Insects and Fungi Associated With Cardus

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INSECTS AND
FUNGI
ASSOCIATED WITH
CARDUUS THISTLES
(COMPOSITAE)
CIRCULATING COPY
AGRICULTURE LIBRARY
UNITED STATES
(C^l) DEPARTMENT OF
AGRICULTURE
TECHNICAL
BULLETIN
NUMBER 1616
PREPARED BY
SCIENCE AND
EDUCATION
ADMINISTRATION
ABSTRACT
Batra, S. W. T., J. R. Coulson, P. H. Dunn, and P. E. Boldt. 1981.
Insects and fungi associated with Carduus thistles (Compositae). U.S. Department of Agriculture, Technical Bulletin No.
1616, 100 pp.
Six Eurasian species of Carduus thistles (Compositae: Cynareae)
are troublesome weeds in North America. They are attacked by
about 340 species of phytophagous insects, including 71 that are
oligophagous on Cynareae. Of these Eurasian insects, 39 were ex
tensively tested for host specificity, and 5 of them were sufficiently
damaging and stenophagous to warrant their release as biological
control agents in North America. They include four beetles: Altica
carduorum Guerin-Meneville, repeatedly released but not estab
lished; Ceutorhynchus litura (F.), established in Canada and
Montana on Cirsium arvense (L.) Scop.; Rhinocyllus conicus (Froelich),
widely established in the United States and Canada and beginning
to reduce Carduus nutans L. populations; Trichosirocalus horridus
(Panzer), established on Carduus nutans in Virginia; and the fly
Urophora stylata (.), established on Cirsium in Canada.
Potentially useful and probably host-specific fungi attacking
Carduus include five species of Puccinia and two species of Uredo.
KEYWORDS: Biological control, Carduus, Cirsium, phytophagous
insects, rust fungi, thistles, weed control, weeds.
A terminal inflorescence of musk thistle (Carduus nutans) with its introduced

biological control agent, the seed-destroying European weevil (Rhinocyllus
conicus).
INSECTS AND
FUNGI
ASSOCIATED WITH
CARDUUS THISTLES
(COMPOSITAE)
by
S. W. T. BATRA, J. R. COULSON,
P. H. DUNN, AND P. E. BOLDT
UNITED STATES
DEPARTMENT OF
AGRICULTURE
TECHNICAL
BULLETIN
NUMBER 1616
PREPARED BY
SCIENCE AND
EDUCATION
ADMINISTRATION
ACKNOWLEDGMENTS
We thank the following colleagues for providing unpublished in
formation included in this publication: P. Harris and M. G. Maw,
Agriculture Canada, Regina, Saskatchewan; G. M. Baloch and H.
Zwblfer, Commonwealth Institute of Biological Control, Rawal
pindi, Pakistan, and Delemont, Switzerland; L. T. Kok and R. D.
Hendrick, Virginia Polytechnic Institute and State University,
Blacksburg; J. D. Hacker, West Virginia Department of
Agriculture, Charleston; S. M. A. Kazmi, Peshawar University,
Pakistan; and L. A. Andres, K. E. Frick, B. Puttier, M. K. McCarty,
and H. L. Parker (retired), Science and Education Administration,
U.S. Department of Agriculture.
CONTENTS
Page
Economic importance, control, and bionomics of Carduus
thistles
Biological control of Carduus thistles
Phytophagous insects associated with Carduus thistles
List of insects
Collembola
Thysanoptera
Dermaptera
Orthoptera
Hemiptera-Heteroptera
Hemiptera-Homoptera
Coleoptera
Lepidoptera
Diptera
Fungi associated with Carduus thistles
List of specific fungi
Uredinales
Ustilaginales
Summary
Literature cited
Index
Fungi
Insects
Plants (Carduus species)
1
5
6
10
10
10
11
11
12
18
22
62
72
81
83
83
85
85
86
96
96
96
99
INSECTS AND
FUNGI
ASSOCIATED WITH
CARDUUS THISTLES
(COMPOSITAE)
By S. W. T. Batra, J. R. Coulson, P. H. Dunn, and P. E. Boldt1
ECONOMIC IMPORTANCE, CONTROL, AND

BIONOMICS OF CARDUUS THISTLES
The genus Carduus (Compositae: Cynareae: Carduinae) includes
species in the subgenera Alfredia and Afrocarduus Kazmi (1963,
1964),2 which are not weedy. The Palaearctic subgenus Carduus in
cludes 70 species, 59 apparent species hybrids, and 60 subspecies,
according to the revision by Kazmi (1964); some species are noxious
weeds. An additional species, four apparent hybrids, and one
subspecies of Eurasian Carduus were included in the "Index
Kewensis" in 1974. Although similar to Cirsium thistles, Carduus
species may readily be distinguished from them by their simple,
not feathery, pappus hairs and lack of resin streaks on involucral
leaves (Kazmi, 1964).
Infestations of some introduced Eurasian species of Carduus
(plumeless thistles) of actual or potential economic importance oc
cur in 18 percent of the counties in the contiguous United States.
Musk or nodding thistles (C. nutans and C. thoermeri)3 occur as
economic infestations in over 10 percent of all counties, primarily
in the Midwest, and appear to be spreading (Dunn, 1976). They oc
cupy about 2 million acres in the United States, and chemical con
trol alone cost about $6.50 per acre in 1976 (M. K. McCarty, pers.
1S. W. T. Batra and J. R. Coulson, Beneficial Insect Introduction Laboratory,
Beltsville Agricultural Research Center, Beltsville, Md. 20705; P. H. Dunn,
Biological Control of Weeds Laboratory, Albany, Calif. 94706; and P. E. Boldt,
Biological Control of Weeds Laboratory, Rome, Italy.
2The year in italic after an author's name refers to Literature Cited, p. 86.
3For complete scientific names with authority or describer, see index (plants).
TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE
commun.). Plumeless thistle (C. acanthoides) is a problem in the Ap

palachian region; Italian thistle (C. pycnocephalus) and slender
flower thistle (C. tenuiflorus) are economically important in Califor
nia; and curled or welted thistle (C. crispus) is common in western
Virginia (Johnson, 1974; Dunn, 1976).
These large, spiny, annual or biennial plants are particularly
troublesome in pastures, where they crowd out forage plants (fig. 1),
and in fallow fields and along railways or roadsides, where mowing
or spraying with herbicides may be uneconomical or impractical,
allowing Carduus thistles to become a seed source for reinfestation
of treated areas. They may invade cultivated areas as the use of the
no-tillage method of cultivation increases.
The Carduus thistles from Eurasia also have become problem
weeds where introduced into other continents. Carduus nutans and
C. thoermeri are important weeds in South America and have re
cently been found in eastern Australia. In New Zealand, C. nutans is
common (Doing et al., 1969). The slender thistles (C. pycnocephalus
and C. tenuiflorus) are the major weeds in sheep pastures in
Tasmania, but they may be effectively controlled by grazing
management (Bendall, 1973, 1974). Carduus pycnocephalus is an im-
PN-7015
Figure 1. Dairy cattle in a typical overgrazed pasture infested by Carduus nutans in
Pennsylvania. Not only are the thistles not eaten, but the cattle do not reach for
the edible plants growing near them.
INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES
portant introduced weed in South Africa (Henderson and Ander

son, 1966); C. nutans also occurs there, where it is used for making
bows (Kazmi, 1964). In Canada, C. crispus, 2N=16, C. nutans (ssp.
nutans, leiophyllus, macrolepis), and C. acanthoides are present, as
well as the hybrid C. x orthocephalus, 2N = 17 = 21 ( = nutans, 2N= 16;
acanthoides, 2N = 22) (Mulligan and Frankton, 1954; Moore and
Frankton, 1974). These hybrids, resembling C. acanthoides, have
become numerous and appear to have been ecologically and
gametically selected for local survival in rocky pastures in Grey
County, Ontario (Moore and Mulligan, 1956, 1959; Mulligan and
Moore, 1961). In Nebraska, "musk" thistles are C. thoermeri (M. K.
McCarty, pers. commun., 1978), not hybrids of C. nutans and C.
acanthoides (Fuller, 1969). The identification and descriptions of
these thistles are discussed by Kazmi (1964), Mulligan (1965), Furrer
and McCarty (1966), Wunderlin (1969), McCarty et al. (1973),
Johnson (1974), and Moore and Frankton (1974).
The readiness with which Carduus species apparently interbreed
could lead to the development of some hybrids that are even more
adaptable and agressive weeds than their parents because of the
enrichment of variation through introgressive hybridization. This
and related problems in other weeds are discussed by Baker (1965).
However, hybridization may also provide an opportunity for con
trol by introducing clones relatively susceptible to natural enemies
or herbicides.
In addition to the known hybrid C. x orthocephalus mentioned
previously, possible European hybrids of the weedy species
already present in the United States and Canada are as follows ac
cording to morphological studies of Kazmi (1964):
Species in North America
C. acanthoides
C. acanthoides
(2N = 54)
C. crispus
C. crispus
C. crispus
C. nutans
C. nutans
C. pycnocephalus
x C. crispus (2N = 16)

x C. pycnocephalus
x C.
x C.
x C.
x C.
x C.
x C.
nutans
tenuiflorus (2N = 54)
thoermeri (2N = 16)
pycnocephalus
tenuiflorus
tenuiflorus
European 'hybrids'
C. x leptocephalus
not named, morphologically
resembles C. acanthoides
C. x dubius
C. x crispo- tenuiflorus
C. x semiperegrinus
C. x pernutanti-pycnocephalus
C. x mixtus
C. x theriottii
More cytogenetic and ecological studies of possible Carduus

hybrids are much needed.
Because of its economic importance, considerable basic research
has been undertaken on the bionomics and control of the musk
thistle in North America (see Feldman et al. (1968), McCarty et al.
(1969), McCarty and Scifres (1969), and McCarty and Hatting (1975)).
It was evidently introduced at ballast dumps on the east coast
(Dunn, 1976) and in Alabama between about 1850 and 1900
(specimens at U.S. Natl. Mus.), and it was considered a weed in the
mid-Atlantic States by the early 1900' s (Furrer and McCarty, 1966).
It rapidly spread, becoming economically important in the Midwest
by 1950 (Furrer and McCarty, 1966).
This adaptable plant is now abundant in North Dakota, Loui
siana, and on both coasts, but the most serious infestations are in
the Cental States (Dunn, 1976). In Nebraska, it is found primarily in
rich, moist lowland, although it will occupy poor soils (McCarty,
1964; McCarty et al., 1973), and in Ontario, it occurs on well-drained
loam or stony pastures (Mulligan and Moore, 1961). In western
Virginia, it is common in the Appalachian region, but it is scarce in
the Piedmont or coastal plain (Johnson, 1974). In Maryland and
south-central Pennsylvania, musk thistle is locally abundant with
C. acanthoides in pastures and on roadsides in the Appalachian
Great Valley. Seeds may have been initially spread by wind,
vehicles, and in soil from infested quarries. According to Hensley
(1973), C. nutans and C. acanthoides are most plentiful in limestone
soils in Virginia. In Ohio, C. nutans is most common where
limestone or dolomite bedrock is less than 6 feet below the surface
(Stuckey and Forsyth, 1971).
In its native Europe, musk thistle is a ruderal pioneer in dis
turbed environments on calcareous, loamy, or sandy soil rich in
nitrogen; however, it may become a problem in overgrazed
pastures. According to Doing et al. (1969), the most significant
characters restricting its distribution are low tolerance for lack or
excess of moisture and of deficient or acid soils and an only
moderately high competitive power.
Musk thistles may be controlled by grazing management in main
taining a dense mat of perennial grasses, but the release of grazing
pressure on annual grassland has resulted in an increase of these
thistles in Australia (Doing et al., 1969). However, in Nebraska,
Feldman et al. (1968) found that musk thistles did not become
established in ungrazed pastures; survival was highest in inten
sively grazed pastures. This thistle may behave as a short-lived
perennial, producing new shoots when grazed or sprayed in the
spring (Doing et al., 1969), and may thus increase difficulty of con
trol. Treatment with the herbicides dicamba (3,6-dichloro-o-anisic
acid) plus 2,4-D (Jensen, 1970; McCarty and Hatting, 1975) or
picloram (4-amino-3,5,6-trichloropicolinic acid) (Feldman et al.,
1968; Jensen, 1970) was fairly effective. Control for 2 years cost
from $6.25 to $14.75 per acre (Jensen, 1970). Mowing musk thistles
within 2 days of the first terminal heads in a population showing

anthesis eliminated production of viable seed from all mowed
stalks, but repeated mowing is necessary owing to differences in
plant maturity (McCarty and Hatting, 1975).
Carduus thistles have been little used by people. The young stalks
are edible and the pappus has been used for papermaking (Moore
and Frankton, 1974). They are not considered important to
vertebrate wildlife in North America (Martin et al., 1961). Seeds of
C. nigrescens are a rich source of pentacyclic triterpenes (Madrigal
et al., 1974); C. albidus was tested but found lacking in ecdysones
(Ganiev, 1975). The fragrant, beautiful flowers of C. acanthoides
(white and purple varieties) and C. nutans are occasionally visited
by insects (Mulligan and Kevan, 1973). At Beltsville, Md., bumble
bees and halictine bees are important pollinators of these plants.
According to Crane (1975), honey bees working Carduus and Cirsium
thistles provide a light, sweet honey of good flavor; Carduus
hamulosus gives a moderate honey yield (26-50 kg per hectare). The
elaiosome-bearing seeds of the myrmechore C. pycnocephalus are
transported by ants (Uphof, 1942).
BIOLOGICAL CONTROL OF CARDUUS THISTLES
Because of the expense of controlling Carduus thistles by mowing
or applying herbicides in areas where they are prevalent, such as
pastures, fallow fields, wasteland, quarries, roadsides, or railways,
biological control by arthropods or pathogens is a reasonable alter
native or adjunct. It has the advantage of being relatively inexpen
sive and harmless to nontarget organisms, and biological control
agents can multiply and spread to adjacent infested areas. The
principles and procedures of biological control have been well
defined and tested with some spectacular successes (see Huffaker
(1956), Wilson (1964), Harris and Zwolfer (1968), Andres (1971), Har
ris (1971), Zwblfer and Harris (1971), Harris (1973a, b), Wapshere
(1974), Andres and Bennett (1975), and Goeden and Louda (1976)).
Carduus thistles are prime candidates for biological control
because (1) it is often impractical or too expensive to mechanically
destroy or spray them; (2) they have been introduced from another
continent, leaving behind their natural enemies; (3) they are usually
biennial; (4) they build up dense populations, maintained for
several years; (5) they are not closely related to major crop plants
except artichoke, sunflower, and safflower; and (6) they have little
value to people or wildlife.
Biological control increases environmental pressure on Carduus
thistles. At best, the natural enemies may eliminate the need for
other control methods over much of the plant's range and form a
sound basis for future weed-management schemes. At least these
natural enemies would augment existing control practices. A com
prehensive list of the natural enemies of Carduus species is a needed
step in the process of selecting promising candidates.
Biological control of Carduus thistles in North America began
with the introduction of the weevil Rhinocyllus conicus (Froelich),
which was first released in Canada in 1968 after 6 years of study
and testing in Europe and North America by scientists of the Com
monwealth Institute of Biological Control, the Canada Department
of Agriculture, and the former Agricultural Research Service (now
a part of the Science and Education Administration (SEA-AR)). It
is beginning to provide a significant degree of control of C. nutans
in Virginia (Kok and Surles, 1975) and in Montana (Hodgson and
Rees, 1976). Rhinocyllus conicus has also been established in Califor
nia to control milk thistle (Silybum marianum (L.) Gaertn.) (Hawkes
et al., 1972) and in New Zealand to control C. nutans (Jessep, 1975).
Assistance in locating sources of approved agents for biological
control of Carduus thistles can be provided by the U.S. Department
of Agriculture's Beneficial Insect Introduction Laboratory,
Building 417, Beltsville Agricultural Research Center-East,
Beltsville, Md. 20705; USDA Biological Control of Weeds
Laboratory, 1050 San Pablo Avenue, Albany, Calif. 94706; and
Agriculture Canada Research Station, Box 440, Regina, Sas
katchewan S4P 3A2.
PHYTOPHAGOUS INSECTS ASSOCIATED WITH

CARDUUS THISTLES
The first step in any project for the biological control of weeds is
to prepare a list of natural enemies associated with the target plant.
A list has been compiled here to give interested workers available
information on the biological control of weeds. It incorporates data
extracted from publications, as well as from reports, letters, and
other unpublished material located in the files of the Insect Iden
tification and Beneficial Insect Introduction Institute (IIBIII),
SEA, USDA, Beltsville, Md., with few exceptions, as of December
31, 1976. 4 Records from museum specimens have not been includ
ed, and the taxonomic literature is not exhaustively cited because
this is not intended as a bibliography. Data on Carduus thistle in4Although most data on which this publication is based were collected prior to
1977, the information is still valid and useful as guidelines for those interested in
the biological control of weeds.
sects not in the Beneficial Insect Introduction Laboratory (BIIL)

files are not summarized here. The compilers do not claim to have
been able to detect or correct all possible inaccuracies in the
sources available.
This list is to serve only as a source of information to aid in fur
ther studies and to make known some of the insects discovered on
Carduus and what studies have been or are being conducted. It is
not intended to serve as a final authority for determining host
plants, geographical ranges, and so forth. Detailed information on
any of the insects or laboratory tests should be obtained from those
laboratories from whose reports or publications the pertinent data
have been extracted. An unpublished preliminary version of this
list, circulated by J. R. Coulson in 1969, has been cited, e.g., in
Goeden (1974, 1976). An index to the Carduus species mentioned in
the list and their associated insects and fungi is included.
Names of insects. The names of insects have been checked by
specialists of the Systematic Entomology Laboratory, SEA-AR.
Plant records.All species of Carduus (as recognized by Kazmi
(1964) and in litt. (1977)) are listed here with the particular insects
with which they have been associated or recorded. This association
may not always indicate a specific host plant relationship, but it
may sometimes be simply a record of the insect casually feeding on
the species in the field or in a laboratory test. In the field records,
the location usually follows the plant species in parentheses. Ex
planations of these records are to be found under Remarks. These
comments refer also to the records on plant genera other than
Carduus. However, in these instances, only the generic name of the
plant is given, and the record may refer to one or more species of
that genus.
Remarks. This section is a brief summary of available informa
tion on the insect from laboratory and field studies.
Sources. The sources of unpublished information are referred to
by the names of the stations reporting the information and the
years of the reports. When data are from many sources, each item is
not specifically attributed to a definite source. If such information
on a specific source is desired, it can be supplied by the authors.
The laboratories that have kindly allowed the use of information
from their unpublished reports for this list include the Com
monwealth Institute of Biological Control stations of Delemont,
Switzerland, and Rawalpindi, Pakistan; the SEA-AR Biological
Control of Weeds laboratories at Rome, Italy, and Albany, Calif.;
the BIIL, Beltsville, Md.; the Biological Control of Weeds Research
Group of Agriculture Canada at Regina, Saskatchewan (formerly at
Belleville, Ontario); and the Department of Entomology, Virginia
Polytechnic Institute and State University, Blacksburg. Data from

reports of several P.L. 480 projects on biological control of weeds
supported by SEA-AR are also included in this list; e.g., projects at
Rawalpindi, Pakistan (A17-ENT-9 and A17-ENT-14 terminated in
1965 and 1970, respectively) and at the Ministry of Agriculture,
Dokki, Egypt (F4-ENT-5 and F4-ENT-16, completed in 1970 and
1974). Much of the data included in these unpublished reports has
since been published, e.g., in Zwblfer (1965a, b) and Baloch et al.
(1971), and efforts have been made here to refer to the published
source. However, some other records from the unpublished reports
have also been included in this list for completeness.
Abbreviations used here are as follows:
General Abbreviations
A - Adults
Addit. - Additional
Agr. - Agriculture
Ann. - Annual
ARS - Agricultural Research
Service (now SEA-AR)
Centr. - Central
CIBC - Commonwealth Institute of Biological
Control
Coll. - Collected
Dept. - Department
Distrib. - Distribution
E. - East, eastern
Et al. - and others
Ft. - Fort
Ident. - Identified, identification
Incl. - Includes, including
In litt. - In correspondence
Inst. - Institute
Is. - Island
Km - Kilometers
L - Larvae
Lab. - Laboratory
Locs. - Locations, localities
Misident. Misidentified, misidentification
Mts. - Mountains
N. - North, northern
Nat. - National
No(s). - Number(s)
Nr. - Near

Pers. comm. - Personal communication
Poss. - Possibly
Prob. - Probably
Rec(s). - Record(s), recorded
Res. - Research
Rpt(s). - Report(s)
S. - South, southern
SEA-AR - Science and Education
Administration-Agricultural
Research
Sp(p). - Species
Spec. - Special
Ssp. - Subspecies
SW. - Southwest, southwestern
Unident. - Unidentified
USDA - United States Department of
Agriculture
Var. - Variety
VPI - Virginia Polytechnic Institute and State
University
Vs. - Versus
W. - West, western
Yr(s). - Year(s)
Locality Abbreviations
Countries
U.S. - United States
USA - United States of America
USSR - Union of Soviet Socialist Republics
Canadian Provinces
Alta. - Alberta
B.C. - British Columbia
Man. - Manitoba
N.B. - New Brunswick
N.S. - Nova Scotia
Ont. - Ontario
Que. - Quebec
Sask. - Saskatchewan
United States
Amer. - America
Calif. - California
Colo. - Colorado
Del. - Delaware
10
United StatesContinued
Ind. - Indiana
Kans. - Kansas
Ky. - Kentucky
La. - Louisiana
Md. - Maryland
Mich. - Michigan
Minn. - Minnesota
Mo. - Missouri
Mont. - Montana
N. Dak. - North Dakota
Nebr. - Nebraska
Nev. - Nevada
N.J. - New Jersey

