biocontrols, especially fungi, may infect humans
(Rippon 1988, Samuels 1996).
li = very harmful (>go% mortality)

According to Orr & Suh (2000), biocontrol product sup- Figure 10.2: Average toxicity of pesticides on biocontrol
arthropods, from data computed by Theiling & Croft (1988).
port in Europe is superior to that in the USA. European
suppliers maintain rigid quality control procedures, and ship
their products by overnight delivery in refrigerated trucks
or in containers with ice packs. Furthermore, European sup- pupal stages were less susceptible than larvae and adults.
pliers provide their customers with extensive tec-hical in- The average toxicity of different groups of pesticides is
formation, and even monitor temperature data and pest presented in Fig 10.2. Theiling & Croft (1988)lumped micro-
populations in areas where biocontrol releases take place. bial biocontrols (viruses, bacteria, fungi, and nematodes) with
Customers are informed in advance of product delivery and microbial fermentation prodlicts (abamectin, streptomycin, etc.)
release dates. In contrast, nearly 50% of USA suppliers into one group, "Microbials." This was the least-toxic group,
shipped dead biocontrols or species other than that which naturally Within "Microbials," Theiling & Croft reported that
was claimed, shipped products in padded envelopes by viruses, bacteria, and fungi were safer to biocontrol arthro-
standard "snail mail," and provided little or no product in- pods than were predatory nematodes and fermentation prod-
formation or instructions (Orr & Suh 2000). The biocontrol ucts. Within the group labelled "Juvenoids," the chitin in-
industry in the USA needs to improve its self-regulation; hibitors were safer than juvenile growth hormones.
perhaps government regulation is required to implement Botanicals and minerals (e.g., Bordeaux mixture, copper,
tighter quality control and product support. sulphur) were moderately toxic to biocontrols. The most toxic
pesticides were synthetic pyrethroids. Of course, within each
BIOCONTROLS & PESTICIDES group we find exceptions,-which is why the Koppert list is
We present the effects of some pesticides on some so valuable.
biocontrol organisms in Table 10.1. For a complete list of Theiling & Croft also presented pesticide "selectivity ra-
pesticides and biocontrol organisms, see the "side effects" tios," calculated by dividing the LDS0of pests by the LD50of
list available on Koppert's web page (www.koppert.nl/ their biocontrols. These selectivity ratios are valuable
english). You will note many dash-marks (-) in Table 10.1, information for IPM. The more selective a pesticide, the better.
indicating unknown data. The Koppert list is updated The most selective pesticides included pirirnicarb and dicofol;
monthly with new information. some of the least selective pesticides were DDT, parathion,
Theiling & Croft (1988) summarized 1000 publications and cypermethrin. Theiling & Croft's literature review re-
citing the effects of pesticides on biocontrol arthropods garding biocontrols and pesticides has been compiled on
(biocontrol insects and mites). Overall, pesticides were less computer diskettes. The database is called NERISK (Natural
toxic to predators than parasitoids. The least susceptible Enemy Risk Assessment, formerly called SELCTV), and is
biocontrols were lacewings and predatory bugs (Miridae, sold by the National Technical Information Service,
Lygaeidae, Anthocoridae). For most biocontrols, egg and Springfield, Virginia.
 "All stlbstances are poisons; there is none zuhich is not a poison. The right dose differentiates a poisonfrom a remedy."
-Paracelsus

Chapter 11: Biorational Chemical Control

Chemical controls are also known as pesticides. Use them permitted in The National List of materials mandated by the
only when all else fails. All pesticides cause side effects. Some Organic Foods Production Act of 1990. But some chemicals
are worse than others. See the warnings below. Some pesticides listed here are restricted by the List, such as nicotine and
can be used on industrial crops (i.e.,fibre hemp) but should sodium nitrate, and eschewed by organic farmers.
never be used on food and drug crops such as seed oil or Some chemicals kill pests, but other chemicals provide
marijuana. us with other options-like repelling pests, or confusing
We define biorational chemicals as naturally-occurring them. Baiting a field with an artificial sex pheromone, for
compounds or synthetic analogues of naturally-occurring instance, confuses males and makes reproduction impossible.
compounds, such as synthetic pyrethroids (Djerassi et al. Repellents drive pests away from plants. Some repellents
1974).Natural means these chemicals occur in nature. It does d o not harm pests, they are simply offensive to pests.
not mean they are safe or belong in baby food. Natural Bordeaux mixture and copper compounds repel insects
chemicals can be quite toxic, naturally toxic. without killing them (these chemicals, however, are lethal to
Most of the biorational chemicals we discuss here are fungi). Garlic extracts are classic olfactory repellents-flying

Table 11.l:
Pesticide formulations. The abbreviations for formulations are used throught the text.

FORMULATION
ABBREV. DESCRIPTION

Aerosol A Pesticide dissolved in volatile solvent and pressurized in a can by a propellant gas like CO,, a
convenient but expensive formulation.

Bait B Pesticide impregnated into a substrate attractive to pests, such as food or sex pheromones.

Dry flowable DF Dry pesticide formulated as granules, to be mixed with water and sprayed; much like a WP, but the
heavier DF granules reduce a person's exposure to airborne particles during handling and mixing.

Dust D Nearly microscopic particles of powder-dry pesticides diluted in a dry carrier; dusts adhere well to
plant surfaces, useful for treating small indoor areas but tend to drift outdoors.
-- -

Emulsifiable EC Petroleum-based liquid plus emulsifier, mix with water and spray; like all sprays, may ineffectively
concentrate roll off plant surfaces. Phytotoxicity hazard usually greater than other liquid formulations, may
leave visible residue on plants.

Flowable FC Solid or semisolid pesticide wet-milled into a pudding-like consistency to be mixed in water and
concentrate sprayed; requires frequent agitation to remain suspended, may clog spray equipment, may leave
visible residue on plants.

Fumigant F Vapour usually stored under pressure in metal bottles as liquified gas; vaporizes when released,
quickly dissipates, dangerous, expensive.

Granule G Prepared by applying liquid pesticides to coarse particles of porous material; granules are like
dusts, but larger so they wind-drift less.

Plant- p2 A new category proposed by the EPA for bioengineered crop plants yielding toxins normally
Pesticide produced by other organisms, such as plants that produce BT-a toxin ordinarily produced by
Bacillus thuringiensis.

Slow-release SR Pesticide embedded in polychlorovinyl resin (e.g., No-Pest Strip@) to slow the rate of volatilization;
other SRs include paint-on pesticides and microencapsulation in semi-permeable membranes.

Soluble SP Soluble powder that dissolves in water to form a true solution; packaged as powders or
powder concentrated solutions; unlike FCs, no further agitation is needed after SPs are dissolved.

Insoluble pesticide mixed with mineral clay into tiny particles <25 ym in diameter (WPs look like Ds
powder but are more concentrated); suspended in water with a surfactant; act like FCs (i.e., sprays may
clog), safer to plants than ECs, may leave visible residue on plants.
190 Hemp Diseases and Pests

Table 11.2: EPA pesticide acute toxicity classification.

LDjO FOR THE RAT LDSoFOR THE RAT LD50FOR THE RAT
CLASS ORAL (mg kg") DERMAL (mg kg") INHALED (mg 1") EYEEFFECTS SKINEFFECTS

I 50 or less 200 or less 0.2 or less corrosive corrosive

opacity not reversible

50-500 200-2000 0.2-2.0 corneal opacity reversible severe irritation at

within 7 days, irritation 72 hours
persisting for 7 days

500-5000 2000-20,000 2.0-20 no corneal opacity, irritation moderate irritation

reversible within 7 days at 72 hours

IV greater than 5000 greater than 20,000 greater than 20 no irritation mild irritation at
72 hours

aphids "smell" garlic extracts sprayed on plants and turn resistance arises by a simple mutation of a single gene.
away before landing. Irritant repellents such as pyrethroids Resistant pests arise soon after systemic pesticides are de-
cause insects to perform exaggerated grooming behaviours; ployed
- . (Agrios 1997).
the insects become stressed and agitated and go away. Ovi- chemicals come in an assortment of formulations, see
position repellents deter females from depositing eggs on Table 11.1. Fumigants are gases. They are very effective but
treated surfaces. THC and CBD act as oviposition repellents they must be used in enclosed areas, they require expensive
against Pieris brnssicae butterflies (Rothschild & Fairbaim 1980). equipment, and most are carcinogenic. sprays a r h ~ i ~ u i d s
Antifeedant chemicals inhibit feeding behaviour, such and much easier to apply, but may be ineffective if they bead
as polygodial and azidarachtin (from the neem tree). Some up and roll off plant surfaces. Spray formulations include
antifeedants are not repellents, so pests remain on plants until emulsifiable concentrates (ECs),flowable concentrates (FCs),
they starve. Antifeedants and repellents can be applied to soluble powders (SPs), dry flowables (DF), and wettable
flowering tops, which drive pests down to older, unharvested powders (WPs). Dusts adhere firmly to plant surfaces but
leaves. On unharvested leaves, the pests can then be killed are susceptible to wind drift during application. Granules
with a localized application of a nasty chemical. are similar to dusts but consist of larger particles, so they
The nasty chemicals are pesticides, which are designed wind-drift less. Granules Dose a serioushazard to birds, whb
to kill pests-not confuse them-and kill them quickly. mistake them for food or grit and eat them. Baits are often
Pesticides work when pests injest them (stomach poisons) formulated as granules but have the power of attraction (e.g.,
or touch them (contact poisons). Stomach poisons only kill food attraction, colour attraction,-sex attractant). plant-
pests that chew and swallow their food. Stomach poisons pesticides are a new category the EPA proposed for plants
are not very effective against insects with piercing-sucking genetically manipulated by recombinant DNA technology
mouth parts (e.g., aphids). Aphids push their stylets right (EPA 1994). Bioengineered plants produce toxins normally
past poisons adhering to plant surfaces. Sucking insects must produced by microbes or other organisms-a grey zone
be controlled with contact poisons. between biocontrol and chemical control (Cook & Qualset
Pesticides are named by their target: miticides 1996).Much remains unknown about bioengineered plants,
(acaricides) kill mites, insecticides kill insects, fungicides yet USA regulatory agencies have approved them, so the
kill moulds, antibiotics kill bacteria, herbicides kill plants, USA has become a large-scale experiment for transgenic field
nematocides kill nematodes, avicides kill birds, and studies. Unpredicted side effects can be ecological, such as
rodentocides kill rodents. Disinfestants eradicate surface the death of monarch butterflies from pollen blown off
organisms, whereas disinfectants work systemically, killing transgenic Bt corn (Losey et al. 1999). Medical side effects
pests and pathogens within plants. also arise, such as changes in the gastrointestinal tract after
r\/Iost older pesticides work on the surface. They do not eating potatoes engineered for insect and nematode
penetrate plants, so they cannot control pests or pathogens resistance (Ewen & Pusztai 1999),and lethal allergic reactions
already established within plant tissue. Surface pesticides from eating transgenic soyabeans (Nordlee et al. 1996).
are frequently called protectant pesticides - they repel The amount-of actiGe ingredient (a.i.) present in a
insects off plants, or prevent fungal spores from germinating. pesticide formulation is listed on the product label. Solid
New pesticides may work deeper than the surface. Sys- formulations list the amount as a percentage of weight. For
temic pesticides can penetrate tissue and redistribute within instance, a 50%WP contains 50% a.i., or 50% pesticide and
plants. Diffusion of pesticides via the xylem is called 50% "inert ingredients." Liquid formulations list the a.i.
L,

apoplastic mobility, and movement cell-to-cell, such as concentration two different i a y s : as a percentage of weight,
through phloem cells, is called symplastic mobility. Thus, or as the number of pounds per gallon (lbs/gal or lb gal-1).
systemic pesticides provide surface protection as well as deep For instance, a 4 lb/gal EC contains 4 pounds a.i. per gallon
and durable eradication of established pests and pathogens. of product. This equals a 40-50°/oEC, depending on specific
The development of systemic pesticides in the 1960s was gravity of the "inert ingredients."
hailed as a breakthrough. But systemics tend to remain in Chemical controls, whether natural or synthetic, can
plant tissues without decomposing, which is not good for cause serious side effects. Technically, there is no such thing
products destined for human consumption. Furthermore, as a pesticide. "Pesticide" implies that a chemical selectively
many systemics kill pests by a "site-specific" action upon kills a pest, leaving everything else alone. Pesticides are really
one metabolic pathway. As a result of this specific action, pest biocides-they harm many living things. They may even harm
 Chapter 11: Biorational Chemical Control 191

plants they are supposed to protect. They often damage or juana are a different story. Few pesticides have been tested
kill nontarget organisms. Honeybees and bald eagles are for their effects when burned and inhaled. Only tobacco re-
famous nontarget victims. Mycorrhizae are less known searchers seem to be interested in these questions, and little
nontarget victims, killed by fungicides as well as insecticides research has been done (Lucas 1975).
and nematocides (Menge 1983). Our final Faustian bargain with pesticides is the rebound
Damage to honeybees can be reduced by spraying crops of resistant pests. In 1950, shortly after DDT was invented,
after sundown. Protecting bald eagles is not so easy. Over 1 only 20 insects were resistant to the new panacea. Since then,
billion pounds of pesticides are applied annually in the USA, the number of resistant pests has risen to 535 insects, 210
which represents 34% of total worldwide application fungal pathogens, and 200 weed species; many of these pests
(Pimentel 1991).A cynic might point out this averages 20 g are resistant to multiple pesticides (Pimentel1991).The search
of pesticide per acre-pretty thin. But persistent pesticides for effective pesticides places farmers on a "pesticide
often become concentrated and "biomagnify" in the food treadmill." Pesticides eventually become counter-
chain. One ppm DDT in pond water might accumulate to productive-Pimentel (1991) argues that pesticide
100 ppm within plankton and accumulate to 1000 ppm in application on USA croplands has grown 337fold since the
plankton-feeding fish. Eagles eat an accumulation of fish and 1940s, yet crops losses from pests have actually increased in
become an endangered species. that time, from 31% to 37%.
Chemicals also affect us. Farmers face acute poisoning Despite these serious caveats, chemicals will continue
as a job hazard. Organophosphates cause the greatest to be used on crops. If chemicals are to be used, we want
number of pesticide poisonings in the USA because of their people to use them correctly, to minimize their toxic side ef-
potency and widespread use. Most are EPA Class I toxins fects. We dedicate this chapter (and Appendix 2) to the spirit
(see Table 11.2).A lethal dose can be absorbed through the of harm reduction.
skin, eyes, lungs, skin, or stomach (don't use your mouth to Some vesticides are less toxic than others. Less-toxic
clear a spray line or prime a siphon!). In one remarkable biorational pesticides are preferable, but these require more
report, parathion spilled on a pair of coveralls hospitalized precise timing to be effective, work more slowly, must be
three workers over a two-week period despite repeated applied more often, a n d pests must be moie closely
laundering (Clifford & Nies 1989). monitored. Nevertheless, biorational pesticides are safer,
Approximately 40,000 people, mostly children, are more ecological, and morally imperative. A classification of
treated-for pesticide poisoning in the USA every year. biorational pesticides is presented in Table 11.3; a selection
Worldwide, pesticides kill 250,000 people annually. To quote of biorational pesticides is presented in Table 11.4.For sources
Cvnthia Westcott (19641,~, "The use of chemicals bv amateurs see the Directory ofleast-Toxic Pest Control Products, published
is hazardous in any event." Always read the label and never annually by BIRC (Bio-Intergral Resource Centre), P.O. Box
act in haste. 7414, Berkeley, California 94707, telephone (510) 524-2567.
Many pesticides are carcinogenic (causing cancer), mu- For more information on pesticides, see the series written by
tagenic (causing gene damage), teratogenic (causing birth Thomson or the annual Farm Chemicals Handbook edited by
defects), or oncogenic (causing tumours). Agricultural Meister.
workers suffer an increased incidence of brain tumours,
testicular cancers, leukaemias and lymphomas. The EPA Table 11.3: Classification of Biorational Pesticides.
ranks pesticides on a multi-tiered scheme adapted from the - -

World Health Organization's International Agency for Natural Non-botanicals

Research on Cancer (IARC). Many pesticides fall in IARC Mineral-based
Group 2B: Sl~bstancesPossibly Carcinoget~icTo H i l n ~ a n s . Carbon-based
Rockwool and saccharin also fall into this category. Botanicals
Some pesticides are n o t acutely poisonous nor Fermented products
carcinogenic, b u t act as female hormones (they are
oestrogenic), and disrupt our endocrine systems. Oestrogenic Synthetic Synthetic botanicals
compounds are especially harmful to developing fetuses. (or semi-synthetic) Pest-growth regulators
and-me-down poisons" absorbed by females Synthetic pheromones
may harm their offspring post iltero, at any time of their
offspring's lives, and may even harm offspring of their
offspring. A male exposed before birth to oestrogenic
compounds may have undescended testicles at birth, a low NONBOTANICALS
sperm count at puberty, testicular cancer in middle age, or Nonbotanical natural pesticides are either mineral-based
prostate cancer as an old man-all from his prenatal exposure or carbon-based. Mineral-based pesticides usually work on
i ~ o l b o r net al. 1996). Certain plastics produce oestrogenic contact. Afew work as stomach poisons, such as once-popu-
effects. Nonylphenol plastics are added to polystyrene and lar but now banned arsenic insecticides (e.g., Paris green)
polyvinyl chloride (PVC) to make these plastics less brittle. and mercurous insecticides (e.g., calomel). Mercurous com-
Bisphenol-A is a d d e d to polycarbonate for stability. pounds also served as fungicides, such as Ceresanm -
Polystyrene, PVC, and polycarbonate are frequently used methoxyethyl mercury chloride. Ceresan was popular in
in hydroponic systems, a n d they leach oestrogenic European Community countries from 1929 until it was
nonylphenol and bisphenol-A into the water (Colborn et al. banned in 1992 due to adverse toxicology (Maude 1996).
1996). In 1992, fourteen European countries agreed to ban Many mineral-based pesticides are permitted on "Certified-
these chemicals by the year 2000. Organic" farms if they come from a mined source, not a
Consumers face unknown exposures; pesticide residues chemical laboratory.
on foodstuffs are a major concerny~heEPA accepts pesticide Bordeaux mixture is a foliar fungicide (introduced by
"tolerances" (maximum allowable residues), then tests Millardet in 1883), b u t also kills bacteria and repels
foodstuffs in random market samplings. If residues exceed caterpillars, beetles, and other insects. Bordeaux is a mix of
tolerance levels, the crop cannot be s d d . Residues in mari- Ca(OH)2 (calcium hydroxide or lime) and CuS04 (copper
192 Hemp Diseases and Pests

Table 11.4: Some biorational pesticides.

GENERIC
NAME US TRADE
NAME(S)
TM OR @ CHEMICAL
TYPE EPA CLASS

abamectin Avid, Vertimec fermented product Ill

boric acid Tim-Bor. Borid. Roachkil mineral-based Ill-IV
calcium cvanamide Cvanamid carbon-based
Cannabis Muaales botanical IV
carbon dioxide co2 carbon-based
castor castor oil botanical Ill
cholecalciferol Muritan, Rampage, Quintox carbon-based I
copper sulphate Basicop, Tribasic, Bordo mineral-based Ill
copper sulphate & lime Bordeaux mixture mineral-based Ill
copper oxychloride Champ F, C.O.C., C.0.C.S mineral-based Ill
cryolite Kyrocide, Prokil mineral-based Ill
cupric hydroxide Kocide 101, Blue Shield mineral-based Ill
cuprous oxide Cuprocide, Perenox mineral-based II
, cyromazine Aurmor, Trigard growth regulator Ill
diatomaceous earth Celite, Celatom mineral-based IV
diflubenzuron Dimilin, Vigilante growth regulator Ill
formaldehyde Formalin carbon-based II
griseofulvin Fulvicin, Grisactin fermented product II
1 hellebore Hellebore botanical Ill
1 horsetail Horsetail botanical IV
hydroprene Gencor growth regulator IV
imidacloprid Confidor, Admire, Provado synthetic botanical Ill
kinoprene Enstar growth regulator II
methoprene Kabat, Minex, Altosid growth regulator IV
neem (azadirachtin) Bioneem, Azatin, Neemayad botanical IV
1 nicotine Black-Leaf 40 botanical II
1 oil Sunsprav, Scalecide carbon-based IV
piperonvl butoxide PBO, Pvbuthrin botanical svneraist Ill
- -

polygodial Polygodial botanical

pyrethroid Pokon, Safer's Bug Killer synthetic botanical Ill
pyrethrum Buhach, Insect powder botanical Ill-IV
quassia Bitterwood botanical IV
red squill Rodine, Dethdiet botanical II-Ill
rotenone Derris, cube botanical 11-111
ryania Ryan 50, Ryanicide botanical Ill
sabadllla Red Devil, Veratran D botanical IV
soap M-Pede, Safer's, Dr. Bronner's carbon-based IV
sodium bicarbonate Baking Soda carbon-based IV
sodium hypochlorite Bleach mineral-based II-Ill
1 sodium nitrate Chilean nitrate carbon-based
I spinosad Conserve, Tracer, Naturalyte fermented product
1 streptomycin Agrimycin 17, Agri-Strep fermented product IV
1 strychnine Strvchnine botanical I
S U ~ D ~ U ~ Cosan, Hexasul mineral-based IV
trimethyl docecatriene Stirrup M pheromone
urea ClandoSan carbon-based
 Chapter 11: Biorational Chemical Control