N.Y. - New York
Okla. - Oklahoma
Oreg. - Oregon
Pa. - Pennsylvania
S. Dak. - South Dakota
Tenn. - Tennessee
Va. - Virginia
W. Va. - West Virginia
Wash. - Washington
Wis. - Wisconsin
Wyo. - Wyoming
LIST OF INSECTS
COLLEMBOLA
Sminthuridae
Unident. sminthurid sp.
Plant recs.: Carduus pycnocephalus (S. Europe)
Remarks:
Occasional adults feeding on leaves
Sources:
Goeden (1974)
THYSANOPTERA
Unident. thrips sp.
Plant recs.: Carduus defloratus (Switzerland); C. nutans (S.
Dak.)
Remarks:
In flowers
Sources:
Delemont Rpt. (1963); Morihara and Balsbaugh
(1976)
Phlaeothripidae
Haplothrips distinguendus (Uzel)
Remarks:
Locally common; adults on flowers;
polyphagous
Sources:
Goeden (1974)
Thripidae
Anaphothrips (Apterothrips) secticornis (Trybom)
Plant recs.: Carduus pycnocephalus (S. Calif.)
Remarks:
Rare; adults feeding on rosette leaves;
polyphagous
Sources:
Goeden (1974)

Frankliniella occidentalis (Pergande)
Remarks:
Rare; adults ectophagous on
polyphagous
Sources:
Goeden (1974)
11
flowers;
DERMAPTERA
Forficulidae
Forficula auricularia L.
Remarks:
Rare; nymphs and adults on leaves and stems;
polyphagous
Sources:
Goeden (1974)
ORTHOPTERA
Gryllidae
Unident. gryllid sp.
Plant recs.: Carduus pycnocephalus (S. Calif.); C. nutans (S.
Dak.)
Remarks:
Rare; nymphs feeding on leaves of rosette
Sources:
Goeden (1974); Morihara and Balsbaugh (1976)
Oecanthus nigricornis Saussure
Plant recs.: Carduus nutans (S. Dak.)
Remarks:
Nymphs and adults ectophagous
Sources:
Morihara and Balsbaugh (1976)
Tettigoniidae
Unident. tettigoniid spp. (3)
Dak.)
Remarks:
Sources:
Acrididae
Unident. acridid spp. (2)
Remarks:
Nymphs and adults ectophagous on leaves
Sources:
Melanoplus sp.
Remarks:
Nymphs and adults ectophagous on leaves;
polyphagous crop pest
Sources:
12
ORTHOPTERA Continued
AcrididaeContinued
Melanoplus bivittatus (Say)
Remarks:
Abundant; nymphs and adults ectophagous on
leaves; polyphagous crop pest
Sources:
Melanoplus differentialis (Thomas)
Remarks:
Abundant; adults ectophagous on leaves;
Sources:
Melanoplus femurrubrum (De Geer)
Remarks:
Abundant; nymphs and adults ectophagous on
Sources:
HEMIPTERA-HETEROPTERA
Miridae
Unident. mirid spp.
Plant recs.: Carduus pycnocephalus (S. Europe); C nutans (S.
Dak.)
Remarks:
Nymphs and adults on foliage
Sources:
Adelphocoris lineolatus (Goeze)
Plant recs.: Carduus crispus (Europe); C. nutans (S. Dak.);
Cirsium
Remarks:
Occasional visitor on Carduus; polyphagous
crop pest
Sources:
Zwblfer (1965a); Morihara and Balsbaugh (1976)
Calocoris norvegicus (Gmelin)
Plant recs.: Carduus crispus (Europe); C. pycnocephalus (S.
Europe); Cirsium
Remarks:
Nymphs and adults from foliage of Carduus;
polyphagous
Sources:
Zwblfer (1965a); Goeden (1974)
Chlamydatus associatus (Uhler)
Remarks:
Adults; ectophagous on Compositae
Sources:
Irbisia sp. nr. J. californica Van Duzee

Remarks:
Sources:
Lopidea sp.
Plant recs.:
Remarks:
Sources:
Lygus hesperus
Plant recs.:
Remarks:
13
Locally common; nymphs and adults ectophagous on flowers, leaves, and stems;
polyphagous
Goeden (1974)
Carduus nutans (S. Dak.)

Adults; uncommon
(Knight)
Carduus pycnocephalus (S. Calif.)
Occasional nymphs and adults ectophagous on
flowers; polyphagous crop pest
Sources:
Goeden (1974)
Lygus lineolaris (Palisot de Beauvois)
Remarks:
Adults ectophagous on stems and flowers;
Sources:
Lygus pratensis (L.)
Plant recs.: Carduus nutans (Europe); Cirsium
Remarks:
Occasional visitor on Carduus; polyphagous
Sources:
Zwblfer (1965a)
Plagiognathus sp. nr. P. confusus Reuter
Remarks:
Adults on flowers
Sources:
Goeden (1974)
Plagiognathus arbustorum (F.)
Remarks:
Occasional visitor on Carduus
Sources:
Zwolfer (1965a)
Plagiognathus obscurus Uhler
Plant recs.: Carduus sp. (Va.); Cirsium (Canada)
Remarks:
Frequent feeder; polyphagous
Sources:
L. T. Kok in litt. (1976); Maw (1976)
Plagiognathus politus Uhler
Remarks:
Eggs, nymphs; ectophagous on Compositae
Sources:
Poecilocapsus lineatus (F.)
Plant recs.: Carduus sp. (Va.); C. nutans (S. Dak.); Cirsium
(Canada)
Remarks:
Nymphs and adults frequent feeders;
polyphagous pest
Sources:
L. T. Kok in litt. (1976); Morihara and
Balsbaugh (1976); Maw (1976)
14
HEMIPTERA-HETEROPTERAOmfmued
MiridaeContinued
Semium hirtum Reuter
Remarks:
Uncommon; adults ectophagous; polyphagous
Sources:
Tingidae
Tingis cardui (L.)
Plant recs.: Carduus defloratus (Switzerland); C. crispus
(Rhine Valley); C. acanthoides (Jura, W.
France); C. pycnocephalus, C. acanthoides
(Europe); Cirsium, Galactites (Europe)
Remarks:
Adults and nymphs found externally on leaves
and stems; literature restricts hosts to CarduusCirsium
Sources:
Delemont Rpts. (1962-63); Zwolfer (1965a);
Eguagie (1974)
Lygaeidae
Unident. lygaeid spp.
Plant recs.: Carduus pycnocephalus (S. Europe); C. nutans (S.
Dak.)
Remarks:
Rare; adults ectophagous on leaves;
polyphagous
Sources:
Geocoris uliginosus (Say)
Remarks:
Rare; adults ectophagous on flowers; on other
Compositae
Sources:
Lygaeus equestris (L.)
Plant recs.: Carduus nutans (Europe); C pycnocephalus (S.
Europe)
Remarks:
Occasional visitor on flowers and leaves;
polyphagous
Sources:
Zwolfer (1965a); Goeden (1974)
Melanocoryphus bicrucis (Say)
Remarks:
Rare; adults ectophagous on flowers and
stems; polyphagous
Sources:
Goeden (1974)
Coreidae
Ortholomus sp.
15
Remarks:
Rare; adults ectophagous
Sources:
Syromastes marginatus (L.) ( = Coreus marginatus)
Remarks:
Occasional visitor
Sources:
Zwblfer [1965a)
Pentatomidae
Aelia sp.
Plant recs.: Carduus crispus (Europe)
Remarks:
Occasional visitor
Sources:
Zwblfer [1965a)
Carpocoris pudicus (Poda)
Plant recs.: Carduus nutans (Europe); Cirsium, Silybum, Cen
taurea
Remarks:
Nymphs and adults found on stem and leaves
Sources:
Zwblfer [1965a)
Carpocoris purpureipennis De Geer (and/or var. pallidus Distant)
Plant recs.: Carduus edelbergii (Pakistan); many other hosts
Remarks:
Polyphagous; "a known pest"; exophytic on
stems and leaves
Sources:
A17-ENT-9 Rpt. [1965); Baloch et al. [1971)
Chlorochroa uhleri St&l
Remarks:
Rare; adults; polyphagous
Sources:
Morihara and Balsbaugh [1976)
Cosmopepla bimaculata (Thomas)
Remarks:
Nymphs, adults on flowers and leaves;
polyphagous
Sources:
Morihara and Balsbaugh [1976)
Dolycoris baccarum (L.)
Plant recs.: Carduus nutans (Europe); Cirsium, Onopordum,
Centaurea, Echinops
Remarks:
Nymphs and adults found on stems and leaves;
polyphagous; attacks crops
Sources:
Zwblfer [1965a)
Dolycoris indicus Stal
Plant recs.: Carduus edelbergii (Pakistan); Carthamus, Xanthium, Calotropis, Cannabis, Cnicus, Salvia,
Ziziphus
Remarks:
Adults suck sap of Carduus and Xanthium;
Sources:
A17-ENT-9 Rpts. [1961, 1965); Baloch et al.
[1968, 1971)
16
HEMIPTERA-HETEROPTERAContmued
PentatomidaeContinued
Eurydema lituriferum Walker
Plant recs.: Carduus edelbergii (Pakistan)
Remarks:
Adults suck sap from leaves of Carduus
Sources:
A17-ENT-9 Rpt. (1965); Baloch et al. (1971)
Eurydema oleraceum (L.)
Remarks:
Occasional visitor on Carduus
Sources:
Zwblfer (1965a)
Eurydema omatum (L.) ( = E. festivum L.)
Remarks:
Exophytic on stems and leaves; polyphagous;
attacks many crops
Sources:
A17-ENT-9 Rpts. (1961, 1965); Baloch et al.
(1971)
Euschistus conspersus Uhler
Remarks:
leaves; polyphagous; pest of Cynara
Sources:
Goeden (1974)
Euschistus euschistoidea (Vollenhoven)
Remarks:
Common; nymphs and adults ectophagous on
flowers and leaves; polyphagous
Sources:
Euschistus tristigmus (Say)
Remarks:
Nymphs and adults ectophagous on flowers
and leaves; attacks other Compositae
Sources:
Evacanthus frepexus Distant
Plant recs.: Carduus edelbergii (Pakistan); many other hosts
Remarks:
Polyphagous crop pest
Sources:
Baloch et al. (1971)
Nezara viridula (L.)
Plant recs.: Carduus edelbergii (Pakistan); C. pycnocephalus
(S. Europe); C. nutans (Va.); Xanthium, Can
nabis, Salvia
Remarks:
Adults feeding (sucking sap) on leaves and
flowers of Carduus; polyphagous crop pest
Sources:
(1968, 1971); Goeden (1974); L. T. Kok in litt.
(1976)
17
Odontotarsus purpureolineatus (Rossi)

Plant recs.: Carduus nutans (Europe)
Remarks:
Occasional visitor
Sources:
Zwblfer (1965a)
Thyanta pallidovirens pallidovirens (Stal)
Remarks:
Rare; adults on stems; polyphagous
Sources:
Goeden (1974)
Piesmatidae
Piesma cinera (Say)
Remarks:
Rare; adults on flowers and leaves;
polyphagous
Sources:
Pyrrhocoridae
Pyrrhocoris apterous L.
Plant recs.: Carduus acanthoides (S. Europe)
Remarks:
Rare; adults on leaves; polyphagous
Sources:
Goeden (1974)
Rhopalidae
Leptocoris sp. nr. L. rubrolineatus Barber
Remarks:
Sources:
Goeden (1974)
Leptocoris trivittatus (Say)
Remarks:
Nymphs, adults ectophagous on leaves, stems,
and flowers; polyphagous crop pest
Sources:
Liorhyssus hyalinus (F.)
Remarks:
Rare; nymphs ectophagous on shoots;
polyphagous
Sources:
Goeden (1974)
Scutelleridae
Unident. scutellerid sp.
Remarks:
Rare; adults eat leaves
Sources:
Goeden (1974)
Eurygaster amerinda amerinda Bliven
Remarks:
Rare; adults ectophagous on leaves;
polyphagous
Sources:
Goeden (1974)
18
HEMIPTERA-HOMOPTERA
Cicadellidae
Aceratagallia sp.
Remarks:
Occasional adults ectophagous on stems,
leaves, and flowers
Sources:
Goeden (1974)
Aceratagallia uhleri (Van Duzee)
Remarks:
Adults; ectophagous
Sources:
Agallia quadripunctata (Provancher)
Plant recs.: Carduus nutans (S. Dak.); Cirsium (Canada)
Remarks:
Rare; adults only; polyphagous
Sources:
Morihara and Balsbaugh (1976); Maw (1976)
Agalliopsis novella Say
Remarks:
Adults; ectophagous; polyphagous
Sources:
Deltocephalus signatifrons (Van Duzee)
Remarks:
Adults; uncommon
Sources:
Elymana sp.
Remarks:
Sources:
Empoasca sp.
Dak.)
Remarks:
Rare; nymphs and adults ectophagous on
leaves
Sources:
Endria inimica (Say)
Remarks:
Nymphs and adults abundant; polyphagous
crop pest; vector of aster yellows and wheat
streak mosaic
Sources:
Euscelis lineolatus Brulle
Remarks:
Sources:
Goeden (1974)
19
Graphocephala coccinea Forster

Remarks:
Sources:
Latalus sp.
Remarks:
Sources:
Macrosteles divisa (Uhler)
Remarks:
Adults ectophagous on Compositae; crop pest
Sources:
Neosteles neglecta Delong and Davidson
Remarks:
Sources:
Paragonia tredecimpunctata Ball
Remarks:
Nymphs and adults common; ectophagous on
leaves and stems; polyphagous
Sources:
Goeden (1974)
Paraphlepsius irroratus (Say)
Remarks:
Adults, nymphs moderately abundant;
polyphagous
Sources:
Psammotettix sp.
Remarks:
Rare; adults
Sources:
Aphididae
Unident. aphid spp.
Plant recs.: Carduus edelbergii (Pakistan); C. nutans
(Europe); C. pycnocephalus (Egypt)
Remarks:
None
Sources:
A17-ENT-9 Rpt. (1961); Zwblfer (1965a); F4ENT-5 Rpt. (1967)
Aphis spp. (2)
Remarks:
Adults and nymphs; ectophagous on stems
Sources:
Aphis craccivora Koch (= A. medicaginis (Koch))
Plant recs.: Carduus pycnocephalus (France, S. Europe);
Tribulus, Cnicus (Pakistan)
20
HEMIPTERA-HOMOPTERAContmued
AphididaeContinued
Remarks:
Sources:
Winter host is Robinia; polyphagous

L. Andres in litt. (1961); A17-ENT-9 Rpt. (1965);
Goeden (1974)
Aphis fabae Scopoli
Remarks:
Occasional nymphs and adults; reproduce on
leaves and stems; polyphagous crop pest
Sources:
Goeden (1974)
Aphis gossypii Glover
Remarks:
Occasional nymphs and adults on flowers,
leaves, and stems; polyphagous crop pest
Sources:
Goeden (1974)
Brachycaudus sp.
Remarks:
Nymphs and adults feeding on leaves of
Carduus
Sources:
A17-ENT-9 Rpt. (1965)
Brachycaudus cardui (L.) ( = Anuraphis cardui, Aphis cardui)
Plant recs.: Carduus pycnocephalus (France, Italy); C. nutans
(Italy); Carduus sp. (Va.); C edelbergii, Cnicus
(Pakistan); Cynara, Cirsium, Senecio
Remarks:
Locally common on flowers and stems; winter
host is Prunus
Sources:
L. Andres in litt. (1961); Baloch et al. (1971);
Goeden (1974); L. T. Kok in litt. (1976)
Brachycaudus helichrysi Kaltenbach ( = Aphis helichrysi (Kalt.))
Plant recs.: Carduus pycnocephalus (Egypt, S. Calif.)
Remarks:
This species severely infests growth of bloom
ing Carduus in Egypt and is preyed on by the
cecidomyiid Phaenobremia aphidivora Rubs.; B.
helichrysi also is known in N. and S. Amer.,
Europe, S. Asia, and N. Africa; winter host is
Prunus; summer hosts are artichoke, sugar
beets, clover, aster, chrysanthemum, iris,
petunia, almond, apricot, peach, plum, and
others
Sources:
F4-ENT-5 Rpts. (1966-67); Goeden (1974)
Capitophorus carduinus (Walker) ( = C. flaveolus (Walk.))
Plant recs.: Carduus nutans, C. acanthoides, C. crispus, Cir
sium (Europe); Cirsium (Canada); Cynara (USA)
Remarks:
Reported on foliage of Carduus; on Cirsium all
year
21
Sources:
Zwblfer (1965a); Maw (1976)
Capitophorus elaeagni (del Guercio) ( = C. braggii Gillette)
Plant recs.: Carduus edelbergii (Pakistan); C. pycnocephalus
(S. Calif.); Cirsium (Canada)
Remarks:
Nymphs and adults feeding on leaves of
Carduus; summer hosts are several Compositae
including Cynara; winter hosts are Elaeagnus
hippophae and Shepherdia
Sources:
A17-ENT-9 Rpt. (1965); Baloch et al. (1971);
Goeden (1974); Maw (1976)
Capitophorus flaveolus Walker
Plant recs.: Carduus acanthoides (Europe); Cirsium
Remarks:
Stenophagous
Sources:
Zwblfer (1965a)
Cerosipha wartenbergi Borner
Plant recs.: Carduus defloratus (Europe)
Remarks:
None
Sources:
Zwblfer (1965a)
Dactynotus aeneus (Hille Ris Lambers)
Plant recs.: Carduus nutans, C. defloratus, C. acanthoides, C.
crispus all year (Europe); Cirsium (Europe)
Remarks:
Nymphs and adults on leaves and stems
Sources:
Zwblfer (1965a)
Dactynotus jaceae (L.) (= Macrosiphum jaceae (L.))
Plant recs.: Carduus pycnocephalus (Italy); Carthamus, Cir
sium, Campanula (Europe, Rhodesia, India,
Egypt)
Remarks:
Nymphs and adults locally common on flowers
and stems; on Centaurea all year
Sources:
L. Andres in litt. (1961); Goeden (1974)
Dactynotus macrosiphon (Hille Ris Lambers)
Plant recs.: Carduus defloratus, C. personatus, Cirsium,
Carlina (Europe)
Remarks:
Nymphs and adults on stems and leaves
Sources:
Zwblfer (1965a)
Rhopalosiphum sp.
Remarks:
Nymphs, adults ectophagous on stems
Sources:
Toxopterina acanthoides (Borner)
Plant recs.: Carduus acanthoides (Europe)
Remarks:
Nymphs and adults on leaves and stems
Sources:
Zwblfer (1965a)
Cercopidae
Unident. cercopid spp.
22
HEMIPTERA-HOMOPTERAOmtmued
Cercopidae Continued
Plant recs.:
Remarks:
Carduus pycnocephalus (S. Calif., S. Europe)

Nymphs locally common to rare on stems; ectophagous
Sources:
Goeden [1974)
Philaenus spumarius (L.)
Remarks:
Nymphs and adults on shoots; polyphagous
Sources:
Goeden (1974)
Delphacidae
Unident. delphacid sp.
Remarks:
Nymphs and adults; ectophagous
Sources:
Dictyopharidae
Orgerius concordus Ball and Hartzell
Remarks:
Rare; adults on shoots; polyphagous
Sources:
Goeden (1974)
Issidae
Danepteryx sp. nr. D. manca Uhler
Remarks:
Sources:
Goeden (1974)
Acanaloniidae
Acanalonia bivittata (Say)
Remarks:
Sources:
Pseudococcidae
Unident. pseudococcid sp.
Remarks:
Rare; nymphs on leaves
Sources:
Goeden (1974)
COLEOPTERA
Staphylinidae
Mesomorphus sp.
Remarks:
Adults; feeding on leaves
Sources:
Melyridae
Unident. melyrid spp.
23

Remarks:
Adults; locally common on flowers
Sources:
Goeden (1974)
Procerallus ( = Acanthocnemus) altivolans Champ.
Remarks:
Adults taken from flowers; no other hosts
known
A17-ENT-9 Rpts. (1962-63, 1965)
Sources:
Phalacridae
Phalacrus sp.
Carduus edelbergii (Pakistan)
Plant recs
Adults found on plant; occasional visitors; no
Remarks:
other host known
Sources:
Nitidulidae
Unident. nitidulid sp.
Carduus acanthoides (France)
Plant recs.
Larvae in flower heads
Remarks:
Delemont Rpt. (1963)
Sources:
Meligethes sp
Remarks:
Adults feeding on flowers; polyphagous; a
polyphagous Meligethes also known from
"thistles" in Europe
Sources:
A17-ENT-9 Rpts. (1962-63, 1965); Baloch et al.
(1971)
Meligethes nigrescens Stephens
Rare; adults endophagous in flowers
Remarks:
Goeden (1974)
Sources:
Oedemeridae
Oedemera sp.
Carduus nutans (Austria); C. crispus (Rhine
Plant recs
Valley)
Remarks:
Ovipositing in stems of C. nutans; adult on
leaves of C. crispus
Sources:
Delemont Rpts. (1962-63)
Mordellidae
Unident. mordellid sp.
Plant recs.: Carduus acanthoides (France); C. crispus (Rhine
Valley); C. nutans, C. tenuiflorus (Europe)
Remarks:
Larvae on leaves of C. acanthoides, in stems of
C. crispus, C. nutans, and C. tenuiflorus
Sources:
Delemont Rpt. (1963); Zwblfer (1965a)
24
COLEOPTERAContinued
MordellidaeContinued
Mordella sp.
Plant recs.:
Remarks:
Carduus nutans (S. Dak.)

Adults; larvae endophagous in leaves and
stems
Sources:
Mordellistena spp.
Plant recs.: Carduus nutans (Rhine Valley, S. Dak.); C.
crispus (Europe); C. pycnocephalus (S. Calif.)
Remarks:
Larvae in stems of C. nutans and listed as en
dophytic in stems of C. crispus; adults occa
sional on leaves, stems, and flowers
Sources:
Delemont Rpt. (1963); Zwblfer (1965a); Goeden
(1974); Morihara and Balsbaugh (1976)
Meloidae
Epicauta atrata (F.)
Plant recs.: Carduus nutans (Sask.)
Remarks:
None
Sources:
M. G. Maw in litt. (1976)
Epicauta ferruginea (Say)
Remarks:
None
Sources:
Nemognatha lutea LeConte
Remarks:
None
Sources:
Alleculidae
Unident. alleculid sp.
Remarks:
Occasional visitor
Sources:
Zwblfer (1965a)
Byturidae
Unident. byturid sp.
Remarks:
Rare; larva
Sources:
Anobiidae
Lasioderma sp.
Remarks:
Larvae common; endophagous in flowers; also
in other Cynareae
Sources:
Goeden (1974)
25
Lasioderma baudii Schilsky

Plant recs.: Carduus pycnocephalus, C. tenuiflorus (Italy);
Cirsium, Centaurea, Cynara, Carthamus
Remarks:
Larvae reared from capitula, feed in seeds;
first identified as L. serricorne
Sources:
Rome Rpts. (1963-64)
Lasioderma redtenbacheri Bach
Plant recs.: Carduus nutans, Onopordum, Centaurea (Europe);
Centaurea (Egypt)
Remarks:
Larvae endophytic in flower heads and buds;
main hosts in Egypt listed as chestnuts;
distrib. Sicily and Egypt
Sources:
Zwblfer (1965a); F4-ENT-5 Rpt. (1967)
Lasioderma serricorne (F.)
Plant recs.: Carduus nutans, C. tenuiflorus, Centaurea, Cynara
(Italy); Centaurea (France, Egypt); Silybum
(Lebanon)
Remarks:
Some earlier recs. (by Rome) are L. bauddii; lar
vae feed in receptacle and are probably second
ary in their attack following weevils and seed
flies; numerous reports of attacking Cynara,
following L. baudii in Italy
Sources:
Rome Rpts. (1963-64); Delemont Rpt. (1965); F4ENT-5 and F4-ENT-16 Rpts. (1967-68, 1976)
Scarabaeidae
Cetonia aurata (L.)
Remarks:
Rare; adults in flowers; polyphagous
Sources:
Goeden (1974)
Oxythyrea sp.
Adults found in flowers; occasional visitor
Remarks:
Sources:
(1971)
Oxythyrea funesta Poda
Plant recs.: Carduus nutans, C. pycnocephalus, Cirsium, Cen
taurea, Onopordum (Europe)
Remarks:
Occasional visitor on thistles; adults common
on flowers; polyphagous
Sources:
Potosia hungarica Herbst
Plant recs.: Carduus nutans, C. acanthoides, Cirsium (Europe)
Remarks:
Occasional visitor on thistles
Sources:
Zwblfer (1965a)
26
COLEOPTERAContmued
ScarabaeidaeContinued
Potosia morio (F.)
Remarks:
Occasional adults feeding on flowers;
polyphagous
Sources:
Goeden (1974)
Tropinota hirta (Poda)
Plant recs.: Carduus nutans (Europe); C. pycnocephalus (S.
Europe)
Remarks:
Rare; occasional visitor; adults ectophagous
on flowers; polyphagous
Sources:
Tropinota squalida Scopoli
Plant recs.: Carduus pycnocephalus (Egypt)
Remarks:
None
Sources:
F4-ENT-5 Rpts. (1967, 1971)
Cerambycidae
Unident. cerambycid spp.
Plant recs.: Carduus nutans, C. pycnocephalus, C. tenuiflorus,
Silybum, Onopordum (adults only, Italy); C.
nutans (larvae, Italy)
Remarks:
Small adults found on these plants with no
associated larvae; large larvae commonly
found boring in pith of larger stems of C.
nutans, but apparently predaceous
Sources:
Rome Rpt. (1963)
Agapanthia sp.
Plant recs.: Carduus crispus (Rhine Valley)
Remarks:
Larvae endophytic in stems
Sources:
Agapanthia cardui (L.)
Plant recs.: Carduus nutans (Rhine Valley); C. nutans, C.
acanthoides, C. tenuiflorus, C. pycnocephalus, Cirsium, Galactites, Onopordum, Centaurea, Cynara
(Europe); ? Centaurea (France, Bulgaria)
Remarks:
Larvae (endophytic in stem) reared only from
Cirsium; other recs. here are adults only (on
foliage); identity of larvae in Centaurea in
France and Bulgaria in question, given as
"possibly cardui"
Sources:
Delemont Rpts. (1963, 1965, 1967); Zwblfer
(1965a); Goeden (1974)
27
Agapanthia dahli Richter

Plant recs.: Carduus nutans (Austria); C. nutans, C. acanthoides, C. pycnocephalus, C. litigiosus, Cirsium,
Arctium (Europe); Carduus pycnocephalus, Centaurea (Egypt)
Remarks:
Larvae (feeding in stem) reared only from
Carduus acanthoides, C. pycnocephalus, and Centaurea; other recs. shown here are adults only
(feeding on foliage); A. dahli studied in lab. at
Delemont; reared in lab. in Egypt from C. pyc
nocephalus and Centaurea; one of main pests of
C. pycnocephalus in Egypt; an economic pest of
sunflower in USSR
Sources:
Delemont Rpt. (1963); CIBC Ann. Rpt. (1963);
Zwblfer [1965a); F4-ENT-5 and F4-ENT-16
Rpts. (1967, 1976); Goeden (1974)
Agapanthia lateralis Ganglbauer
Plant recs.: Carduus pycnocephalus, Centaurea, Silybum
(Egypt)
Remarks:
One of main pests of C. pycnocephalus in Egypt;
reared in lab. in Egypt from C. pycnocephalus
and Centaurea; larvae feed in stems
Sources:
F4-ENT-5 and F4-ENT-16 Rpts. (1967-68, 1976)
Agapanthia villosoviridescens De Geer
Plant recs.: Carduus nutans, C. pycnocephalus, Cirsium,
Onopordum, Echinops (Europe)
Remarks:
Larvae (feeding in stems) recorded only from
Cirsium and Echinops; other recs. here are
adults only, eating stems or foliage; literature
rec. hosts are Angelica, Aconitum, Eupatorium,
Heracleum, Senecio, and Cirsium
Sources:
Delemont Rpts. (1962-63); Zwblfer (1965a);
Goeden (1974)
Agapanthia violacea (.)
Plant recs.: Carduus pycnocephalus, Centaurea (Europe)
Remarks:
Occasional visitor (adults)
Sources:
Zwblfer (1965a)
Purpuricenus kaehleri (L.)
Remarks:
Rare; adults ectophagous on flowers;
polyphagous
Sources:
Goeden (1974)
28
COLEOPTERAContinued
Chrysomelidae
Unident. chrysomelid spp.
Plant recs.: Carduus pycnocephalus and/or C. tenuiflorus
(Calif.)
Remarks:
Feeding habits unknown
Sources:
Albany Rpt. (1963)
Altica sp. nr. A. viridicyanea Baly (= Haltica sp.)
Remarks:
Adults feeding on foliage; possibly
oligophagous; morphologically different from
European Altica spp.
Sources:
Altica carduorum Guerin-Meneville
Plant recs.: Carduus pycnocephalus, C. nutans, C. crispus, C.
tenuiflorus, C. defloratus, C. personatus, C. acanthoides, Cirsium, Silybum, Onopordum, Cnicus,
Arctium, Xeranthemum, Cynara, Echinops,
Helianthus, Chrysanthemum, Carthamus, Lactuca, flax (Europe and lab. recs. - see remarks)
Remarks:
Adults and larvae skeletonize leaves of host,
desiccating plant incl. roots; believed to be
host specific to Cirsium, with Carduus and
Silybum being marginal hosts, probably not in
nature
Field recs. (adults only); Cirsium arvense (L.)
Scop. (Switzerland, France, Spain), C. vulgare
(Savi) Ten. (Spain), and Carduus pycnocephalus
(Europe)
Other distrib. (from museum specimens):
Germany, Czechoslovakia, Yugoslavia,
Greece, Albania, Italy, Sicily, Sardinia, Cor
sica, Cyprus, ?Austria, and ?Asiatic USSR
History: First discovered in Swiss Valais,
May 1961; shipped to Canada, 1962-64; Work
ing Group on Biological Control of Weeds ap
proved release in Canada, Mar. 1963, in U.S.,
1966; released in Canada, 1963-70; released in
U.S., 1966-72; insect studied and released to
control Canada thistle (Cirsium arvense)
Results of lab. screening tests (D = Delemont,
B = Belleville, A = Albany): Adults fed regu
larly on Carduus nutans (D), C. crispus (D), C.
tenuiflorus (D), C. acanthoides (B), C. sp. prob.
29
tenuiflorus (A), C. pycnocephalus (A), Cirsium (D,

B, A), Silybum (D, B, A), and occasionally on
Carduus defloratus (D), C. personatus (D), Cnicus
(D), Arctium (D, B), Xeranthemum (B),
Onopordum (D, B), Centaurea (B, A), Helianthus
(B), Chrysanthemum (B), Lactuca (A), and
especially the new tender growth of Carthamus
(A) and Cynara (A); larvae fed and developed
successfully only on Carduus acanthoides (B), C.
pycnocephalus (A), Cirsium (D, B, A), and
Silybum (D, B); fed occasionally, but could not
develop on Cnicus (D), Xeranthemum (D), Cynara
(D, A), Centaurea (B), Onopordum (B), Echinops
(B) , Carthamus (A), and flax; eggs deposited
only on Cirsium (D, B, A) and Silybum (D, ?B,
A); detailed results of specificity tests in Har
ris
distrib. and ecology in Zwblfer
(i9<556)
Releases (A = adults, L = larvae) against
Cirsium arvense: Canada - 21 A in Ont. and 100
A and L in Alta. in 1963; 600 A and L in Ont. (2
Iocs.), and 100 A and L in B.C. (82 A and L sent
to Albany for lab.) in 1964; 149 A in Alta., 72 A
in B.C., and 1,460 A and L in Ont. (2 Iocs.) in
1965; 60 A in N.S. and 35 A ( + 482 L in field
cage study) in Ont. in 1966; 367 ?A ( + 64 in field
cage study) in B.C. and 490 ?A in Ont. in 1967;
1,147 A in B.C. (2 Iocs.), 1,160 A and 107 L in
Alta., and 291 A in Ont. in 1968; 5,534 A in
B.C. (2 Iocs.) and 170 A in Alta. in 1969; and
1,363 A (from lab. culture and France); in Ont.
(2 Iocs.) in 1970
United States - 1,000 A in Calif. (4 Iocs.), 400
A in Idaho, 200 A in Oreg., 200 A in Wash., and
in Mont. (640 A sent, but prob. only 200 re
leased, 2 Iocs.) in 1966; 1,700 A in Calif. (Ft.
Bragg area) and 175 in Idaho (in field cage) in
1967; 500 A in Calif. (Ft. Bragg field cage), 150
A in Colo. (2 Iocs.), 200 A in Idaho, 100 A in
Wash., 200 A in Minn., 200 A in Wis., 200 A in
Ind., 200 A in Del., and 200 A(+ some larvae
reared in lab.) in N.J. in 1968; unknown No. A
in Calif. (Ft. Bragg area) in 1969; 500 A and 242
L in S. Dak. (field cages) and 360 L in Md. (field

cages) in 1970; 160 A in Md. and 65 sent to S.
Dak. in 1971; and 52 A sent to S. Dak. in 1972
(S. Dak. and Md. releases from French
material)
Recovery data: Little survival in Canada,
not established on Cirsium in B.C., Alta., Ont.,
or N.S. by 1975 and studies discontinued;
possible reasons are predation and low sum
mer temperatures; no overwintering observed
at Ft. Bragg, Calif., release site; no known
establishment elsewhere in U.S.; in Canada, it
survived only when protected by field cages
from predation; the mites Leptus sp. nr. curtipes
Schweitzer and Sphaerolophus sp. ate the eggs
(Peschken et al., 1971); over 2,100 adults were
released in 1969 at 3 sites in England (Claridge
et al., 1970); poor survival in cages was at
tributed to the cool climate (Baker et al., 1972);
in S. Dak., establishment was limited by the
hot, dry summers and predation by Lebia
viridis Say and Harpalus pennsylvanicus De Geer
(Schaber et al. 1975)
Sources:
Delemont Rpts. (1961-65, 1967); Belleville Res.
Inst. Res. Rpts. (1962-63); CIBC Ann. Rpts.
(1963-64, 1969, 1970); Karny (1963); Albany
Rpts. (1964-72); Harris (1964, 1975); Zwblfer
(1965a, b); R. B. Hawkes in litt. (1966); William
son (1966-70); Canada Dept. Agr. Res. Inst.
Res. Rpt. (1966); Zwblfer and Harris (1966);
ARS Lab., Moorestown, N.J., Rpt. (1968);
Zwblfer (1969, 1970); Zwblfer and Pattullo
(1970); Peschken et al. (1970); Andres and Davis
(1973); Goeden (1974); Maw (1976)
Aphthona pygmaea Kutschera
Remarks:
Rare; adults on leaves
Sources:
Goeden (1974)
Arima marginata (F.)
Plant recs.: Carduus nutans, Centaurea (France); other hosts
from literature: Carduus, Cirsium, Onopordum,
Arctium, Asteraceae, Heliantheae, Anthemideae, Calenduleae, Centaureinae, Lactuca
(lab. tests)
Remarks:
Adults and larvae on leaves

Sources:
31
Grandi (1952); Heikertinger (1954); Zwblfer

(1969)
Cassida deflorata Suffrian
Plant recs.: Carduus tenuiflorus, C. personatus, C. pycnocephalus, C. defloratus, C. nutans, C. crispus,
Cnicus, Arctium, Cynara, Cirsium, Onopordum,
Silybum, Centaurea, Chrysanthemum, Echinops,
Xeranthemum, Carthamus, Carlina, Tagetes,
Leontodon, Erigeron (Europe and in lab. tests see remarks)
Remarks:
Field recs.: Carduus sp., C. tenuiflorus, C. pycnocephalus (S. Europe), Cynara, Silybum (France),
Cirsium (Switzerland), and Cynara (Italy);
adults and larvae on all species here except on
C. tenuiflorus (adults and eggs only) and on
Cirsium in Switzerland (adults only); identity of
"sp. prob. deflorata" in Italy on Cynara in ques
tion
Results of lab. tests (Delemont): Adults fed
heavily or regularly on foliage of Carduus per
sonatus, C. defloratus, C. nutans, C. tenuiflorus,
C. crispus, Arctium, Cynara, Onopordum,
Silybum, Cirsium, Centaurea, and rather incon
sistently or occasionally on Cnicus, Chrysan
themum, Carthamus, Echinops, Xeranthemum,
and Carlina; larvae fed regularly on Carduus
(same 5 spp. as above), Cnicus, Carthamus, Arc
tium, Cynara, Onopordum, Silybum, Centaurea,
and inconsistently or occasionally on Echinops,
Xeranthemum, Tagetes, and Leontodon; adults
show high feeding indices in Cynara and
Onopordum; cannot be considered in biological
control program because of feeding on Cynara
Sources:
Delemont Rpts. (1961-63); Zwblfer (1965a, 1969);
Zwolfer and Eichhorn (1966); Goeden (1974)
Cassida rubiginosa Mueller
Plant recs.: Carduus pycnocephalus, C. acanthoides, C.
defloratus, C. crispus, C. nutans, C. personatus, C.
tenuiflorus, Cirsium, Onopordum, Arctium, Cen
taurea, Carthamus, Cynara, Silybum, Lappa,
Tanacetum, Xeranthemum, Cnicus, Echinops,
Helianthus, Erigeron, Solidago, Aster, Carlina,
Taraxacum (Europe, N. Amer., and in lab. tests
- see remarks)
32
COLEOPTERAContinued
ChrysomelidaeContinued
Remarks:
Sources:
Adults and larvae skeletonize and severely

damage leaves of hosts; prefer Cirsium, Carduus, and Silybum, but also feed on Cynara, Carthamus, and other plants
Field recs.: Adults and larvae from Carduus
nutans and C. pycnocephalus (Europe), C.
thoermeri and C. nutans (abundant in N. Va.,
Md., Pa., 1975), C. crispus (Rhine Valley), C. personatus (Europe), C. acanthoides (Swabian Jura),
C. defloratus (Switzerland), Carduus (Canada),
Cirsium (Switzerland, Canada, France, Swa
bian Jura, Rhine Valley), and Onopordum, Arc
tium, and Centaurea (Europe); adults only from
Carduus acanthoides and Onopordum (France),
Carduus nutans and Cirsium (Austria), and
Carduus defloratus and C. personatus (Europe);
unknown stages from C. pycnocephalus (France,
Italy), Cirsium, Lappa, and Silybum (N. Amer.)
Other distrib.: Germany, England, Ont.
(found 1927), Que. (1940), N.Y. (1945), and N.J.
(1962)
Results of lab. screening tests (Delemont,
1962-63): Adults fed regularly on Carduus per
sonatus, C. defloratus, C. nutans, C. tenuiflorus,
C. crispus, Cirsium, Silybum, Onopordum,
Cynara, Arctium, Centaurea, Carthamus, and oc
casionally (or nibbled) on Xeranthemum,
Cnicus, Echinops, and Helianthus; larvae fed
and developed on same 5 Carduus spp. in adult
test plus Cirsium, Silybum, Onopordum, Cynara,
Arctium, Centaurea, and fed but did not develop
on Carthamus, Cnicus, Echinops, and Erigeron;
oviposition obtained on Carduus defloratus, C.
nutans, Cirsium, Silybum, Onopordum, Cynara,
Arctium, Centaurea, Carthamus, Cnicus, Helian
thus, Erigeron, Solidago, Aster, Carlina, and
Taraxacum (fate of eggs not studied in this test);
parasitism may prevent high populations in
parts of N. Amer.
L. Andres in litt. (1961); Delemont Rpts.
(1962-63, 1967); Rome Rpt. (1963); Zwblfer
33
(1965a, 1969); Zwblfer and Eichhorn (1966);

Mohr (1966); Zwblfer and Pattullo (1970); S. W.
T. Batra and L. T. Kok in litt. (1976); Maw (1976)
Cassida sp. nr. C. vibex (L.)
Plant recs.: Carduus sp. (Austria, Switzerland); C. acanthoides (Switzerland); C. personatus (France); C.
edelbergii (Pakistan)
Remarks:
Eggs and larvae found on plants in Europe;
adults feeding on leaves of C. edelbergii in
Pakistan
Sources:
Delemont Rpts. (1962-63); A17-ENT-9 Rpt.
(1965); Zwblfer and Eichhorn (1966); Baloch et
al. (1971)
Cassida vibex (L.)
Plant recs.: Carduus defloratus, C. crispus, C. pycnocephalus,
C. tenuiflorus, C. nutans, C. personatus, Cirsium,
Centaurea, Arctium, Onopordum, Silybum, Serratula, Cnicus, Carthamus, Cynara, Tanacetum,
Xeranthemum, Echinops, Carlina, Helianthus,
Chrysanthemum, Solidago, Erigeron, Aster,
Taraxacum, Lactuca (Europe and in lab. tests see remarks)
Remarks:
Adults and larvae skeletonize the leaves of the
host; a potential pest of Cynara and Carthamus
defloratus and Cirsium (Switzerland, France,
Austria, Rhine Valley), Onopordum and Arctium
(France), and Centaurea and Serratula (Europe);
adults only from Carduus crispus, C. pyc
nocephalus, C. personatus, C. tenuiflorus, and
Silybum (Europe)
Results of lab. tests (Delemont, 1962-63) for
Cirsium arvense: Adults fed heavily or regularly
on Carduus nutans, C. defloratus, C. tenuiflorus,
C. crispus, C. personatus, Cirsium, Silybum, Cen
taurea, Onopordum, Cynara, Carthamus, Arctium,
Cnicus, and occasionally on Xeranthemum,
Echinops, Carlina, and Helianthus; larvae fed
and developed on Carduus (same 5 spp. as for
adult test), Cirsium, Silybum, Centaurea, Onopor
dum, Cynara, Carthamus, Cnicus, and nibbled
(but did not develop) on Carlina and
Chrysanthemum; oviposition obtained (fate of
eggs not studied) on Carduus defloratus, Cirsium,

Silybum, Centaurea, Onopordum, Cynara, Arc
tium, Cnicus, Xeranthemum, Echinops, Chrysan
themum, Solidago, Erigeron, Aster, Taraxacum,
and Lactuca
Sources:
Delemont Rpts. (1961, 1963); Zwblfer (1965a,
1969); Zwblfer and Eichhorn (1966); Mohr (1966)
Chaetocnema confinis Crotch
Remarks:
Adults ectophagous on leaves; polyphagous
crop pest
Sources:
Chrysomela banksi F.
Plant recs.: Carduus pycnocephalus (S. Europe); Compositae,
Labiatae
Remarks:
Rare; adults on leaves; prob. polyphagous
Sources:
Zwblfer (1969); Goeden (1974)
Chrysomela fuliginosa Olivier
Plant recs.: Centaurea (Europe); Carduus nutans, Cirsium,
Cynara, Cnicus, Carthamus, Centaurea (lab. tests)
Remarks:
Adults and larvae on leaves
Sources:
Zwblfer (1969)
Crepidodera ferruginea Scopoli
Plant recs.: Carduus defloratus, C. personatus, Cirsium,
Carlina, other Compositae (Europe)
Remarks:
These recs. are adults only on foliage; larvae
generally attack plant stem base
Sources:
Zwblfer (1965a, 1969)
Crepidodera transversa Marsham
Plant recs.: Carduus defloratus, Cirsium, Carlina (Austria,
Switzerland)
Remarks:
These recs. are adults only on foliage; oviposition and screening tests conducted with this
sp. at Delemont in 1962 and 1963
Sources:
Delemont Rpts. (1962-63)
Cryptocephalus sericeus (L.)
Remarks:
Occasional visitor
Sources:
Zwblfer (1965a)
Diabrotica undecimpunctata undecimpunctata Mannerheim
Remarks:
Occasional adults; poss. breeding on flowers,
leaves, and stems; polyphagous crop pest
Sources:
Goeden (1974)
35
Diachus auratus (F.)