sulphate) in water. Only the copper is fungitoxic; the lime contact activity against mites and thrips. Available as a 96%D
serves as a "safener" to protect plants. Preparations of or 96%WP, applied with a duster or sprayer. Cryolite is per-
Bordeaux are labelled as a ratio of copper sulphate (in sistent in dry weather. Do not combine cryolite with lime or
pounds) and quick lime (in pounds) to water (in gallons). other alkali. Some phytotoxicity of corn and fruit trees occurs
The most popular mixture is 4:4:50 (= 8:8:100). Bordeaux is in damp climates. Nontoxic to birds and bees, moderate tox-
widely available in SP or WP formulations. To make Bor- icity to fish. Natural (organic-approved) cryolite is mined in
deaux, suspend copper sulphate snow (not fixed copper) in Greenland, synthetic sources are manufactured in the USA
half the water, suspend quick lime in the other half, then and other countries.
mix the two solutions. Spray immediately after mixing. Diatomaceous earth (DE) is a contact insecticide, often
Agitate the mix frequently to keep it from settling in the spray combined with pyrethrins. DE tears microscopic holes in the
tank. Metcalf et al. (1962) said a gallon of Bordeaux covers surfaces of soft-bodied insects, and slices between the
1000 square feet of leaf surface. ~ersistenceis long. Bordeaux exoskeleton plates of hard-bodied insects. In hot, dry weather
causes some phytotoxicity in plants, especially on seedlings the injured insects rapidly dehydrate and die. DE is
when applied in cool, wet weather. To reduce phytotoxicity, formulated as a talc-like D. It is persistent in dry weather.
decrease copper or increase lime (mix 2:6:100 or 8:24:100), DE can irritate eyes. It should not be applied to drug crops
and spray on a warm, dry day. Bordeaux is slightly toxic to destined for inhalation, because DE is a serious respiratory
honeybees, practically nontoxic to mammals, but highly toxic hazard.
to fish. The RE1 is 48 hours. Sodium bicarbonate, NaHC03 (baking soda) works
Boric acid (H3B03), is a stomach and contact poison well against powdery mildew. Spray a 0.5% solution of
useful against roaches, earwigs, crickets, and ants. Available baking soda (15 ml per 4 1 or 3 tsp/gallon). Some growers
as a 98%D, 18%G, a n d 1-5%B formulations. These double the concentration. Sodium bicarbonate works better
formulations are persistent in dry conditions, but wash out when combined with light horticultural oil (also 15 ml per
rapidly after rainfall. Boric acid is phytotoxic. Borax is not 4 1). Potassit~rnbicarbonate (Kaligreena, Armicarb loo@)can
dangerous to honeybees or birds. The dust irritates eyes (but be used the same way as sodium bicarbonate. Nontoxic, low
boric acid sol~itionis used as an eyewash). ~ o d i t i hborate persistence, little ph)totoxicity.
(Na,B,O,,. 4H,O) is a mineral borate used on wood products. Sodium hypochlorite, a 5% solution of NaOCl is sold
Copper is a fungicide, useful against grey mould, leaf as household bleach. Bleach disinfects viruses, bacteria, fungi,
spots, blights, anthracnose, powdery mildews, downy and small arthropods from equipment, glasshouse walls, and
mildews, and also some foliar bacterial diseases. Fixed the air. Asolution of one part household bleach in nine parts
copper is less soluble than the copper used in Bordeaux water (0.5% NaOCl) is used as a seed soak or soil drench
mixture, so it is less phytotoxic, but also less effective. It is against fungi. Undiluted bleach is toxic to plants and caustic
not toxic to birds and bees, but very toxic to fish, and acts as to our eyes and skin.
a eye and skin irritant to mammals. The RE1 for all copper Sulphur (sulfur) is applied as a 98%D (fungicidal against
compounds is 48 hours. Copper can bioaccumulate in food rusts and powdery mildews) or a 30-92%WP (miticidal, but
chains. Below we present several useful inorganic salts of not against spider mites). Mix and spray at a rate of 600 g
fixed copper. Collectively, copper compounds are the second sulphur per 100 1water. Available from many manufacturers.
most widely-selling fungicides, behind only mancozeb Sulphur is the fourth most-popular fungicide, behind
(Hewitt 1998). mancozeb, copper and chlorothalonil (Hewitt 1998).Sulphur
Copper sulplmte (CuS04*5H20)is the most popular form persists on plants until washout, then it becokes a
of fixed copper. It is often called "tribasic copper." Available component of the soil environment. Sulphur can injure
as a 1-82%WP, 6-8%FC, or 7%D. Residuals last five to 14 plants, especially in hot, dry weather. DO-not apply if the
days. temperature is expected to exceed 30°C. Sulphur mixed with
Copper oxychloride (Cu2Cl(OH)3)is sold as a 45-85%WP lime kills more insects, but this mixture causes greater
and 10-25%D. A sulphated version (copper oxychloride phytotoxicity. Honeybees, birds, and fish are safe. sulphur
stilpllate), is available as a 50-53%WP and 3-15%D. Copper -- dust is irritating to eyes and lungs; ingested sulphur acts as
oxychloride is more popular in Europe, copper oxychloride a laxative. The RE1 is 24 hours.
sulphate is more popular in the USA. Water, full-strength, loo%, controls some pests and
C~cprichydroxide (CU(OH)~) is sold as a 21-77OloWP and pathogens. Parker (1913b) washed 50-70% of spider mites
3-55%D. This formulation dissolves in water better than off hops plants by directing a strong but fine stream of cold
most fixed coppers, causing less clogging of spray nozzles. water at undersides of leaves. This also destroys their
Ctiprous oxide (Cu20) is sold as a 19-80%WP, 64%EC, webbings. Gentle misting works if predatory mites (e.g.,
60-65%FC, 4-83%D, and G. It is less phytotoxic than other Phytoseiultrs persirnilis) are present-the mist slows spider
copper compounds, but more toxic to us. Residual period mites yet speeds the performance of predators.
and reapplication interval is seven to ten days. Cuprous Unfortunately, spraying water increases the likelihood of grey
oxide is also used as a seed treatment. mould in flowering plants.
Clay microparticles, modified from kaolin, can be
sprayed onto crops in a thin, watery slurry. The material CARBON-BASED NON-BOTANICALS
sticks to plants and dries to a white powdery film. The The other category of nonbotanical chemicals is carbon-
microscopic clay particles attach to the bodies of insects. The based pesticides. We define carbon-based pesticides as
microparticles repel small, thin-skinned pests, such as mites, naturally-occurring compounds that contain carbon but are
aphids, thrips, and leafhoppers. The coating does not not extracted from plants. Some are carcinogenic and not
interfere with photosynthesis; sunlight diffuses into leaves permitted in "Certified Organic" operations.
with little reduction of light. Clay microparticles are nontoxic Calcium cyanamide, CaCN,, also known as lime
and do not harm predatory biocontrol insects. nitrogen, is made from heating lime and coal. It has been
Cryolite, sodium aluminofluoride (Na3A11F6)works as used since the turn of the century as a nitrogen fertilizer, a
a stomach poison against insects with chewing mouthparts weed killer, and a fungicide. Available as a G, mix 562 kg
(flea beetles, weevils, and some caterpillars). It also has some ha-1(500 lbs/acre) in soil to kill soil pathogens. It is phytotoxic
194 Hemp Diseases and Pests

and must be applied in late autumn or very early spring, but temperatures are cool. Follow each soap spray after several
is broken down into urea by seeding time. hours by a water rinse, especially in warm weather. Soap is
Carbon dioxide is a fumigant gas that kills nearly any- not selective and kills many beneficial insects. Soap kills
thing not green. Seal plastic bags over plants and inflate bags predatory mites but their eggs survive, so soap and preda-
with the gas for several hours. C 0 2 is useful against recalci- tory mites can work together. Soap can be used on medical
trant insects and fungi. High concentrations are lethal to Cannabis up to a week before harvest without any- distaste-
humans, so use caution in growrooms. ful residues discerned on finished dry flowers.
Cholecalciferol, a sterol hormone, is used as a rat poi- Sodium nitrate, NaN03, kills damping off fungi and
son, with an oral LDS0of 43 mg kg-'. Available as a 0.075%B. nematodes (which act synergistically and should be eradi-
It causes hypercalcemia and kills rats in two to four days. A cated together). ~ v a i l a b l efrom many manufacturers as a G.
lethal dose can be consumed i n a single feeding or Mix into topsoil at a rate of 135-270 g m-2 (4-8 oz/ yard>).It is
accumulated in smaller, multiple feedin& unfortunately, phytotoxic but not persistent; mix into soil several weeks
dogs and cats may be more sensitive to cholecalciferol than beiore setting seed. -
rats. In humans, the RDA of cholecalciferol (vitamin D3) is Urea, CO(NH2)2, kills some damping off fungi and
10 pg. Vitamin D2, ergocalciferol, is also sold as a rodenticide. nematodes like sodium nitrate, and is also available as a G.
Formaldehyde is carcinogenic and not used much Mix 540 g m-2 (16 ozlyard2) into topsoil several weeks before
anymore. It works as a liquid/fumigant, lethal to damping- sowing seed. Clandosano combines urea with chitin from
off oomycetes, fungi, bacteria, insects, nematodes, and ground-up crab shells. Chitin is a polymer of the amino sugar
viruses. Formalin is a 38% solution of formaldehyde; N-acetyl-glucosamine (NAG is a building block of cartilage
paraformaldehyde is a solid polymer. Using formaldehyde in humans). Chitosan is a nonacetylated form of chitin. Crab
as a seed disinfectant began in the 1890s, to eliminate shell chitin encourages the gydwth of nematicidal soil
seedborne fungi. It has also been used as a soil drench, or organisms when mixed into soil at a rate of 2-4 kg per 10 m2.
mixed into porous materials (oat hulls, sawdust) and sprinkled in However, chitin may cause phytotoxicity if mixed into soil
field furrowsto kill soil organisms.Formaldehyderapidly oxidizes at levels exceeding 1% (Mian et al. 1982).
to formic acid. It is phytotoxic, toxic to honeybees, fish, and
- ~
Human urine repels many vertebrats, such as deer and
mammals. The vapour irritates eyes and lungs. other varmints. Mark your territory.
Oil was traditionally sprayed on trees in winter as a dor-
mant oil to kill aphid eggs and mites. Dormant oil is heavy
(with a 50% distillation temperature or "flashpoint" of BOTANICAL POISONS
2460°F) and herbicidal to herbaceous plants. Improved re- Many companion plants mentioned in Chapter 10, such
fining techniques have produced lighter horticultural oils as garlic, retain their repellency after they are harvested and
(summer oils) that can be sprayed on sensitive foliage. Light dried. They can be crushed up, mixed with water, and
"hort oils" flash as low as 412°F (oils below 400°F do not kill sprayed on plants as bug repellents. Repellent sprays are
insects). Hort oils are ranked by their UR rating (unsulpho- commercially available (Guardians a garlic extract; Hot Pep-
nated residue, the percentage of oil that is free of phytotoxic per Wax9 a Capsiczrm extract).
residues after flashing)-look for 92% or greater. Oil gravity "High powered" botanicals do more than repel-they
("10unsaturated hydrocarbons) is also important-look for kill. They act as contact or stomach poisons. Some botanicals
32(%) or less. are extrenzely lethal and not permitted on Certified-Organic
Hort oil is derived from animals (fish oil), vegetables farms, such as nicotine and strychnine.
(seed oil), or minerals (petroleum). The oil plugs up spiracles In their simplest form, botanical pesticides are simply
(breathing tubes). It suffocates mites, whiteflies, aphids, dried plants, ground to a fine powder. The powders can be
mealybugs, scales, and other soft bodied insects. Hort oil also used full-strength or diluted in a carrier such as clay or talc.
kills many beneficial insects, especially immatures (adults Pesticidal extracts can be produced by soaking fresh or dried
may escape the nozzle). Hort ~ i l r a r e l ~ c a u s phytoxicity
es if plants in a solvent, such as water or alcohol. After a while
sprayed in a 1% solution. A 3% solution may cause dam- the solid material is filtered out, leaving the liquid extract.
age-always test-spray. To reduce phytotoxicity, crops should Extracts can be sprayed full-strength on crop plants, boiled
be well-watered and then sprayed on a warm day (21-38°C) down to liquid concentrates, distilled to pure pesticidal
with low humidity (<40% RH), so oil can kill insects and chemicals, or applied to clay for use as dusts.
then evaporate quickly. Dried oil is inactive. Hort oil can be Rosenthal(1999) described a home-made botanical brew
used against fungi if mixed with 0.5% baking soda (15 ml he sprayed on marijuana plants. To 1 1 (1 quart) of boiled
per 4 1 or 3 tsp/gallon). Do not mix with sulphur. Oil can water add 30 ml (2 tablespoons) of the following: ground
also be used to form soap (see below). cinnamon (Cinnanzonum zeylanicz~m),ground chili pepper
Soap is oil combined with sodium or potassium alkali. (Capsicumannuum), ground black pepper (Pipernigri~nz),mint
Most soaps sold today consist of potassium oleate (the po- or peppermint leaves (Mentha species), fresh crushed garlic
tassium salt of oleic acid, a fatty acid). Soap suffocates many (Allium sativunz), fresh chopped onion (Allillrn cepa), and or-
insects (but not their eggs). Safer's@Insecticidal Soap causes ange peel (Citrt~ssinensis). When the mixture cools but is
98.69/0mortality in aphids and 91% mortality in two-spotted still warm, add 500 m l ( 2 cups) isopropyl alcohol, 250 m l ( l
spider mites (Puritch 1982). Dr. Bronner's Soap@also works cup) of strong coffee, and 125 ml (0.5 cup) low-fat milk).
well (mix 2.5 ml per 4 1 or 0.5 tsp/gallon). The USDA is mak- Strain through a fine sieve, add 30 ml liquid soap, and add
ing soap from tobacco extract that kills sweetpotato enough water to make 2 1 of pesticide.
whiteflies. A 1% solution of M-Pede@soap enhances the ac- Some botanical poisons have been used for centuries,
tivity of synthetic miticides (abamectin, dicofol) and insecti- such as neem and ganja in India, pyrethrum in the Middle
cides (diazinon, malathion), according to the manufacturer. East, and rotenone in South America-all described below.
Soaps work best if mixed in soft water; soften the water New commercial products are being developed from cala-
if hardness exceeds 300 ppm (or 17.5grains per gallon). Plants mus (Acorlrscalamus),basil (Ocimum basilia~m),big sagebrush
should be sprayed at five-day intervals. Soaps lose activity (Artemisia tridentata), chilcuan (Heliopsislongipes),and mamey
after drying, s o s p r a y early in the morning w h e n (Mammea anzericana).
 Chapter 11: Bioraticma1 Chemical Control 195

Cannabis flowers, leaves, seeds, and their extracts have insects if sprayed directly on them. Limonene irritates the
been used as repellents or pesticides against insects eyes, mucous membranes, and skin.
(Culpepper 1814, King 1854, Riley & Howard 1892, Indian Neem is extracted from seeds and foliage of the Indian
Hemp Drugs Commission 1894, MacIndoo & Stevers 1924, neem tree (Azadirachta indica) or the chinaberry tree (Melia
Metzger & Grant 1932, Chopra et al. 1941, Bouquet 1950, azedarach).The primary active ingredient, azadirachtin, is a
Abrol & Chopra 1963, Reznik & Imbs 1965, Fenili & steroid-like triterpenoid. Neem is a broad-spectrum agent
Pegazzano 1974, Khare et al. 1974, Stratii 1976, Pakhomov & that mimics insect growth hormones, so it primarily kills
Potushanskii 1977, Rothschild et al. 1977, Rothschild & immature insects. Neem also serves as an antifeedant and
Fairbairn 1980, Prakash et al. 1987, Kashyap et al. 1992, Bajpai repellent. Poisoned larvae fail to complete the moulting proc-
& Sharma 1992, Jalees et al. 1993, Sharma et al. 1997), mites ess and die slowly (typically three to 14 days) but neem de-
(Reznik & Imbs 1965, Fenili & Pegazzano 1974, Surina & ters larvae from feeding almost immediately (unlike roten-
Stolbov 1981), bacteria (Ferenczy 1956, Ferenczy et al. 1958, one). Neem works best against caterpillars, but also controls
Schultz & Haffner 1959, Zelepukha 1960, Bel'tyukova 1962, immature whiteflies, leafminers, fungus gnats, mealybugs,
Radosevic et al. 1962, Zelepukha et al. 1963, Gal et al. 1969, leafhoppers, and some thrips and beetles. It is less effective
Veliky & Genest 1972, Veliky & Latta 1974, Farkas & against aphids and some grasshoppers. Neem is available
Andrassy 1976, Klingeren & Ham 1976, Fournier et al. 1978, as an EC from several manufactures at different concentra-
Braut-Boucher et al. 1985, Vijai et al. 1993, Krebs & Jaggi 1999), tions. Azatinn is the most concentrated (mix 4 ml per 4 1 or
fungi (Vysots'kyi 1962, Bram & Brachet 1976, Dahiya & Jain 3/4 tsp /gallon).
1977, Misra & Dixit 1979, Pandey 1982, Gupta & Singh 1983, Neem breaks down rapidly in standing water, so don't
McPartland 1984, Singh & Pathak 1984, Kaushal &Paul 1989, mix more than needed. Diluting water should be acidic (pH
Grewal1989, Upadhyaya & Gupta 1990, Krebs & Jaggi 1999), 3-7). Use a spreader/sticker. Spray when humidity is high
nematodes (Kir'yanova & Krall 1971, Haseeb et al. 1978, (morning or evening). Neem only persists four to eight days
Vijayalakshmi et al. 1979, Goswami & Vijayalakshmi 1986, on plant surfaces in bright sunlight. It persists longer, for
Grewal 1989, Mojumder et al. 1989, Kok et al. 1994, Mateeva several weeks, when plants absorb neem systemically, when
1995), protozoans (Nok et al. 1994), "worms" (Parkinson it is applied as a soil drench (Mordue & Blackwell 1993).Some
1640, Culpeper 1814, Pliny 1950), and weeds (Stupnicka- neem oil products leave a sticky residue on plants. Under
Rodzynkiewicz 1970, Srivastava & Das 1974, Muminovic laboratory conditions, normal doses of neem (<20 ppm aza-
1991). dirachtin) do not harm most beneficial insects ord due &
Many terpenoids in Cannabis inhibit acetylcholineste- Blackwell 1993). Neem does not harm foraging honeybees,
rase, including limonene, limonene oxide, acterpinene, y earthworms, fish or birds; in mammals it irritates eyes.
terpinene, terpinen-4-01, carvacrol, I- and d-carvone, 1,8-ci- Neem oil, a clarified hydrophobic extract of neem seeds
neole, p-cymene, fenchone, pulegone, and pulegone-1,2- (which does not contain azadirachtin), is marketed as
epoxide (McPartland & Pruitt 1999). These cholinergic ter- Neemguardm and TriactB. Neem oil has fungicidal, insecti-
penoids paralyse and kill insects the same way as malathion cidal, and miticidal activity. Locke et al. (1993) found that a
and other synthetic organophosphate pesticides, although foliar spray of 1%Neemguard protected plants against pow-
plant terpenoids are 4.4- to 17.1-fold less potent than dery mildews and rust fungi better than benomyl (but
malathion (Ryan & Byrne 1988). Research by Rothschild et benomyl worked better at eradicating established infections).
01. (1977) suggests that cannabinoids are insecticidal. Neem oil kills aphids and mites more effectively than regular
Cannabinoids are safe to mammals; the oral LD.j of THC in neem. It may also harm beneficial insects (Cherim 1998).
mice is >21,600 mg kg-' (Loewe 1946),safer thanneem. Work- Nicotine, derived from tobacco (Nicotiana tabaciim), first
ing with THC may, however, elicit an allergic contact served as an insect control in the 1690s. It is useful against
dermatitis (Watson et al. 1983). aphids, mites, whiteflies, young borers, thrips, young beetles
Castor, from seeds of Xicilziis comn~tinis,suppresses and bugs, and other soft-bodied insects. Nicotine works as a
nematode populations when mixed in the soil. Seed oil and stomach poison, contact poison, and by inhalation (no
leaf extracts have been used against a variety of insects surprise there). There are two forms -free nicotine alkaloid
(Jacobsen 1990). Persistence and phytoxicity unknown. and conjugated nicotine salts.
Creosote is distilled from wood tar or coal tar, useful as Nicotine alkaloid is volatile and serves as a fumigant. Years
a wound treatment against fungi. It is a benzene compound, ago, the New York Agricultural Society suggested placing a
quite carcinogenic. A synthetic derivative, dinitro-o-cresol, box over infested plants and smoking aphids to death with
is also used. Both are best avoided. a cup of burning tobacco. Today this can be accomplished
Hellebore, a hot water decoction of rhizomes from sev- with a "smoke generator" formulation. Tobacco dust is also
eral Veratriim species, acts as a stomach poison against many available (0.5% nicotine alkaloid). Hot water can extract the
insects. Available as a D. Hellebore loses potency quickly free alkaloid, but do not boil it. Parker (1913a) killed over
after exposure to air and sunlight; short residual. It is toxic 95% of hops aphids (Phorodon humiili) with two types of "to-
to livestock. bacco tea": 1) he soaked 11.3 kg of "tobacco waste" in 378 1
Horsetail, from Eqiiisettim hycmale, kills epiphytic fungi water (25 lbs/100 gallons); 2) he soaked 6.1 kg of commer-
and powdery mildews. Boil 40 g of dried leaves and stems cial blackleaf tobacco in 378 1 water (13.5 lbs/100 gallons).
in 4 l water (1.5 oz/gallon) for 20 minutes (Yepsen 1976). Free nicotine alkaloid is very unstable, has a very short half-
Cool, strain, and spray on plants. life, and is not very popular.
Limonene is a terpenoid distilled from citrus peel oil. It Nicotine stilphate (NS) is the most common conjugated
also occurs in Cannabis. Its mode of action is similar to pyre- salt. It is sold as a 40% aqueous solution. Parker (1913a)mixed
thrum (Thomson 1998). Limonene, particularly d-limonene, 130 ml of 40% NS in 378 1 water (4.4 oz/100 gallons), and
repels and paralyzes insects; this activity is synergized by killed 99.9% of hops aphids (Plrorodon htimiili). The potency
PBO. Available as a 37%EC, and sold in aerosol cans of NS increases when it is combined with an alkaline activa-
(DeMizea) and shampoos for ridding house pets of fleas and tor (e.g., soap or calcium caseinate). Parker (1913a) mixed
ticks. Limonene is volatile and evaporates rapidly from 1.8 kg soap in 378 1 water (4 lbs/100 gallons). The alkaline
treated surfaces. It harms honeybees and other beneficial activator reverts NS to a free nicotine alkaloid. The half-life
196 Hemp Diselases and Pests