Remarks:
Adults on leaves; polyphagous
Sources:
Enneamera variabilis Baly
Remarks:
Adults exophytic on stems and leaves
Sources:
Epithrix cucumeris (Harris)
Remarks:
Adults on leaves; polyphagous crop pest
Sources:
Galeruca pomonae (Scopoli)
Plant recs.: Carduus crispus, C. nutans, C. defloratus, C. personatus (in lab. tests only); Centaurea, Cirsium,
Carlina (Europe)
Remarks:
Adults and larvae feed on leaves of host, often
causing defoliation; Centaurea believed to be
main host plant
Field recs.: Larvae reared from Centaurea
(Switzerland); adults and larvae found from
Cirsium and Carlina (Europe)
on Carduus crispus, Centaurea, Cirsium, Arctium,
Cnicus, and occasionally on Carduus nutans and
C. defloratus; larvae fed heavily or regularly on
C. nutans, C. personatus, Achillea, Silybum, Cen
taurea, Cnicus, Cynara, Cirsium, and occa
sionally on Carduus defloratus
Sources:
Delemont Rpts. (1962-63); Zwblfer, (1965a,
1969); Mohr (1966)
Galeruca tanaceti (L.)
Plant recs.: Carduus nutans (Europe); C. defloratus (in lab.
tests only); Achillea (Switzerland and in lab.
tests); Cirsium (Switzerland, France, Swabian
Jura, and in lab. tests); Centaurea (Europe and
in lab. tests); Rudbeckia, Solidago, Erigeron,
Aster, Chrysanthemum, Xeranthemum, Silybum,
Helianthus (in lab. tests only)
Remarks:
Adults and larvae feed externally on foliage;
apparently relatively polyphagous; these field
recs. are all of adults and larvae except
Carduus nutans (adults only)

Results of lab. tests (Delemont, 196263) for
on Carduus defloratus, Rudbeckia, Achillea, Cir
sium, Centaurea, Solidago, Erigeron, Aster,
Chrysanthemum, Xeranthemum, Silybum, and oc
casionally on Helianthus
Sources:
Heikertinger (1954); Delemont Rpts. (1962-63);
Zwblfer (1965a, 1969)
Hoplasoma sexmaculata Hope
Remarks:
Adults feeding on leaves
Sources:
Lachnea cylindrica LeConte
Remarks:
Rare; adults ectophagous on flowers;
polyphagous
Sources:
Goeden (1974)
Lema cirsicola Chujo ( = L. "cyanella, " misident.)
Plant recs.: Carduus nutans, Silybum, Cynara, Carthamus,
Centaurea, Helianthus (in lab. tests only);
Cirsium (Japan and in lab. tests)
Remarks:
Coll. from Cirsium in Japan, as "cyanella"
Results of lab. tests (Albany, 1971-72) for
Cirsium arvense and C. vulgare, early tests as
"cyanella": Adults fed readily on Cirsium and
Silybum spp., and very much less on Carduus
nutans, Centaurea, Cynara, Carthamus, and
Helianthus; adults oviposited on Cirsium,
Silybum, and Carduus nutans; some minimal lar
val feeding on C. nutans; since coll. only from
Cirsium in shaded forest stands, value of this
sp. as control for Cirsium arvense questionable
Sources:
Albany Rpts. (1971-73)
Lema cyanella L. ( = L. puncticollis Curtis)
Plant recs.: Carduus defloratus, Silybum, Onopordum (in lab.
tests only); Cirsium (France, Switzerland, and
in lab. tests)
Remarks:
Adults and larvae feed on leaves of Cirsium
In lab. tests (Delemont, 1962-63, 1967 as
puncticollis, rugicollis; 1964 as puncticollis,
ruficollis, rugicollis; and 1967 as cyanella) for
Cirsium arvense: Adults fed regularly on
Cirsium, Silybum, Carduus defloratus, and occa
sionally on Onopordum; host plants apparently
37
limited to Carduus, Cirsium, and Silybum; also

fed on Cirsium arvense, C. occidentale, and
Carduus nutans; being tested in Sask., Canada
(1976)
Sources:
Delemont Rpts. (1962, 1964, 1967); CIBC Ann.
Rpts. (1963, 1964); Zwblfer, (1965a, 1969);
Zwblfer and Harris (1966); Zwblfer and Pattullo
(1970); P. Harris pers. comm. (1976)
Leptinotarsa decemlineata (Say)
Remarks:
Adults ectophagous on leaves; polyphagous
crop pest
Sources:
Longitarsus Itestaceus (Melsheimer)
Remarks:
Adults on leaves; feeds on other thistles
Sources:
Merista quadrifasciata Hope
Plant recs.: Carduus edelbergii, Cnicus (Pakistan)
Remarks:
Adults feeding on leaves; no other host plants
known
Sources:
Monolepta signata Olivier
Remarks:
Adults feeding on leaves; polyphagous on
crops
Sources:
Oulema melanopus (L.) ( = Lema melanopus L.)
Plant recs.: Carduus tenuiflorus (France); C. pycnocephalus
(S. Europe); Centaurea (Europe)
Remarks:
Occasional visitor (adult); pest of grains in
Europe and N. Amer.
Sources:
Delemont Rpt. (1962); Zwblfer (1965a); Goeden
(1974)
Psylliodes sp.
Remarks:
Occasional visitor
Sources:
Zwblfer (1965a)
Psylliodes chalcomera (Illiger)
Plant recs.: Carduus acanthoides (Germany); C. nutans and
C. pycnocephalus (Germany, Italy, and in lab.
tests); C. crispus (Italy); Cynara, Carthamus, Cen
taurea (in lab. tests only)
38
COLEOPTERAContinued
Remarks:
Adults feed on leaves, eggs laid on soil, larvae
bore in growing tips of host plants at the
crown; of these field recs., only Italian recs. on
Carduus nutans and C. pycnocephalus are
definitely adults and larvae; rec. from C.
crispus is adult only
Results of lab. tests (Rome, 1966-71) for
Carduus spp.: Adults fed on Carduus nutans, C.
pycnocephalus, Cynara, Carthamus, and
Centaurea; oviposition obtained on Carduus
nutans, C. pycnocephalus, Cirsium, Silybum, and
Cynara; 1st instar larvae successfully transfer
red to Carduus spp., Silybum, and Cirsium;
adults produced only on C. nutans, mature lar
vae on Carduus and Cynara; introduced in
quarantine to USA (Albany, Calif.) in 1972
(600), 1973 (3,362), and 1974 (3,600)
In lab. tests (Albany, 1972-74) adults fed on
4 Cirsium spp.; oviposition and larval develop
ment on Carduus nutans, Centaurea, and Cynara;
some pupation and adult emergence on Cynara;
this insect rejected as biocontrol agent (Albany
Rpt., 1974); gregarine parasite, in 79 percent of
sample, influences oviposition behavior;
Microctonus sp. also parasitic (Rome Rpt., 1971)
Sources:
Heikertinger (1954); K. E. Frick in litt. (1966);
Rome Rpts. (1966-74); Albany Rpts. (1972,
1974); Goeden (1974); Dunn and Rizza (1976)
Psylliodes punctulata Melsheimer
Remarks:
None
Sources:
Sphaeroderma rubidum Graells (= S. "rubrum")
Plant recs.: Carduus pycnocephalus, C. crispus, C. personatus,
Cirsium, Cynara, Galactites, Onopordum, Serratula, Centaurea, Arctium, Carthamus (Europe
and in lab. tests - see remarks)
Remarks:
Adults feed externally on leaves; larvae mine
leaves of host plants
Field recs.: Adults only (ident. questioned in
all cases) from Cirsium (Rhine Valley),
Centaurea, Galactites, Onopordum, and Carduus
39
pycnocephalus (S. France or N. Spain); larvae

and adults from Serratula, Cynara, and
Centaurea (Europe); stage unknown from
Cynara (Italy)
Results of lab. tests (Delemont, 1963, 1965)
for Cirsium arvense: Adults fed heavily or
regularly on Centaurea, Carthamus, Arctium,
Cynara, Onopordum, Cirsium, Carduus crispus,
and C. personatus; Centaurea is apparently main
host, but Cynara and Carthamus are heavily
damaged; therefore not considered for
biological control
Sources:
Delemont Rpts. (1963-65); Rome Rpt. (1963);
Zwblfer (1965a, 1969); Mohr (1966); Goeden
(1974)
Sphaeroderma testaceum Weise
Plant recs.: Carduus nutans, C. crispus, C. personatus, C.
defloratus, Cirsium, Carlina, Onopordum, Silybum
(Europe and in lab. tests - see remarks)
Remarks:
Adults feed on foliage; larvae mine leaves of
host plants; apparently prefers Carduus, Cir
sium, and Silybum; appears to be a promising
biological control candidate
Field recs.: Adults (only) from Carduus
defloratus, C. nutans, C. personatus, Carlina
(Switzerland), Carduus crispus (Rhine Valley,
poss. ovipositing), Cirsium (Switzerland,
France), and Onopordum (Europe); larvae from
Carduus nutans and Cirsium (Europe)
on Carduus nutans, C. crispus, C. personatus, C.
defloratus, and Cirsium; some feeding on
Onopordum and Silybum
Sources:
Delemont Rpts. (1962-63, 1967); Zwblfer (1965a,
1969); Zwblfer and Harris (1966); Zwblfer and
Pattullo (1970)
Systena blanda Melsheimer
Remarks:
Rare; adults feeding on leaves and stems;
Sources:
Goeden (1974)
Systena elongata (F.)
40
COLEOPTERA Continued
Remarks:
Sources:
Systena frontalis (F.)
Remarks:
Adults on leaves; polyphagous
Sources:
Curculionidae
Unident. curculionid spp.
Plant recs.: Carduus acanthoides (Austria, Switzerland); C.
nutans (Italy)
Remarks:
Feeding in roots and flower heads; larvae in
crowns
Delemont Rpt. (1962-63); Rome Rpt. (1963)
Sources:
Acalles sp.
Carduus pycnocephalus (Sicily)
Plant recs.
Remarks:
None
Sources:
L. Andres in litt. (1961)
Acalles diocletianus (Germar)
Plant recs.: Carduus Inigrescens, Silybum (Europe)
Occasional visitor
Remarks:
Zwblfer (1965a)
Sources:
Apion spp.
Plant recs.: Carduus pycnocephalus (Sicily, Italy, France); C.
acanthoides (Austria, Switzerland); C. crispus
(Rhine Valley); C. defloratus (Switzerland);
Carduus spp. (Sicily, Switzerland); Cirsium
(Switzerland, France, Rhine Valley); Cynara
(France); Centaurea (Europe)
Remarks:
Adults attack crowns and stems, larvae bur
row in pith and pupate in cortical layer of C.
pycnocephalus; adults on, and larvae boring in,
stems of C. acanthoides; adults in stems of C.
crispus; larvae in stems of C. defloratus; adults
on Carduus sp., Cynara, and Centaurea; adults
on, and larvae boring in, stems of Cirsium;
although often recorded as "numerous" or
"abundant," damage attributed to these
beetles is "little" or none
Sources:
Rome Rpt. (1960); L. Andres in litt. (1961); Dele
mont Rpts. (1962-63, 1965); Goeden (1974)
41
Apion (Ceratapion) sp. nr. A. armatum Gerstaecker

Plant recs.: Carduus pycnocephalus, C. tenuiflorus, C. nutans,
Centaurea (Italy)
Remarks:
Most common weevil attacking Carduus crowns
in Italy (up to 65 larvae found in single large
crown); damage to host is negligible; same sp.
or closely related sp. also feeds on pith of
stems; these plant recs. are rearing recs.
Sources:
Rome Rpt. (1963)
Apion (Ceratapion) alliariae Herbst
Plant recs.: Carduus tenuiflorus, C. pycnocephalus (France);
Centaurea (Yugoslavia, Italy); Onopordum
(Italy)
Remarks:
Reared from Centaurea in Yugoslavia; other
recs. here are of adults on foliage
Sources:
Zwblfer (1965a)', Albany Rpt. (1966)
Apion (Ceratapion) carduorum Kirby (and ssp. galactitis Wencker)
Plant recs.: Carduus pycnocephalus, C. personatus, C. nutans,
?C. acanthoides, Cirsium (Europe - see
remarks); ssp. galactitis C. nutans, C. pyc
nocephalus, C. tenuiflorus (France, Italy, or
Sicily; reared); Galactites, Cynara
Remarks:
Adults only reared from Carduus nutans (larvae
also, but identity questioned), C. pyc
nocephalus, and C. personatus (all from Europe);
larvae reared from Cirsium (Europe); adults
from Cirsium (Switzerland, Germany); adults
(France) and larvae (Europe, identity of both
recs. questioned) from Carduus acanthoides;
adults on foliage, larvae within stems
Sources:
Delemont Rpt. (1963); Zwblfer (1965a); Albany
Rpt. (1966)
Apion (Ceratapion) onopordi Kirby
Plant recs.: Carduus nutans, C. acanthoides, C. crispus, C.
pycnocephalus, Carduus sp., Cirsium, Centaurea,
Onopordum, Silybum (Europe - see remarks)
Remarks:
Adults on foliage, larvae in stems (and
crowns); larvae reared from Carduus acan
thoides and C. nutans (Germany), C. pyc
nocephalus and Carduus sp. (France), Onopordum
and Silybum (Italy), and Cirsium (identity not
verified) and Centaurea (Europe); adults (only)

recorded from foliage of Carduus crispus
(Europe) and C. acanthoides (France, ident. not
verified); screening tests for A. onopordi vs.
Centaurea at Delemont (1965)
Sources:
Delemont Rpts. (1963, 1965); CIBC Ann. Rpt.
[1965); Zwblfer (1965a); Albany Rpt. (1966);
Goeden (1974)
Apion (Ceratapion) pisi F.
Plant recs.: Carduus nutans (Italy)
Remarks:
One specimen "probably" reared from C.
nutans
Sources:
Albany Rpt. (1966)
Apion (Pseudapion) fulvirostre Gyllenhall
Remarks:
Occasional visitor
Sources:
Zwblfer (1965a)
Brachynotus sp.
Remarks:
Rare; adults and larvae on leaves
Sources:
Goeden (1974)
Ceutorhynchus sp.
Remarks:
Adult feeding on Carduus in Pakistan, poss.
only an occasional visitor; different from
Ceutorhynchus spp. of Europe
Sources:
(1971)
Ceutorhynchus sp. nr. C. litura (F.)
Plant recs.: Carduus acanthoides, C. nutans (E. Austria)
Remarks:
Eggs and larvae common in rosettes of C. acan
thoides; poss. an ecotype of Ceutorhynchus litura
preferring Carduus over Cirsium
Sources:
Ceutorhynchus (Hadroplontus) litura (F.)
Plant recs.: Carduus defloratus, C. nutans, C. tenuiflorus, C.
personatus, C. acanthoides, C. crispus, Cirsium,
Silybum, Onopordum, Centaurea, Cynara, Cnicus,
Aster, Carthamus, Echinops, Xeranthemum, Lactuca (Europe and in lab. tests - see remarks)
Remarks:
Adults feed on foliage; larvae endophytic in
leaves, stems, root collars, and occasionally in
roots; hosts limited to Cirsium, Carduus, and
Silybum
43
defloratus (Switzerland), Cirsium (Switzerland,

Sweden, France, Germany), and Carduus nutans
and C. crispus (France)
History: Screening tests at Delemont,
1961-66; shipments made to Canada, 1963-67;
Working Group on Biological Control of
Weeds approved release in Canada, 1965;
small field cage study, 1965; first field release,
1967; insect studied and released to control
Canada thistle (Cirsium arvense)
Results of lab. screening tests (D = Delemont,
B = Belleville): Adults fed regularly on Carduus
defloratus (D), C. nutans (D), C. tenuiflorus (D), C.
personatus (D), C. acanthoides (B), Cirsium (D, B),
Silybum (D, B), and occasionally ate Onopordum
(D, B), Centaurea (D), Cynara (D, B), Cnicus (D),
Aster (D), Carthamus (D), Echinops (D), Lactuca
(D, B), and Xeranthemum (D); larvae fed and
developed only on Carduus defloratus (D) and
Cirsium (D), and fed but did not develop on
Silybum (D); eggs deposited in C. defloratus (D)
and Cirsium and Silybum (D) only
Releases in Canada on Cirsium arvense: 22 A
in field cage study in Ont. in 1965; 270 A
released in Ont. (2 Iocs.) in 1967; 70 A in Sask.
in 1973; 100 A in Sask. in 1974; 69 A in B.C.,
57A in Alta, and 41 A in Sask. in 1975; 56 A in
Sask. in 1976; cleared for release on Cirsium
arvense in U.S. by the Working Group on
Biological Control of Weeds in 1969; 25 Cana
dian collected adults sent to SEA-AR lab.,
Albany, Calif., in 1970 (rearing attempt failed);
adults (2,900 + ) collected on Cirsium arvense
and Carduus nutans sent to Albany from Ger
many and Switzerland in 1971-75
Releases in U.S. on Cirsium arvense: 154 A in
S. Dak. and 100 A in Idaho in 1972; 231 A in
Idaho, 125 A in Mont., and 192 A in Calif. in
1973; 410 A in Calif., 200 A in Wash., 400 A in
Idaho, and 150 A in Colo, in 1974; 280 A in
Idaho and 150 A in Md. in 1975; rearing at
tempted in 1975 in N.J. by N.J. Dept. Agr.; lar
vae from Mont, released in Md. in 1978 and
1979

Recovery data: Recovered at 1967 Ont.
release site in 1968 and spreading at one site in
Ont. (established on Cirsium); established and
increasing in Sask. by 1976; established in
Mont.; some evidence of potential establish
ment in Idaho and Calif.; 1967 release popula
tion at Belleville, Ont., has caused decline of C.
arvense shoots from 28.8 to 0.24/m2 in density
Parasite: C. litura may disseminate the this
tle rust Puccinia punctiformis (Str.) Kohl, in
Ont.; the fly Phaonia trimaculata (Bouche) is a
European enemy as is the protozoan Nosema
sp.
Sources:
Delemont Rpts. (1961-67); CIBC Ann. Rpts.
(1963-67, 1974); Williamson (1963-65, 1966-70);
Zwblfer (1964, 1965a); Zwblfer and Harris (1966);
Canada Dept. Agr. Res. Inst., Belleville, Rpts.
(1967-73); Albany Rpts. (1969-75); Zwblfer and
Pattullo (1970); Peschken (1971); Peschken and
Beecher (1973); P. H. Dunn in litt. (1978)
Ceutorhynchus (Hadroplontus) trimaculatus (F.)
Plant recs.: Carduus nutans, C. pycnocephalus, C. tenuiflorus,
C. myriacanthus, Galactites, Cirsium (Europe);
Carduus acanthoides, Cirsium, Cynara, Silybum,
Onopordum, others (in lab. tests - see remarks)
Remarks:
Adults feed on foliage, oviposit on new grow
ing tips; larvae bore into crown itself, feed
gregariously
nutans, C. pycnocephalus, and Galactites (Italy),
C. pycnocephalus (S. France, N. Spain), Carduus
myriacanthus (Algeria), and C. tenuiflorus and
Cirsium (Europe)
Tests for Carduus spp. at SEA-AR lab.,
Rome, 1968-73: In tests, adults fed on Carduus
pycnocephalus, C. nutans, C. acanthoides, Cir
sium, Cynara, Galactites, Silybum, Onopordum,
Echinops, and nibbled other plants; oviposited
on Carduus spp., Cirsium, Cynara, Silybum, and
Galactites; larvae developed to adult on Carduus
nutans, C. acanthoides, Cirsium, Cynara, Onopor
dum, and Galactites
Field tests to study biology and survival on
Cynara begun at Rome, 1975: Adults (1,400 + )
45
sent to SEA-AR lab., Albany, Calif., to test for

potential release against Carduus tenuiflorus, C.
nutans, and C. pycnocephalus, 1970-73; tests un
successful, few adults, no egg production;
some adult feeding on Carduus nutans, Cirsium
spp., and Cynara; in 1975, adults (900 + ) sent
from Rome lab. to VPI for initiation of testing
of C. trimaculatus at that lab.
Sources:
Rome Rpts. (1964, 1966, 1968-70, 1975); Delemont Rpt. (1965); Zwblfer (1965a); K. E. Frick in
litt. (1966, 1969); Zwblfer and Harris (1966);
Albany Rpts. (1970-73); Goeden (1974); Goeden
et al. (1974); Maw (1976)
Chlorophanus viridis (L.)
Plant recs.: Cirsium (Austria, Rhine Valley); Carduus
tenuiflorus, C. crispus, C. personatus, Aster,
Chrysanthemum, Tussilago, Rudbeckia, Achillea,
Senecio, Calendula, Echinops, Silybum, Centaurea, Taraxacum, Sonchus, Inula, Cnicus,
Cynara (in lab. tests only)
Remarks:
In lab. tests (Delemont, 1963), sp. fed heavily
or regularly on all these listed plants except
last three, on which only nibbling occurred;
polyphagous sp. of no biological control in
terest
Sources:
Cleonus piger Scopoli
Plant recs.: Carduus nutans, C. acanthoides, C. pycnocepha
lus, C. defloratus, C. crispus, C. edelbergii, Cir
sium, Arctium, Onopordum, Silybum, Centaurea,
Cnicus, Rumex, Inula, Zinnia, Rudbeckia, Carthamus, Cynara, Taraxacum, Lactuca, Sonchus,
Leontodon, Carlina, Cicer, Cannabis, Ziziphus,
Tanacetum (Europe, Asia, N. Amer., and in lab.
tests - see remarks)
Remarks:
A crown and root borer; adults feed on foliage;
larvae bore into basal part of roots and also at
tack rosettes of blooming host plant; larvae in
flict damage, causing stunting and premature
dying; reported as an economic pest of Cynara
in Italy, will feed on wide range of Compositae
Field hosts: Carduus nutans (Italy, Europe), C.
edelbergii (Pakistan, Kashmir), C. acanthoides

(Europe), Cirsium (Pakistan, Switzerland, Italy,
N. Amer.), Cnicus and Rumex (Pakistan; stage
unknown), Silybum (Europe; adults only,
Pakistan), and Centaurea (USSR - Caucasus);
N. Amer. distrib.: N.Y. (since 1929), Pa. (1956),
Mich. (1974), and Ont. and Que. on Cirsium
arvense (1940, 1971)
Results of lab. tests (Delemont 1963, 1968):
Adults fed heavily or regularly on Carduus
nutans, C. defloratus, C. crispus, Inula, Zinnia,
Rudbeckia, Cnicus, Carthamus, Arctium, Cynara,
Onopordum, Silybum, Cirsium, Centaurea, Tarax
acum, Lactuca, Sonchus, Leontodon, and occa
sionally on Tanacetum; found breeding on
Carduus nutans, Cirsium, and Carlina
Sources:
Mellini (1951a); Anderson (1956); A17-ENT-9
Rpts. (1960-62, 1965); Delemont Rpts. (1962-63,
1968); Rome Rpt. (1963); Zwblfer (1965a);
Peschken (1971); Harris and Zwblfer (1971);
Baloch et al. (1971); Goeden (1974); Maw (1976);
D. Whitehead, pers. comm. (1977)
Conotrachelus sp.
Remarks:
Rare; adults on leaves; polyphagous crop pest
Sources:
Donus crinitus (Boheman)
Remarks:
Occasional larvae and adults eat leaves
Sources:
Goeden (1974)
Hypera spp.
Plant recs.: Carduus nutans (Italy, Europe); C. pycnocephalus
(Italy, S. Calif.)
Remarks:
Occasional visitor on C. nutans in Europe,
locally common; larvae and adults eat leaves
in Calif.
Sources:
L. Andres in litt. (1961); Zwblfer (1965a); Goeden
(1974)
Hypera comata Boheman
Plant recs.: Carduus nutans, Cirsium (Switzerland);
Tussilago, Aconitum, Chaerophyllum (literature);
Solidago, Erigeron, Aster, Zinnia, Achillea,
Chrysanthemum, Tussilago, Calendula, Echinops,
Arctium, Silybum, Cirsium, Lactuca (in lab. tests)

Remarks:
47
Field rec. on C. nutans was of larvae; in lab.

tests, species found to have wide host range
and thus of no interest for biological control
Sources:
Hypera postica Gyllenhall (= Phytonomus variabilis Herbst)
Plant recs.: Carduus edelbergii, Cynoglossum, Medicago,
Polygonum, Ziziphus (Pakistan); Cirsium
(Canada)
Remarks:
Larvae feeding on leaves of C. nutans; polyphagous; pest of alfalfa
Sources:
(1971); Maw (1976)
Larinus spp.
Plant recs.: Carduus defloratus (Europe); C. crassifolius, C.
acanthoides (Austria); C. pycnocephalus, C.
nutans (Italy); C. chrysacanthus (Yugoslavia)
Remarks:
Larvae (only) found in flower heads of first of
these three listed plants; stage of insect found
on last three of these recs. is unknown
Sources:
L. Andres in litt. (1961); Delemont Rpt. (1962);
Zwblfer (1965a); Zwblfer et al. (1971)
Larinus australis Capimont
Plant recs.: Centaurea (Austria, France, Switzerland,
Yugoslavia, Italy, Spain, Hungary); Carduus
defloratus, C. nutans, C. tenuiflorus, C. personatus, Centaurea, Arctium, Cirsium, Silybum,
Cynara, Onopordum, Carthamus, Cnicus, Lactuca
(in lab. tests)
Remarks:
In lab. tests, L. australis adults fed heavily on
Centaurea and regularly on all plants listed
here except Lactuca and 4 Carduus spp., on
which only slight feeding occurred; listed as
meriting further study for biological control of
Centaurea
Sources:
Delemont Rpts. (1963-64); Rome Rpt. (1963);
Larinus brevis (Herbst) (= L. sanctaebalmae Alberville)
Plant recs.: Carlina (Switzerland, Austria, Germany);
Atractylis (France); Carduus nutans, C. personatus, Carlina, Rudbeckia, Tussilago, Echinops,
Cirsium, Cynara, Onopordum, Centaurea (in lab.
tests)
Remarks:
Adults feed on young leaves; larvae feed
within receptacle, destroying ovaries and

seeds; species prefers Carlina; specimens from
Carlina vulgaris differ from C. acaulis (ident.
questioned)
In lab. tests: L. brevis adults fed heavily on
Carlina or regularly on all these listed plants
Recorded parasites: Exeristes roborator (F.),
Tetrastichus sp., and braconid sp.
Sources:
Zwblfer et al. (1971)
Larinus carlinae Olivier ( = L. planus Germar)
Plant recs.: Carduus crispus, C. pycnocephalus, C. acanthoides, C. nutans, C. nigrescens, C. litigiosus, C.
tenuiflorus, C. personatus, C. defloratus, Galactites, Cirsium, Centaurea, Carlina, Arctium,
Cynara, Onopordum, Silybum, Zinnia, Erigeron,
Helianthus, Echinops, Xeranthemum, Cnicus,
Carthamus (Europe and lab. recs. - see
remarks)
Remarks:
Adults feed on foliage, damage not particu
larly noticeable; larvae feed on flower buds,
destroying ovarioles, etc.; feeding preference:
Carduinae and some Centaureinae, indices
high on Silybum-Cirsium-Carduus
crispus (Rhine Valley, Europe), C. pycnocephalus
and C. acanthoides (Europe, France, Italy),
Galactites (Europe), and Cirsium (France,
Switzerland, Austria); adults (only) from
Carduus acanthoides, C. nutans, C. nigrescens, C.
litigiosus, and Centaurea (Europe); larvae from
Carduus tenuiflorus, Centaurea, and Carlina;
other distrib. S. Germany and Caucasus
heavily or regularly on Carduus nutans, C.
tenuiflorus, C. crispus, C. personatus, Silybum,
Cirsium, Centaurea, Arctium, Cynara, Onopor
dum, Zinnia, and occasionally or irregularly on
Erigeron, Helianthus, Echinops, Xeranthemum,
Cnicus, Carthamus, and Carduus defloratus; suc
cessful oviposition obtained on Cirsium and
Carduus defloratus; successful development of
larvae observed on Carduus crispus, C.
defloratus, and Cirsium; small number of adults
shipped to Canada in 1962 and 1964 for testing
49
Recorded parasites: "Ephialtes sp." (France)

and Bracon sp.
Sources:
Delemont Rpts. (1961-64); L. Andres in litt.
(1961); Rome Rpt. (1963); CIBC Ann. Rpt. (1964);
Larinus cynarae F.
Plant recs.: Carduus pycnocephalus, C. nutans, Cirsium,
Silybum, Onopordum, Cynara (Europe)
Remarks:
Adults and larvae oligophagous; in flowers
Sources:
Zwblfer et al. (1971)
Larinus flavescens Germar
Plant recs.: Carduus chrysacanthus (Yugoslavia); Silybum,
Centaurea, Carthamus, Echinops, other plants (S.
Europe, N. Africa)
Remarks:
Adults rec. in flower heads of Carduus; larvae
in flower heads of other plants
Sources:
F4-ENT-5 Rpt. (1969); Zwblfer et al. (1971); F4ENT-16 Rpt. (1974)
Larinus jacae F.
Plant recs.: Carduus acanthoides, C. nutans, C. personatus,,
C. pycnocephalus,, C. tenuiflorus, C. crispus, C.
defloratus, C. chrysacanthus, Cirsium,, Galactites,
Silybum, Cynara, Centaurea, Carthamus, Onopor
dum, Arctium, Carlina, Echinops, Chrysan
themum, Zinnia, Erigeron, Aster, Achillea, Xeranthemum (Europe and lab. recs. - see remarks)
Remarks:
Adults feed on host leaves and peduncles
without causing visible damage; larvae (1-3)
feed in capitulum; subtribe Carduinae pre
ferred
acanthoides (France, Austria, Europe), C. nutans
(Austria, Italy, Europe), C. personatus (France),
C. pycnocephalus, C. tenuiflorus (Italy), and
Cirsium, Centaurea, Galactites (Europe); adults
(only) from Cynara (Italy, 1 "resting"); stage
unknown from Carduus chrysacanthus (Yugo
slavia); distrib. also in SW Asia to Turkestan;
not observed in Rhine Valley or Swiss Valais
crispus, C. defloratus, C. tenuiflorus, C. per
sonatus, Cirsium, Silybum, Cynara, Zinnia,
Echinops, Chrysanthemum, Carlina, Arctium,
50

Onopordum, Centaurea, Carthamus, and occa
sionally on Erigeron, Aster, Achillea, and
Xeranthemum', successful larval development
on Carduus acanthoides, C. nutans, and C. personatus
Recorded parasites: Tetrastichus sp., Bracon
urinator F., and Exeristes roborator (F.)
Sources:
Mellini (1951b); L. A. Andres in litt. (1961); Delemont Rpts. (1963-64); Rome Rpt. (1963); Zwblfer
(1965a); Zwblfer et al. (1971)
Larinus scolymi Olivier
Plant recs.: Carduus pycnocephalus (Italy, France); C. nutans
(Europe); Cynara (Italy, France, Europe);
Cirsium (Europe); Onopordum (Europe);
Echinops (Algeria)
Remarks:
Recorded as pest of Cynara in Italy and S.
France; all these other recs. are of adults only
Sources:
L. A. Andres in litt. (1961); Rome Rpt. (1963);
Delemont Rpt. (1964); Zwolfer (1965a); Zwolfer
et al. (1971)
Larinus sturnus Schaller
Plant recs.: Carduus acanthoides, C. nutans, C. crispus, C.
nigrescens, C. personatus, C. defloratus, C.
tenuiflorus, Cirsium, Centaurea, Arctium,
Silybum, Cynara, Onopordum, Carthamus,
Echinops, Rudbeckia, Erigeron, Zinnia, Xeran
themum, Cnicus, Aster, Carlina (Europe and lab.
Remarks:
Adults feed on foliage and peduncle; larvae
mine receptacle destroying ovarioles
nutans (Swiss Alps, French Rhine Valley,
Italy), C. acanthoides (centr. Europe), C.
nigrescens (France), C. personatus (Europe),
Cirsium (Swiss Alps, French Rhine Valley,
Italy), Centaurea (French Rhine Valley, Italy),
and Arctium (Austria); adults (only) from
Carduus crispus (Switzerland); distrib. also E.
Asia and N. Africa (from literature) and N.
Yugoslavia (Cynara given as host in literature);
Austrian population is only Larinus found in
Europe to attack Arctium; adults from Austria
prefer Arctium over Carduus or Centaurea; those
bred from Carduus (from Rhine Valley) pre
51
ferred Carduus to Arctium or Centaurea, and

those from Centaurea (from Rhine Valley)
preferred Centaurea over Carduus or Arctium
Results of lab. tests (Delemont): Adults
(Rhine Valley Carduus population) fed heavily
or regularly on Carduus nutans (also in lab.
tests at SEA-AR lab. in Rome), C. crispus, C.
personatus, C. tenuiflorus, C. defloratus, Cirsium,
Centaurea, Silybum, Cynara (also at Rome),
Onopordum, Carthamus, Echinops, Arctium,
Rudbeckia, and occasionally on Erigeron, Zin
nia, Xeranthemum, Cnicus, Aster, and Carlina;
oviposition obtained on Carduus defloratus, Cir
sium, Centaurea, and Arctium (by Arctium
population only); larvae bred successfully
from Arctium (Austrian population only),
Cirsium, Carduus nutans, and Centaurea
Recorded parasites: Bracon sp. and
chalcidoids; a possible biocontrol agent for
Centaurea or Carduus
Sources:
Delemont Rpts. (1962, 1964); Rome Rpt. (1963);
Zwblfer (1965a); Zwblfer et al. (1971); CIBC Rpt.
(1971)
Larinus turbinatus Gyllenhall
Plant recs.: Carduus nutans, C. pycnocephalus, C. crispus, C.
personatus, C. tenuiflorus, C. defloratus, Cirsium,
Onopordum, Galactites, Centaurea, Serratula,
Echinops, Cnicus, Arctium, Cynara, Silybum,
Tanacetum, Zinnia, Xeranthemum, Carlina,
Senecio, Leontodon (Europe and/or in lab. tests
- see remarks)
Remarks:
Adults feed on leaves, stems, buds, and
peduncles of hosts; larvae in flower heads
seem restricted to Cirsium
Field recs.: Adults and larvae from Cirsium
(Germany, France, Switzerland, Austria,
Yugoslavia, Italy) and Carduus nutans and
Silybum (France); larvae (only) from Serratula
(Austria); adults (only) from C. nutans
(Austria), C. pycnocephalus (France, Italy),
Onopordum (Italy), Galactites (Italy), and
Galactites and Centaurea (Europe)
Sources:
crispus, C. personatus, C. tenuiflorus, Cirsium,

Echinops, Cnicus, Arctium, Cynara, Onopordum,
Silybum, Centaurea, Tanacetum, and occasion
ally (nibbled) on Carduus defloratus, Leontodon,
Senecio, Carlina, Xeranthemum, and Zinnia;
oviposition obtained only on Cirsium
Recorded parasites: Bracon sp. and
chalcidoids; small shipments of L. turbinatus
adults made to Canada in 1962 and 1965 for
tests
H. L. Parker in litt. (1957); Delemont Rpts.
(1961, 1964); Rome Rpt. (1963); CIBC Ann. Rpt.
(1964); Zwblfer (1965a); Zwblfer et al. (1971)
Larinus ursus F.
Plant recs.: Carduus nutans, Centaurea, Carlina (S. Europe)
Remarks:
Adults rec. on Carduus and Centaurea; larvae on
Carlina; in flower heads
Sources:
Zwblfer (1965a); Zwblfer et al. (1971 )
Leptomias jekeli Faust
Plant recs.: Carduus edelbergii, Cnicus, Dipsacus, Abies
(Pakistan)
Remarks:
Adults feeding on leaves
Sources:
Leptomias stolickzae Faust
Remarks:
Adults feeding on leaves; no other hosts known
Sources:
Lixus spp.
Plant recs.: Carduus pycnocephalus, C. nutans (Italy,
France); C. crispus (Rhine Valley)
Remarks:
Larvae in flower head of C. crispus
Sources:
Rome Rpts. (1960-61); L. A. Andres in litt.
(1961); Delemont Rpt. (1963)
Lixus algirus (L.)
Plant recs.: Carduus nutans, C. litigiosus, C. acanthoides, C.
pycnocephalus, C. defloratus, C. personatus, C.
crispus, C. tenuiflorus, Cirsium, Onopordum,
Galactites, Cnicus, Silybum, Malva, Althea, Vicia,
Centaurea, Sonchus, Carthamus, Arctium,
Cynara, Rudbeckia, Zinnia, Calendula, Xeran
themum, Chrysanthemum, Senecio, Echinops,
Carlina (Europe, Pakistan, and/or in lab. tests
- see remarks)

Remarks:
53
Adults feed on leaves and oviposit in stems of

hosts; found to be rather polyphagous (see
following remarks); a "potential pest" in
Europe and a "known pest" in Pakistan
Field recs.: Ovipositing adults and larvae
from Cirsium (Switzerland, France, Rhine
Valley), Carduus nutans (Austria, Europe), and
C. litigiosus (France); adults (only) from
Onopordum and Galactites (France), Carduus
acanthoides, C. pycnocephalus, and Silybum
(Europe), and Cnicus (Pakistan); further
literature recs. (adults only) from Centaurea,
Malva, Althea, and Vicia (damage reported to
flowers and foliage of vetch); larvae recorded
from 'Malvaceae' in literature
heavily or regularly on Cirsium, Silybum, Car
duus crispus, C. defloratus, C. nutans, C. personatus, Sonchus, Carthamus, Arctium, Cynara,
Centaurea, Rudbeckia, Onopordum, and occa
sionally on Zinnia, Calendula, Xeranthemum,
Cnicus, Chrysanthemum, Senecio, Echinops, and
Carlina; of no interest in biological control
Sources:
Delemont Rpts. (1962, 1964); A17-ENT-9 Rpt.
(1965); Zwblfer (1965a); Goeden (1974)
Lixus (Lixochelus) cardui Olivier
Plant recs.: Onopordum (Austria, Europe); Carduus nutans
(Austria, Italy?); C. pycnocephalus (Europe,
Italy?); C. acanthoides, Cirsium (Europe);
Silybum (Syria); Carduus defloratus, C.
tenuiflorus, C. crispus, Arctium, Cynara, Silybum,
Centaurea, Carlina (in lab. tests only)
Remarks:
Larvae found breeding in field only on
Onopordum; other field recs. listed here are of
adults only; Italian recs. are of L. "prob.
cardui"
heavily or regularly on Onopordum, Cirsium,
Carduus defloratus, C. nutans, C. tenuiflorus, C.
crispus, Arctium, Cynara, Silybum, Centaurea,
and occasionally on Carlina
Sources:
Delemont Rpt. (1963); Rome Rpt. (1963); Zwblfer
(1965a); F4-ENT-16 Rpt. (1976)
54
COLEOPTERAContinued
CurculionidaeContinued
Lixus sp. nr. L. elongatus Goeze (or L. rufitarsis Boheman)
Plant recs.: Carduus pycnocephalus (Lebanon, Syria); C.
tenuiflorus, C. pycnocephalus, C. nutans, C.
litigiosus, C. defloratus, Galactites, Cirsium,
Silybum, Cynara, Onopordum, Xeranthemum,
Helianthus (Europe and/or in lab. tests - see
remarks)
Remarks:
Field recs.: Ovipositing adults from Carduus
tenuiflorus, C. pycnocephalus, and C. litigiosus
(France); larvae observed breeding only on C.
tenuiflorus; adults (France) and larvae (Rhine
Valley) from Cirsium; adults from Carduus
nutans and Galictites (Europe)
heavily or regularly on Carduus tenuiflorus, C.
nutans, Cirsium, Silybum, Onopordum, Cynara,
and Xeranthemum; in early tests, also fed on
Carduus defloratus and Helianthus; 130 adults
sent to Canada in 1964
Sources:
Delemont Rpts. (1962-64); CIBC Ann. Rpt.
(1964); Zwblfer (1965a); F4-ENT-16 Rpt. (1974)
Lixus (Lixochelus) elongatus Goeze
Plant recs.: Carduus nutans, C. acanthoides, C. crispus, C.
pycnocephalus, C. Inigrescens, C. defloratus, C.
tenuiflorus, C. personatus, Cirsium, Silybum,
Onopordum, Centaurea, Cynara, Arctium, Carlina,
Xeranthemum, Erigeron (Europe and/or in lab.
Remarks:
Adults feed on foliage, often skeletonizing
host; larvae feed in root collars and in lower
parts of stem; feeding restricted enough to war
rant further testing for biological control of
Carduus spp. or Cirsium vulgare (not C. arvense)
acanthoides (France, Europe), C. nutans (Italy,
Austria, Rhine Valley, Europe), C. crispus
(Europe), and Cirsium (Rhine Valley,
Switzerland, Austria, France); adults (only)
from Carduus pycnocephalus (Italy) and C.
nigrescens, Silybum, and Onopordum (Europe)
heavily or regularly on Cirsium, Silybum, Car
55
duus crispus, C. personatus, C. tenuiflorus, C.

nutans, C. defloratus, Centaurea, and occa
sionally on Xeranthemum, Carlina, Arctium,
Cynara, Onopordum, and Erigeron; oviposition
obtained only on Carduus and Cirsium; ship
ment of 110-120 adults made to Canada in
1963; judged unsuitable for biological control
of Cirsium arvense in Canada, larvae do not sur
vive on this species
Sources:
Mellini (1951a); L. A. Andres in litt. (1961); Delemont Rpts. (1962-63); Belleville Res. Inst. Rpt.
(1962-63); CIBC Ann. Rpt. (1963); Rome Rpt.
(1963); Zwblfer (1965a); Goeden (1974)
Lixus junci Boheman
Plant recs.: Carduus nutans, Cirsium (Europe)
Remarks:
Occasional visitor on C. nutans
Sources:
Zwblfer (1965a)
Lixus sp. prob. L. scolopax Boheman
Plant recs.: Carduus pycnocephalus, C. nutans, Silybum,
Cynara (Italy)
Remarks:
Adult feeding and oviposition observed on C.
nutans only
Sources:
Rome Rpt. (1963); Goeden (1974)
Lixus spartii Olivier
Remarks:
Occasional visitor
Sources:
Zwblfer (1965a)
Otiorhynchus sp.
Plant recs.: Carduus personatus (Europe)
Remarks:
Occasional visitor
Sources:
Zwblfer (1965a)
Pseudocleonus grammicus Panzer
Plant recs.: Centaurea (Yugoslavia, France); Carlina
(Europe); Carduus personatus, Cirsium, Silybum,
Onopordum, Arctium, Carthamus, Echinops,
Tanacetum, Chrysanthemum, Inula (in lab. tests
only)
Remarks:
Field recs. on Centaurea are of larvae found in
the roots; recs. on Carlina are of adults on
foliage
Results of lab. tests (Delemont) for Cirsium:
Adults (from Carlina) fed regularly on Carduus
personatus (but not on 3 other spp. of Carduus),
Centaurea, Cirsium, Silybum, Onopordum, Arc

tium, Carthamus, Carlina, Echinops, Tanacetum,
Inula, and Chrysanthemum
Sources:
Delemont Rpts. (1963, 1965); CIBC Ann. Rpts.
(1963, 1965); Zwblfer (1965a)
Rhinocyllus conicus (Froelich)
Plant recs.: Carduus nutans, C. acanthoides, C. personatus, C.
pycnocephalus, C. nigrescens, C. crispus, C.
litigiosus, C. tenuiflorus, Cirsium, Silybum,
Onopordum, Centaurea, Cnicus, Galactites,
Xeranthemum, Carlina, Tagetes, Carthamus,
Helianthus, Cichorium, Cynara, Lactuca, rye,
mangel, apple, radish (Europe and/or in lab.
Remarks:
Adults gnaw peripheral tissue of stems,
peduncles, and leaves; feeding is usually of lit
tle importance, less pronounced than that of
Larinus spp.; eggs laid externally on flower
bud; larvae (1-16 per flower (200 larvae per
flower in Mont.)) mine receptacle, 3-4 larvae
prevent seed formation; host range CarduusCirsium-Silybum (occasionally Onopordum) is
believed to be highly stable, i.e., no tendency of
R. conicus to form "races" attacking other
plants
nutans (Switzerland, Austria, Italy, Europe),
rarely C. acanthoides (France), C. personatus
(Austria, Switzerland), C. pycnocephalus
(France, Spain, Italy), C. nigrescens (France), C.
crispus (Switzerland, France), mainly in large
flowered species of Carduus and Cirsium
(France, Switzerland, Germany, Austria),
Silybum (France, Italy), Onopordum (France),
and Galactites; adults (only) from Carduus
litigiosus (France), Xeranthemum, Centaurea,
Cnicus, and Carlina (Europe)
Other distrib.: N. Africa, S. England, E.
Europe (Hungary, Poland, Czechoslovakia,
Romania), and USSR (Ukraine, Caucasus,
Kazakhistan, Altai)
Recorded European parasites: Tetrastichus
sp., Bracon urinator ., Aritranis ( = Holocryptus)
nigripes (Gravenhorst), and an egg parasite
Results of lab. screening tests (all tests at
57
Delemont except a few at Belleville