of NS, without an alkaline activator, is 4.5 days. Phytotoxic- and other aphids. For small batches, he mixed 140 g chips
ity is rare. Nicotine kills honeybees if sprayed directly on per 4 1 (5.3 oz/gallon). Chips should be soaked for 24 hours
them. So spray late, after bees have retired for the evening. then boiled for two hours. Warning: the quality of chips may
Nicotine on plants repels honeybees before they are harmed. vary. Yepsen (1976) potentized quassia by adding larkspur
Nicotine is moderately toxic to fish and birds, and very toxic seeds. Mixing quassia with potassium soap also enhances
to mammals (LD5@ = 50-60 mg kg-I), so it is prohibited from its activity. Quassia supposedly spares ladybeetles and bees,
most Certified Organic farms. High dosages of nicotine cause and is nontoxic to mammals. It has substituted for hops in
tremors, convulsions, paralysis, then death. beer manufacturing (Meister 1998).
Oil from peanut, safflower, citrus, corn, soyabean or sun- Red squill, Urginea ntaritima, is an onion-like plant from
flower, kills small insects. Vegetable oil consists primarily of the Mediterranean region. Bulb extracts and dried powder
fatty acids and glycerides. See more information in the de- have been used for rodent control since the 13th century
scription of carbon-based pesticides, above. (Godfrey 1995). Red squill extract has an oral LDjo of 500
Pawpaw extracts, from the pawpaw blowtorch tree, mg kg-1 in rats; the purified active ingredient, scillirnside,
Asinzina triloba, have been applied as a soil drench to kill soil has an oral LDSoof 0.43 mg kg-' in female rats. It affects the
nematodes (Meister 1998). cardiovascular and central nervous systems, causing con-
Physcion, extracted from a European weed, giant vulsions and death after 24-48 hours. Some countries ban
knotweed (Reynoufria sachalinensis) is marketed as a foliar red squill as a cruel poison. The extract is very bitter; when
spray (Milsanaa) for protecting fruits and vegetables against administered to nontarget animals, red squill usually induces
the powdery mildew fungus Sphaerothecafiiliginia. Physcion vomiting (thus it is considered relatively safe).
works by inducing plant resistance via phytoalexin produc- Rotenone has been used since ancient times to kill fish.
tion; technically it is not a fungicide. Physcion takes one or The Chinese discovered rotenone's insecticidal qualities,
two days to induce resistance, weekly treatments may be which attracted Western attention around 1848.Rotenone is
needed. a resin extracted from Peruvian cube root (Lonchocaupusspe-
Polygodial, derived from Warblirgia and Polygonunz cies) or, less frequently these days, from Malaysian err is or
plants, works as a feeding deterrent against aphids (e.g., Tephrosia species. It was the most popular pesticide in the
Myzlis persicne), armyworms (Spodoptera species), and USA until DDT in the 1940s. Rotenone works best against
budworms (Heliothis species). A 0.1% solution of polygodial insects with chewing mouthparts, such as budworms, cut-
sprayed on plant surfaces kills less than 20% of these pests, worms and beetles, but it also works against thrips, aphids,
but it deters them from feeding on treated surfaces, so they and occasional mites. Formulations include a D, 1-59/oWP,
migrate elsewhere. Polygodial is a sesquiterpene dialdehyde and 1.1 lb/gal EC. Unfortunately, rotenone has a slow onset
and unstable in sunlight; its half-life on plant surfaces is short. of action-fast-feeding insects may consume up to 30 times
Pyrethrum, a contact insecticide, kills aphids and their lethal dose before rotenone affects them. Consider add-
whiteflies best, and also works against young bugs, beetles, ing pyrethrum to rotenone-fast-acting pyrethrum slows
and caterpillars. Frank (1988) killed spider mites with a py- insect feeding rates, allowing rotenone time to kill. Roten-
rethrum-PBO product, Holiday Fogger". Pyrethrum is a pow- one persists longer than most botanicals. It breaks down
der produced by grinding dried flowers of Chysanthenzlinz after three to ten days on plant surfaces in bright sunlight,
plants, predominantly C. cinerariifolium. The active ingredi- quicker in the presence of soaps or lime. Phytotoxicity is rare.
ents in pyrethrum are collectively called pyretkrins (e.g., Rotenone is highly toxic to honeybees, fish, and mammals
pyrethrins, cinerins, and jasmolins). Synthetic pyrethrins are (rotenone is more toxic to humans than malathion). Chronic
collectively called pyrethroids (e.g., permethrin a n d use causes skin irritation, and possibly liver or kidney dam-
cypermethrin). Pyrethrum originated in the Middle East and age. Some organic growers refuse to use rotenone; the Neth-
was introduced into the USA around 1870.It is now our most erlands banned its use in 1980.Rats given chronic, low doses
popular botanical pesticide. of rotenone develop a selective apoptosis of nigrostriatal
Formulations include pyrethrum powder (powdered dopaminergic neurons, a syndrome akin to Parkinson's dis-
flowers, 1% pyrethrins), pyrethrin dust (20-60%D, mixed ease (Friedrich 1999).
with clay or talc carriers), pyrethrin sprays (1-20%EC, 1- Ryania is extracted from roots and stems of Ryania
20°/0WP), and aerosol products. Many formulations contain speciosa (Jacobsen 1990). It is a slow-acting stomach poison,
the synergists PBO or MGK 264. Disadvantages include high used against European corn borers, hemp borers, other
cost and pyrethrum's rapid deterioration after exposure to caterpillars, and thrips. Ryania was "discovered" at Rutgers,
air (about one day). Sunlight accelerates its deterioration, so by researchers screening extracts of plants used by
apply in early evening or during cloudy weather. To slow Amazonian natives. It works better in hot weather, whereas
degradation, combine pyrethrum with Pheast", a lepidop- sabadilla works better in cool weather (Ellis & Bradley 1992).
teran feeding stimulant. Do not mix pyrethrum in alkaline Available as a D or WP. Ryania is relatively on
water; it is not compatible with Bordeaux, lime, and soaps. treated surfaces. The residual remains toxic much longer than
Pyrethrum exerts a rapid paralytic action and can "knock pyrethrum; it persists almost as long as rotenone. Phytotoxic-
down" flying insects, but they may recover. Some insect re- ity rare. Ryania is moderately toxic to mammals, fish, and birds.
sistance has appeared. Pyrethrum is not phytotoxic. It kills Sabadilla, an alkaloid like ryania, is extracted from
honeybees if sprayed directly on them (foraging bees who seeds of a New World lilly (Schoenocalilonofficinale) and Eu-
encounter previously-sprayed plants are repelled but not ropean white hellebore (Veratrunz album). It serves as a con-
killed). Pyrethrum is highly toxic to fish. Mammals safely tact and stomach insecticide against bugs, thrips, beetle
ingest pyrethrum (it is rapidly hydrolysed by stomach ac- grubs, caterpillars, and grasshoppers. Yepsen (1976)claimed
ids); pyrethrum is more toxic to humans when inhaled. Some sabadilla becomes more powerful with storage; heating it to
people develop skin allergies when handling it. 75-80°C for four hours also markedly activates it. Applied
Quassia is extracted from the wood of bitterwood as a D or F. Once dusted, sabadilla breaks down quickly in
(Qliassia aniara) and the lowly tree-of-heaven (Ailanthus sunlight (about two days). It is highly toxic to honeybees.
altissirna). Parker (1913a) chopped up 1.4 kg quassia chips The dust irritates mucous membranes and may cause vio-
and 1.8 kg soap into 378 1water to kill 96% of Phorodon hllmtlli lent sneezing.
 Chapter 11: Biorational Chemical Control 197

Strychnine, derived from seeds of Strychnos nux-uonzica, nAChrs. Spinosad works as a contact and stomach insecticide
is formulated as a 0.5-1%B rodenticide. It causes violent against caterpillars, as well as thrips, beetles, and leafminer
muscular spasms in birds and mammals, killing them within maggots. Spinosad is used on fruits, vegetables, and cotton.
30 minutes. Strychnine is not humane, and intensely poi- It degrades rapidly from UV light (persistence up to seven
sonous to humans. days), has no phytotoxicity, is safe to beneficial organisms
Wild buffalo gourd, roots of Cucirrbita foetidissima, is (except some adult wasp parasitoids), and exhibits low tox-
toxic to cucumber beetles. Grind roots into powder, suspend icity to mammals, fish, and birds.
in soapy water, and spray on plants. Tetracycline is an antibiotic produced by several Strep-
tomyces species, useful against bacteria and phytoplasmas.
Asemisynthetic derivative, oxytetracycline (Terramycin"),is
FERMENTATION PRODUCTS more popular for controlling plant diseases. The RE1 is 24
Natural metabolites oozed by fungi and actinomycetes hours.
have many uses. These are not botanicals, because the
organisms that produced them are not plants. The most
popular products of fermentation are antibiotics, used in SYNTHETIC ANALOGUES
human medicine and agriculture. These products should be Here we discuss synthetic chemicals that are consid-
used sparingly, because widespread antibiotic use increases ered biorational, because they closely mimic naturally-oc-
pressure upon bacterial populations to select for resistant curring compounds. They include synthetic botanicals,
mutants. Resistant genes may pass from plant bacteria to pheromones, and growth regulators.
human pathogens. This passage of genes is accomplished
by bacterial conjugation, plasmid exchange, or DNA scav- SYNTHETIC BOTANICALS
enging. Cinnamaldehyde (Cinnamite",Valero")is a new contact
Abamectin is a mix of auermectin Bla and auermectin Blb, insecticide and miticide, a synthetic oil of cinnamon
produced by an actinomycete, Streptomyces auernzitilis. Syn- (Cinnamonurn zeylanict~m).It is currently registered for use
thetic derivatives are called ivermectins, related products on fruits, vegetables, and hops, against aphids and mites.
include etrlairlectin and milbenzectin. Abamectin is very po- Cinnamaldehyde, however, is a broad spectrum agent capa-
tent against russet mites and spider mites on hops, and kills ble of killing most insects, including beneficials. It also kills
fire ants, leafminers, and nematodes. It can be diluted to con- powdery mildew. Cinnamaldehyde provides rapid knock-
centrations as low as 0.01 p p m for spraying on plants. down of spider mites and has a short half-life (beneficials
Abamectin is not truly systemic, but has translalninar actiu- can be reintroduced after 24 hours). Some phytotoxicity may
ity in some plants (especially in young leaves), meaning it is occur at the recommended 2% concentration; test on a small
absorbed from exterior surfaces to internal leaf parts. It is number of plants for potential damage.
available as a 0.15 lb/gal EC (mix 1 ml per 4 1 or 1/4 tea- Imidacloprid is a chlorinated derivative of nicotine. It
spoon/gallon); be sure to add a wetting agent. Abamectin activates nicotinic acetylcholine receptors (nAChRs), like
degrades readily in the environment (half life eight to 12 nicotine. But unlike nicotine, imidacloprid has greater affin-
hours when exposed to sunlight). It does not bioacc~mulate. ity for insect nAChRs than mammal nAChRs, s o
The RE1 is 24 hours. Repeat application may be needed in imidacloprid is safer. It acts as a contact insecticide, most
seven to ten days. Abamectin works poorly in cold weather effectively against insects with sucking mouthparts, such as
conditions. High concentrations are toxic to mammals, fish, aphids, whiteflies, and leafhoppers. It is also used against
and honeybees (nontoxic to birds). Abamectin may harm thrips, leaf beetles, and stem borers. Imidacloprid does not
immature biocontrol organisms. The translaminar absorp- harm spider mites. It is applied as a foliar spray, soil drench,
tion protects beneficial insects, while phytophagous insects or seed treatment; available as a 10%WP, 35%FC, and 1-5%G.
are exposed to the pesticide. Abamectin is restricted is some Imidacloprid is less toxic to mammals than nicotine (LD5, =
localities. Several products related to abamectin are now 2000 mg kg-I), and serves as a flea spray for pets. It can kill
sold-eman~ectin and milbemectin. predatory biocontrols, such as Chrysoperla, Coccinella, and
Griseofulvin, a fungicide produced by the fungus Peni- Orills species, as well as bees, but does not affect beneficial
cllliilrn griseofirlvilm, is used to treat human fungal infections mites and spiders (Elbert et al. 1998). Plants absorb it sys-
and may cause urticaria in sensitive individuals. Brian et al. temically and persistently, up to 70 days. When imidacloprid
(1951) eliminated Botrytis cinerea and Alternaria solani by is used against Phorodon htrnzuli in hops, a pre-harvest inter-
watering vegetables with a 100 mg 1-1solution of griseoful- val of 28 days is required. The RE1 is 24 hours. Imidacloprid
vin. It is systemic. is expensive and Bayer keeps increasing its price New nico-
~ t r e d t o m ~ c i nan, antibiotic secreted by Streptomyces tine derivatives are coming: acetamiprid and thiamethoxam.
griseus, kills bacteria (Psetldomonas and Xanthomonas species) Piperonyl butoxide (PBO) is derived from sesame
and some oomycetous fungi (Hewitt 1998 reported activity (Sesamt~nzindicum). PBO inhibits the microsomal detoxifica-
against Pse~ldoperonosporalu~rnuliin hops). Available as tion of many compounds, including insecticides and THC
streptomycin or streptomycin sulphate, as a 0.1-0.2%D, 8- (Gill &Jones1972).PBO usually serves as a synergist, mixed
62%WP.It is mixed 200 ppm and applied as a soil eradicant with insecticides to enhance their impact. PBO is present in
or foliar spray on tobacco and many other crops. It works most pyrethrum products, and is also mixed with rotenone,
systemically. The RE1 is 12 hours. Some phytotoxicity (chlo- ryania, sabadilla, and some oils. PBO may be carcinogenic.
rotic flecking) may occur-reduce this by spraying in slow- Synthetic pyrethroids serve as rapid-acting, broad-spec-
drying conditions, like at night. Streptomycin is nontoxic to trum, contact insecticides against aphids, whiteflies, bugs,
birds and mammals, and slightly toxic to fish. Applicators beetles, and caterpillars. Resistance has arisen in some aphids
may develop allergenic reactions. and spider mites; these pest populations explode after
Spinosad is produced u n d e r fermentation by pyrethroids are used, because pyrethroids kill off their preda-
Saccharopolyspora spinosa, an actinomycete. It activates tors (almost all biocontrol insects are killed by pyrethroids).
nicotinic acetylcholine receptors (nAChRs), b u t with At last count, 33 different pyrethroids were available in many
remarkable selectivity for insect nAChRs and not mammal formulations and concentrations. Pyrethroids are more toxic
198 Hemp Diseases and Pests

to mammals and more persistent in the environment than agents. A variety of pheromone traps have been designed
their natural cousins. synthetic pyrethroids are known to (Howse et al. 1998), a selection is shown in Fig 11.1.Delta
disrupt the endocrine system (Colborn et al. 1996). Many traps and wing traps utilize non-drying adhesives to hold
pyrethroids are mixed with piperonyl butoxide, a possible trapped insects; they are sufficient for most pests. For larger
carcinogen. pests, such as nocthrid moths and Japanese beetles, funnel
Pernzetkriti is commonly sold as a 0.2% aerosol, easy to traps or bucket traps are recommended. These have a one-
apply. Also available as a 37%EC, 25%WP, and 0.5%D. way entrance with no exit, and may be laced with poisons
~ e i m e t h r i nbreaks down quickly on sun-exposed plants (Lure N Kill" traps). Most pheromone lures should be re-
(half-life 4.6 days); in soil the half-life is longer (three to six placed every four to six weeks, depending on temperature.
weeks). The RE1 is 24 hours. To delay degradation, combine Traps should be checked daily and dead pests must be re-
permethrin with PheastB, a lepidopteran feeding stimulant. moved before their bodies saturate the trap.
Permethrin is rarely phytotoxic. It is extremely lethal to hon- In addition to monitoring pests, sex pheromones can
eybees if sprayed directly on them, so spray in the evening be applied for direct control of pests, by mass-trapping or
(bees find dried permethrin very irritating, so it repels them "confusing" pests. Scattering about 500 sex pheromone drops
from sprayed plants before they absorb a lethal dose). per ha will saturate the field in a "pheromone fog." Male
Permethrin is highly toxic to fish, nontoxic to birds, low tox- moths become confused and overwhelmed, leaving the fe-
icity to most mammals (very toxic to cats), irritating to the male's eggs unfertilized. Hemp researchers have used the
eyes and skin. A tingling of nerves in the fingers may arise "confusion" tactic against European corn borers, cutworm,
from prolonged occupational exposure. armyworms, budworms, and hemp moths (Nagy 1979).
Deltaniethrin is formulated w i t h polybutene as Small rubber septums, hollow-fibre "twistees," and other
ThripsticksE. The mix is sticky-paint it on sticks and place simple devices can be impregnated with pheomones and
sticks in soil between plants. Thrips hop on the Thripsticks, serve as "controlled release dispensers," lasting for several
stick, and die. It is also available as a 2.5 lb/gal EC, but not weeks.
in the USA. It is considered the most potent pyrethroid. As an added benefit, sex pheromones may also attract
Less common pyrethroids include allethrin, bifenthrin, biocontrol organisms to the field. Lewis et al. (1982) showed
cyfluthrin (10-20%WP, used on hops), cypermethrin (10- that parasitization of budworm eggs by Trichogramma wasps
20%EC, used on tobacco), esfetzvalerate (10-30%EC, used on doubled in fields treated with budworm pheromones, com-
hops), fenpropathrin, plzenothrin (sl~mithrin),resmetkrin, pared to untreated fields.
tefluthrin (a systemic pyrethroid), and tralometkritz (used on Aggregation pheromones attract pests to traps where
tobacco). Some are sold unrestricted in aerosol cans (e.g., they can be removed from the environment. Japanese beetle
Raida Flying Insect Killer), while others are restricted-use traps, for instance, contain Japonilure" (a pheromone bait,
pesticides (e.g., bifenthrin). Many of the older pyrethroids which attracts males) and geraniol (a food attractant, which
are less toxic and degrade quickly (e.g.,resmethrin), whereas predominantly attracts females). These chemicals lure the
the newer pyrethroids tend to be more persistent (e.g., pests into a cul-de-sac. The cul-de-sac contains malathion
permethrin). which kills them. This strategy is called the attract-annihilate
tactic (Foster & Harris 1997).
SEMIOCHEMICALS Alarm pheromones or dispersal agents such as P
Semiochemicals are chemical messengers which can be farnesene cause Myzus persicae and other aphids to drop to
used to disrupt pest behaviour or growth. They are also called the ground, where they may be eaten by predators (Howse
growth and reproduction regulators (GRRs). Synthetic et al. 1998).pfarnesene signals other pests to disperse across
semiochemicals mimic natural compounds produced by the surfaces of plants. Dispersal increases pest exposure to
pests. GRRs are called "third generation pesticides" by some pesticides or biocontrol organisms. But dispersal agents don't
researchers. Semiochemicals fall into two groups, always work as planned (Dombrowski et al. 1996). Interest-
pheromones and allelochemicals.
Pheromones are chemical signals emitted by animals,
and act on others of the same species, such as reproductive
pheromones, alarm pheromones, a n d aggregation
pheromones. Allelochemicals are emitted by animals and
plants, and act on other species, such as allomones (chemi-
cals which favour the emitter, including juvenoids,
antifeedants, repellents, and seed germination inhibitors),
and kairomones (chemicals which favour the receiver, such
as feeding stimulants).
Reproductive (sex) pheromones are emitted by insects
to attract the opposite sex for mating. In most cases, females
emit the pheromones and males follow. Night-flying insects,
such as nocturnal moths, make the greatest use of these odour
signals (Howse et al. 1998). Sex pheromones are scented al-
dehydes or esters, usually 14-16 carbon atoms long. Syn-
thetic sex pheromones for dozens of insects and mites are
now commercially available, and described in Chapter 4.
Sex pheromones are primarily used for pest monitor-
ing. When pheromones lure males into traps, this indicates
females are mating in the area and eggs are on their way. Figure 11.1: Examples of pheromone trap designs.
Traps monitor the timing and intensity of pest invasions. A. Tent or Diamond trap, B. Vertical sticky trap, C. Wing
Pheromones can lure males onto Tanglefoot@(a non-drying trap with spacers, D. Delta or Jackson trap, E. Funnel or
adhesive), or into traps containing pesticides or biocontrol Heliothis trap, F. Bucket trap (McPartland).
 Chapter 11: Biorational Chemical Control 199

ingly, the glandular trichomes of Cannabis produce P- is not effective against Homopteran pests. Flwcyclozuron and
farnesene (Nigam et al. 1965, Hood et al. 1973, Hood & Barry dinfentkiuron work against insects as well as spider mites;
1978, Ross & ElSohly 1996, Mediavilla & Steinemann 1997). they are applied to many crops outside the USA.
Unfortunately, Cannabis also produces pcaryophyllene, Teflubenztlron is used against European corn borers,
which inhibits the pheromone activity of pfarnesene (Pickett budworms, grasshoppers, whiteflies, and many others.
et al. 1992). Hexythiazox is a highly selective growth hormone that
Juvenoids or juvenile growth hormones are produced kills mite eggs; immature mites are rarely killed and adult
by insect larvae for purposes of moulting and growth. Plants mites are immune (eggs laid by treated adult females, how-
produce allelochemicals which mimic insect hormones. The ever, are not fertile). Hexythiazox only affects mites in the
neem tree, for instance, produces azadirachtin, which arrests mite family Tetranychidae (e.g., Cannabis pests Tetranychns
larval growth, prevent pests from maturing, and kills them and Eutetraizychus), so beneficial mites are not harmed, but
before they can reproduce. Aculops pests are not harmed, either. Hexythiazox and a re-
We now have synthetic juvenoids. Applying these lated compound, clofentezine, are used outside the USA on
semiochemicals to a crop results in a gradtlal reduction of fruits, vegetables, cotton, and other crops. Formulated as a
pest populations. Juvenoids work slowly. Most juvenoids 10-50%WP and 10 Ib/gal EC. Hexythiazox has a long re-
have little effect against adults, except to render adults ster- sidual on plants. Non-phytotoxic, honeybees and insect
ile. Juvenoids work well against insects with short life cy- predators are not affected, nontoxic to birds, slightly toxic to
cles and high reproductive rates, such as aphids and other mammals > 5000 mg kg-1).
Homopterans. When first introduced in 1967,juvenoids were Cowpea trypsin inhibitors (CpTI), interfere with the di-
considered "resistance proof." Unfortunately, insects have gestion of plant proteins by insects. A CpTI gene has been
developed resistance to their own hormone mimics (Gould bioengineered into other crops to control various insects, in-
1991). Some selective juvenoids exhibit great toxicity to tar- cluding Heliothis, Spodoptera, Diabrotica, and Triboliilm spe-
get species, and zero toxicity to other organisms. Other GRRs cies (Hilder et al. 1987). CpTIs have little effect against
interfere with insect immune systems, rendering pests de- phloem-feeding pests, such as leafhoppers, planthoppers,
fenceless against bacteria (Reuveni 1995). and some aphids. These sap sucking pests can be controlled
Methoprene works best against aphids, leafhoppers, cat- with a group of proteins called lectins.
erpillars, beetles, flies (including leafminers), and dog fleas. Lectins can bind the surface glycoproteins of two cells
Formulated as a 1-5 lb/gal EC and an aerosol. Methoprene together; they are also called agglutinins. Lectins bind up
has a half-life of two days in water, two weeks on tobacco insect gut epithelium and block nutrient uptake. The cDNA
leaves in sunlight, and persists longer on tobacco in storage. for a lectin from snowdrops, called Galaiztklls niualis aggluti-
Nontoxic to honeybees, birds, and mammals. Moderately nin (GNA) has been engineered into potatoes and other
toxic to fish. plants. Unfortunately, a diet of these transgenic potatoes also
Kinoprene specifically kills Homopterans (aphids and causes changes in rat gut mucosa (Ewen & Pusztai 1999),
whiteflies) and has little effect on most beneficial predators and GNA binds together human blood cells (Fenton et al.
and parasites (Godfrey 1995). It has a very short half-life on 1999).
foliage; it is currently limited to use on ornamentals. Feeding stimulants encourage pest feeding behaviour,
Hydroprene is currently limited to use against cock- so thev are kairomones. But kairomones can be used to trick
roaches, applied indoors. It is very effective against M y z u s pests, by mixing the feeding stimulants with pesticides or
persicae, but has not been registered for this application biocontrols (e.g., Bt or NPV), or adding them to attract-
(Godfrey 1995). annihilate traps. Feeding stimulants range from simple car-
Fetzoxycarb is a carbamate that acts as an insect growth bohydrates to complex triterpenoid compounds such as
regulator. It is used on fruit trees and grapes against aphids cucurbitacin. Pheasts, a lepidopteran feeding stimulant, is a
and scales, and has ovicidal activity against a Grayholita spe- mix of nutritional yeast, flour, vegetable oil, and
cies (Godfrey 1995). Regrettably, fenoxycarb harms the lar- polysaccharides, formatted as a wettable powder. It also
vae of lacewings, lady beetles, and Aphidoletes aphidimyza. slows the photodegradation of Bt and NPV, and masks the
Furthermore, pet use (to control fleas) has been curtailed repellency of synthetic pyrethroids.
due to possible carcinogenic effects on dogs. Pests themselves produce kairomones, which attract
Pyriproxyfen is a flea control for dogs, used agricultur- their natural enemies, such as parasitic wasps. Pest sex
ally against aphids, whiteflies, scales, and thrips on fruit and pheromones, mentioned above, act in this fashion. Plants also
vegetable crops. Unfortunately, pyriproxyfen may kill im- produce volatile chemicals which attract the natural enemies
mature Encarsin fornzosa biocontrols inside whitefly pupae; of their pests, described at length in Chapter 3 under the
but it does not harm adult E. forinosn wasps, or predators section on insects.
such as Orizis and Macroloplz~lsspecies. Feeding repellents and oviposition repellents are natu-
Buyyofezin is also used against whiteflies, mealybugs, rally produced by plants, and we now have synthetic ver-
and scales, on rice, vegetables, and fruit trees. Cyromazine sions. These are described earlier in this chapter.
controls dipterous leafminers in vegetable crops.
Dlf2ubenzuron is technically not a juvenoid but a chitin
inhibitor. It is used against armyworms but disrupts
PESTICIDE REGULATION
moulting in many insects. The pupal stages of parasitoids IN THE USA
within affected insects are not killed. Formulated as a Pesticides are regulated in the USA under the Federal
25%WP, 2-4%FC. Half-life is one week on plant surfaces. Insecticide, Fungicide and Rodenticide Act of 1988 (FIFRA
The RE1 is 24 hours. Diflubenzuron is highly toxic to crusta- '88), as amended by the Food Quality Protection Act of 1996.
ceans, nontoxic to birds and bees, and slightly toxic to mam- About 800 pesticidal active ingredients are made into 19,000
mals. Restricted-use. registered pesticide products. The EPA, USDA, FDA, and
Several new chitin inhibitors have appeared: Triflt~mi~ron OSHA enforce pesticide regulations. FIFRA '88 limits acces-
is used against caterpillars, beetles, and flies in Europe. sibility to certain pesticides. These "restricted-use" chemi-
C h l o ~ f l i ~ r n z ~ lworks
r o n well against caterpillars in Europe, it cals can only be purchased by persons certified by the state.
200 Hemp Diseases and Pests