(B)): Adults fed heavily or regularly on
Carduus nutans, C. crispus, C. personatus, C.
acanthoides (also B), Cirsium (also B), Silybum,
Centaurea, and C. tenuiflorus (last 3 in 1 test
only, "occasionally" in 2d test), and occa
sionally Tagetes, Xeranthemum, Onopordum,
Carthamus, Cnicus, Helianthus, Cichorium,
Cynara, Lactuca, and (at B only) rye, mangel,
apple, and radish; normal (4.5 eggs per female)
oviposition obtained only on Carduus nutans, C.
personatus, and Cirsium; atypical oviposition
occasionally (0.3 egg per female) obtained on
other (unpalatable) test plants (Carthamus,
Helianthus, Cnicus, and Cynara); successful lar
val development only in Cirsium, Carduus
nutans, and C. personatus (successful larval
transference only into last one); in lab., prefers
Silybum to Cirsium for oviposition (Albany
Rpt., 1970)
History: First studied in 1962 by CIBC
lab., Delemont; tested at Delemont and
Belleville, 1962-68; U.S. Working Group on
Biological Control of Weeds approved release
in Canada, July 1968, and in U.S., Feb. 1969;
adults collected and shipped by CIBC, Dele
mont, to Canada in 1964 (390), 1967-68
(2,500 + ), 1969-70 (many thousand), and to
VPI, Blacksburg, Va., and SEA-AR lab.,
Albany, Calif., in 1969-73 (several thousand);
adults also collected and shipped by SEA-AR
lab., Rome, Italy, to Albany in 1969, 1971-75
(many thousand) and to VPI in 1971 (1,400)
Releases: First released by Canada Agr. on
Carduus nutans in Sask. (400) and C. acanthoides
in Ont. (370) in 1968; additional Canadian
releases on C. nutans in Sask. (3,795) and in
Ont. at 3 sites (4,952) in 1969; on C. acanthoides
in Ont. at 3 sites (5,275) and C. nutans in Ont. at
2 sites (450) and on both spp. in Ont. (700) in
1970; released on C. nutans in Ont. (10,100),
Sask. (120), and Man. at 2 sites (4,500) in 1974;
and in Ont. at 2 sites (2,185) in 1975 (it was also
released in New Zealand in 1975)

First released in U.S. by VPI on C. nutans, C.
thoermeri, and C. acanthoides in Va. at 14 sites
(2,000) and by SEA-AR, Albany, on C. nutans
in Nebr. (60), Mo., Calif. (12), and Mont. (1,805)
in 1969; addit. U.S. releases on C. nutans, C.
thoermeri, and C. acanthoides in Va. at 9 sites
(5,950) in 1970; on Silybum marianum in Calif.
(1,141) and on C. nutans in Mont. (667) in 1971;
on S. marianum in Calif. at 5 sites (2,062) and on
C. nutans in Nebr. at 2 sites (663), Mont. (492),
and Idaho (800) in 1972; on C. nutans in Nebr. at
2 sites (750), S. Dak. (1,000), Kans. (725), and
Mont. (425); on S. marianum in Calif. at 4 sites
(1,143) in 1973; on C. nutans in Calif. (440),
Nebr. (440), Idaho (440), Colo. (440), and Nev.;
on S. marianum in Calif. (370) in 1974; and on C.
pycnocephalus in Calif. at 6 sites (3,069) in 1975
Recoveries and establishments: Well estab
lished in Sask. by 1974, seed production
reduced by 63 percent at 1969 site, and
recolonization begun in 1974; by 1976, 85 per
cent of seeds destroyed at 1 site in Sask.; also
reported established in Ont.; in Va., R. conicus
is well established at several sites on Carduus
nutans and C. thoermeri, but it is less effective
against C. acanthoides; at 1 Va. site, C. nutans
was reduced by 95 percent 6 yrs. after initial
release of 100 adults; dispersal was 1.6 km
each year and 32 km 6 yrs. after release
recolonization begun in 1972; at a 1971 Silybum
release site in Calif., 67 percent of flower heads
were infested by 1972, later increasing to 94
percent in 1973; it became established at 5 of
10 release sites in S. Calif. (Goeden and Ricker,
1977); it also attacks C. pycnocephalus in Calif.;
in Mont., beetles at 5 1969-73 release sites had
spread over an area of 1,280 km2 by 1974, at
tacking C. nutans and Cirsium arvense;
established in Colo, in 1976
Recolonizations: Over 20,000 adults col
lected from Va. establishments for recoloniza
tion in Va., W. Va., and Tenn. in 1972; reported
established, infesting 70-80 percent of flower
heads of Carduus crispus 3 yrs. after release in
Sources:
59
W. Va.; further recolonization on C. crispus and

C. nutans in W. Va. made in 1975; 20,000 adults
collected annually in 1973-75 for redistribu
tion against C. nutans and C. acanthoides in Va.;
additional beetles collected in Va. for release
in Md. (757), N.J. (600), and Pa. (70) against C.
nutans and C. thoermeri in 1975, and Md. (416)
against C. nutans and Pa. (297) against C.
nutans, C. thoermeri, and C. acanthoides in 1976;
established in Md. and Pa. in 1978. In Canada,
collections for recolonization were made in
Sask., Que. (10,100), and Man. (4,500) in 1974;
establishments reported in both Que. and Man.
in 1975; in Calif., beetles were collected at
Silybum establishments for recolonization on
S. marianum at new Calif. sites in 1974 (2,016 at
4 sites) and 1975 (365); in 1975, collections were
made at establishments in Mont, for release on
C. nutans in Calif. (500), Colo. (500), Idaho
(1,000), Iowa (500), Minn. (500), N. Dak. (500), S.
Dak. (500), Utah (1,000), Wash. (500), and Wyo.
(500); collections also made in Va. in 1975 for
releases in Kans. (375), Ky. (430), La. (350), Mo.
(500 established and spread 2 mi by 1976),
Okla. (388 established in 1977), and Tenn. (500)
Mortality factors: In Va., larval and egg
mortality of 68 percent was caused by weather,
crowding, and parasitism by Aliolis curculionis
(Fitch), Bracon mellitor Say, and a parasite of
Lepidoptera, Campoplex polychrisidis Viereck;
in Calif., R. conicus was destroyed by
Hippodamia sp., Raphidia adnixa Hagan,
spiders, pyemotid mites, rodents, and cattle
grazing thistles (Goeden and Ricker, 1977)
Mellini (1951b); Delemont Rpts. (1962-64,
1966-69, 1971); Zwblfer (1963, 1965a, 1967);
Rome Rpts. (1963, 1969, 1971-75); CIBC Ann.
Rpts. (1964, 1966-70); Williamson (1966-73); H.
Zwblfer in litt. (1967); Albany Rpts. (1969-75);
Zwblfer and Pattullo (1970); Harris and Zwblfer
(1971); R. D. Hendrick in litt. (1971); Hawkes et
al. (1972); Andres and Davis (1973); Canada
Agr. Res. Branch Rpt. (1974); Goeden (1974);
Goeden and Ricker (1974); Surles et al. (1974);

Kok (1974, 1975b); P. Harris in litt. (1975); Kok
and Surles (1975); Surles and Kok (1975, 1976,
1977); Surles (1975); Surles et al. (1975); Hacker
(1975); Batra in litt. (1975-76); Surles (1975); M.
G. Maw in litt. (1976); Letendre et al. (1976);
Hodgson and Rees (1976)
Rhodobaenus tredecimpunctatus (Illiger)
Plant recs.: Carduus sp. (Va.)
Remarks:
Feeding
Sources:
L. T. Kok in litt. (1976)
Sitona callosus Gyllenhall
Plant recs.: Carduus edelbergii, Cnicus (Pakistan); alfalfa
(USSR)
Remarks:
Adults feeding on leaves; "a known pest"
Sources:
Alimdzhanov (1941); A17-ENT-9 Rpt. (1965);
Tanymecus pallia t us F.
Plant recs.: Carduus acanthoides (Austria); Cirsium (France,
Austria, Europe); Vitis, Chrysanthemum (also in
lab. test); Beta, cereals (in literature); Solidago,
Erigeron, Aster, Helianthus, Zinnia, Achillea,
Tussilago, Calendula, Xeranthemum, Carthamus,
Arctium, Cynara, Onopordum, Silybum, Centaurea, Taraxacum, Lactuca (all in lab. tests
only)
Remarks:
These field recs. are of adults only; adults feed
on foliage; occasionally found listed in
literature as pest of Vitis, Chrysanthemum,
Beta, and cereals
In lab. tests (Delemont): Adults fed on all
these plants except Carduus (Vitis, Beta, and
cereals not incl. in tests); of no interest in
biological control; broad feeding range
Sources:
Delemont Rpts. (1962-63); Zwblfer (1965b)
Trichosirocalus horridus (Panzer) (= Ceuthorrhynchidius horridus
(Panzer))
Plant recs.: Carduus nutans, C. pycnocephalus (Italy); C.
tenuiflorus, C. nutans, C. acanthoides, C. crispus
(Europe); Galactites, Cirsium (Italy); Onopordum
(Europe); Cynara, other plants (lab. tests only)
Remarks:
Adults in lab. feed at base of leaves,
ovipositing on tips of new growth; larvae mine
and bore to base of growing leaves, then feed
Sources:
61
on crowns (12-18 larvae per crown not uncom

mon); reared from Carduus nutans, C. pycnocephalus, Galactites, and Cirsium; adults and
larvae found on Carduus acanthoides and
Onopordum; adults rare on new stands of C.
nutans in Europe
Screening tests conducted at Delemont,
1964, and Rome (preliminary, 1964):
Hexamermis brevis (nematode) reared from C.
horridus larvae from crowns of Carduus nutans
(Italy)
Culture and studies conducted at Rome lab.,
1968-70: Adults fed on Carduus nutans, C. acan
thoides, Cirsium, Galactites, Cynara, and other
plants; oviposited on these plants, Carduus pycnocephalus, and other plants
Adults (6,800 + ) collected in Italy and sent
for final testing in quarantine at VPI, 1970-75;
host-specificity tests at VPI: 1st instar larvae
tested on 35 plant species; larvae were able to
complete development in Carduus nutans, C.
acanthoides, and Cirsium vulgare, not in Cynara
and Carthamus; adults and larvae fed on these
and other plants to lesser degree (Ward et al.,
1974; Kok, 1975a); biology studies (Kok et al.,
1975); European distrib. (Zwblfer and Harris,
1966); U.S. Working Group on Biological Con
trol approved release in U.S. in 1974; released
in Va. in 1975, established by 1978 (L. T. Kok,
pers. comm., 1976, 1979); adults (490 + ) sent to
Canada by CIBC-Delemont in 1975; released
in Sask. (87 in 2 sites) in 1975; bred success
fully (Harris, 1975); testing begun at SEA-AR
lab., Albany, Calif., in 1974-78, confirmed
safety to Cynara (release in Kans., N. Dak., S.
Dak., Mo., Wyo., Nebr., Mont, planned for
1978-79)
Rome Rpts. [1964-66, 1968-73); Zwblfer
(1965a); K. E. Frick in litt. (1966); Zwblfer and
Harris (1966); Frick (1969); Ward et al. (1974);
Kok (1975a); Kok et al. (1975); Harris (1975);
CIBC Ann. Rpt. (1975); L. T. Kok, pers. comm.
(1976)
62
COLEOPTERA Continued
Buprestidae
Unidentified buprestid sp.
Remarks:
Rare; endophagous larva
Sources:
Tenebrionidae
Unident. tenebrionid sp.
Remarks:
Rare; adults feeding on leaves
Sources:
Goeden (1974)
Lathridiidae
Macrophthalmus sp.
Remarks:
Rare; adults endophagous in
oligophagous
Sources:
Goeden (1974)
flowers;
LEPIDOPTERA
Unident. microlepidoptera
Plant recs.: Carduus crassifolius (Austria)
Remarks:
Larvae found in 0.7 percent of flower heads in
area
Sources:
Arctiidae
Unident. arctiid sp.
Remarks:
Rare; larvae on leaves
Sources:
Goeden (1974)
Estigmene acrea (Drury)
Plant recs.: Carduus pycnocephalus (Calif.)
Remarks:
Rare; larvae on leaves; economic pest
Sources:
Goeden (1974)
Nymphalidae
Cynthia cardui (L.) ( = Vanessa cardui)
Plant recs.: Carduus edelbergii (Pakistan); C. nutans
(Austria, Canada); C. acanthoides (France); C.
pycnocephalus (Egypt, Calif.); C. pycnocephalus
and/or C. tenuiflorus (Calif.); Carduus spp. (U.S.,
Canada); Cynara (Italy); Centaurea (Pakistan,
Egypt); Carthamus, Cnicus, Astragalus, Silybum,
Urtica (Pakistan); Cirsium (Switzerland, France,
Canada); Onopordum (France, Austria); Carlina
(Europe); many other hosts

Remarks:
Sources:
63
Larvae feed externally on foliage; polyphagous; a pest of Cynara in France; pest of

sunflower, soybean, and other crops; recorded
parasites: Apanteles spp. (Pakistan,
Switzerland)
Huffaker (1956); A17-ENT-9 Rpts. (1961, 1965)
Delemont Rpts. (1962-63); CIBC Spec. Rpt
(1962); Rome Rpt. (1963); Albany Rpt. (1963)
Zwblfer (1965a); F4-ENT-5 Rpts. (1966-67); Har
ris and Zwblfer (1971); Baloch et al. (1971)
Goeden (1974); L. T. Kok in litt. (1976); Maw
(1976); Morihara and Balsbaugh (1976)
Noctuidae
Unident. noctuid sp.
Remarks:
Rare; larvae ectophagous on leaves
Sources:
Goeden (1974)
Autographa gamma (L.) ( = Phytometra gamma)
Plant recs.: Carduus pycnocephalus, Centaurea (Egypt, S.
Europe)
Remarks:
Rare; larvae on leaves; polyphagous crop pest
Sources:
F-4-ENT-5 Rpts. (1966-67, 1969); Goeden (1974)
Eublemma parva (Hubner) ( = Porphyrinia parva)
Plant recs.: Carduus edelbergii (Pakistan); Xanthium
(Bangladesh, Pakistan); Carthamus, Blumea, In
ula (Pakistan); Centaurea (Pakistan, Egypt);
Ziziphus (China); Tagetes, Conyza (in lab. tests
only)
Remarks:
Larvae bore into burs of Xanthium, feed on
seeds, flowers, and tender shoots; larvae
feeding in flowers and fruits of C. edelbergii;
larvae infesting drying heads of Centaurea in
Egypt; in 1960, 12 shipments (3,285 larvae) of
E. parva sent from CIBC Pakistan lab. to
Queensland, Australia, as breeding stock for
release vs. Xanthium; E. parva found to
damage Carthamus in Pakistan (1961-62); E.
parva found to be previously established in
Australia
Results of lab. tests (Rawalpindi): Larvae fed
to pupation on Tagetes, Conyza, and Carthamus;
oviposition obtained on Carthamus
Recorded parasites (Pakistan): Brachymeria
sp. nr. aegyptiaca Masi, Apanteles sp. nr. stanton

(Ashmead), Eurytoma sp., Netelia sp., and
Paraholaspis sp.
Sources:
CIBC Ann. Rpts. (1959-62, 1966-67); A17-ENT9 Rpts. (1960-65); Ghani (1963); Tsao and Wang
(1965); A17-ENT-14 Rpts. (1966-67); F4-ENT-5
and F4-ENT-16 Rpts. (1966-67, 1974); Baloch et
al. (1968, 1971)
Eublemma purpurina (Denis and Schiffermuller) ( = Thalpochares
purpurina)
Plant recs.: Carduus nutans (Italy); Cynara (lab.); Cirsium
(Europe)
Remarks:
Larvae in petioles, crown, and buds
Sources:
Mellini (1951a); Zwblfer (1965a); Rome Rpt.
(1975)
Euxoa spp. (= Agrotis spp.)
Plant recs.: Carduus pycnocephalus (Egypt, Calif.); Carduus
sp. (Va.)
Remarks:
Common feeder on foliage; larvae on leaves in
Calif. "prob.-E. tesselata (Harris)"
Sources:
F4-ENT-5 Rpts. (1967, 1969); Goeden (1974); L.
T. Kok in litt. (1976)
Euxoa segetum Schiffermuller
Remarks:
Larvae feeding in flowers; exophytic on stems
and leaves; polyphagous crop pest
Sources:
Heliothis armigera (Hiibner)
Plant recs.: Carduus edelbergii, Xanthium, Carthamus,
Datura, Chrozophora, Loranthus, Trichodesma,
Hypericum (Pakistan); many others
Remarks:
Larvae destroy flowers of Carduus nutans and
Carthamus; feed on fruits of Xanthium and on
leaves of Loranthus; polyphagous; an economic
pest
Sources:
CIBC Ann. Rpts. (1959, 1961); A17-ENT-9 Rpts.
(1960-61, 1964-65); Baloch et al. (1968, 1971)
Lycanades purpurea (Grote)
Remarks:
Larvae occasionally feeding on leaves
Sources:
Goeden (1974)
Papaipema arctivorans Hampson
Remarks:
Common feeder
Sources:
65
Papaipema nebris (GuenSe)

Remarks:
Larvae in stems; polyphagous crop pest
Sources:
Peridroma saucia (Hiibner)
Remarks:
Larvae found rarely feeding on leaves;
economic sp.
Sources:
Goeden (1974)
Geometridae
Unident. geometrid spp.
Plant recs.: Carduus acanthoides (Austria); C. defloratus
(Switzerland); C. pycnocephalus (S. Europe); C.
nutans (S. Dak.)
Remarks:
Larvae in flower heads; on leaves
Sources:
Delemont Rpts. (1962-63); Goeden (1974);
Eupithecia sp.
Remarks:
Common feeder
Sources:
Pyralidae
Unident. phycitine sp.
Plant recs.: Carduus personatus (centr. Europe)
Remarks:
Larvae endophytic in flower heads
Sources:
Zwblfer (1965a)
Unident. pyralid spp.
Plant recs.: Carduus pycnocephalus and/or C. tenuiflorus (S.
Europe, Calif.)
Remarks:
Larvae boring into stems at leaf axils in Calif.;
locally common in Europe; ectophagous on
leaves; another sp. "probably Myclopsis sp."
endophagous in flowers of C. pycnocephalus in
Calif.
Sources:
Albany Rpt. (1963); Goeden (1974)
Homoeosoma sp. ( = Phycitodes sp.)
Plant recs.
Carduus acanthoides (France); C. nutans
(Switzerland)
Remarks:
Larvae attacking flower heads
Sources:
Delemont Rpt. (1963); CIBC Ann Rpt. (1971)
Homoeosoma binaevella (Hiibner) (= Phycitodes binaevella)
Plant recs.: Carduus nutans (Europe); C. edelbergii (Paki
stan); C. acanthoides, C. personatus (Europe);

Cnicus, Echinops (Pakistan); Cirsium (France,
Europe); Centaurea, Anthemis, Senecio (Europe)
Remarks:
Larvae endophytic in capitula; 3 generations
per year; low incidence of attack generally in
lab.; fed on Carduus and Cirsium spp. only, and
not the 5 other Compositae tested; considered
of possible use in control of Carduus nutans
Sources:
A17-ENT-9 Rpts. (1962-65); Rawalpindi Rpts.
(1962, 1966, 1968); Delemont Rpt. (1963); Zwblfer
(1965a); A17-ENT-14 Rpts. (1966, 1968); CIBC
Ann. Rpt. (1970); Baloch et al. (1971)
Homoeosoma electellum (Hulst)
Plant recs.: Carduus nutans (Sask., S. Dak.)
Remarks:
Larvae in flowers; pest of sunflowers
Sources:
M. G. Maw in litt. (1976); Morihara and
Balsbaugh (1976)
Homoeosoma nebulellum (Denis and Schiffermiiller)
Plant recs.: Carduus acanthoides (Austria); C. crispus (Rhine
Valley); C. nutans (Europe); Cirsium, Silybum,
Centaurea (Europe)
Remarks:
Sources:
Myelois sp.
Plant recs.: Carduus nutans (Rhine Valley)
Remarks:
Sources:
Myelois cribrumella Hiibner (= M. "cribrella")
Plant recs.: Carduus nutans (Rhine Valley, Europe); C. acan
thoides (Europe); Cirsium (Germany, France,
England, Switzerland, Austria); Onopordum
(Austria); Centaurea, Arctium, Carthamus
(Europe); Jurinea, Echium (literature); Aster,
Helianthus, Senecio, Silybum, Chrysanthemum (in
lab. oviposition tests only)
Remarks:
Larvae endophytic in stems; also in flower
heads of Cirsium
Results of lab. tests (Del&mont, for Cirsium):
Oviposition obtained on Aster, Helianthus,
Chrysanthemum, Senecio, Silybum, Centaurea,
Arctium, and especially on Onopordum and
Cirsium (no Carduus incl. in tests); found to have
a wide host range, dropped from further con
sideration in biological control

Sources:
67
Mellini (1951a); Delemont Rpts. (1962-63);

Zwblfer; (1965a)
Noctuelia floralis (Hiibner)
Plant recs.: Carduus pycnocephalus (Egypt)
Remarks:
Rare
Sources:
F4-ENT-5 Rpt. (1967)
Ostrinia kasmirica (Moore)
Remarks:
In stems
Sources:
Ostrinia nubilalis (Hiibner) (= Pyrausta nubilalis)
Plant recs.: Carduus edelbergii (Pakistan) occasionally
Cnicus (Pakistan)
Remarks:
Larvae in stems; this "strain" (of 0. nubilalis
from Carduus nutans) was not observed on corn;
Ephialtes sp. and Chlorocytus sp. parasites of 0.
nubilalis on Cnicus; this sp. is a common crop
pest
Sources:
(1971)
Phycita sp.
Remarks:
Larvae feeding on leaves
Sources:
Pyrausta silhetalis Guen6e
Remarks:
Larvae bore in stems; no other hosts known
Sources:
Olethreutidae (= Eucosmidae)
lEpiblemma sp.
Plant recs.: Carduus pycnocephalus (Europe)
Remarks:
Larvae endophagous in stems
Sources:
Goeden (1974)
Epiblemma sp. nr. E. scutulana (Denis and Schiffermuller)
Plant recs.: Carduus nutans, Cirsium (Italy)
Remarks:
Larvae mine stems and flower heads; damage
to capitula of C. nutans reported to be severe
Sources:
P. H. Dunn in litt. (1966); Rome Rpt. (1975)
Epiblemma scutulana (Denis and Schiffermuller) (= E. pflugiana
(Haworth))
Plant recs.: Carduus acanthoides (France); C. nutans
(Europe); Cirsium (France as E. pflugiana and
Europe as E. 1'pflugiana); Carlina, Centaurea
(Europe); other Compositae (in lab. tests)
68
LEPIDOPTERAContinued
OlethreutidaeContinued
Remarks:
Larvae rare; in stems of Carduus nutans and C.
acanthoides (Europe)
Results of lab. tests (Delemont): Larvae
developed successfully to live pupae only on
Cirsium; fed to various degrees with some lar
val development (no live pupae) on Centaurea
(fed heavily), Aster, Echinops, Cynara, Onopordum, Silybum, Cnicus, Zinnia, and Lactuca (no
Carduus incl. in these tests)
This sp. was first discovered (first known as
E. pflugiana) and tested for biological control of
Cirsium at Delemont in 1961; small shipment of
pupae (as E. scutulana) sent to Canada in 1962;
recorded tachinid parasite: Actia lamia
(Meigen)
Sources:
Delemont Rpts. (1961-63); Zwblfer (1965a)
Eucosma sp. (? = cana Haworth)
Plant recs.: Carduus personatus (Austria); Cirsium, Centaurea
(Europe)
Larvae endophytic in flower heads of these
Remarks:
hosts
Sources:
IGrapholitha sp.
Plant recs.:
Remarks:
Sources:
Herpystes sp. ( =
Plant recs.:
Remarks:
Sources:
Carduus acanthoides (France)

Larvae in stems
"Acroclita sp.")
Cuscuta (Bangladesh, Pakistan); Carduus, many
other plants (in lab. tests only)
Main enemy of Cuscuta in Pakistan, attacking
fruits and vines; Carduus edelbergii and other
plants received slight feeding by larvae in lab.
tests; larvae completed development on
several economic plants; not likely of use in
biological control of Cuscuta
A17-ENT-9 Rpts. (1961-65); A17-ENT-14 Rpts.
(1967-68); Rawalpindi Rpts. (1967-68); CIBC
Ann. Rpt. (1967)
Pterophoridae
Platyptilia carduidactyla (Riley)

Remarks:
69
Larvae in stems; attacks other Compositae, in

cluding Cynara
Sources:
Tortricidae
Unident. tortricid spp.
Plant recs.: Carduus acanthoides (France); C. nutans (S. Dak.)
Remarks:
Delemont Rpt. (1962); Morihara and Balsbaugh
Sources:
(1976)
Unident. cneph asiine sp.
Occasional visitor
Remarks:
Zwblfer (1965a)
Sources:
Archips sp.
Larvae exophytic on stems and leaves;
Remarks:
Sources:
Clepsis strigana (Hiibner)
Plant recs.: Carduus nutans (Austria)
Larvae on leaves
Remarks:
Sources:
Cnephasia spp.
Plant recs.: Carduus acanthoides (Austria); C defloratus
(France); C. nutans (Europe)
Larvae on C. acanthoides and C. defloratus; occa
Remarks:
sional visitor on C. nutans
Sources:
Cnephasiella incertana (Treitschke)
Listed as occasional visitor on C. nutans
Remarks:
Sources:
Zwblfer (1965a)
Cochylidae(= Phaloniidae)
Unident. cochylid sp.
Remarks:
Larvae endophytic in flower heads and buds
Sources:
Zwblfer (1965a)
Cochylis dubitana (Hiibner)
Plant recs.: Carduus nutans, C. acanthoides, Arctium
(Europe); Circium, Centaurea
Remarks:
Sources:
Hannemann (1964); Zwblfer (1965a)
Cochylis sp. nr. C. posterana (Zeller) or C. hybridella (Hiibner)
70
LEPIDOPTERAContmued
CochylidaeContinued
Remarks:
Larvae bore into flowers for feeding; no other
hosts known; of possible interest as a
biological control agent; efforts to study
biology in lab. (Rawalpindi) unsuccessful;
more study warranted
Sources:
A17-ENT-9 Rpts. (1962-63, 1965); A17-ENT-14
Rpt. (1966); Rawalpindi Rpts. (1966, 1968);
Baloch et al. (1971); Baloch and Khan (1973)
Cochylis posterana (Zeller) (= Phalonia posterana)
Plant recs.: Carduus nutans (Rhine Valley, Europe); C. acanthoides (France, Europe); C. personatus
(Europe); C. crispus (Europe); C. pycnocephalus
(S. Europe, France); Cirsium (Austria, France,
Switzerland, Europe); Centaurea (Austria,
Europe); Arctium, Onopordum, Serratula, Carlina
(Europe)
Remarks:
Larvae reared from flowers; endophytic
Sources:
CIBC Ann. Rpt. (1969); Goeden (1974)
Euxanthoides straminea (Denis and Schiffermiiller)
Plant recs.: Centaurea (S. Europe); Carduus nutans, Carthamus (lab. tests)
Remarks:
Oviposited on Carduus nutans and other Compositae in lab. tests
Sources:
Delemont Rpt. (1970); CIBC Ann. Rpt. (1970)
Cosmopterigidae
Pyroderces argyrogrammos Zeller
Plant recs.: Carduus nutans (Upper Rhine Valley, Austria);
C. pycnocephalus (S. Europe); C. crispus (Upper
Rhine Valley); Centaurea (?France, Swit
zerland, Romania, Egypt); Silybum, Onopor
dum, Carlina (Europe); Silybum (Egypt)
Remarks:
Sources:
Delemont Rpts. (1962-63, 1965-66); CIBC Ann.
Rpt. (1965); Zwblfer (1965a); F4-ENT-5 and F4ENT-16 Rpts. (1966-68, 1976); Goeden (1974)
Gelechiidae
Scrobipalpa acuminatella Sircom (= Lita acuminatella)
Plant recs.: Carduus nutans, Cirsium, Onopordum, Serratula,
Centaurea (literature recs., Europe)
Remarks:
Larvae mining leaves
Sources:
Schutze (1931); Zwblfer (1965a)
71
Oecophoridae
Agonopteryx arenella (Denis and Schiffermiiller)
Plant recs.: Carduus nutans, C. personatus, Cirsium (Europe)
Remarks:
Larvae reared externally on stems and leaves;
reported mining shoots of Cirsium in
Switzerland
Sources:
Agonopteryx propinquella (Treitschke)
Plant recs.: Carduus acanthoides (Switzerland); Cirsium,
Onopordum, Serratula (Europe)
Remarks:
Larvae mine leaves; ident. of larvae from C.
acanthoides not verified
Sources:
Agonopteryx subpropinquella Stainton
Plant recs.: Carduus nutans, C. pycnocephalus, Cirsium, Centaurea, Cynara, Onopordum (Europe)
Remarks:
Locally common; larvae mining leaves
Sources:
Schutze (1931); Delemont Rpt. (1964); Zwblfer
(1965a); Goeden (1974)
Endrosis sarcitrella (L.) (= E. lactaeella Schiffermiiller)
Plant recs.: Carduus acanthoides (Europe); Cirsium (Rhine
Valley, France); Centaurea (Europe)
Remarks:
Larvae feed in flower heads, destroy seeds;
listed as occasional visitor on C. acanthoides;
recorded in literature on mushrooms, fruits,
flour, etc.; of no interest in biological control
Sources:
Glyphipterygidae
Choreutis bjerkandrella Thunberg
Plant recs.: Carduus crispus, Carlina (Europe)
Remarks:
Larvae mining leaves of C. crispus; found exter
nally on leaves of Carlina
Sources:
Schutze (1931); Zwblfer (1965a)
Coleophoridae
IColeophora sp.
Plant recs.: Carduus defloratus (Switzerland)
Remarks:
Case (only) found on plant
Sources:
Coleophora laeripennis Wocke
Plant recs.: Carduus defloratus (Europe)
Remarks:
Larvae mine leaves; European literature recs.
for C. aeripennis = Arctium, Serratula, and
Centaurea
Sources:
Zwblfer (1965a)
72
LEPIDOPTERAContinued
ColeophoridaeContinued
Coleophora therinella Tengstrom
Plant recs.: Carduus nutans, Cirsium, Carlina (Europe)
Remarks:
Larvae mining leaves
Sources:
Zwblfer (1965a)
DIPTERA
Cecidomyiidae
Unident. cecidomyiid spp.
Plant recs.: Carduus nutans, C. defloratus, C. acanthoides, C.
crispus (Europe); C. nutans (S. Dak.); C. pycnocephalus (Egypt)
Remarks:
Larvae endophytic in buds, galls, and flower
heads
Sources:
Ross and Hedicke (1927); Zwblfer (1965a); F4ENT-5 Rpt. (1970); Morihara and Balsbaugh
(1976)
Jaapiella cirsiicola Rubsaamen
Plant recs.: Carduus crispus, Cirsium (Europe)
Remarks:
Larvae endophytic in (and deforming) flower
heads
Sources:
Buhr (1964); Zwblfer (1965)
Syrphidae
Cheilosia sp.
Plant recs.: Carduus crispus (Switzerland); C. acanthoides, C.
personatus, C. sp. (Austria); C. nutans (Italy,
France, Europe); C. tenuiflorus, C. pycnocephalus, Cirsium (Italy)
Remarks:
Generic identity of some of these larval recs.
originally questioned; larvae (usually solitary)
mine stems, roots, and sometimes buds and
crowns, damaging 10 percent of rosettes; rec.
from Carduus sp. in Austria was as leaf miner;
only specimen reared (from many larvae) in
Italy was identified as Cheilosia chrysocoma
(from C. nutans)
Sources:
Delemont Rpts. (1962-63); Rome Rpts. (1965,
1974-75); Zwblfer (1965a); K. E. Frick in litt.
(1966)
Cheilosia chrysocoma (Meigen)
Plant recs.: Carduus nutans (Italy); C. crispus (Europe)
Remarks:
Larvae (usually solitary) in stems and crowns;
only specimen reared to adult (from many syr
73
phid larvae found on Carduus spp.) in Italy was

identified as C. chrysocoma (from C. nutans); C.
chrysocoma chosen for study for possible use in
biological control of Carduus (Rome)
Sources:
Brauer (1883); K. E. Frick in litt. (1966); Rome
Rpt. (1975)
Cheilosia cynocephala Loew
Remarks:
None
Sources:
Brauer (1883); K. E. Frick in litt. (1966); Rome
Rpt. (1975)
Cheilosia flavicornis (.)
Plant recs.: Carduus crispus, ICnicus (Europe)
Remarks:
None
Sources:
Brauer (1883); K. E. Frick in litt. (1966)
Cheilosia grossa Fallen
Plant recs.: Carduus pycnocephalus, C. nutans, C. crispus,
Cnicus (Europe)
Remarks:
Adults feed on pollen in early spring
Sources:
Mellini (1951a); Goeden (1974); Rome Rpt. (1975);
F. C. Thompson in litt. (1976)
Cheilosia mutabilis (Fallen)
Remarks:
None
Sources:
F. C. Thompson in litt. (1976)
Cheilosia variabilis (Panzer)
Plant recs.: Carduus nutans (Italy); Cirsium, Scrophularia
(Europe)
Remarks:
Larvae in stems
Sources:
Rome Rpt. (1975)
Platystomatidae
Platystoma sp.
Remarks:
Occasional visitor
Sources:
Zwolfer (1965a)
Platystoma seminationis F.
Remarks:
has been reared from mushrooms
Sources:
Zwblfer (1965a)
Tanyderidae
Unident. tanyderid sp.
Remarks:
Rare
Sources:
74
DIPTERAContinued
Tephritidae ( = Trypetidae)
Unident. tephritid spp.
Plant recs.: Carduus defloratus, C. acanthoides (Switzerland);
C. nutans (Austria); C. tenuiflorus (France); C.
pycnocephalus (S. Calif.)
Remarks:
Larvae found in flower heads in all these recs.;
addit. rec. of unident. adult on C. defloratus; lar
vae on C. acanthoides in 1 rec. said to be
gregarious
Sources:
Delemont Rpts. (1962-63); Goeden (1974)
Acanthiophilus eluta (Meigen) (= A. helianthi (Rossi))
Plant recs.: Carduus pycnocephalus (France, Italy); C.
edelbergii (Pakistan); C personatus (Austria);
Cirsium (France, Europe); Centaurea (France,
Pakistan, Italy, Romania, Bulgaria, USSR
(Caucasus), Egypt, Syria, Lebanon); Carthamus
(India, Pakistan); Carlina (Austria); Cynara,
Onopordum (Italy); Serratula, Xeranthemum,
Leuzea (Europe); Cnicus, Echinops (Pakistan);
Silybum (Egypt, Pakistan, Lebanon)
Remarks:
Larvae reared from flower heads, tend to be
gregarious; viable seed production reduced;
polyphagous within Cynareae; major pest of
Carthamus in India and Pakistan; most impor
tant enemy of Centaurea in Europe and Egypt;
some oviposition tests conducted on Centaurea
(at Delemont)
Sources:
Rome Rpts. (1959, 1963); L. A. Andres in litt.
(1961); A17-ENT-9 Rpts. (1961-62, 1964-65);
CIBC Ann. Rpts. (1961, 1965-66); Delemont
Rpts. (1963, 1965-68); Zwblfer (1965a); F4-ENT5 and F4-ENT-16 Rpts. (1966-68, 1976); Baloch
et al. (1971); Zwolfer (1974)
Chaetostomella cylindrica (Robineau-Desvoidy)(= C. onotrophes
Loew)
Plant recs.: Carduus nutans, C. crispus (Europe); Cirsium,
Serratula (Austria, Europe); Centaurea,
Microlonchus (Europe)
Remarks:
Larvae reared from flower heads
Sources:
Euaresta bella (Loew)

Remarks:
75
Endophagous in flowers; normally in Ambrosia

seeds
Sources:
Orellia winthemi Meigen
Plant recs.: Carduus crispus (Britain, Rhine Valley); C.
crassifolius (Austria); ?C. acanthoides (Europe)
Remarks:
Larvae in flower heads; identity of larvae from
Carduus defloratus and C. acanthoides not
verified
Sources:
Delemont Rpt. (1963); Zwblfer (1965a, 1974)
Paracantha culta (Wiedemann)
Remarks:
Endophagous; polyphagous
Sources:
Tephritis sp.
Carduus edelbergii, Cirsium (Pakistan)
Plant recs
Six generations per year in unopened flower
Remarks:
heads; did not attack other Compositae tested
in lab.
Sources:
CIBC Ann. Rpt. (1969)
Tephritis cardualis Hardy
Plant recs.: Carduus edelbergii, Cirsium spp. (Pakistan)
Remarks:
Endophytic in flowers
Sources:
Rawalpindi Rpt. (1968); Baloch et al. (1971)
Tephritis heiseri Frauenfeld
Plant recs.: Carduus nutans (Europe); C. edelbergii
(Pakistan); C. acanthoides (Europe, ?Austria); C.
defloratus (Europe); Cirsium, Cnicus (Pakistan)
Remarks:
Larvae reared from flower heads; larvae of
Austrian rec. from C. acanthoides not reared;
biological studies conducted at Rawalpindi:
Lab. oviposition obtained on C. nutans but not
on 7 other spp. of Compositae; recorded
parasites (Pakistan): Paraholaspis sp. and
Tetrastichus sp.
Sources:
Seguy (1934); A17-ENT-Rpts. (1962-63, 1965);
Delemont Rpt. (1963); CIBC Ann. Rpts.
(1965-68); Zwblfer (1965a); A17-ENT-14 Rpts.
(1966-68); Rawalpindi Rpts. (1966-68); Baloch
et al. (1971)
Tephritis hyoscyami (L.)
Plant recs.: Carduus personatus (Austria, France); C. uncinatus (USSR); C. nutans (Europe); C. crispus

(Sweden, Britain); ?C. acanthoides (Austria);
Carduus sp. (Sweden)
Remarks:
Larvae reared from flower heads; larvae from
C. acanthoides not reared, may be Tephritis
heiseri; formed galls in Carduus sp.
Sources:
Seguy (1934); Delemont Rpt. (1963); Zwblfer
(1965a, 1974); A17-ENT-9 Rpt. (1965)
Terellia serratulae (L.)
Plant recs.: Carduus nutans (Europe, Britain); C. defloratus
(Switzerland, Europe); C. crassifolius (Austria);
C. acanthoides (Europe); C. tenuiflorus (France);
C. edelbergii (Pakistan); Cirsium (Europe,
Pakistan); Onopordum, Galactites, Centaurea
(Europe); Cnicus (Pakistan)
Remarks:
Larvae attack seeds and tissue of flower heads
of Carduus; found to reduce both seed produc
tion and seed viability of C. edelbergii (in
Pakistan)
Field and lab. studies conducted at
Rawalpindi: Regular oviposition and suc
cessful larval development obtained only on
Carduus edelbergii and Cnicus; in early test,
some oviposition obtained on following plants
(in isolation): Cynara, Conyza, Centaurea,
Cousinia, Acroclinum, Gerbera, Gaillardia, and in
later test on Calendula; no larval feeding seen
on these plants; recorded parasites (Pakistan):
Paraholaspis cothurnata Masi and Tetrastichus
sp.
Sources:
Seguy (1934); A17-ENT-9 Rpts. (1961-65); CIBC
Ann. Rpts. (1961, 1963, 1965-68); Delemont
Rpts. (1961-64, 1966); Zwblfer (1965a); A17ENT-14 Rpts. (1966-67); Rawalpindi Rpts.
(1966-68); Baloch et al. (1971); Baloch and Khan
(1973); Zwblfer (1974)
Urophora sp. nr. U. approximata (Hering)
Plant recs.: Carduus defloratus (Switzerland)
Remarks:
Larvae endophytic in flower heads and/or
buds; true U. approximata rec. (Europe) only
from Cirsium
Sources:
Hering (1938); Zwblfer (1965a, 1974)
Urophora cardui (L.)
Plant recs.: Cirsium arvense only (Europe); Carduus nutans
(in lab. tests)
77
Remarks:
Larvae endophytic in stems