Spraying any registered pesticide in a manner inconsistent

with its labelling is illegal. No restricted-use pesticides cited in
PESTICIDE SAFETY
this text are registered for use on Cannabis. Penalties for using Pesticide containers m u s t be labelled. Those
pesticides in a manner inconsistent with labelling range up emblazoned with "Danger/PoisonU are extremely toxic, cat-
to US$1000 per offence. egorized by the EPA as Class I poisons (see Table 11.2).Sim-
One of the mandates of FIFRA '88 is the reregistration ply tasting these chemicals can be lethal. A "Warning" label
of all pesticides. The EPA must retest pesticides that were marks pesticides of moderate toxicity (EPA Class 11); a tea-
registered years ago when requirements were less stringent spoon to a tablespoon will kill. Chemicals carrying the sig-
than they are today. Thus, some chemicals used today will nal word "Caution" (EPA Class I11 & IV) are also lethal, but
be banned tomorrow-so far the reregistration rejection rate at bigger doses. For a list of labels, see www.cdms.net.
has been 45% (Hembra 1993). The EPA has suspended sale
of DDT, BHC, mercuric and arsenic compounds, mirex, vi- W h e n zuorking zuith pesticides, follozv these common-sense pre-
nyl chloride, and has curtailed use of other pesticides, such cautions:
as dieldrin, aldrin, chlordane, endrin, and heptachlor. 1. Read the pesticide label and follow instructions.
Cynics claim the EPA's testing of synthetic chemicals is 2. Handle all used spraying/dusting equipment with as
biased-the EPA tests chemicals in cancer-prone B6C3F1 much caution as the pesticide itself.
mice. Captan, for instance, induced carcinomas in three of 3. Store pesticides and equipment in lockable storage away
47 mice fed more tlzan tlieir weight in pesticides (Abelson 1994). from children and animals, and away from water, food,
When the dose was reduced to haytheir weight in captan, no and feed.
females developed cancer, suggesting a threshold level is 4. Keep pesticides in original containers. Discard unlabelled
required for carcinoma development. Nevertheless, captan pesticide containers; do not guess at contents. Dispose
may be banned despite the fact that it is relatively nontoxic of pesticide wastes properly. Triple-rinse empty pesticide
to humans and decomposes quickly (it is rarely detected on containers before throwing them away or recycling them.
food or in ground water). Meanwhile, coffee remains unre- Do not reuse containers in the home.
stricted. Coffee contains caffeine, which is a neurotoxin 5. Prepare pesticides and spray equipment in a protected
(McPartland & Mitchell 1997). Coffee also contains caffeic area away from children and animals, where spills can
acid, a compound nzilclz nzore carcinogenic than captan, and the be cleaned up easily. Clean up spilled chemicals with
8 mg of caffeic acid in a single cup of coffee is 1000 times absorbent materials such as sawdust. Do not hose down
greater than the amount of captan an average American re- the area. Decontaminate most pesticides with household
ceives in food residues per year (Ames et al. 1995). bleach and hydrated lime.
Registration of pesticides for new crops requires an av- 6. Do not combine two pesticides unless you are certain
erage of ten years of expensive research, costing $50 million they are compatible (e.g., mixing the insecticide pyre-
or more (Abelson 1994). This price keeps many low-cost thrum with a lime/sulphur fungicide deactivates the
botanicals from being registered. Rotenone, for instance, may former). To avoid cross-contamination of pesticides, use
lose its registration because no rotenone suppliers can af- separate labelled sprayers for each class of pesticide (fun-
ford the cost of EPA reregistration. Pesticide manufacturers gicide, insecticide, herbicide, etc.).
will not get involved unless predicted sales exceed the cost 7. Do not work alone when handling dangerous pesticides,
of registration. For minor crops neglected by private indus- especially in enclosed areas like storage sheds, prepara-
try, the USDA established the Interregional Research Project tion areas, or glasshouses and growrooms.
No. 4 ("IR-4") Program. Submitting an IR-4 Pesticide Clear- 8. Always wear protective clothing, including long-sleeved
ance Request permits the experimental use of pesticides on shirts and long-legged pants made from tightly woven
unregistered minor crops. "Minor" is defined as any crop cotton or hemp fabric. Protect your scalp with a water-
grown on less than 121,400 ha (300,000 acres). In the USA, proof, brimmed hood or hat. Rubber or neoprene foot-
this definition includes kenaf and hops, two beneficiaries of wear should fit over (or replace) leather or fabric shoes.
the IR-4 program; hopefully in the near future it will also Wash or discard protective clothing immediately follow-
include hemp. The National IR-4 Program is headquartered ing use.
at Rutgers University, New Jersey. 9. Some pesticides require additional Personal Protective
Organic farmers restrict their use of pesticides to The Equipment (PPE): an apron, coveralls, or chemical-resist-
National List mandated by the Organic Foods Production Act ant suit (especially when pouring and mixing chemicals
of 1990. The National List approves the use of some syn- or cleaning pesticide equipment). Gloves should be made
thetic materials, such as synthesized versions of C02, bleach, of polyvinyl chloride or rubber (nitrile, neoprene, etc.),
insect pheromones, micronutrient fertilizers, fish emulsions unlined (lining fabric absorbs pesticides), and extend to
(which contain synthetic p H stabilizers), petroleum oils and mid-forearm. Protect your eyes with safety glasses, gog-
soaps, and plastic mulch and row covers. The List disapproves gles, or a full-face shield. Wear a well-fitting dust mask
the use of some natural substances, such as strychnine. or respirator. Take steps to avoid heat illness when wear-
Conventional pesticide regulations may change with ing respirators and spray suits.
implementation of the Food Quality Protection Act of 1996. 10. Never eat, drink, or smoke while working with pesti-
This new law gutted the Delaney "zero cancer risk" clause cides or in treated areas. Wash your hands before using
with a compromise defined as "reasonable certainty of no the toilet. Avoid inhaling pesticide fumes; if pesticides
harm." The law passed because it will consider, for the first drift nearby, get away. If eyes or skin become contami-
time, children's increased vulnerability to pesticides. Presi- nated, quickly flush with large volumes of water. Remove
dent Clinton presented it to the public, saying, "I call this contaminated clothing; handle clothing cautiously; wash
the Peace of Mind Act, because parents will know that the in hot water with maximum detergent. Wash work
foods children eat are safe." But two weeks later Congress clothes separate from the family laundry. Be sure that
killed funding for the Pesticide Data Program, the very pro- cleaners know clothes are contaminated and dangerous.
gram the EPA needs to set new pesticide standards for chil- 11. Know the symptoms of pesticide poisoning-irritated
dren (Colby 1996). skin/throat/eyes, nausea, vomiting, dizziness, headache,
 Chapter 11: Biorational Chemical Control 201

Table 11.5: Personal Protective Equipment (PPE) recommended for some Botanicals and Biocontrols.**

ACTIVE RECOMMENDED
INGREDIENT PPE

Bacillus species Gloves (latex is sufficient), dust mask, goggles, LSS*, LLP*

Copper Gloves (latex or rubber), dust mask, goggles for mixing powders or safety glasses for mixing
liquids, coveralls or LSS, LLP

Diatomacious earth Goggles, dust mask

Neem Safety glasses, dust mask, LSS, LLP

Oil, dormant or Safety glasses, mask

horticultural

Pyrethrins Gloves (latex or rubber), safety glasses, dust mask or respirator, coveralls or LSS, LLP

Rotenone Gloves (latex or rubber), goggles for powders, safety glasses for liquids, dust mask, LSS, LLP

Ryania Safety glasses, dust mask, LSS, LLP

Sabadilla Gloves (latex or rubber), safety glasses, dust mask, LSS, LLP

Soaps Gloves (latex or rubber), safety glasses, respirator, LSS, LLP

Sulphur Gloves (latex or rubber), goggles for powders or safety glasses for liquids, dust mask (if using
micro-sul formulation, use respirator with HEPA filter), coveralls or LSS, LLP

**Usethese recommendations only in the absence of specific label requirements. Chart adapted from John Berino.
*LSS=long-sleevedshirt, LLP=long-legged pants with shoes and socks.

trouble breathing, muscle pains, pinpoint pupils. Post 1. The WPS is label-driven. Requirements are pesticide-
telephone numbers of the local hospital and in the USA, specific and listed on labels. Both general-use and re-
the national Centre for Poison Control, 1-301-443-6260. stricted-use pesticides are covered by WPS '92. If you are
using a pesticide with labelling that refers to the Worker
THE WORKER PROTECTION STANDARD Protection Standard, you must comply with WPS '92.
Most of the common-sense precautions listed above Many home-use pesticides and biocontrols are exempt.
are now law. The EPArevised its Worker Protection Standard But we recommend using precautions even in the absence
in 1992 (WPS '92). Employers who hire agricultural workers of specific label requirements (see Table 11.5).
must meet new federal guidelines for pesticide safety. (Some 2. All workers potentially exposed to pesticides must be
states have additional requirements.) The provisions in WPS trained in pesticide safety-not just pesticide applicators.
'92 are complicated. Employers should contact their regional The EPA provides a checklist of required topics, includ-
EPA office or a state agricultural extension agent for full in- ing the precautions described above.
formation regarding compliance. In summary: 3. Before spraying, a fact sheet must be posted that describes
the pesticide (its EPAnumber and active ingredients), the
area treated, date of application, and the RE1 (restricted-
entry interval or re-entry interval). In general, pesticides
Table 11.6: Methods of applying pesticides to soil. classified as Class I poisons (Table 11. 2) have an RE1 of
48 hours, the Class 11s have an RE1 of 24 hours, Class 111s
METHOD DESCRIPTION and IVs have an RE1 of 12 hours. Fact sheets and other
pesticide information must be displayed at a central lo-
Broadcast Pesticide broadcasted on the surface cation. Some pesticides also require "Keep Out" signs be
and mixed into soil before sowing seed posted during the REI.
4. Information on pesticide safety and emergency assistance
Furrow Pesticide selectively poured into soil must also be displayed at a central location.
behind plough or drill, before sowing seed 5. Establish decontamination stations within 400 m (1/4
mile) of all agricultural workers. These should be mo-
Root zone Pesticide mixed into soil of intended root bile, so field workers can clean up before meals and at
zone, before transplanting seedlings the end of the workday. Supply water for routine and
emergency washing, plenty of soap, and paper towels.
Irrigation Pesticide added to trickle irrigation system 6. Workers applying pesticides require additional training,
decontamination supplies, and personal protective equip-
Seed coating Seeds coated with pesticide using an ment (PPE). PPE must be clean and in operating condi-
adhesive tion, worn and used correctly, and washed or replaced as
needed.
202 Hemp Diseases and Pests

RE1 posters and warning signs, pesticide fact sheets, of the compounds he tested are no longer available. Two
worker training manuals, compliance record books, PPE effective compounds still in use are copper oxide, 12 g per
equipment, and decontamination equipment can be pur- kg seeds (9 ounces per bushel), and thiram (1.4-2.8 g kg-' or
chased at farm supply stores. For one-stop shopping, con- 1-2 oz/bushel).
tact Gempler's for their master catalogue (1-800-382-8473, The trouble with dust treatment is poor adhesion to
web site http:/ /www.gemplers.com). seeds, resulting in the environmental and health hazards of
loose pesticide dust in seed bags. Spraying seeds with a fast-
drying liquid pesticide results in better adherence. But the
PESTICIDE APPLICATION liquid may adhere irregularly, with too little on some seeds
Pesticides can be applied directly on plants, or applied and heavy phytotoxic loads on other seeds. "Film coating" is a
near plants. Pesticides are applied on plants as foliar treat- technology that provides good adherence and uniform coating.
ments, seed treatments, or onto roots as soil drenches. Off- A second approach to seed treatment is seed immer-
plant applications include pest baits, pest barriers, and pre- sion-steeping seeds in liquids or fumigant gases for vari-
season soil treatments. Off-plant applications are safest for ous periods of time, at ambient or elevated temperatures.
Cannabis consumers, but are still hard on the environment. Many hemp researchers have utilized this method. Soaking
seeds in plain hot water eradicates viruses (see Chapter 9,
BAITS, BARRIERS & SOIL TREATMENTS method 11).To eradicate bacteria, Kotev & Georgieva (1969)
Most baits are sold ready-to-use. Examples include soaked hemp seeds in 0.3% formalin. Booth (1971) soaked
pheromone baits for moths, food baits for cutworms, Japa- seeds in 0.1% formalin to eliminate fusaria. But for how long?
nese beetle traps, and rodent baits. Tanglefoot@is a popular Chinese growers soak seeds in formalin for one hour.
barrier, useful against crawling insects. Soil treatments can Soaking seeds in 10% household bleach for one minute
be applied by any method in Table 11.6. Furrow, root zone, kills many moulds. After immersion in bleach, rinse seeds
and seed coating methods are most commonly used. in water at least three times before drying or planting. Mishra
(1987)tested seven fungicides on Cannabis seeds-benomyl,
SEED TREATMENT carboxin, thiram, captan, zineb, and PCNB. Benomyl and
Treating seeds with pesticides may be the best way to carboxin were mixed at a rate of 100 mg active ingredient
control some hemp diseases (Patschke et nl. 1997). Planting (a.i.)per litre water, and the other fungicides were mixed at
pesticide-coated seeds is much easier on the environment 1000 mg a.i. per litre. Mishra soaked seeds for 15-20 min-
than applying soil drenches. Pesticides work at different lev- utes. In contrast, Maude (1990) soaked seeds in 0.2% thiram
els within seeds. Some only kill pests and pathogens on the at 25-30°C for six to 24 hours. Obviously, more research needs
surface of seeds. Other pesticides penetrate seeds and elimi- to be done. Ferri (1961b) soaked hemp seeds in 0.2% sul-
nate deep-seated pathogens. Some deeply penetrating sys- phuric acid for up to 30 minutes, or in 90% isopropyl alco-
tenzic pesticides persist in young seedlings, protecting them hol for ten minutes, apparently without seed damage.
from pests and pathogens lurking in soil or air. Maude (1996) Kryachko et al. (1965) killed European corn borers
noted that metalaxyl was detected for up to 20 days in the overwintering in stored seeds by fumigating with methyl
tissues of pea seedlings grown from treated seeds; benomyl bromide. Another popular seed fumigant was chloropicrin,
lasted up to four weeks in lightly-dosed seeds and up to nine but Migge (1921) noted chloropicrin reduces hemp seed ger-
weeks in heavily-dosed seeds; carboxin only lasted a few mination by at least 30%.
days before oxidizing into nontoxic sulphoxide. Pesticides
applied to seeds are long gone by harvest time. FOLIAR TREATMENT
Seeds may be coated with adhesive dusts, sprayed with Pesticides may be applied directly on plants as dusts
slurries, or immersed in solutions (see Table 11.1). The ma- or sprays. Be sure to treat the undersides of leaves, where
chinery used for coating seeds is reviewed and illustrated many foliar pests congregate.
by Maude (1996).Seeds are dusted in drum mixers or auger Dusts adhere firmly to plant surfaces, particularly if
mixers. Mixers can be modified to apply liquid formulations electrostatically charged or applied to dew-covered plants.
(WP and FC). Improved machinery for spraying seeds with Dusts are more susceptible to zuind drift than are sprays. Do
pesticides include perforated drztms (drum mixers with tiny not dust when wind speed exceeds 5 mph. (Sprays tolerate
holes which draw warmed air to speed drying),jztidized beds breezes up to 12 mph.)
(seeds are tossed in the air by vertical columns of forced hot Sprays are easier to apply to undersides of leaves than
air while being sprayed with fluids), spinning disk techniques dusts, but spray droplets often roll off plant surfaces, espe-
(fluid is poured onto a spinning disk which breaks the fluid cially off plants covered with trichomes such as Cannabis.
into fin; droplets by centrifugal force-seeds fall past the Spray adjuvants must be added to improve the effective-
disc through a peripheral curtain of pesticide spray), and ness of spray applications. There are many types of spray
film-coating (applying pesticides mixed with special binders adjuvants (over 150 adjuvants are registered in 17 catego-
and adhesives-often polysaccharides or synthetic poly- ries for use in Washington state). Here are a few:
mers-to create a durable film coating). Spreaders (i.e.,zuettingagents)keep sprays from bead-
Toole et al. (1960) reported good viability when seeds ing up and rolling off leaves. They reduce the surface ten-
were dusted with calcium chloride and stored for 14 years. sion of liquids, so the sprayed droplets spread out and hold
Maude (1996) mixed 1.5-5 g of thiram (a fungicide dust) per onto sprayed surfaces. High-rounded drops on leaf surfaces
kg seeds. In comparison, old mercurial dusts like calomel indicate the need for a spreader. Flat drops that slide off
and Ceresan required 250-500 g per kg seeds (Maude 1996). leaves indicate too much spreader. There are nonionic, ani-
Many hemp researchers recommended mercurial dusts to onic, and cationic spreaders. Nonionics are the most common.
control damping-off fungi (Flachs 1936, Robinson 1943a, They do not ionize in water, so they do not react with most
Wilsie & Reddy 1946, Andren 1947, Ferri 1961b, Kotev & pesticides (but they tend to remain as residues on plant sur-
Georgieva 1969). Robinson (1943a) tested the germination faces). Two common nonionic spreaders are alkyl-aryl-poly-
rate of hemp seeds treated with eight different pesticides, at oxy-ethylenate (AAPOE) and alcohol-poly-oxy-ethylenate
various concentrations. He presented copio~tsdata, but most (APOE).Triton AG-44M" is a popular nonionic spreader. Mix
 Chapter 11: Biorational Chemical Control 203

0.25 ml of 50% Triton solution per 50 ml pesticide solution. Spray equipment:

About 10% of spreaders fall into the anionic category; they 1. Aerosol can: prepackaged, very convenient, but high cost.
are ionized with a strong negative charge. Examples include 2. Trigger-pump sprayer: inexpensive, reusable, Windex-
fatty acids and linear alkyl sulphonate (LAS). Cationic spread- type hand sprayers work best with water-soluble pesti-
ers (ionized with a strong positive charge) are rarely used. cides for small problems. They are too small and tiring
Stickers provide an adhesive effect after the spray has for large p r d e n s .
dried on the plant. Otherwise, the pesticide would quickly 3. Slide-pump sprayer: similar to the trigger-pump sprayer,
wash off from heavy dew, irrigation, or rain, as well as from but generates pressure via a telescoping plunger. More
wind erosion and leaf abrasion. Stickers work by three applicable to larger gardens. Usually limited to 1 noz-
mechanisms: 1)increasing adhesion, 2) slowing evaporation, zle, and most sprayers only discharge during half of the
and 3) providing a waterproof coating. Some stickers are pump cycle. They are leaky and often contaminate the
also spreaders, such as AAPOE, LAS, fatty acids, and some applicator.
Triton formulations. These spreader-stickers utilize the first 4. Compression sprayer: a manual pump compresses air
two mechanisms. Waterproofing stickers include latex, above liquid in a tank. Air pressure forces the liquid out
polyethylene, resins, and menthene polymers. through openings at the bottom of the tank to a hand-
Extender adjuvants (also called stabilizing agents) held nozzle. The tank can be carried in the other hand or
protect spray residues against UV radiation and heat, which in a backpack. Pressure slowly discharges as pesticides
degrade many pesticides. Ennllsifiableoil adjuvants enhance are applied, requiring frequent repumping. These
the penetration of sprays into waxy plant surfaces. They are sprayers clog easily, and require frequent cleaning and
also called plant penetrants or translocntors. Acfiuator is a maintenance to prevent corrosion of working parts.
general term indicating any adjuvant that increases the 5. Power compressed-air sprayer: a gas or battery-powered
effectiveness of a pesticide; all of the aforementioned engine pumps air pressure into a tank, and the rest works
adjuvants are activators. like a compression sprayer. Usually mounted on a trac-
Buffers include acidijying adjuvants and softening tor. Higher maintenance costs.
agents. Many pesticides require mixing in water that is 6. Hydraulic sprayer: engine drives a hydraulic pump that
slightly acidic (optimal p H = 6.0) and soft. The most common draws liquid up from a tank and forces it under pres-
acidifying adjuvant is phosphoric acid. Muriatic acid and sure through multiple discharge tubes (spray booms) and
vinegar may not be suitable. Hard water contains metal ions out any number of nozzles. Usually mounted on a trac-
(Ca, Fe, Mg, etc.) which precipitate many pesticides tor-xpensive maintenance. Nozzles come as cones, flat
(especially fatty acids and soaps) into a scum, curd, or "ring fans, whirl chambers, and many other configurations.
around the tub." Water hardness is measured in ppm or Dz~stsand granz~les:
grains; if water hardness exceeds 300 ppm or 18 grains, 7. Bulb or bag duster: squeeze by hand to propel dust onto
softening may be required. Softening agents include Spray- plants. Simple, easy, and many are reusable.
Aide%,BlendexE, and Triton AG-44Ma. 8. Hand-crank dust or granule applicator: dust or granules
Defoamer adjuvants keep foam from forming in spray are stored in a hopper, beneath which spins a fan, pow-
tanks. The most common defoamer is a silicone/carbon poly- ered by a hand-crank. Air currents or the fan itself dis-
mer called dimethylpolysiloxane. Drift retardants keep sprays charge the granules. Hole size at the bottom of the hop-
from evaporating into ultrafine droplets that float away. Cotn- per regulates application rate, and fan speed controls the
patibility agents facilitate the mixing of pesticides to keep them range of discharge. The granules often contaminate the
from separating or curdling in spray tanks. Feeding attractants applicator.
andfecding stinullants encourage pests to eat pesticides (e.g., 9. Power duster: similar to the above, but an engine gener-
Coax3, Gustol'a).Their ingredients consist of sugars, starches, ates compressed air to blow dust. usualiy tractor-
seed oils, and proteins. Some attractants/stimulants use mounted, generating considerable drift and collateral
extracts of the crop to which they are applied (Hunter-Fujita contamination (Fig 11.2).
et n l . 1998). 10. Power granule applicator: like the power duster, but
spreads granules, much less drift.
FOLIAR APPLICATION EQUIPMENT
Many pesticide application devices are sold at
hardware stores and garden centres. Or contact Gempler's
for their master catalogue (1-800-382-8473,web site http://
www.gemp1ers.com). Selecting equipment adapted to your
individual needs makes the choice less overwhelming.
Liquid pesticides can be atomized into different drop-
let sizes. The smaller the droplet the better the coverage, and
smaller droplets are more toxic to insects (Thacker et al. 1995).
But smaller droplets are more susceptible to drift. Fogs and
smokes are the smallest, with a droplet size of <5 pm diam-
eter. Fogs and smokes almost act as fumigants-they cover
all plant surfaces (including the undersides of leaves). Fogs
are produced by ultrasonic or thermal fogging machines,
and smokes are generated by combustion. Aerosol droplets
(5-50 pm) are produced by blasting pressurized air over a
liquid to atomize the spray. Mists (50-100 pm) also require
high pressure. Fine sprays (100-400 pm) and coarse sprays
(A00 pm) can be generated at low pressures.

Here is a partial list of foliar application equipment, with Figure 11.2: Tractor-drawn duster used in the Ukraine
comments: (from Lesik 1958).
204 Hemp Diseases and Pests

Many soil pesticides are applied per ha or acre. To con- (18 inches), only one hollow-cone spray nozzle is required
vert to square-foot gardening, refer to conversion factors per row. Spraying rows 90 cm apart (36 inches) at a pressure
provided in the front of this book. Foliar pesticides need not of 4.2 kg cm-2 (60 pounds/inch2), while moving 6 km h-1 (4
be measured in gallons per ha, but simply applied until "run miles/hour) will use 65 1ha-1 (7 gallons/acre). For tall plants,
off." For calibration and use of tractor-drawn power equip- use three hollow cone nozzles per row, same pressure, same
ment, refer to Pimentel et al. (1991) or Matthews (1992) for tractor speed, and apply 196 1 ha-1 (21 gallons/acre). Always
information concerning droplet size, nozzle types, pump position the centre nozzle 15-30 cm (6-12 inches) above plant
capacity and other aspects of application technology. height. The two side nozzles should be positioned on either
Here is an example of calibration: Using tractor-drawn side of the row, low to the plants, with the nozzles angled
power sprayers, the amount of insecticide applied per ha (or upward-so the sprays hit pests that congregate on under-
acre) depends on age of the crop. For plants under 45 cm tall sides of leaves.
 OM!/Golden Rule to keep out of trouble: never do two stupid things at the same time. "
-Frank LeCase

Appendix 1: Synthetic Chemicals

According to Metcalf et al. (1962), synthetic pesticides were structures. The major groups include chlorinated hydrocar-
first used in 1892.They run a gamut from analogues of natu- bons, organophosphates, carbamates, dipyridilium herbi-
rally-occurring pyrethroids to mutant DDT. Much of our cides, and the most popular group, "miscellaneous." Syn-
information on pesticide toxicity was gleaned from ExToxNet thetic pyrethroids and synthetic pheromones are considered
(http:/ /ace.ace.orst.edu/info/extoxnet/ghindex.html), a co- "biorational" and treated in Chapter 11. A selection of syn-
operative project sponsored by Oregon State University and thetic pesticides is provided in ~ p p e n d i xTable 1.For more
several other agricultural universities in the USA. For a da- information on pesticides, see the series by W.T. Thomson or
tabase of current pesticide labels with full text, updated on the annual buyer's guide edited by R.T. Meister, Farm Chen~i-
a daily basis, see http://www.cdms.net. cals Handbook.
No pesticides have been labelled in the USA for use on
cannabis. Using pesticides in a manner inconsistent with their
labelling brings penalties, u p to US$1000 per offence. CHLORINATED
European hemp researchers have tested many synthetic
chemicals on fibre crops. Synthetic chemicals, however,
HYDROCARBONS
Chlorinated hydrocarbons are also called
should not be used o n crops destined for h u m a n
organochlorines. Many are relatively safe to mammals and
consumption, such as seed oil or medical marijuana. We say
rarely cause acute poisoning in humans (the antidote is
this knowing authors of marijuana books recommended
atropine). But their long residual (i.e., resistance to
using chemicals in the past. For instance, Rosenthal (1998)
degradation), initially a desired feature, makes them
recommended spraying marijuana with a tree nursery
environmentally dangerous. Some have been banned, such
product, Wilt-Prufa, which suffocates spider mites. Wilt-
as DDT, chlordane, and dieldrin. Spraying hemp fibres with
Pruf's generic name is polyvinyl chloride, the same as Saran
pentachlorophenol (PCP) is now illegal (Bocsa & Karus 1997).
Wrap. Not a desirable product to smoke!
In 1993 the American Public Health Association urged a
Chemical toxicity depends onset and setting. Many syn-
phaseout of all chlorinated organic compounds.
thetic chemicals are systemic. Systemics are particularly dan-
Chloropicrin (CC13N02)was developed as tear gas but
gerous on plants destined for human consumption. The
found use as an agricultural fumigant in 1919, manufactured
poisons remain in plant tissues without breaking down. For
in the USA by Great Lakes Chemical Corp. Soil fumigation
instance aldicarb, a systemic pesticide used to treat spider
with chloropicrin is expensive, technically difficult, and
mites on hops, produces a peak concentration in hops flowers
generally nasty-but it eliminates "thick-skinned" sclerotia-
60 days after application (Duke 1985).
producing fungi such as Sclerotillni rolfsii. It provides better
Many new pesticides are desirable because they are very
control of soil fungi, oomycetes, and bacteria than methyl
selectiz~e.They kill pests while sparing beneficial organisms
bromide, whereas methyl bromide is better against
and other innocent bystanders. Many selective pesticides
nematodes, insects, a n d weed seeds (Agrios 1997).
described in this chapter are used in the USA on hops and
Chloropicrin reduces Cannabis seed germination by at least
tobacco: Hops are related to Cannabis (so phytotoxicity data
30% (Miege 1921). Restricted-use.
regarding hops should be applicable to hemp), and tobacco
Dicofol is a miticide used against Tetranychus urticae, T
pesticides have been tested for their by-products of
cinnabarinl~s,and Brevipalpils obovatl~son hops, although
combustion.
many mite populations are now resistant. Dicofol is mite-
Placing synthetics in baited traps is a relatively safe ap-
specific-it spares beneficial insects (but kills most predatious
plication of pesticides. Baited traps (such as Japanese beetle
mites) and is nearly nontoxic to humans. Frank (1988)
traps) keep the chemicals off plants, and keeps chemicals
considered dicofol harmful and did not recommend it; in
containerized for effective disposal. Using synthetics for seed
1986 dicofol supplies were contaminated by DDT. Made by
tveatilrents (coating seeds with pesticides before they are
Rohm & Hass, and is available as a D, 35%WP, and 18.5-
planted) is also relatively safe-little pesticide is introduced
42%EC. Its residual on plants is two weeks, with a soil half-
into the environment, and the pesticide is long gone by the
life of 60 days. The RE1 is 12 hours. Dicofol is not toxic to
time crops are harvested (see discussion in Chapter 12).
plants and honeybees, slightly toxic to birds, and highly toxic
Pesticides come in a variety of formulations (see Table
to fish. It is carcinogenic to male (not female) mice, and may
11.1), including aerosols (abbreviated A), baits (B), dusts (D),
disrupt the endocrine system (Colborn et al. 1996).Strangely,
errllllsifiable coilceiztrates (EC), flowable conceiztrates (FC),
Wu et al. (1978) showed that Papaver poppies sprayed with
filrnigants (F), granules (G), slow release (SR), soluble powders
dicofol produced higher concentrations of opiates; all other
(SP), and wettable poiuders (WP). The amount of active
pesticides caused decreases in opiate production.
ingredient (a.i.) in solid formulations is presented as a
Pentachloronitrobenzene (PCNB), a fungicide, has been
percentage of weight. For example, a 50%WP contains 50%
used as a seed treatment or soil drench, applied to Cannabis
a.i. by weight (the remaining 50% consists of "inactive
at planting time to control hemp canker and southern blight
ingredients"). Liquid formulations are expressed as either a
(Ashok 1995). Mishra (1987) did not find PCNB helpful
percentage of weight or as the number of pounds a.i. per
against Rhizoctonia disease. Made by Olin and Uniroyal, and
gallon of product. For example, 4 lb/gal EC contains 4
is available as a 10%G, 2 lb/gal EC, and 35-75%WP. PCNB is
pounds of a.i. per gallon, which equals about 45%EC.
persistent in soil (five to ten months). It is slightly phytotoxic,
Synthetic pesticides are classified by their chemical
205
206 Hemp Diseases and Pests

Appendix Table 1: Some synthetic chemicals.

USA TRADE NAME(S) EPA

GENERIC
NAME OR 03 CHEMICAL
TYPE ACUTE TO XI CITY^

acephate Orthene, Ortho-12420 organophosphate II

aldicarb Temik carbamate I
4-aminopyridine Avitrol 200 miscellaneous I
benomyl Benlate, Bavistin carbamate IV
carbaryl Sevin carbamate Ill
carboxin Vitavax amide Ill
chloropicrin Larvacide chlorinated hydrocarbon II
chlorothalonil Daconil2787, Bravo organophosphate II
chlorpyrifos Dursban, Raid Hornet organophosphate II
diazinon Spectracide organophosphate II
dicofol Kelthane chlorinated hydrocarbon IV
dichlorvos (DDVP) No-Pest Strip, Vapona organophosphate II
dienochlor Pentac organophosphate II I
dimethoate Dexol, Cygon, Dimate organophosphate II
etridiazole TruBan, Terrazole carbamate II
fenbutatin oxide Vendex, Torque miscellaneous Ill
ferbam Carbamate carbamate IV
fosetyl-Al Aliette organophosphate IV
iprodione Chipco, Roval amide IV
malathion Malathion organophosphate Ill
maneb Dithane M-22 carbamate Ill
mancozeb Dithane M-45, Manzate 200 carbamate Ill
metalaxyl Ridomil, Subdue miscellaneous Ill
metaldehyde Slugit, Deadline, Metason miscellaneous Ill
metam-sodium Vapam carbamate Ill
methomyl Kipsin, Lannate carbamate Ill
methyl bromide Brom-0-Gas, Ozoneholer miscellaneous I
parathion, ethyl Parathion, Thiophos organophosphate I
parathion, methyl Metaphos, Penncap-M organophosphate I
pentachloronitro- benzene PCNB, Brassicol, Terraclor chlorinated hydrocarbon Ill
pirimicarb Pirimor carbamate II
propargite Comite, Ornamite chlorinated hydrocarbon I
pyridaben Sanmite, Dinomite miscellaneous Ill
sodium fluoroacetate Compound-1080 miscellaneous I
thiram Arason, Tersan carbamate Ill
tolclofos-methyl Rizolex, Risolex, Basilex organophosphate Ill
triadimefon Bayleton, Fung-Away miscellaneous Ill
triforine Funginex miscellaneous I
vinclozolin Ronilan amide IV
warfarin Coumadin, d-Con miscellaneous II
zineb Dithane 2-78, Parzate carbamate Ill
for a description of EPA acute toxicity, see Table 11.2.
 Appendix I : Synthetic Chemicals 207

slightly toxic to honeybees and birds, and highly toxic to nearly as persistent on plants. The RE1 is 48 hours. Nontoxic
fish. Restricted-use. Kotev & Georgieva (1969) used a related to plants and bees, somewhat toxic to birds and mammals
compound, Germisan@(hexachlorocyclohexane), to disinfect (especially eye and skin irritation), and highly toxic to fish.
hemp seeds of Psei~donzonasinfections. These compounds Chlorpyrifos has agricultural uses against hemp borers,
may disrupt the endocrine system (Colborn et al. 1996). cutworms, flea beetles, termites, flies, and even nematodes
Propargite is a miticide useful against the same pests i n field, fruit, hops, a n d vegetable crops. Made by
as dicofol (see above), as well as mites resistant to dicofol. It DowElanco, and is available as a 15-50%WP, 1-4 lb/gal EC,
is used on hops and many fruits and vegetables, and works and 30%FC. Short residual on foliage (ten to 14 days), but
best when the daily temperature is above 21°C. Kac (1976) persists longer in soil (half-life of 60-120 days). The RE1 is 24
sprayed it in Slovenian hemp fields. Propargite is specific; it hours. Chlorpyrifos can be phytotoxic. It was used safely on
does not affect insects (beneficial or otherwise), and spares hemp by Sandru (1976) against Grapholita delineana. It is
predatory mites (Helle & Sabelis 1985). Made by Uniroyal extremely poisonous to honeybees, very toxic to fish and
as a 30%WP and 6 lb/gal EC, currently suspended in the birds, and moderately toxic to humans. General and
USA and Canada pending results of additional research. restricted uses.
Propargite is moderately persistent in soil. It is slightly Diazinon is an insecticide first marketed in 1953. Frank
phytotoxic (especially to young plants), very toxic to fish, (1988) considered it the best control of scale insects, flea
slightly toxic to birds, slightly toxic to mammals by ingestion beetles, and whiteflies; he also used it against aphids,
and harmful to eyes. mealybugs, spider mites, leafhoppers, and caterpillars. It
works very well against hemp borers (Sandru 1975, Peteanu
1980) and leafminers. Made by Novartis (formerly Ciba-
ORGANOPHOSPHATES Geigy), and is available as a 5%D, 5-14%G, and 25%EC, and
The organophosphates, like chlorinated hydrocarbons, 40-50%WP. Residual activity one or two weeks on plants and
act as contact or stomach poisons. But unlike chlorinated two months in soil. The RE1 is 24 hours. Frank & Rosenthal
hydrocarbons, organophosphates degrade quickly in the (1978) recommended a safety period of 35 days between
environment. Lucas (1975) monitored the degradation of application of diazinon and harvesting of treated plants.
organophosphates on tobacco plants: Ten days after spraying Phytotoxicity is rare. Honeybees are poisoned. Birds are quite
parathion, diazinon, and malathion, only malathion residues susceptible to diazinon poisoning (especially the G formula-
exceeded 3 ppm. The malathion was gone after another ten tion, which they mistake for food).
days. On the dark side, organophosphates are far more Dichlorvos (DDVP) is an insecticide with contact and
poisonous to mammals than chlorinated hydrocarbons. vapour action (i.e., poison gas), used since the 1950s.
Parathion, a common insecticide, is half-strength Sarin, a Physicians prescribe dichlorvos for the treatment of
chemical warfare nerve gas. The antidote for both is atropine schistosomiasis. Wang et al. (1987)killed hemp sawfly larvae
or pralidoxime chloride. Organophosphates also cause with a dichlorvos spray Marijuana growers eliminated spider
delayed neurobehavioural effects. Fryday et al. (1994) ex- mites and thrips by hanging No-Pest@resin strips in their
posed house s p a r r o w s to chronic, low levels of growrooms (Alexander 1982,1988b). Made by Shell, and is
organophosphates. The sparrows' feeding behaviour became available as a 1-4 lb/gal EC, 10%A, 0.5%B and 20%
clumsy, they dropped more Catznabis seeds than unexposed impregnated resin strips. The EPA estimates that resin strips
sparrows, and they lost weight. pose a 1-in-50 cancer risk if used over a lifetime (Journal of
Conversely, low levels of organosphosphates are the American Medical Associati071 260:965).
prescribed for the treatment of Alzheimer's disease (e.g., Dienochlor, a selective miticide, was introduced in the
tacrine, donepezil). Metrifonate, which is metabolized into 1960s but Tetranychus tirticae remains uniquely susceptible
dichlorvos (described below), is being tested for the to it. Treated mites do not die for one to three days (but stop
treatment of dementia (Anonymous 1998). feeding within hours), so dienochlor is relatively slow-acting.
Organosphosphates inhibit acetylcholinesterase-a Made by Sandoz as a 50%FC. Dienochlor is less toxic than
mechanism lethal to insects (Ryan & Byrne 1988) but malathion and decomposes quickly (twice as fast as dicofol),
beneficial to Alzheimer's patients. Many terpenoids in especially in direct sunlight or temperatures above 130°C.It
Cannabis also inhibit acetylcholinesterase (McPartland & is not phytotoxic, does not harm beneficial insects (but is
Pruitt 1999). toxic to predaceous mites), is nontoxic to honeybees and
Acepliate is a frequently used systemic insecticide; Frank birds, but highly toxic to fish. General use.
(1988) applied a foliar spray against spider mites (sprayed Dimethoate is a systemic insecticide recommended by
three times at seven-day intervals) a n d thrips (two Frank (1988) against sap-sucking insects such as aphids,
applications seven days apart); he also sprayed aphids, whiteflies, thrips, and planthoppers in hops and vegetable
whiteflies, mealybugs, scale insects, and caterpillars. Kac crops. Made by BASF Corp. as an A, D, 10%G, 2-4 lb/gal
(1976) applied acephate against mites in Slovenian hemp EC, and 25-50%WP. Dimethoate degrades in 1 5 4 5 days on
fields. It has been used against flea beetles, leafminers, and plants and four to 16 days in soil. The RE1 is 24 hours. Some
grasshoppers in hops, tobacco, fruit, and vegetable crops. phytotoxicity occurs. Dimethoate is highly toxic to
Made by Chevron, and is available as a 0.25%A, 2%D, honeybees, fish, and birds.
50?'WP, and 75-90%SP. Acephate dissipates rapidly with Fosetyl-A1 selectively kills oomycetes such as Pythil~m
residual activity of ten to 15 days on plants and one to ten (damping off disease) and Psel~doperonospora(downy mildew)
days in soil. Highly toxic to bees, moderately toxic to birds, in hops, applied as a foliar spray, root dip, or soil drench. It
fish, and mammals. also treats bacterial diseases caused by Pset~domonassyringae
Chlorothalonil is a broad-spectrum foliar nonsystemic and Xanthonzonas species. Fosetyl-A1was the first product to
fungicide recommended by Frank (1988) to control mould be successfully reregistered under FIFRA'88. Made by Rh6ne-
and rot (not effective against Pytlziiinz damping off). It is the Poulenc, and is available as a 15%G and 80%WP. It is
third most-popular fungicide, behind mancozeb and copper moderately persistent in plants (systemically absorbed), but
(Hewitt 1998). Chlorothalonil works well against Septoria decomposes quickly in soil. The RE1 is 12 hours. It is not
species. Made by ISK Biotech since the mid-1960s, as an 11- phytotoxic, and nearly nontoxic to honeybees, fish, and birds.
409/oFC or 75%WP. Persistence in soil is 30-90 days, and
208 Hemp Diseases and Pests

Malathion, an insecticide and miticide, was first syn- Aldicarb is an extremely lethal systemic miticide, insec-
thesized in 1950. Sandru (1976) found it effective against ticide, and nematicide (especially against cyst nematodes).
hemp borers. Mountain Girl (1998) used it against spider Made by RhBne-Poulenc, and is only available as a 15G.
mites. Frank & Rosenthal (1978) applied it as a foliar spray Aldicarb is persistent, with residues in hops flowers peak-
against spider mites, cucumber beetles, and thrips, or as a ing 60 days after application (Duke 1985).Residual in soil is
soil drench against fungus gnats and other soil pests. Frank ten weeks. It is not phytotoxic, repellent to honeybees, mod-
(1988) also recommended it against aphids, whiteflies, erately toxic to fish, and extremely toxic to birds and mam-
mealybugs, scale insects, leafhoppers, and flea beetles. mals. Restricted-use.
Unfortunately, many pests are now resistant to malathion, Benomyl is a systemic fungicide, first available in 1968.
especially spider mites and whiteflies. It still works well Foliar sprays control grey mould, hemp canker, and powdery
against aphids a n d their a n t masters, h e m p borers, mildew. Soil drenches control southern blight (Ashok 1995).
budworms, cutworms, beetles, weevils, leafminers, Mishra (1987) soaked seeds and root-dipped seedlings in
leafhoppers, root maggots, grasshoppers, crickets, and benomyl to kill Rhizoctonia solani. Seed soaks also prevent
fungus gnats. Made by Rh6ne-Poulenc, and is available as a Fzisaritim and Macrophomina diseases in other crops (Maude
D, G, 5-10%B, 4-10 lb/gal EC, and 25-50%WP. The average 1996).Benomyl distorts P-tubulin, preventing fungal mitosis.
half-life for malathion is six days; residues persist longer in Because of this "site-specific" action, resistance arises by a
plants with high lipid contents (e.g., seeds). The RE1 is 24 simple mutation of a single gene. Benomyl-resistant fungi
hours. Malathion is highly toxic to honeybees, moderately appeared just two years after the fungicide became available
toxic to birds, and exhibits a range of toxicities to fish- (Hewitt 1998).To inhibit resistance, combine benomyl with
slightly toxic to goldfish, but very toxic to trout. In mammals, a nonsystemic fungicide, such as iprodione or captan (Agrios
malathion is less acutely toxic than aspirin (LDS0levels 1997). Made by DuPont, available as a 50%WP. Persistent
between 1375-10,700 mg kg-', versus 1200 mg kg-' for aspirin). on plants, residual six to 12 months in soil. The RE1 is 24
Parathion (ethyl) kills insects and careless farmers. It is hours. Benomyl is not directly toxic to beneficial predatory
much more toxic than malathion and possibly carcinogenic mites (Pkytoseilil~~s species), but may cause sterility in them
to rats; in 1992 its use was voluntarily restricted to field crops (Hussey & Scopes 1985). Benomyl is not phytotoxic, nearly
(cereals and soyabeans). Ethyl parathion has been used as a nontoxic to honeybees and mammals, slightly toxic to birds,
foliar spray and soil drench against beetles, weevils, and moderately toxic to fish. General use.
leafminers, leafhoppers, mealybugs, scales, and grasshoppers Carbaryl is the most commonly-used carbamate
on hops and other crops. Made by Cheminova and others, insecticide worldwide, useful against flea beetles, root grubs,
and is available as a D, 10%G, 2-8 lb/gal EC, and 15-25%WP. grasshoppers, and many other insects. Wanget al. (1987) used
It is nonpersistent. It is not phytotoxic, moderately toxic to it to kill hemp sawfly larvae. Frank & Rosenthal (1978)
fish, and extremely toxic to birds, bees, and mammals. recommend a 35 day safety period between carbaryl
Restricted-use. application and harvest of treated marijuana plants. Half-
Parathion (methyl) is less acutely lethal to mammals life is three to ten days on plants, seven to 28 days in soil.
than ethyl parathion, but it may cause chronic nerve damage The RE1 is 24 hours. Made by RhBne-Poulenc, and is available
in humans, especially children. It dissipates so rapidly its as a 5-10%D, 5-10%B, 4%FC, 50%WP, and 22-40%SP. It is
insecticidal efficacy is questioned-half-life on foliage is a not phytotoxic, moderately toxic to birds and fish, very toxic
couple hours. Nevertheless, the RE1 is 48 hours. No to honeybees, nephrotoxic to mammals (although carbaryl
persistence in soil. It is used on hops against hops aphids, is used in some flea shampoos), and kills earthworms. Re-
requiring a 15-day pre-harvest interval. Bosca & Karus (1997) peated use may stimulate spider mite infestations.
controlled Psylliodes atteizl~ataand Grapholita delineana with Etridiazole is a systemic fungicide recommended by
methyl parathion. Made by Cheminova and others, and is Frank (1988). It cures damping off caused by Pytkilim in
available as a 2-8 lb/gal EC and 20-40%WP. It is not cotton, cucurbits, tomatoes, and other crops, applied as a
phytotoxic but extremely toxic to bees, fish, and animals that soil drench or seed treatment. Made by Uniroyal as a 2.5%D,
eat fish. Restricted-use. 5-8%G, 30-40%WP, and 4 lb/gal EC. Moderately persistent
Tolclofos-methyl is a nonsystemic fungicide, applied as in soil. Phytotoxicity is rare. It is moderately toxic to fish,
a seed treatment and soil drench to protect against soilborne and slightly toxic to birds. General use.
diseases caused by Rlzizoctonia solani and Sclerotinm rolfsii. Ferbam is a sulphuric fungicide containing iron, useful
Its half-life on sunlit surfaces is two days, but it persists longer against seedling and foliage diseases in tobacco. Made by
in soil. Dippenaar et al. (1996) used it on hemp seedlings in UCB Chemical Corp., and has been available since 1931 as a
South Africa, without much success. Made by Sumitomo and 1-25%D and 3-98%WP. Ferbam is not phytotoxic, nontoxic
others, available as a 5%G, 10-20%D, 20%EC, and 50-75%WP. to honeybees, and slightly toxic to fish and birds. It leaves a
It is not phytotoxic, practically nontoxic to birds and black spray residue on foliage, and is irritating to the nose
and throat. The RE1 is 24 hours.
mammals, but very toxic to fish.
Mancozeb is a fungicidal mix of maneb and zineb, with
benefits and problems common to both compounds (see
CARBAMATES below). Since mancozeb has gone off patent, it currently out-
Most carbamates, like organophosphates, are very poi- sells all other fungicides (Hewit 1998). Mancozeb has
sonous (antidote: atropine) but biodegrade rapidly in the en- controlled yellow leaf spot in hemp, hops, and tobacco. It
vironment. Some sulphur-containing dithiocarbamates (e.g., should not be applied within 77 days of harvest, which limits
maneb, mancozeb, thiram) biodegrade into ethylene thiourea, its use to seed treatment and foliar treatment of seedlings.
a potent carcinogen. Lucas (1975) reports ethylene thiourea Made by Rohm & Haas and others, and is available as a D,
is not found in cigarette smoke generated from carbamate- G, 37-75%FC, and 80%WP. Residual is moderately persistent,
treated tobacco. To confuse the issue, some dithiocarbamates half-life four to eight weeks in soil. The RE1 is 24 hours. It is
may have medical applications, such as deactivating HIV not phytotoxic, with low toxicity to bees, slightly toxic to
viruses (Schreck et al. 1992). Nevertheless, sulphur-contain- birds, and moderately toxic to fish.
ing carbamates should not be used on plants destined for Maneb is a sulphur- a n d manganese-containing
human consumption, fungicide, used as a foliar spray or seed treatment. B6sca &
 Appendix 1: Synthetic Chemicals 209

Karus (1997)recommended maneb against grey mould and (e.g., iprodione and vinclozolin, two dicarboximide amides),
rust fungi; it controls a wider spectrum of foliar diseases others are absorbed systemically (e.g., carboxin, a
than any other single fungicide (Thomson 1997). Made by phenylamide). They are useful for seed treatments. Some
Rohm & Haas and others since 1950, and is available as a amides may be contaminated with dioxins (a group of about
1-~OYOD, 4%FC, and 70-80%WP. It is moderately persistent, 75 notoriously toxic chemicals).
with a half-life of four to eight weeks in soil. Maneb is not Carboxin is a systemic fungicide. Mishra (1987) soaked
phytotoxic, with low toxicity to birds and bees, but highly seeds and root-dipped seedlings in carboxin to kill Rhizoctonia
toxic to fish. General use. solatzi, but carboxin did not work as well as benomyl. Seed
Metam-sodium is a very effective nematicide and fun- treatments also control Sclerotil~mrofsii, Macrophomina
gicide, applied as a soil drench or as a fumigant injected into phaseolina, and Fl~sarit~m species (Maude 1996). Made by
the soil. Made by Stauffer, and is available as a soluble Uniroyal, and is available as a 25-75%WP and 3%FC.
concentrate. Metam-sodium is very phytotoxic, requiring an Carboxin has a half-life of 24 h in soil, but longer in plants. It
interval of two or three weeks between soil application and is not phytotoxic, nontoxic to honeybees, slightly toxic to
crop planting. It degrades to methyl isothiocyanate, with a birds, and very toxic to fish. It causes eye irritation.
residual 21-60 days. It is nontoxic to bees, moderately toxic ~ ~ r o d i o is
n ea fungicide used as a foliar spray and seed
to fish, and very toxic to birds. Restricted-use. treatment against Botrytis, Sclerotinia, Rhizoctonia,
Methomyl is a broad-spectrum systemic insecticide used Macrophomina, Alternaria, and many other fungi (Maude
on tobacco a i d hops to control spider mites, budworms, and 1996). Made by RhBne-Poulenc, in 50%WP and 41%FC
flea beetles. Made by DuPont, and is available as a 1-2 lb/ formulations. Residues on plants have a half-life of seven
gal SC and 90%WP. Methomyl takes three to six weeks to days, with a persistent half-life of 20-120 days in soil. The
degrade in soil, with a half-life of three to five days on plants. RE1 is 12 hours. Iprodione is not phytotoxic, nontoxic to
It is not phytotoxic, highly toxic to honeybees, birds, and honeybees, slightly toxic to birds, but very toxic to fish. See
biocontrol insects, and moderately toxic to fish. Restricted-use. comments under vinclozolin, below.
Pebulate is a systemic herbicide used against grasses Vinclozolin is a fungicide used as a foliar spray against
and some broadleaf weeds. Zabrodskii (1968),Tarasov (1971), sclerotia-producing fungi, such as Botytis and Sclerotinia
and Tarchokov (1975) tested pebulate and other herbicides species, and a seed treatment against Macrophovzina phaseolina
(trifluralin, linurom, dalapon, etc.) for use in hemp against (Maude 1996). De Meijer et al. (1995) sprayed hemp fields
An~bvosla,Polyxont~m,Setaria, Echinochloa, Chenopodiuna, with vinclozolin and iprodion, at rates of 500 g ha-1. They
Ainilrantlzl~s,and Raphanl~sweed species. Made by ~ e n e c a , alternated the two fungicides, spraying at 14-day intervals
in G and EC formulations. It is slightly toxic to honeybees, from June through August (after canopy closure and during
moderately toxic to mammals, and very toxic to fish. Holland's high-humidity summer). The treatment reduced
Pirimicnrb is a selective aphidicide, useful against orga- grey mould, but did not significantly increase crop yield-
nophosphate-resistant Myzus persicae on hops. It is nontoxic 11.2 t ha-1 for control plots versus 12.3 t ha-1 for treated plots
to most beneficial insects. Pirimicarb has fumigant (Van der Werf 1994). De Meijer et al. (1995) also sprayed
properties. It is translocated by xylem (taken up by plant vinclozolin and iprodion on fibre crops suffering from hemp
roots), with a relatively short residual. It was voluntarily canker. This treatment did not significantly improve crop
withdrawn in the USA because of marketing problems (even yield (Van der Werf 1994). Made by BASF as a 50%WP.
in the 1970s farmers rejected selective pesticides in favour Residues on plants have a half-life of seven days, and are
of broad-spectrum pesticides), but pirimicarb may be making moderately persistent in soil. The RE1 is 12 hours. Vinclozolin
a comeback. Made by Zeneca, and is available as an A, 5 lb/ is not phitotoxic, nontoxic to honeybees, slightly toxic to
gal EC, and 50%WP. It is not . phytotoxic,
.
nontoxic to birds, but very toxic to fish. It blocks testosterone receptors,
honeybees and fish. which may cause feminization in males (Colborn et al. 1996).
Thiram, a sulphuric fungicide used since 1931, is the
methyl analogue of disulfuram (Antabuse"). It is used against
damping off fungi, especially Pythit~mspecies, Rhizoctonia
solaili, and Botytis cinerea, applied as a seed treatment or
BIPYRIDILIUM HERBICIDES
soil drench. It is less effective against Fllsariilrn (Maude 1996). These herbicides act on contact, they are not translocated.
Thiram also exhibits deer-repellent activity, and has been Paraquat is a notorious example. Bipyridiliums cause harm
used in sunscreen ointments. Made by DuPont and others, if inhaled or splashed on the skin and kill if swallowed.
and is available as a 60%D, 2-5%G, 4%FC, and 3-90%WP. Symptoms do not arise until after treatment can be effective.
Thiram's half-life in soil is one or two weeks, the RE1 is 24
hours. Thiram is not phytotoxic, nontoxic to honeybees, but
highly toxic to fish. In mammals it is more toxic inhaled than HETEROCYCLIC COMPOUNDS
ingested. This group includes captan and captafol, nonsystemic
Zineb is a sulphur- and zinc-containing fungicide, fungicides introduced in 1952, and commonly found in home
applied to foliage or seeds. Bosca & Karus (1997) fruit-tree spray mixtures. Clarke (1981)used captan to protect
recommended it against rust fungi. Mishra (1987) did not clone cuttings from root rot (made by Zeneca as a 50-
find zineb helpful for controlling Rkizoctonia disease. Made 80%WP). High doses are carcinogenic; Frank (1988)
by DuPont and others since 1943, and is available as a 3- discouraged the use of captan. Web scuttlebutt suggests that
15%D, 4%FC, and 1-75%WP. Residual one or two weeks on the seed treatment used by French hemp producers to protect
foliage. Zineb is more phytotoxic than maneb. It is nontoxic seeds against damping off fungi may contain captan (captan
to honeybees, and more toxic to fish than to birds. also repels seed-eating birds). The usual dose is 0.5-6 g a.i.
per kg seed. Captan's half-life in soil ranges one to ten days,
residues on plants drop to 150 ppm after 27 days and cannot
be detected after 40 days. The RE1 is 96 hours. Captan is
AMIDES moderately toxic to birds, bees and mammals (EPA class I1
Amides are ammonia derivatives, fairlv new due to eye irritation), but highly toxic to fish.
agrochemicals. Some are nonsystemic or weakly systemic
210 Hemp Diseases and Pests

MISCELLANEOUS ing off fungi and seeds of weeds and broomrape (Orobanche
4-aminopyridine is a bird repellent formulated as a grain ranzosa). Restricted-use.
bait. It causes victims to emit distress calls, scaring off the Pymetrozine is a systemicpyridine azomethine, Fulfill@,
flock. The avicide Starlicide" kills starlings and blackbirds, Relay@,and Sterling@in the USA, for use on vegetables,
yet spares seed-eating sparrows and pigeons. All avicides tobacco, and other crops. It selectively controls sucking
require permits, and you run the risk of killing untargeted insects (aphids, whiteflies, planthoppers) by blocking their
birds. Overdoses kill mammals too. Restricted-use. stylets so they die from starvation about 48 h posttreatment
Fenbutatin oxide, a nonsystemic miticide organo-tin (Fuog et al. 1998).Made by Novartis (formerly Ciba-Geigy),
compound, is applied to hops, glasshouse crops, and fruit half-life in soil is two to 29 days. Nontoxic to mammals (EPA
trees. It is rotated with pyrethrins against spider mites in class 111), birds, fish, earthworms, bees. It is very safe to
Holland and Canada (High Times #257 p. 8,1997). Made by biocontrols-5x to 50x safer than pirimicarb to Oriiis,
DuPont and Shell, and is available as a 50%WP. Relatively Chysoperla, and Coccinella species (Fuog et al. 1998).
persistent in soil. The RE1 is 48 hours. Fenbutatin oxide is Pyridaben is a pyridozinone compound. It controls
nontoxic to honeybees. It is less toxic to predator mites than mites and sucking insects (aphids, whiteflies, leafhoppers,
to spider mites, so it has been used in IPM. It can be and thrips) on vegetables and fruit trees. It is not effective
phyfotoxic, is toxic to birds, and very toxic to fish. Organo- against caterpillars, grubs, and maggots. Pyridaben has little
tin compounds may disrupt the endocrine system (Colborn effect on predators, which makes it useful in mite
et al. 1996). management. Made by Nissan Chemical Industries as a
Fenaznquin is a broad spectrum miticide. It is a 20%WP and 15%EC. Moderately persistent. Slight toxicity
quinazoline with a novel mode of action (mitochondria1 to mammals and birds.
inhibition), used o n fruit a n d grape crops. Made by S o d i u m fluoroacetate is an odourless, tasteless,
DowElanco, in EC and SC formulations. Fenazaquin is not intensely poisonous rodenticide. It does not cause cancer, it
phytotoxic, does not affect beneficial insects, and is safe to just kills. Formulated as a B. It is extremely toxic to warm-
mammals and birds. blooded animals. Restricted-use.
Metaldehyde, an aldehyde, serves as a molluscicide. It Triadimefon is a new systemic triazole fungicide. It
causes slugs to secrete a heavy trail of mucus, so they inhibits sterol production in powdery mildews (exhibiting
dehydrate and die. Death comes sooner in warm weather. curative properties on established infections), with less
At higher doses metaldehyde also acts as a neurotoxin. Made activity against other fungi. It is used on hops, tobacco, and
by Chevron, a n d is available as a 1-20%B a n d 4G. vegetable crops, as a foliar spray and seed treatment. Made
Metaldehyde has a short half-life in soil. It is phytotoxic by Bayer a n d Miles as a 1-50%FC a n d 25-50%WP.
(apply bait near plants, not on them), nontoxic to honeybees, Triadimefon's half-life is one to four weeks on foliage. The
nontoxic to fish, and moderately toxic to birds and mammals. RE1 is 24 hours. It can be phytotoxic, is nontoxic to bees and
Restricted-use. birds, and toxic to fish.
Metalaxyl, an alanine ester, is a widely-used systemic Trifon'ne is a systemic fungicide recommended by Frank
pesticide. It selectively kills oomycetes such as Pythium (cause (1988).It controls powdery mildew and other foliar diseases
of damping off disease) and Pse~~doperonospora (cause of on hops, tobacco, and fruit trees. It has curative properties
downy mildew) in hops and tobacco. Made by Novartis on established infections. Triforine also suppresses spider
(formerly Ciba-Geigy), and is available as a D, 2%G, 25%WP, mites. It is a derivative of piperazine (a deworming drug).
and 2 lb/gal EC. Metalaxyl is applied as a seed treatment or Made by American Cyanamid as a 6.5-18%EC and 50%WP.
soil drench, with a residual of 70 days in soil. The RE1 is 12 Half-life of triforine in soil is approximately three weeks. It
hours. Metalaxyl may also be applied to foliage if combined causes little phytotoxicity, is nontoxic to honeybees and birds,
with another broad-action fungicide such as mancozeb or moderately toxic to fish, but highly toxic to mammals, with
chlorothalonil (to inhibit the rise of resistant organisms). moderate inhalation toxicity. Restricted-use.
Metalaxyl is not phytotoxic, nontoxic to honeybees and birds, Warfarin, CI9Hl5NaO4,is a coumarin derivative
and moderately toxic to fish. used as a rodenticide. Made bv Hacco and others as a 0.13%B
Methyl bromide, a deadly fumigant, is particularly dan- or in paraffin blocks; formulations over 3% are restricted.
gerous because it is odourless. The EPArecently listed it as a Many rodents in North America are now resistant to
class I ozone-depleting substance, 120 times-more potent warfarin. This has led to the develooment of
than chlorofluorocarbon-111. Thus, methyl bromide is being "superwarfarins" such as brodifacoum, which is occasionally
phased out to a total ban by January 1,2005. Kryachko et al. mixed with marijuana (LaRosa el al. 1997). Warfarin is
(1965) used it to kill hemp borers overwintering in stored nontoxic to honeybees, and very toxic to warm-blooded
seed. Methyl bromide has also been used as a soil fumigant animals. In human medicine it serves as an anticoagulant.
to kill nematodes, and at higher concentrations kills damp- The antidote is Vitamin K1.
 " W e zuill eitherfind a way or make one."
-Hannibal

Appendix 2: A Dichotomous Key of Diseases and Pests

This dichotomous key includes over 90 of the most common pests and diseases of Cannabis, which collectively
cause approximately 99% of crop losses. To save the forest from the trees, rarely-encountered organisms are
omitted from the key. These organisms can still be identified by referring to Chapters 4-6.
To begin using this key, the user must first turn to the section concerning the part of the plant showing signs
and symptoms (I. Seedlings and seeds in soil; 11. Roots; 111. Lower stems and crowns; IV. Upper stems and
branches; V. Leaves; VI. Flowers; VII. Entire plants). Every level of the key presents the user with two (sometimes
three) choices marked by the same letter. Trace key elements down to a specific disease or pest. Check the index
to find further descriptions of your tentative diagnosis. Illustrations are often provided to further evaluate the
accuracy of your diagnosis.

I. Seeds and seedlings

Al.Seed does not germinate, no seedlings appear
B1. Seed no longer present in soil
C1. Soil site disturbed .........................................................................................................
Birds, Rodents
C2.No soil disturbance seen .........................................................................
Root maggots, Cutworms
B,. Seed present
C1. Seed gnawed or other mechanical damage .............................................
Root maggots, Rodents
C,. Seed covered with fuzz ..........................................................................................
Damping off fungi
C3. Seed appears normal. ...........................................................................................
Drought, Poor seed
Az. Seed germinates and produces a seedling
B,. Above-ground parts show signs of feeding damage or insects present
C1. Stem completely severed from root
Dl. Severed seedling lying nearby .......................................................
Cutworms, Crickets, Slugs
D,. Severed seedling absent. ..................................................................................
Birds, Mammals
Cz.Plant upright but with holes in leaves or other insect symptoms
Dl. Leaves or cotyledons with holes and notched edges
El. Holes small, leaf edges rarely notched ............................................................
Flea beetles
El. Large holes and notched edges
Fl. Leaping insect with large rear legs .......................................................
Grasshoppers
F,. Caterpillars present on plant or nearby in soil
GI. Pink-skinned, often spin webs ......................................................
Hemp borers
GZ Grey or brown cuticle, rarely spin webs. .................Cutworms, Armyworms
DZ.NOmissing pieces but webbing or honeydew present
El. Webbing present with tiny red or green mites .............................................
Spider mites
E2,No webbing, much honeydew, pear-shaped insects .......................
...............Aphids
212 Hemp Diseases and Pests

B,. Seedling obviously sick but no feeding damage evident on above-ground parts
C,. Seedling wilted and/or toppled
Dl. Brown discolouration of root or stem near soil line
El. Roots discoloured and gnawed ....................................................................
Root maggots
E,. Root and/or stem discolouration, no insect damage ........................
Damping off fungi
D,. No root discolouration
El. Fungal hyphae present microscopically .............................................Damping off fungi
E,. No hyphae present
Fl. Bacteria present in sectioned tissue .......................................................
Bacterial wilt
F,. No bacteria present .......................................................................Drought, Frost
C,. Seedling upright but cotyledons and leaves discoloured
Dl. Discoloured areas light in colour, produce tiny, black-dot fruiting bodies
El. Discoloured areas greenish-yellow, form pycnidia ...............................Yellow leaf spot
E2. Discoloured areas grey, form acervuli. ..........................................................Anthracnose
D,. Discoloured areas dark, no fruiting bodies present
El. Cotyledons and true leaves covered with brown spores. ..........................Brown blight
E,. Cotyledons or leaves covered with dark green spores .......................................................
F1. Primarily cotyledons, spores two-celled and oval ...................Trichothecium spot
F,. Primarily true leaves, spores 4-5 celled, elongate ........................Curvularia blight

11. Roots (not including crown, see I11 below)

A,. Insect signs and symptoms (gnawed roots or insects present)
B,. Root damage an extention of stalk infestations
C1.Insect a green to brown caterpillar. .................................................................
assorted stem borers
C2.Insect a pale beetle grub with large head .............................................. Longhorn beetle grubs
B,. Root damage alone, no stem feeding
C1. Insect a 6-legged pale-bodied beetle grub
Dl. Grub fat, assuming a C-shape when disturbed ...........................................
White root grubs
D,. Grub cylindrical, resembles a caterpillar (but no prolegs) .........................
Flea beetle grubs
C,. Insect not a beetle grub
Dl. Insect a slow-moving legless maggot
El. Maggot spike-shaped with pointy head, <7 mm long ............................ Root maggots
EZ Maggot cylindrical or thread-like
F1. Maggot pink-grey in colour, up to 35 mm in length ................Crane fly maggots
F,. Maggot white, thread-like, 3 mm or smaller ........................Fungus gnat maggots
D2.Insect a quick-moving 6-legged ant or termite
El. Soft-bodied, white, almost always underground ..............................................
Termites
E,. Hard-bodied, red or black, often above ground ........................................................
Ants
 Appendix 2: A Dichotomous Key of Diseases and Pests 213

AZ.NO signs or symptoms of insects

B1.Abnormal swellings (galls, knots) and bushy-appearing root
C1.Deformed sections with small round cysts (<1mm) ..............................................
Cyst nematode
C2.Large irregular galls contain nematodes but no cysts ...................................
Root knot nematode
C3.Parasitic plant attached to deformed roots ...................................................
Branched broomrape
BZ Roots discoloured, rotten in appearance
C1. Sclerotia present
Dl. Mycelium white, sclerotia black, oblong, 5-12 mm long .................................Hemp canker
D,. Mycelium pale brown, sclerotia black, spherical, 1 mm dia. ......................Southern blight
D,. Mycelium forms funicles, sclerotia yellow-brown ...........................................
Texas root rot
CZ.Sclerotia not present
Dl. Root stele dark brown, no spores seen in culture ..................................Rhizoctonia root rot
D2.Root stele reddish, spores present in culture .........................................Fusarium root rot

111. Lower stalk, including crown (stalk at soil level)

Al. Stalks swollen into galls, puncture holes often present, frass or insect visible
B1. Insect a caterpillar with 6 true legs in addition to 8 prolegs
C1. Caterpillar pale brown with dark head ......................
.............................European corn borer
CZ.Caterpillar darker brown, white, green or red-violet ........................................
Other stem borers
BZ. Insect a pale white beetle grub with 6 true legs, no prolegs
C1.Grubs <7 mm long, plump, assume a C-shape when disturbed ............. Weevil, curculio grubs
C,. Grubs >7 mm, cylindrical, with enlarged heads .......................................
Longhorn beetle grubs
A>.No swollen stalks, puncture holes, or other signs and symptoms of insects
B1. Epidermis shredded, sclerotia present on surface or in pith
C1.Mycelium white, sclerotia oblong, 5-12 mm in length ............................................ Hemp canker
C2.Mycelium pale brown, sclerotia nearly spherical, 1mm dia. ..............................Southern blight
B> No sclerotia
CI. Crown turns dark brown, epidermis shredded ...........................................
Rhizoctonia sore shin
CZ.Crown dark brown, epidermis intact, root stele discoloured red ...................Fusarium foot rot
C3.Crown with beige-coloured, lumpy, cancer-like growths ...........................................
Crown gall
214 Hemp Diseases and Pests

IV. Upper stem and branches

A1. Stem galls, puncture holes, frass or insect visible
B,. Insect on surface of stem, feeding externally
C1. Insect sedentary or slow moving
Dl. Insect covered in froth of spittle ...............................................................................
Spittlebugs
D2.Insect flattened, often with waxy filaments
El. Filaments disappear with age, insects brown, round ......................................... Scales
E,. Filaments persist, insects white, slowly mobile ...........................................
Mealybugs
Cz Insect quickly leaps away when spotted
Dl. Insect small, spindle-shaped, with fringed wings. ................................................................Thrips
D2.Insect flattened shield shape, wings leather-like ........................................
Tarnished plant bugs
B2.Larvae feed within stem gall
C1.Insect a 3-legged pale-bodied beetle larva
Dl. Grubs <7 mm long, plump, assume C-shape when disturbed ........ Weevil, curculio grubs
D2.Grubs >7 mm, cylindrical, with enlarged heads ...............................Longhorn beetle grubs
D3. Grubs different than above .................................................................
Assorted boring beetles
C2.Larvae with prolegs (caterpillars) or no legs (maggots)
Dl. Small (<7mm long) legless maggots ..................................................................
Nettle midges
D2.Larva a caterpillar
El. Pale brown bristly body with dark head, 10-20 mm long ......... European corn borers
E2.Plump pinkish-white body <I0 mm long with dark head .......................Hemp borers
E3. Caterpillar darker brown, white, green or red-violet ........................
Other stem borers
A?. No galls, puncture holes, frass or insect visible
B,. Stem or branch epidermis sunken or discoloured
C1. Canker covered with a mat of mycelium and spores
Dl. Spores grey, round ....................................................................................................
Grey mould
DZ.Spores white or light pink, ellipsoid .......................................................................
Pink rot
D,. Spores green, oblong ..............................................................................
Cladosporium canker
C2.Canker not covered with mycelium, usually light brown
Dl. Fruiting bodies present in older lesions
El. Initial fruiting body a simple pycnidium
F1. Fungus only produces pycnidia in culture .............................................
Yellow spot
F> Other spore stages sometimes form in culture ................Brown spot, Twig blight
E2.Initial fruiting body an acervulus or perithecium
F,. Grey to pink acervuli bristling with setae ....................................... Anthracnose
F2.Perithecia with long cylindrical ascospores .............................. Ophiobolus canker
D,. No fruiting bodies
El. Fungal spores canoe-shaped .................................................. Fusarium stem canker
E2.No fungal elements, bacteria seen under microscope .......................Striatura ulcerosa
 Appendix 2: A Dichotomous Key of Diseases and Pests 215

B-. Epidermis ilormal but stem misshapened and often spongy

CI. Grub or caterpillar present in pith .................................................................. go to section IV AlB:
C?.No insects present
Dl. Microscopic nematodes present ....................................................................... Stem nematode
DZ.No nematode present
El. Stem tissue discoloured red ...........................................................Fusarium stem canker
E-. No discolouration, plant stunted ................................................... Nutrient deficiency

V. Leaves
A,. Insect damage (leaf cutting, puncture wiounds) or other signs (honeydew, webbing)
B,. Leaf skeletonized or holes present and/or notched leaf edges
C,. Leaf skeletonization
Dl. Skeletonization alone
El.Insect a conspicuous green and bronze beetle ....................................... Japanese beetles
E?. Insect a hyperactive, small, black beetle ..................................................Flea beetles
E3. Insect a white larva <I2 mm long, resembles a caterpillar ..................... Sawfly larvae
D2.Skeletonization followed by large holes and notched edges
El. Only leaves attacked ......................................................................Leaf-eating caterpillars
E2. Other plant parts also attacked, webbing often present
F1. Caterpillars bore into stems and branches ....................... Stem-boring caterpillars
F2. Caterpillars attack flowering tops ............................................... Bollworms, Borers
C2.No skeletonization, only leaf holes and/or notched edges
D,. Caterpillars present
El. Individuals usually solitary .......................................................... Leaf-eating caterpillars
E?. Individuals gregarious ...................................................................................Armyworms
Dz. Insect not a caterpillar
El. Insect with large hind legs, conspicuous.................................................... Grasshoppers
E:. Insect a less-conspicuous beetle
F1. Only edges notched, beetle with long snout. ......................... Weevils & Curculios
F2.Notched edges and leaf holes, no long snout ...................Flea beetles, leaf beetles
B2. Tiny puncture wounds, rasp marks, or subsurface tracks-no leaf holes or skeletonization
C,. Wounds punctate (often appear as white specks), plants often wilted
Dl. Honeydew present, insects <3 mm long
El.Webbing produced by small red, green or brown mites .... Spider mites, Other mites
E2. NOwebbing present
FI. Insects usually green, pear-shaped .................................................................Aphids
F2. Insects resemble tiny white moths. ............................................................. Whiteflies
D2. Little or no honeydew, insects >3 mm long
El. Insects flattened shield-shaped, often slow moving ....................................... Plant bugs
Ep Insects elongated, quickly leap or fly away ................................................ Leafhoppers
216 Hemp Diseases and Pests

CZ.Wounds linear, on surface or subepidermal

Dl. Short rasp marks on surface, sometimes oozing sap ....................................................
Thrips
D2.Epidermis intact, white or brown tracks beneath surface .................................... Leafminers
A,. NO insects or feeding damage but leaf spots or other discolourations present
B1. Leaf spots with distinct edges
C1. Spots white, yellow, light-green to light-brown
Dl. Fruiting bodies present in older lesions
El. Fruiting bodies pycnidia or perithecia
F,. Spots yellow, large, irregular, with scattered pycnidia ................. Yellow leaf spot
F,. Spots white, large, irreg., pycnidia in concentric circles .................White leaf spot
F3. Spots brown, small, circular, often breaking apart
GI. Only pseudothecia with ascospores present ................................Pepper spot
G2.Pycnidia, rarely with perithecia present .................................
Brown leaf spot
E2.Fruiting bodies other than pycnidia or perithecia
F,. Grey to pink acervuli bristling with setae ............................................
Anthracnose
F2.Bright yellow or orange fruiting bodies. .............................................................
Rust
FS. Black pin cushion-like sporodochia ...............................................
Black dot disease
D2.No fruiting bodies in older lesions
El. Fungal spores present
F,. Spores borne mostly on upper leaf surface
GI. Leaf lesions often with chlorotic ring ..........................................
Brown blight
G2.Leaf lesions without chlorotic ring .............................Stemphylium leaf spot
F,. Spores mostly on underside of leaf
GI. Spores emerge from stomates only on undersides. .................Down mildew
GZ. Spores borne in tufts, mostly on undersides ...........................Olive leaf spot
E2.NO fungal spores or mycelium present
F1. Bacteria present in sectioned material. ..............................................
Bacterial blight
Fz. No bacteria present ............................................................................
Abiotic diseases
C2.Leaf spots dark, nearly black
Dl. Fruiting bodies present within fungal stroma .............................................................
Tar spot
D2.No fruiting bodies present, spores borne on leaf surface ......................Cladosporium spot
B,. Leaf lesions or discolourations without distinct edges
C1.Mycelium on leaf surface
Dl. Mycelium white to grey mycelium, sometimes light pink
El. Mostly on upper surfaces. .....................................................................
Powdery mildew
EZ On either surface, even growing on branches .....................................................
Pink rot
D,. Black mycelium, either surface
El. Insects present, leaf surface often sticky .......................................................
Sooty mould
E2.NOinsects present, not sticky .......................................................................
Black mildew
 Appendix 2: A Dichotomous Key of Diseases and Pests 217

CZ.Leaf lesion without mycelium, consists of chlorosis or necrosis in streaks, mosaics

Dl. Bacteria seen in sectioned tissue .......................................................................
Bacterial blight
D,. Bacteria not seen
El. Leaves exhibit phyllody (distorted growth) or rosettes . Mycoplasma-like-organisms
E2. Little distorted growth, mostly leaf chlorosis
F,. Chlorosis in streaks or chevrons ...............................................Hemp streak virus
F,. Chlorosis in small rings or mosaics ................................................
Mosaic viruses
F3. Chlorosis without a pattern ...................................................
Nutrient deficiencies

VI. Flowers and developing seeds

A,. Male flowers
B,. Petioles swell into galls, flowers wilt ....................................................................................
Gall midges
B,. Petioles normal, flowers covered with mycelium
C1.Mycelium produces oval, grey spores. ..........................................................................
Grey mould
C2.Mycelium produces elongated brown spores. ...........................................................
Brown blight
A?.Female flowers and developing seeds
B,. Webbing, honeydew, leaf holes or insects present
C,. Holes or edges notched in flowers, fan leaves, and seeds
Dl. No webbing, holes small, insect a small black beetle ........................................... Flea beetles
D2. Webbing and caterpillars present
El. Larvae pink, immature seeds partially shelled ..................................................
Hemp borers
E2. Larvae light brown, seeds partially shelled .........................................European corn borers
E3.Larvae green or dark brown, whole seed sometimes eaten .................................
Budworms
CZ.No holes in flowers or fan leaves, no shelled seeds
Dl. Webbing present with minute red or green mites ..............................................
Spider mites
D,. No webbing
El. Insects <2.5 mm long, pear-shaped, with honeydew .........................................
Aphids
EZ Insects conspicuous, 15 mm long, flattened shield-shape ..................Green stink bugs
B2. NOsigns or symptoms of insects
C,. Wilted flowers, no other symptoms
Dl. Whole plant wilted. ...........................................................................................
go to section VII
D,. Only apical bud or occasional branches wilted ...........................................
go to section IV
C,. Discoloured flowers with/without wilting
Dl. Fungal mycelium and spores present
....................................................
El. Fan leaves wilted and grey spores present Grey mould
E,. Less conspicuous mould with brown spores present ................................. Brown blight
D2.No signs of mycelium or spores
El. Flowers and fan leaves with chlorotic markings .............Viruses, Nutritional diseases
E2.Flowers turn black overnight .......................................................................
Frost damage
218 Hemp Diseases and Pests

VII. Whole plant

A,. Wilt (sometimes one-sided) leaves evenly chlorotic or normal colour
8,.Stems discoloured internally (xylem or pith)
C1.Pith peppered with microsclerotia. .............................................................................
Charcoal rot
Cz.No microslerotia but xylem discoloured
Dl. Reddish-brown discolouration, spores canoe-shaped ...................................Fusarium wilt
D,. Brown discolouration, spores round to oval .............................................
Verticillium wilt
B2. NOdiscolouration of stem tissue
C,. Roots abnormal ...........................................................................................................
go to section I1
C2.Roots normal
Dl. Bacteria in sectioned plant tissue .......................................................................Bacterial wilt
DZ.No bacteria seen ......................................................................................
Drought, Herbicides
A2.Systemic chlorosis and necrosis
B,. Chlorosis gradual, plants often stunted
C1.Insects or nematodes found upon close inspection of plant
Dl. Insects on undersides of leaves .........................................................................
o to section V
D2.Insects or nematodes on roots ..........................................................................
go to section I1
CZ.No insects. ....................................................................................................
Air pollution, Nutrients
B2. Chlorosis and necrosis sudden ..................................................................................
Toxins, Herbicides
 " A l l concepts, theories, and general ideas are thin and ineffecttlal tlnless they are grounded i n
the concrete reality of things which specifically enter into our lives and which zue steadily deal with." -John Dewey
" N o ideas but i n things. " -William Carlos Williams

Appendix 3: Conversion Factors

Both English and Metric measures are used in this text, depending on the system used by original authors.
Use charts and equations below to convert:

length temperature weight (mass)

lnches cm O F OC kilograms x 2.2046 = pounds lb x 0.4536 = kg
grams x 0.354 = ounces oz x 28.25 = g
milligrams x 0.001 = grams g x 1000 = mg
ounces x 0.0625 = pounds lb x 16 = oz
2000 lbs = 1short ton (U.S.) = 0.89 long ton (U.K.) = 0.91metric tonne
- N -

volume & capacity

litres x 1.057 = quarts qt x 0.946 = 1
millilitres x 0.2 = teaspoons tsp x 5 = ml
cubic centimetre x 0.06 = cubic inch in3 x 16.4 = cm3
- -- bushel x 35.2 = litres 1x 0.0284 = Bu

area
square metres x 10.76 = square feet ft2 x 0.093 = m2
~l~
-
- square miles x 640 = acres acre x 0.0016 = Mi2
1 -; hectares x 2.5 = acres acre x 0.4047 = ha
l w x l 9
I-= - - * =-""- length
i -
-- micrometres x 0.001 = millimetre mm x 1000 = pm
1 ---

_
: a = millimetres x 0.04 = inches in x 25.4 = mm
--
- metres x 3.28 = feet ft x 0.305 = m
" ---
-- Oz==zz
I
kilometres x 0.621 = miles Mix 1.61 = km
3
I Z- A -
A - assorted
1Lumen = 1 footcandle = 10.76 footmetres = 10.76 Lux
1Watt per square foot = 10.76 Watts per square metre
, --: 1.123 kg hectare-1= 1lb/acre
E-1.x
1
- - -
z-
-
-
-
1 0

2 The above values

11.24 litres ha-1 = 1 gallon/acre
1kg cm-2 = 13.3 lb/in2
-
' -
Y *-
d =
are approximate. 1bar = 0.99 atmosphere = 14.5 lb/in2
@

i -- Exact conversions:
"C = (OF-32) x 5/9
OF = ("C x 9/5) + 32
 "An absolutely clear arld exhaustive i~riderstandingo f a n y single tlzing in the world
w o u l d iniply a perfect cornpreherzsion o f e v e q t h i n g else."
-Arthur Schopenhaurer

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ill~tograplliceDelineata No. 55. Berolini, Italy. 509 tabulis. treatment on Caiznabis sativa pollen viability. Plant Cell,
Wollen~veberHW, Reinking OA. 1935. Die Fl~sarien.Verlag Tisslle G.. Orgail Cultlrre 47:189-194.
Paul Parey, Berlin. 355 pp. Zubrin R. 1981. The fungus that destroys pot. War on Drugs
Wood A . 1989. Irlsrcts of Eco~zoinicInlportance: a ckeckllst of Action Reporter, June 1981:61-62. LaRouche Publ. Co.
preferreii iiaines. CABI, Wallingford, UK. 150 pp. Zuskin E, Valic F, Bouhuys A. 1976. Byssinosis and airway
\Vu FF, Dobberstein RH, Morris RW. 1978. The effects of responses due to exposure to textile dust. Lung
selected fungicides and insecticides on growth and 154:17-24.
thebaine production of Papoiler br~lcteatl~in. Lloydia Zycha H, Siepmann R, Linnemann G. 1969. Ml~corales.J .
41 :355-360. Cramer, Lehre, Germany. 355 pp.
Yarxvood CE. 1973. "Erysiphales," pp. 71-86 in The Fiingi: Zylinski T. 1958. Nalrka o Wl6kizle. Centralny Instytut
A11 Ad;~alzcrdTreatise, Vol. 4A. Ainsworth GC, Sparrow Informacji Naukowo-technicznej i Ekonomicznej
FK, Sussrnan AS, eds. Academic Press, NY. 621 pp. Warszawa. 681 pp.
 Index

Note: page numbers in italics refer to figures and tables

abamectin 197 aphids 32-34 beet armyworm 55

Acalyit1ii1a ~lttilt(l67 biocontrol 35-38 beet webworm 59
acephate 207 biorational chemical control beetles see flea beetles; leaf beetles;
Aclzrrontla a t r ~ p o s(death's head moth) 38-39 longhorn beetles; tumbling
60 cultural/mechanical control 34-35 flower beetles
Aclzeta doitzestzclrs (cricket) 87 Aphis species 32-34 Ber7zisia species (whitefly) 39-40
Aciilops cannabicola (hemp russet mite) Aphytis nzelinus 83 benomyl 208
30-31 Apion species 71 bertha armyworm 55
Ailnlza 21iplrticfata (two-spotted lady- apterae, defined 31 bhang aphid 33
beetle) 29, 36 arabis mosaic virus (ArMV) 143 'big-eyed bug' 42
Accr~i~lclrz catlt~abls 123 Arc/zytas rnarrnoratlis 52 bindweed 147
Aeoloflrrlps ii~teriiledllis 64 Arctia caja (garden tiger moth) 58 bio-dynamic farming 2
aerosol formulation 189 armyworms 54,55,56 biocontrol organisms 179,185-187
A ~ i l ; ~ m ~ t l i z m p e c73~es biocontrol 56-57 application rates 184-1 85
A;;riotc.s 11il~atlrs (llned click beetle) 91 chemical control 57 autocidal control 184
Agrol~acter~liiii radlobactcr 147 cultural/mechanical control 55 companion plants 183
A g r o b i l c t ~ ~ r ztllnlefaciens
~~ri~ 146 Arthriizilrm p/zaeosperrni~m 131 en~rironmentaleffects 187
ACyroi7iyzar~ptails77 Arthrobotys species 141 epiphyte antagonists 182
Agrotis scgetliiiz cytoplasmic poly- Arthropoda 20-23 microbes 182
hedrosis virus (AsCPV) 57 Aschelminthes 18-20 parasi toids 182
Agrotis segctl~ingranulosis virus Aschersonia aleyrodis 43 pesticide toxicity 183
(AsGV) 57 ascospores, defined 17 predators 181-182
Agrotls species (cutworms) 54-55 Asochy ta species 104-1 06 refuge/habitats for 179
nlatae, defined 31 Aspergillus species 172 release strategies 181
Albllgo specles 123 Athclia species 124 trap crops 184
aldicarb 208 Atracfotnorpha cretli~lata 87 viruses 13
Alc70dlara b1111zc~at~7 86 Atta species (leafcutter ants) 84 biopulping organisms 165
alfalfa mosaic virus (AMV) 143 Aliveobaszdizinz pzillz~la~zs
129 biorational chemicals 189, 192
allelochemicals 18, 148, 198 autocidal pest control 184 botanical poisons 194-197
allelopaths 148 Azltograplzn gaitzrna (silver Y-moth) 58 classification 191
Altc7rrlarra specles 114-115 Aves 23 fermentation products 197
Altica species 66 Ayurveda 170 formulations 189,190
altitude 11 azadirachtin 195 non-botanicals 190-194
Arnblys~~~lrs species 62-63 synthetic analogues 197-1'39
Airzbrosln tr2frda (ragweed) 147 bipyridilium herbicides 209
amides 209 Bacill~tscereils 100 birds 152
4-aminopyridine 210 Bacillzis yenetrans 141 black bean aphid 32
A1il;7cloitryces ~ ] I I Z ~ L ~ ~112LQ~IS Baczl~iispopilliae 70 black cutworm 54
A I I ~ ~ ~ ~ ~ ~ S L ' I83I L ~ O C O C C I Bacillz~ssubtilis 100, 103 black dot 124
anamorph, deflned 17 Bacilllis thiiringiensis (Bt) 15, 47-48 black mildew 117-118
Aiia,vlles lol~.76 transgenic 48 blight
Anglospermae 18 var. azzaiuai 48 bacterial 144-145
Anlmalia 18-23 var. israelensis 90 brown 114-115
Annelida 20 var. klirstaki 47 Curvularia 129
At~oplolzoraglabripcnrzls 72 var. rrlorrisorzi 48 defined 15
antenna1 tubercles, deflned 31 var. tenebriotzis 67 southern 116-117
At1 tllocorrs izrmorlrn~64 bacteria 4-6, 15 twig 118-119
anthracnose 121-122 bacterial diseases 144-147 Xanthomonas 145
antifeedant pesticide, defined 190 bait, defined 189 bollworm 51
antl-marijuana blocontrol 8 baited traps 205 Bordeaux mixture 191-192
ants 84 banker plants 184 borers 50
i l ~ ~ h ~ l r r al~doirli~lal~s
llls 38 basidio rot 124 see also European corn borers;
Aplllill~isspecles 37-38 basidiospores, defined 17 hemp borers
Ayl~iciolctcsap/llillitlyza 37, 183 Beazizleria species 43, 70 boric acid 193
245
246 Index

boron 157 growth requirements 10,164 conversion factors 219

botanical poisons 194-197 as pest control 194 Conuolz7zilzrs arzlerzsis (bindweed) 147
registration 200 carboxin 209 copper 157,193
synthetic 197-198 carmine spider mite 25-26 corn borers see European corn borers
Bofi yospllnerrn species 118-119,131 caryophyllene oxide 3 cornicles, defined 31
Botrytrs clizeren 93-95, 99, 169 castor 195 Corynebacteriiirn fascians 165
B~rrrlet~~'llnhorfen5ls (garden spring- caterpillars see leaf-eating caterpillars C o s s ~ cosszis
~s (goat moth) 50
tall) 91 cauliflower weevil 70 Cofesin niargirzventris 57
Br~lclyslnspecies 89 Cernin bycidae 72-73 cotton aphid 33
branched broomrape 138 C ~ r n n i s ~rnenes
is 65 cotton bollworm 51-52
BI eej~ynlplrsspecies (privet mites) 31 cerci, defined 20 cottonycushion scale 81
broomrapes 150-151 Cercosporn cannabis 113 crane flies 89-90
brown blight 114-115 Celitor/zyncl~zlsspecies 70-71 creosote 195
brown fleck disease 114-116 Clznctocneirza species 66 crickets 87
brown leaf spot 103-106 Clzactonzrzlin species 126-127 crop damage triangle 1-2
Bryophyta 17-18 chafers, European 68 crop rotations 159, 177
budworms 51-53 charcoal rot 112-113 crown gall 146
biocontrol 52-53 Charles, Vera 7-8 crown rot, defined 15
chemical control 53-53 C!zelolzlrs species 53 cryolite 193
cultural/mechanical control 52 Chinese medicine 170 Cryptolaernlis inontrolrzieri (mealybug
~nfestatlonseveritv index 52 chitin inhibitors 199 destroyer) 82
buffalo treehopper 80 chitosan 194 cucumber mosaic virus (CMV) 143
buffer (spray adjuxlant) 202 chlamydospores, defined 17 curculios 70-72
burdock borer 51 Chlooaltis consyersn (sprinkled locust) Curvularia blight 129
Blrrkhol~l~rin cepacin 100, 141 86 Czisclita species (dodder) 148-149
byssinosis 168 chloride 157 cutworms 54-56
chlorinated hydrocarbons 205-207 biocontrol 56-57
Chlorophyta 16 chemical control 57
cabbage curculio 71 chlorosis, defined 13 cultural /mechanical control
cabbage maggot 85 chlorothalonil 207 55-56
cabbage moth 58 chlorpurifos 207 Cyli~zdrosporizrnzcannabinn 131
calcium 156, 160 cholecalciferol 194 Cyretepistonrnzls castanells 71
calcium cyanamide 193 Clrondracris rose0 (citrus locust) 87 cyst nematodes 138,139
Calzotllrips z i ~ d i c ~(Indian
~s bean thrips) Chordata 23 cystolith trichomes 22
61 chloropicrin 205
Cnlocc~risI I O ~ Z I ~ ' ~ (potato
~ ~ C ~ I Sbug) 75 Chrysomelidae 65-67
Cnlystcya scpilrin (bindweed) 137 Clzrysoperla species (lacewings) 35-36, DDT 205
Cnill/l~rlilprl111ci~ln (clearwinged 183 Dacizzrsa sibi~ica 78
grasshopper) 86 Cicadellidae 79-80 Dactylaria species 141
Clzi~di~lac,lec>ylli/a 173 Cicadas 80 damping off 97-100
cnnker, defined 15 cinnamaldehyde 197 control 99-100
cannabinoids 3,15,22-23,195 citrus locust 87 death's head moth 60
C171.lllflb15 Cladosporlirrrz species 120-121 degree days 6
antibacterial compounds 15 Clazlibacter xyli 47 Delia species (root maggots) 85
antifungal compounds 16 clay microparticles 193 Delphastzis pirsillzis 42
consumer hazards 174 clay, defined 10 deltamethrin 198
disease researchers 6-8 claybacked cutworm 55 Dendroplzorna nzarconii 118
genetics 1, 165 clearwinged grasshopper 86 Derneocoris brevis 64
insect repellents/pesticides click beetle grubs (wireworms) 91 Dewey, Lyster Hoxie 7-8
22-23,193-197 climate 164-165 Diabrotica species 67
medicinal uses 170 Clonostnclrys rosea 95 diapause, defined 21
occupational hazards 168 Clostridizrnl species 168 Dinpromoupha pallens 67
resistance in 1-2, 176-177 Cneplzm~1ainterjectana 59 diatomaceous earth 16,193
seed srr hemp seed cobalt 60 irradiation 174 diazinon 207
taxonomy 3-4,170-171 Coccinella sepfei?zpzrnctala 37 dichlorvos (DDVP) 207
trichomes 22,170-171,185 Cocciizella zindecllnpzlncfata (eleven- Dichroplzis ~~zaciilipeiznis87
Cnnizaliis nj$rnilrcn 4 spotted ladybeetle) 37 dicofol 205
Ccrllllnbis iniiica 3 cockroaches 87 Dicyrna pzrluinata 114
Cnnrlnbis rlrilcrnlis 3-4 coexistence 2 Didyrnilirn claz~zrs 132
Cnn~znbissntieln 3 Colletotr~clz~rm species 121-122 dieback, defined 15
canopy closure, defined 147 common hairy caterpillar 59 dienochlor 207
captan 209 common stalk borer 50-51 Diglyphzrs isaea 78
carbarnates 208-209 companion plants 184 dikaryotic, defined 17
carbaryl 208 conidia, defined 17 disease, defined 1
carbon dioxide Coniothyri 1rn1 17zinitans 97 Dityleizchzis dipsnci (stem nematode)
excess 164 convergent ladybeetle 29/36 139-140
 Index

dodder 148-149 feeding stimulants 199 griseofulvin 197

Dolycoris indiczts (Indian stink bug) 75 Feltiella acarisziga 29-30 growth requirements
dot moth 58 fenazaquin 210 atmosphere 10
downy mildew 106-107 fenbutatin oxide 210 climate 164-165
drought 164 ferbam 208 ecology 11
dry flowable, defined 189 fermentation products 197 light 9-10
dust formulation, defined 189 fertilizers 161-163 moisture 9
Dysdcrci~scilzglrlntiis 76 flavescent leafhopper 80 temperatures 6,9
flax noctuid 52 see also nutrients; soil
flea beetles 65-67 grubs see white root grubs
earwigs 91-92 flowable concentrate, defined 189 Gryllidae 87
ecological relationships 3 flower flies 89 Gryllotalpa species 87
ectoparasites, defined 20 Forficula aziricztlaria (earwig) 91-92 Grylliis species 87
Ediunrdslann rosac 80 formaldehyde 194 Gymnospermae 18
eelworms see nematodes Fosetyl-A1 207 Gypotzana octolineata 80
eleven-spotted ladybeetle 37 Frankinielln occidentalis (western flower
ELISA 4,6 thrips) 61-62
Emponscn species (leafhoppers) 79-80 Franklinotlzrips vespiformis 64 hail damage 165
emulsifiable concentrate, defined 189 frost damage 164 hairy caterpillar, common 58
Encnrsiaforn~osn41-42 Fulgoroidea 80 Harnzonia axyridis (multicoloured
pesticide effects 183 fumigant, defined 189 Asian ladybeetle) 82
release strategies 181 fungi 16-17,18,171-173 heavy metals 164
Ellcarsin l~rtroln42 identification 4-6 Helicoverpa armigera stunt virus 53
E tzdocylytn excrescens 51 taxonomy of Cannabis-associated Helicoverpa species (bollworms) 51-52
endoparasites 19 Heliocotylendizts species 140
migratory 20 see also named fi~ngi Heliothis viriplaca (flax noctuid) 52
sedentary 20 fungus gnats 89-90 Heliothrips lzaemorrhoidalis (greenhouse
endophytes 16 Fzrsariztm species 107-111 thrips) 61
Enterobncter cloacae 171 damping off 100 hellebore 195
entomology, terminology 20 foot/root rot 108-109 Hemiptera 73-76
Entonzoyht/zora species 38, 65/88 human toxicity 108 hemispherical scale 82
enzyme-linked immunosorbent assays nonpathogenic 111 hemp bagworm 52
(ELISA) 4,6 stem canker 107-108 hemp borers 48-49
Epliestin klz17ielln 165 wilt 109-111 hemp canker 96-97
Ep~coccir~~i nigrzirn 124 hemp dagger moth 59
Epilnnclznn dodecostigma 67 hemp fibre 167-168
epiphyte antagonists 182 Galendromiis species 28 hemp flea beetle 65-66
epiphytes, defined 16 gall midges 85-86 hemp longhorn beetle 72
E retrnocerlrs erenzici~s42 gamma moth 57 hemp louse 33
Eriuinin trnclieiphiln 146 garden tiger moth 58 hemp mosaic virus (HMV) 143-144
Ery rzin neonpliidis 38 Geisha distinctissima (planthopper) 80 hemp russet mite 30-31
Er~ysrplzeconzrnzinzs 111 genetic engineering 1-2 hemp sawfly 90
escape cropping 176 genetics 165 hemp seed
essential oil 3 Geocovis pz~nctipes42-43 deterioration 169-170
ethylene oxide (sterilizing agent) glandular trichomes 22,168-169 oil from 169
174 glasshouse leafhopper 79 pesticide treatments 202
ethylene thiourea (carcinogen) 208 Gliocladizim roseiim 95-96 sterilization of 178
etridiazole 208 Glomits in traradices 100 storage of 169-170
Eucclnforia brynnz 53 goat moth 50 hemp streak virus (HSV) 143
European chafers 68 Goetheana shakespearei 65 hemp weevil 71
European corn borers 44-45/47 granule, defined 189 herbicide damage 164
biocontrol 4 5 4 8 Graphocephala coccinea (redbanded herbicides 164
chemical control 48 leafhopper) 79 heterocyclic pesticides 209
cultural/mechanical control 45 Grapholita delineana 49 Heterc7dern species (cyst nematodes)
European fruit lecanium 81 Grapholita interstictana 49 138,139
Eirseizls lzibisci 63 Grapholita tristrigana 49 Hrteron!~c/~~rs nrator 69
.
Elitetrnnyclll~s orientalis (oriental mite) grasshoppers 86/87 Hcterorlznbdif is species 69
31 control 88-89 Hippoiiniiiii7 con~~t~rgriis (con\-ergent
extender (spray adjuvant) 202 green lacewing 35 ladybeetle) 29, 36
extinction 187 green peach aphid 32 Hirsl~tellnrilossilit711sis 141
greenhouse thrips 61 Hirslltelln fllollrpso,lii 30
greenhouse whitefly 39 fTistoplns~i~n cnpsirlflf ~ L I I I 171
facultative parasites 16 grey mould 93-96 Hon~eopro~~c~~inflrs 17lli011!71 30
facultative saprophytes 16 biocontrol 95-96 honeybees 191
false chinch bug 74 chemical control 96 honeydew 21
fasciation 165 cultural / mechanical control 95 hops aphid 33
 Index

hops cyst nematode 138 whitefly control 42 Melanopliis biuittatzis (two-striped

hops flea beetle 66 Lasiodiplodia tkeobromae 132 grasshopper) 86
horsetail 195 latitude 11 Melanoplzis sang1 inipes
hover flies 89 leaf beetles 68-69 entomopoxvirus (MsEPV) 88
humus 11 leaf spot hlelanospora cannabis 125
Hy~~ienoscyplzzls hc)rbar~rnl132 defined 15 MeIoidogyne species (root-knot nema-
hyperparasites, defined 16 olive 113-114 todes) 103,137-138
hyphae, defined 16 white 128 Melolon thn species (Europeanchafers) 68
H!y[~/za?lfria C I I Y ~ ~ Y59
I Wisconsin 146 Mesosei~tlz~s longipes 28
Hypoasp is r~liles90 yellow 101-102 xnetalaxyl 210
H Y ~ O I ~ I ( YS YL L~ 'IS~ O ~ I (gold
~ O Z ~ dust
S wee- leaf tissue analysis (LTA) 155 metaldehyde 210
vil) 71 leaf-eating caterpillars 57-60 metamorphosis, defined 21
leafhoppers 79-81 metam-sodium 209
leafminers 77-79 Mefaphyczrs helvollrs 83-84
1ass11sindic11s 80 biological control 78-79 Metar!zizilim anisopliae 38, 63
Icer!ycl p11rc11asi(cottonycushion scale) chemical control 79 methomyl 209
81 cultural/mechanical control 78 methvl bromide 210
imidacloprid 197 L~ptomastixdactylopii 83 metrifonate 207
imperfect fungi, defined 17 Leptosp/zaerin species 132-133 microbial pesticides 182
inbreeding 165 Lt.ytosplzaerzili~zaspecies 128-129, 133 Microdiplodia abromozlii 133
incipient wilting 20 Lrptospora r~rbella 133 micronutrients, defined 10
Indian bean thrips 61 Lez~eillrilatalirica 111-112 Micropeltopsis can~zabis 133
Indian stink bug 75-76 light 9-10 Microplit is croceipes 53
injury, defined 1 limonene 22,195 mildew
insects 20-23 Lindoris loplzantkae 83 black 117-11 8
feeding 21-22 lined click beetle 91 downy 106-107
fungal diseases 16 Liocoris tri~~zist~ilatiis
76 powdery 111-112
identification and monitoring 4 Lirionryzn species 77, 78 millipedes 151
life cycle 20-21 locusts 86-89 mites see hemp russet mite; oriental
orders associated with Cannabis long-tailed mealybug 81 mite; privet mites; spider
21 longhorn beetles 72-73 mites; ta ma mite
terminology 20 Loxostecge sticticalis (beet webworm) 59 Mollusca 20
toxin protection mechanisms 23 lucerne (alfalfa) mosaic virus (AMV) molybdenum 157
viruses affecting 13 143 'Monday syndrome' 166
see also rlauzed lnsccts lucerne flea 91 Monera 15
instar, defined 21 Lyd~llathornpsonii 48 Monolepta dichroa 67
integrated pest management (IPM) Lygiidineolaris (tarnished plant bug) Mordellistena species 72
2-3,6 74 mosaic, defined 13
monitoring Lycgzis species 74 moult, defined 21
environmental 6 Mucor species 172
pests/disease 4-6 multivoltine, defined 21
post-intervention 6 Macrocentrzls ancyclivorzis 50 mycelium, defined 16
Iphiseilis degenenlns 63 Macrolophz~scaligilzoszis 43 mycoplasma-like organisms (MLOs)
IPM see integrated pest management macronutrients, defined 10 15,146-147
iprodione 209 Macropliomina pliaseolina mycorrhizae 16,130-131
iron 156-1 57 charcoal rot 112-113 Myrotlzecizrm species 133, 141
Iwanowsky, Dmitri 13,142 damping off 98,99 Myxomycophyta 16
Macropsis cannabis 80 Myzzls persicae (green peach aphid) 32
magnesium 156,160
1a1111i('11a bdle~liicn 132 Maladera kolosericea 68
Japanese beetle 67, 68, 69 Malanclzm persicariae (dot moth) 57 needle nematode 140
Java root knot nematode 138 malathion 208 neem 195
jul~enoids199 Manzestm species 55,56,58 nematodes 18-20,137-142
mammals 23,152-153 biological control 20, 141
mancozeb 208 chemical control 141-142
Kirchner, Oskar 7 maneb 208-209 cultural/mechanical control
Klcbsiella ptmvtoniae 171 manganese 156 140-141
Ma~ztisreligiosa (praying mantid) 181 cyst 138-139
marigold 141 differential diagnosis 140
lacewings 35-36 marijuana thrips 61 identification 6
ladybeetles marsh beetle 73 needle 140
aphid control 36 mealybug destroyer 82-83 pesticide effects 183
beneficial 29 mealybugs 81-84 root knot 137-138
mealybug/scalc control 83 Melarnpsora cannabis 123 root lesion 140
pesticide effects 183 Melnnogronzyza 11 rticivora (nettle stem 139-140
spider mite control 28-29 midge) 85 white root grub control 70
 Index

Nernnfopllthora g?/noplzila 141 parasites, facultative 16 pillbugs 151

Neoseirlllrs species 27-28, 62-63 parasitoids 3,182 pinene 22
Ncozygitcs parz?ispora 66 parathion 208 pink rot 119-120
nettle midge 85 ~artizenolecanizlnzcorni (European fruit piperonyl butoxide (PBO) 197
Nczara z~iridz~la (southern green stink lecanium) 81 pirate bugs 63-64
bug) 73, 74 pathogens, defined 1 pirimicarb 209
nicotine 195 Pasfeltria penetrans 141 plant bugs 73-76
nitrogen 155-156 pawpaw extracts 196 plant lice see aphids
Nornzlrn~arileyi 53 PCP (pentachlorophenol) 205 plant-pesticides 190
Northern root knot nematode 137 pebulate 209 defined 189
Nost'r~zaspecies 48, 70, 88 Penicillizlm species 133, 172 Plantae 17-18
nuclear polyhedrosis viruses (NPV) pentachloronitrobenzene (PCNB) planthoppers 80
53,55-56,59 205-206 plants
nutrients 10 pepper spot 128-129 allelopathic 148
deficiencies 6,155-157 Periconia byssoiides 134 parasitic 148-150
in fertilizers 163 Peristenlis digonezltis 76 tissue analysis 155,162
fertilizing field crops 161-162 permethrin 198 weeds 18,147-148
fertilizing glasshouse soils 162-163 Persoon's broomrape 150 Platap Iecfn consangzlis (hemp dagger
functions of 12 Pestalotiopsis species 134 moth) 59
overfertilization 157 pesticides Podagrica aerafa 66
plant analysis 155 application methods 202-204 Podibiig 59
released by retting 163 categories 190 Podiszls nzaczllizlentris 60
soil balancing 159-161 foliar treaments 202-203 pollutants 163-164
and soil pH 1 0 , l l formulations 189,190 polygodial 196
toxicity 155 plant 190 Polyg(~rzz~m c o n u o l z ~ z ~(bindweed)
l~~s 147
Nysitls rricac (false chinch bug) 74 regulation 199-200 polyphagous, defined 19
residues 191 polyvinyl chloride 205
resistance rebound 191 Popillin japonica (Japanese beetle) 67,
obligate parasites 16 safety 200,201 68,69
obligate saprophytes 16 seed treatments 202 potassium 156
occlusion bodies 13 side effects 190-191 potato bug 75
Odorztoterrnes obesrls (termite) 84 soil application 202 potato leafhopper 79
oestrogenic compounds 191 synthetic chemicals 205-210 powdery mildew 111-112
oils 194, 196 toxicity to biocontrol organisms Pratylencliiis penetrans (root lesion
olive leaf spot 113-114 183,187 nematode) 140
onion thrips 61 pests 2 praying mantid 181
Ooerzc~jrt~ls s~ibr~zctalliczis
76 associated with Cannabis 14 privet mites 31
Oomycota 16 autocidal control 184 propargite 207
O y n t r ~ r msablilos~rrrr 67 control 175-179 Protoctista 15-16
Oplri(7bolzls stem canker 125-126 identification and monitoring Protozoa 16,151
Opills pallipcs79 4-6 Psezrdaiilacaspis pentagons (white peach
Orbiln llrteola 134 screens 178-179 scale) 82
organic farming 2-3 traps 178 Pseiidoacfeon species 84
organic matter 11,158-159 PFR (Paecilomyces fz~nzosorosezis) 43 Psc~idocerccsporacarlnablna 113
organochlorines 205-207 pH, soil 10,ll Pseudococczis longispinzis (long-tailed
organophosphates 191,207-208 Phasrnarlzabditis herrnaplzrod ita mealybug) 81
oriental mite 31 151-152 Pstwdonzonas species
Orills species (pirate bugs) 63-64 pheromones 198 for biocontrol 100,109, 141,
Orobanclze species (broomrapes) Plzilaenzrs spzrrnarizis (spittlebug) 80 173
150-151 Phonza species 104-106 pathogenic 144-146
Ost rinia nil bilalis (European corn Phomopsis species 127-128 Pselrdoperonospora species 16, 106
borer) 44-45 Phomopsis stem canker 127-128 Psyche carzrzabinella (hemp bagworm)
Ostrinia scapulalis 45 Phorodon species (aphids) 33 52
Ozllcmn rrlelanoya 67 phosphorus 156 Psylliodes species (flea beetles)
overwatering 164 phylloplane, defined 15 65-66
O x y tlirips cannal7cnisis (marijuana phylloplane fungi 129-130 purslane sawfly 90-91
thrips) 61 Phyllotreta nenzortlm 66 Pyemotes fritici 84
Phyrnatotrichopsis ornniuora 125 pymetrozine 210
physcion 196 pyrethroids 197
'P-squared', spider mite control 27 Phytomonas 151 pyrethrum 196
paddy cutworm 54-55 Phytornyza /zorficola 77 pyridaben 210
Pa~cilor~zyces
fiimosorosezls (PFR) 43 Plzytophthora infestans 97 Pyrrhocoris apterzrs 73
Pnpnipeina species (boring caterpillars) phytoplasmas 15,146-147 Pythillnz species 16,97-100
50-51 Phytoseizilzls macroyilis 27
Parnlongidortis nzaximzls (needle nema- Phytoseiz~lzispersimilis 27, 183
tode) 140 Pichia gziillierrnondii 173 quassia 196
 Index

red boot 125 silver Y-moth 58 Streptococczrs species 171

red squill 196 silverleaf whitefly 40 Streptomyces species 100
redbanded leafhopper 79 Simpleton's Key 5 streptomycin 197
RE1 (restricted-entry interval) 201 Sitona species 71 striatura ulcerosa 145
relative humidity 9 slime moulds 16 strychnine 196
repellent, defined 189 slugs 20,151-152 stunt virus 52
resistance 1-2,176-177 Smintlzzrrils uiridis (lucerne flea) 91 stylet, defined 19
retting 163,167-168 snails 20,151-152 sugar beet cyst nematode 138
Rllabdospora cannabinn 134 soap 194 sulphur 156,161,193
Rhinonclis pericarpllrs (hemp weevil) sodium bicarbonate 193 sun scald 165
71 sodium fluoroacetate 210 sustainable agriculture 2
Rhizoctonia species 98,99,102-104 sodium hypochlorite 193 sweetpotato whitefly 40
rhizoplane, defined 15 sodium nitrate 194 Sycantrs collaris 76
Rlzizoy 11s species 173 soil 10-11,12 symphylans ,151
Rl~yzobil~sventralis 83 balancing 159-161 Synacra pa~lperi 90
Ricnnia japonicn 80 components 10-11,159 syrphid flies 89
ring spot, defined 13 cultural/mechanical pest control systemic pesticides 190,205
Rodolio cardinalis (vedalia) 82 175-177
root knot nematodes 137-138 field crop fertilization 161-162
root lesion nematodes 140 glasshouse 162-163 T-2toxins (trichothecenes) 108
root maggots 85-86 pesticide application 202 ta ma mite 31
Rosellil~ianecatrix 134 pH 10,11 Tagetes species 141
rosette, defined 13 Solenopsis genlinata (red fire ant) 84 Tnlaromyces flaz~zls 123
rot, defined 15 soluble powder, defined 189 Tanacetum uzrlgare (common tansy) 60
rotenone 196 southern blight 116-117 tarnished plant bug 74
Rotylend~risspecies 140 southern green stink bug 73,74 teleomorph, defined 17
roundworms see nematodes southern root knot nematode 137 temperatures 6
R~irizinadecollnta 152 sowbugs 151 termites 84
rust 123 Splzaeria cannabis 7,131 terpenoids 3,16,22-23,195
ryania 196 Splzaerotheca maczrlavis 111 tetracycline 197
spider mite destroyers 28-29 tetrahydrocannabinol (THC)
spider mites 25-26 insect protection mechanisms
sabadilla 196 biological control 26-30 23
sachet 62 biorational chemical control 30 toxicity in mammals 23,195
Sacc/zaropolyspora spinosa 57 cultural / mechanical control 26 Tetranydzus species (spider mites) 25
Saissetia cofiae (hemispherical scale) spiders 151 Tettigonia species 87
82 Spilosoma obliqzra (common hairy cater- Texas root rot 125
Snlnloilella rnl/enclzen 171 pillar) 59 THC see tetrahydrocannabinol
salt, excess 163 spinosad 197 thiram 209
sand, defined 10 spiracles, defined 20 T/zermoactinomyces species 171
sanitation 175 spittlebug 79,80 Tkerodiplosis persicae 29
saprophytes 16 Spudoptera species (armyworms/cut- Thripobiils serniltrtezrs 64-65
Saran Wrap 205 worms) 54-55 thrips 60-62
sawflies 90-91 Sporidesnlitinz sclerotiuorirm 97 biocontrol 62-65
scales 81-84 spray adjuvants 202 chemical control 65
Scambus pteroylzori 72 spreader (wetting agent) 202 cultural/mechanical control 62
scarab beetle grubs 68-70 springtails 91 Thrips tabaci (onion thrips) 61
Sckiffnerlila cannabis 117, 118 sprinkled locust 86 Thyestes gebleri (hemp longnorn
Scllizocerella pilicornis (purslane Stac/zybotrys Iobirlata 168 beetle) 73
sawfly) 90-91 stalk borer, common 50 Tibetan medicine 173
Scirtes japoniczis (marsh beetle) 73 Steinernema species 56,67,69-70,89, tiger moth, garden 58
Scl~rotiniasclerotior~ir~~
96-97,171 90 Tipkia species 70
sclerotium, defined 17 stem nematodes 139-140 Tipzlla species (crane flies) 89,90
Sclcrotil~ilzrolfsil 117 Stemylzyliunt species (leaf/stem spot) tobacco broomrape 150
Sczrtcllorlelrza species 140 115-116 tobacco mosaic virus 142
Sczitigcrella irrznlaczrlnta (symphylan) Stenocralzlls qial~dainirs80 tobacco thrips 60
151 Stetlzorlrs species (spider mite destroy- tobacco whitefly 39-40
seed see hemp seed ers) 28-29 tolclofos-methyl 208
seedcorn maggot 85 sticker (spray adjuvant) 202 Torzlla lzerbarzrm 135
selectivity ratios 187 Sfictoceplzala bi~bal~rs
(buffalo treehop- toxins 163-164
selenium 157 per) 80 trace elements 11,12
semiochemicals 198-199 stink grasshopper 87 Tracheophyta 18
septa, defined 16 storage diseases/pests 171-173 trap plants 184
Septoria species (yellow leaf spot) biocontrol 173 triadimefon 210
101-102 chemical control 174 Trialezrrodes z~aporariorirm(greenhouse
silt, defined 10 cultural/mechanical control 173 whitefly) 39
 Index

Tric/ziocnnzpzls cat171nbis(hemp sawfly) vaporizer devices 174 white peach scale 82

YO Vavilov, Nikolai 7,8 white root grubs 68-70
Trichodrnna species 96, 100, 103, 122, vedalia 82 whiteflies 39-40
123,135 Verticilliz~nzalbo-atrilm 122 biocontrol 40-44
Trrc7zo~yralrzirzaspecies 46-47, 49-50, Verticillirrm clzlanrydosporiz~n~141 wild buffalo gourd 197
181 Verticilliz~rrldaliliae 122 wildfire 145-146
trichomes 22,170-171,185 Verticilliilm lecanii 38, 43, 61, 183 wireworms 91
Tricllopoda pcnnipes 76 Verticillium wilt 122-123 Wisconsin leaf spot 146
trichothecenes 108 vinclozolin 209 witch's broom 13
Trlcllothecilinl rosellm 119-120 viral diseases 13,142-144 woolybears 58
triforine 210 affecting insects 13 Worker Protection Standard (WPS)
Trlssolcus basalis 76 biocontrol 13 201-202
tumbling flower beetles 73 identification 6
twig blight 118-119 symptoms of 13
t.rt.0-spotted ladybeetle 29/36 viroids 13 Xanthomonas blight 145
two-spotted spider mite 25 viruses 13,14 Xanthoproclzilis faiitzzrs 71
two-striped grasshopper 86 visual scales 4 Xe~zorhabdllsnematophiliis 15
T y p l ~ l o d r o ~ ~canlzab~s
lz~s (ta ma mite) 33 von Schweinitz, Lewis David 7

yellow leaf spot 101-102

ultra~rioletlight 10 warfarin 210
univoltine, defined 21 water
urea 194 growth requirements 9 zearalenone 108
U re do kriegeria na 123 as pest control 193 Zetzellia mali 30
~irine,human 194 water potential 9 Zeirzera rtlzlltistrigata 50
Llrc~lcllcotljaceae 33 weed plants 18,147-148 zinc 156
Llrorrzyces i ~ z c o n s p i c z123
~~~~ weevils 70-72 zineb 209
western flower thrips 61-62 zoochory, defined 22
wettable powder, defined 189 zygospores, defined 17
white leaf spot 128