In lab. tests: Will oviposit on Cirsium vulgare
and Carduus acanthoides if Cirsium arvense is ab
sent; some larval development in Carduus
nutans
Galls sent to Canada from Europe for C.
arvense control: 1,400 (1969), 750 (1973), and
1,080 (1974); U.S. Working Group on Biological
Control of Weeds approved release in Canada
(1974) and in U.S. (1977); releases (against C.
arvense): 21A in B.C. and 574 in Sask. (3 sites) in
1974; 96 in Que., 81 in Ont., 98 in B.C., and 374
in Sask. (3 sites) in 1975; 951 in Sask. (5 sites),
472 in Alta. (2 sites), 199 in B.C., 52 in Ont., 451
in Que. (4 sites), and 191 in N.B. (2 sites) in
1976
Sources:
Delemont Rpts. (1961-67, 1970); Zwblfer (1965a);
Peschken (1971); Peschken and Harris (1975);
Harris (1975)
Urophora 1eriolepidis (Loew)
Plant recs.: Carduus nutans (Rhine Valley); Cirsium, Centaurea (Europe)
Remarks:
Larvae doubtfully identified as U. eriolepidis
found in flower heads of C. nutans in associa
tion with larvae of Urophora solstitialis; larvae
definitely identified as U. (or Euribia) eriolepidis
reared only from Cirsium (Germany, Europe)
Sources:
Ross (1927); Delemont Rpts. (1963-64); Zwblfer
(1965a)
Urophora ? solstitialis (L.)
Plant recs.: Carduus nutans (France, Italy, Rhine Valley); C.
acanthoides (France); C. pycnocephalus (S.
Europe); C. crispus (Rhine Valley); C. defloratus
(Switzerland); C. edelbergii (Pakistan); C. personatus (Europe); ^Arctium, ICarthamus,
Carlina, Centaurea (Europe)
Remarks:
Ident. often questioned; larvae from galls in
flower receptacles; feed on seeds
Sources:
Ross (1927); Seguy (1934); Mellini (1952); H. L.
Parker in litt. (1957); Delemont Rpts. (1963-64);
Zwblfer (1965a); L. A. Andres in litt. (1966);
Rawalpindi Rpt. (1968); CIBC Ann. Rpt. (1971);
Baloch et al. (1971); Goeden (1974)
78
DIPTERA Continued
TephritidaeContinued
Urophora stylata (F.)
Plant recs.: Carduus acanthoides, C. personata, C. nutans
(Europe); C. edelbergii, Cnicus (Pakistan);
Cirsium, Senecio (Europe)
Remarks:
European specimens mostly from Cirsium;
Pakistan specimens from Carduus edelbergii,
Cnicus, and Cirsium
Results of lab. oviposition tests (Delemont,
1966-68): Regular oviposition obtained only on
Cirsium, occasionally on Carduus acanthoides,
Arctium, Onopordum, and on Centaurea when ar
tificially stimulated (Rawalpindi, 1966-68).
Oviposition obtained only on Carduus edelbergii
and Cnicus
Shipments of 750 galls containing U. stylata
larvae made in 1964-67 from Del6mont to
Belleville, Ont., for testing as control agent of
Cirsium vulgare; U.S. Working Group on
Biological Control of Weeds approved release
in Canada in 1973
Releases (against Cirsium vulgare): 1,227
adults released in B.C. (4 Iocs.) in 1973,
established, and populations increasing by
1975; 123 larvae released in Que. in 1976;
Pakistan population possible biological con
trol agent for Carduus nutans and/or Cnicus
wallichii
Sources:
Seguy (1934); Delemont Rpts. (1961-68); A17ENT-9 Rpts. (1963-65); CIBC Ann. Rpts.
(1964-67, 1973-74); Zwblfer (1965a, 1974);
Baloch et al. (1971); Williamson (1974); Harris
(1975)
Xyphosia miliaria (Schrank)
Remarks:
Sources:
SSguy (1934); Delemont Rpts. (1961-65);
Zwblfer (1965a, 1974)
Drosophilidae
Scaptomyza pallida (Zetterstedt) (= Parascaptomyza pallida)
Remarks:
Larvae mine leaves; no other host known
79
Sources:
A17-ENT-9 Rpts. (1964-65); Baloch et al. (1971)
Chloropidae
Oscinella spp.
Remarks:
Common; endophagous in stems
Sources:
Polyodaspis sp.
Remarks:
Pupae found in flowers; status unknown
Sources:
Agromyzidae
Unident. agromyzid spp.
Plant recs.: Carduus crispus (Rhine Valley); C. pycnocephalus
(Egypt)
Remarks:
Adults
Sources:
Delemont Rpt. (1962); F4-ENT-5 Rpt. (1969)
Agromyza apfelbecki Strobl
Plant recs.: Carduus acanthoides (Hungary); Cynara (Europe)
Remarks:
Leaf miner; pest of Cynara
Sources:
Spencer (1973)
Cerodontha dorsalis (Loew)
Remarks:
In leaves; polyphagous
Sources:
Liriomyza sp. nr. L. eupatorii Kaltenbach
Remarks:
Rare; larvae endophytic in leaves
Sources:
Goeden (1974)
Melanagromyza sp. (poss. M. aeneoventris (Fall6n)
Plant recs.: Carduus crispus (Rhine Valley); C. acanthoides
(Europe); Cirsium (Europe)
Remarks:
Larvae found in stems of C. crispus
Sources:
(1965a)
Melanagromyza aeneoventris (Fallen) (= M. cirsii (Rondani))
Plant recs.: Carduus nutans, C. acanthoides (Europe); C.
edelbergii (Pakistan); Carduus sp. (Argentina);
Cirsium, Centaurea, Senecio, other Compositae
(Europe)
Remarks:
Larvae mine within the pith of stems; pupae
parasitized by Tetrastichus sp.
Sources:
Seguy (1934); Delemont Rpt. (1962); A17-ENT19 (1962, 1965); A7-ENT-9 Rpts. (1964-65);
Rome Rpt. (1965); Baloch et al. (1971)
80
DIPTERAContinued
Agromyzidae Continued
Napomyza lateralis (Fallen)
Plant recs.: Carduus nutans, C. pycnocephalus (Italy);
Centaurea, Calendula (Europe)
Remarks:
Common feeder on Compositae
Sources:
Spencer (1973)
Phytomyza sp.
Remarks:
Occasional larvae; endophagous in leaves
Sources:
Goeden (1974)
Phytomyza affinis Fall6n
Plant recs.: Carduus acanthoides (Finland); Cirsium, Onopordum, Serratula (Europe)
Remarks:
Leaves mined
Sources:
(1965a)
Phytomyza cardui Hering
Plant recs.: Carduus crispus, Cirsium (France)
Larvae in leaf galls
Remarks:
Sources:
Buhr (1964); Zwblfer (1965a)
Phytomyza cirsii Hendel
Plant recs.: Carduus (Finland, Britain); Cirsium, Serratula
(Europe)
Remarks:
Leaves mined
Sources:
Seguy (1934); Parmenter (1952)
Phytomyza farfarella Hendel ( = P. atricornis Meigen)
Plant recs.: Carduus personatus (France); C. crispus
(Czechoslovakia); C. pycnocephalus (U.S.,
Canada); Cirsium, Onopordum, Silybum, Xeranthemum (Europe); Xanthium, Carthamus, Salvia,
Centaurea, Cannabis, Conyza, Datura, Silybum,
Trichodesma (Pakistan); tCentaurea (Egypt);
many other hosts
Remarks:
Larvae mine leaves; occasional visitor on these
hosts in Europe
Sources:
Frick (1959); A17-ENT-9 Rpts. (1961, 1965);
Delemont Rpts. (1962-63); Zwblfer (1965a); F4ENT-5 Rpts. (1966-68); Baloch et al. (1968)
Phytomyza syngenesiae (Hardy) (= P. "atricornis Meigen" in
part)
Plant recs.: Carduus crispus (Sweden); several other Com
positae (rarely other plants) (Europe, N. Amer.,
Australia, New Zealand)

Remarks:
81
Leaf miner; pest of lettuce and chrysan

themum
Spencer (1973)
Sources:
Anthomyiidae
Unident. dipterous larvae, prob. Pegomya terebrans (Rondani)
(= P. nigricornis (Strobl))
Plant recs.: Carduus pycnocephalus, Cynara (Italy)
Remarks:
Larvae mine leaves; larvae from Cynara ident.
as P. terebrans
Sources:
Rome Rpt. (1965); Goeden (1974)
Hylemya spp.
Remarks:
Endophagous
Sources:
Pegomya carduorum Huckett (= P. bicolor (Wiedemann) )
Plant recs.: Carduus pycnocephalus, Cirsium (Calif.)
Remarks:
Common; larvae mine leaves; reared
Sources:
Albany Rpts. (1968-69); Goeden (1974); G.
Steyskal in litt. (1977)
Pegomya fodiens (Hendel) (= Carduiphila fodiens)
Plant recs.: Carduus crassifolius (Europe)
Remarks:
Mine leaves
Sources:
G. Steyskal in litt. (1977)
Pegomya steini Hendel
Plant recs.: Carduus crispus (Britain)
Remarks:
None
Sources:
Niblett (1957)
FUNGI ASSOCIATED WITH CARDUUS THISTLES

The use of plant pathogens for weed control has been reviewed by
Wilson (1969) and Hasan (1974). It would be desirable to introduce
highly specific fungi to control Carduus thistles with no danger to
other plants. Smuts and some rust strains are notably species,
subspecies, variety, or even race specific. The successful establish
ment of an Italian strain of Puccinia chondrillinaBubak & Sydow to
control the Eurasian skeletonweed (Chondrilla juncea L.) in
Australia was discussed by Hasan (1971).
Carduus thistles are attacked by a variety of relatively
nonspecific and specific fungi, according to information extracted
from herbarium specimens and the host or fungus index in the Na
tional Fungus Collections, Plant Protection Institute, Beltsville
82
Agricultural Research Center,

nonspecific fungi are as follows:
Beltsville,
Md.
Recorded
Ascochyta phomoides Saccardo

Bremia lactucae Regel
B. tulasnei (Hoffman) Sydow
Cercosporella carduicola P. Brunaud
Cladosporium herbarum Link ex Fries
Erysiphe cichoracearum DC. ex Merat
Fusicladium aronici Saccardo
F. schnablianum Allescher
Heteropatella umbilicata (Persoon) Saccardo
Leptosphaeria carduina Passerini
L. clivensis (Berkeley & Broome) Saccardo
L. purpurea Rehm
Leveillula compositarum Golovin forma cardui (Jaczewski) Golovin
L. taurica (Leveille) Arnaud
Macrosporium commune Rabenhorst
Oidium spp.
0. conspicua Fautrey & Lambette
Ophiobolus acuminatus (Sowerby) Duby
O. pellitus (Fuckel) Saccardo
Ovularia vossiana Thiimen
Passalora schnabliana (Allescher) Petrak
Phoma albicans Roberge & Desmazieres
P. pycnocephali Passerini
Phragmidium personatae Allescher
Phyllosticta personatae Allescher
Pleospora albicans Roberge & Desmazieres
P. penicillus (Schumacher) Fuckel
Pseudohelotium micaceum (Persoon) Saccardo
Ramularia cardui Karsten
Rhabdospora cirsii Karsten
Sclerotinia sclerotiorum (Libert) DeBary
Scolecotrichum cardui Schroeter
Septoria associata Bubak & Kabat
S. cardui Fossi
Venturia pellita Hazslinszky
In addition, Carduus glaucus on limestone is associated with a
mycorrhizal fungus in Poland, according to records in the files of
the National Fungus Collections. The obligate parasitic fungus
Albugo trogapogi (Persoon) Schroeter is associated with and may be
host specific on Carduus crispus in Denmark and the Netherlands.
Carduus micropterus broteri was attacked by the lettuce mosaic
virus in Portugal, and a phloem necrosis virus infected C. nutans at
Beltsville. Md.
83
The rusts and smuts in the following list are specific to Carduus
thistles or contain specific races or biotypes attacking various
species or varieties of Carduus. Because many rusts are highly
specific to certain varieties or races of the host, these fungi seem
most promising as potential biological control agents, particularly
since they have not yet been reported from North America.
LIST OF SPECIFIC FUNGI

UREDINALES
Puccinia cardui-pycnocephali H. Sydow & P. Sydow
Plantrecs.:
Carduus pycnocephalus (Italy, Spain,
Yugoslavia, Portugal, Britain, Turkey, Asia
Minor, Iran, Cyprus, New Zealand, Azerbaidzhan SSR, Turkmen SSR, North Africa,
Scotland, Palestine); C. tenuiflorus (Spain, Por
tugal, New Zealand, France, Morocco); C. ammophilus (as C. reuterianus) (Spain); C. seminudus
(Caucasus USSR); C. spachianus var. duriaei
Boiss. & Reut. (Morocco); C. clavulatus (Canary
Is.); C. ballii, C. myriacanthus (Morocco)
Remarks:
States II (uredia) and III (telia) on living leaves
and stems; plants persisted but disease disap
peared after several years (Switzerland); oc
curs in June (Portugal)
Sources:
Sydow and Sydow (1904); Natl. Fungus Coll.
Puccinia carduorum Jacky (= P. hieracii (Roehling) Martius; =
Aecidium cardui Sowerby; = Uredo flosculosorum Albertini & Schweinitz; = P. flosculosorum Winter; = P. cardui-carlinoides Petrak;
= P. compositarum Schlechtendahl (in part) )
Plant recs.:
Carduus nutans (France, USSR, Spain, Britain,
Germany, Hungary, Lithuanian SSR,
Pakistan, Kirghiz SSR, Switzerland, India,
Romania); C. crispus (France, Switzerland,
Romania, USSR, Spain, Siberia, Japan, Korea,
Lithuanian SSR (as Uredo flosculosorum),
Uruguay, Hungary, Finland (Lapland), Nor
way, Sweden, Germany, Altai Mts. - USSR,
Czechoslovakia, Scotland, Europe (as Puccinia
hieracii, Uredo hieracii) ); C. cornius (Greece (as C.
armatus), Bulgaria); C. kerneri (Europe); C.
defloratus (Switzerland, France, Austria (as
Aecidium cardui), Europe (as Puccinia
84
flosculosorum) ); C. getulus (N. Africa); C.

crassifolius (as C. glaucus) (Czechoslovakia); C.
hamulosus (Hungary); C. leptocladus, C.
myriacanthus, C. macrocephalus, C. meonanthus
(Morocco); C. candicans (Europe); C. personatus
(France, Hungary, Germany, Bulgaria,
Switzerland); C. ammophilus (as C. reuterianus)
(Spain, Balearic Is.); C. tenuiflorus (Portugal,
Spain, France); C. onopordioides (Bulgaria); C.
6a//w (Morocco); C. acanthoides (Europe (as
Puccinia flosculosorum), France, Germany,
Czechoslovakia, Ukrainian SSR); C. pycnocephalus (Turkmen SSR (as P. hieracii),
Europe, Cyprus, Spain, Iran, Turkey,
Turkmen SSR, Uzbek SSR); C. gayanus (Spain
(as C. carpetanus) ); C. thoermeri (Africa (as C.
leiophyllus) ); C. chrysacanthus (France (as
Puccinia carduicarlinoides) ); C. collinus
(Romania, Hungary); C. kerneri (as C. scardicus)
(Bulgaria); C. (Alfredia) acantholepis and C. (A.)
m'fea (Kirghiz SSR); C. (A.) cernua (Kirghiz
SSR, Siberia); C. chevallieri (N. Africa); (?)
Silybum marianum (L.) Gaertn. (Portugal)
Remarks:
States II and III on leaves and branches in
July and Aug. (Europe, C. nutans); II and III
common on stems and leaves in July-Oct.
(Europe, C. crispus, C. personatus); June-Oct.
(Switzerland, C. defloratus)
Sources:
Sydow and Sydow (1904); Savulescu (1955);
Natl. Fungus Coll.
Puccinia centaureae DC. (= P. centaureae Fuckel; = P. calcitrapae
DC; = P. scabiosae P. Magnus; = P. jaceae Otth; = P. inquinans
Wallroth; = Epitea jaceae Otth; = Uredo flosculosorum var.
centaureae Wettstein)
Plant recs.:
Carduus crispus (Norway, Finland); C. clavulatus
(Canary Is.); C. pycnocephalus (Iraq, Canary Is.,
as Puccinia centaureae forma asperae Cast, on C.
pycnocephalus as C. tenuifolius); also literature
records from Centaurea spp.
Remarks:
States II and III on C. clavulatus (Canary Is.)
Sources:
Puccinia cnici-oleracei Persoon ex Desmazieres (= P. cardui
Plowright; = P. spectabilis Otth)
85
Plant recs.:
Carduus (Afrocarduus) kikuyorum (Kenya); C.

{Afrocarduus) sp. (Uganda); ? C. crispus (Ger
many); Cirsium spp. (Europe); Carthamus
(France); Lupsia (Canary Is.); Artemisia (China)
Remarks:
In living leaves; probably several spp.
Sources:
Puccinia galatica Sydow
Plant recs.:
Carduus pycnocephalus var. albidus (Asia
Minor); C. arabicus marmoratus (Europe); C.
nutans (Sinkiang, China)
Remarks:
On living leaves
Sources:
USTILAGINALES
Ustilago cardui F. de Waldheim (= U. reessiana Kuhn)
Plant recs.:
Carduus acanthoides (as U. reessiana), C. acanthoides, C. nutans (N. Europe); C. defloratus
(Switzerland); C. personata (USSR, Europe); C.
crispus (USSR - Altai Mts.); C. thoermeri
(Europe)
Remarks:
In flower heads; destroys capitulum
Sources:
Savulescu (1955); Natl. Fungus Coll.
Ustilago violacea S. F. Gray (= U. violacea Persoon)
Plant recs.:
Carduus acanthoides (Germany)
Remarks:
None
Sources:
Natl. Fungus Coll.
SUMMARY
Six Eurasian species of Carduus thistles have become established
in North America, where they are troublesome weeds in pastures
and rights-of-way. A search for biological control agents in Eurasia
and North America has shown that Carduus spp. are attacked by
about 340 species of phytophagous insects, including 71 that are
oligophagous on Cynareae. The host range of 39 Eurasian species
was determined by laboratory testing (30 Coleoptera, 5
Lepidoptera, and 4 Diptera). Five of them were sufficiently damag
ing and host specific to warrant their release in North America.
The results are as follows:
(1) Altica carduorum Guerin-M6neville, repeatedly released
against Canada thistle (Cirsium arvense (L.) Scop.) but not yet
established in North America.
86
(2) Ceutorhynchus litura (F.), first released against Cirsium arvense

and established in Sask., Ont., Idaho, Mont., and Calif., with some
control of the host.
(3) Rhinocyllus conicus (Froelich), first released in 1968, estab
lished on Carduus in Calif., Va., W. Va., Colo., Mo., Mont., Pa., Md.,
Okla., Ont., Sask., Man., Que., and on Silybum marianum (L.) Gaertn.
in Calif., with significant localized control of Carduus in Va., Mo.,
and Mont, and of Silybum in Calif.
(4) Trichosirocalus horridus (Panzer), first released against Carduus
in 1975 in Canada and Va., establishment in Va. (1978); releases
planned in Kans., Mont., Nebr., N. Dak., S. Dak., and Mo. for
1978-79.
(5) Urophora stylata (F.), first released against Cirsium vulgare
(Savi) Ten. in 1973 and established in B.C.
In addition, Cassida rubiginosa Mueller, which had been
laboratory tested, was found to have been adventively established
on Carduus and Cirsium in northeastern North America since 1927.
Carduus thistles are also attacked by 42 species of fungi; of these,
7 species of Puccinia and Ustilago may be host specific and are
worthy of further investigation.
LITERATURE CITED
Alimdzhanov, R.
1941.
THE BIOLOGY OF THE NODULE WEEVILS UNDER CONDITIONS OF IR
RIGATED lucerne. Akad. Nauk Uzbek. SSR Izv. 4: 64-70.
Anderson, D. M.
1956.
NOTES ON CLEONUS PIGER (SCOP.) IN THE UNITED STATES (COLEOPTERA,
curculionidaei. Coleopterists' Bui. 10 (6): 81-85.
Andres, L. A.
1971.
the suppression of weeds with insects. Tall Timbers Conf. Ecol.
Anim. Control by Habitat Mangt. Proc. 1971: 185-195.
and Bennett, F. D.
1975.
and Davis,
biological
C. J. control of aquatic weeds. Ann. Rev. Ent. 20: 31-46.
1973.
THE BIOLOGICAL CONTROL OF WEEDS WITH INSECTS IN THE UNITED

states. Commonwealth Inst. Biol. Control Misc. Pub. 6: 1 1-28.
Baker, C. R. B., Blackman, R. L., and Claridge, M. F.
1972.
STUDIES ON HALTICA CARDUORUM GUERIN (COLEOPTERA:
CHRYSOMELIDAE) AN ALIEN BEETLE RELEASED IN BRITAIN AS A CON
TRIBUTION TO THE BIOLOGICAL CONTROL OF CREEPING THISTLE, CIR
SIUM ARVENSE (L.) SCOP. J. Appl. Ecol. 9 (3): 819-830.
Baker, H. G.
1965.
CHARACTERISTICS AND MODES OF ORIGIN OF WEEDS. In Baker, H. G.,
and Stebbins, G. L., eds., The Genetics of Colonizing Species, pp.
147-172. Academic Press, New York.
87
Baloch, G. M., and Khan, A. G.

1973.
BIOLOGICAL CONTROL OF CARDUUS SPP. II. PHENOLOGY, BIOLOGY AND
HOST-SPECIFICITY OF TERELLIA SERRATULAE L. (DIPT.:
trypetidae). Commonwealth Inst. Biol. Control Tech. Bui. 16:
11-22.
Khan, A. G., and Mushtaque, M.
1971.
BIOLOGICAL CONTROL OF CARDUUS SPP. I. INSECTS ASSOCIATED WITH
these in west Pakistan. Commonwealth Inst. Biol. Control Tech.
Mohyuddin,
Bui. 14:51-58.
A. I., and Ghani, M. A.
1968.
XANTHIUM STRUMARIUM L. INSECTS AND OTHER ORGANISMS

associated with it in west Pakistan. Commonwealth Inst. Biol.
Control Tech. Bui. 10: 103-112.
Bendall.G. M.
1973.
THE CONTROL OF SLENDER THISTLE, CARDUUS PYCNOCEPHALUS L. AND
C. TENUIFLORUS CURT. (COMPOSITAE), IN PASTURE BY GRAZING MANAGE
MENT. Austral. J. Agr. Res. 24: 831-837.
1974.
Brauer, F.
1883.
SLENDER THISTLES IN PASTURE; CONTROL BY GRAZING

management. Tasmanian J. Agr. 45 (1): 62-63.
DIE ZWEIFLUGLER DES KAISERLICHEN MUSEUMS ZU WIEN. III. K. Akad.
der Wiss. Wien, Math. -Nat. Denkschr. 47: 1-100.
Buhr.H.
1964.
BESTIMMUNGSTABELLEN DER GALLEN (ZOO- UND PHYTOCECIDIEN) AN

PFLANZEN MITTEL- UND NORDEUROPAS. Band I, 761 pp. Jena.
Claridge, M. F., Blackman, R. L., and Baker, C. R. B.
1970.
HALTICA CARDUORUM INTRODUCED INTO BRITAIN AS A POTENTIAL CON
TROL AGENT FOR CREEPING THISTLE CIRSIUM ARVENSE (L.)
scop. Entomologist 103: 210-212.
Crane, E.
1975.
HONEY: A comprehensive survey. 608 pp. Crane, Russak and
Co., Inc., New York.
Doing, H., Biddiscombe, E. F., and Knedlhans, S.
1969.
ECOLOGY AND DISTRIBUTION OF THE CARDUUS NUTANS GROUP (NODDING
thistles) in Australia. Vegetatio 17 (6):313-351.
Dunn, P. H.
1976.
DISTRIBUTION OF THE WEEDY THISTLES. CARDUUS NUTANS, C. ACAN
THOSES, C. PYCNOCEPHALUS AND C. CRISPUS IN THE UNITED
and Rizza,
states.
A. Weed Sci. 24:518-524.
1976.
Eguagie, W. E.
1974.
BIONOMICS OF PSYLLIODES CHALCOMERA, A CANDIDATE FOR

biological control OF musk thistle. Amer. Ent. Soc. Ann. 69 (3):
395-398.
BIONOMICS OF THE SPEAR THISTLE LACE BUG, TINGIS CARDUI

(HETEROPTERA: tingidae). J. Nat. Hist. 8: 621-629.
Feldman, I., McCarty, M. K., and Scifres, C. J.
1968.
ECOLOGICAL AND CONTROL STUDIES WITH MUSK THISTLE. Weed Sci.
16: 1-4.
88
Frick. K. E.
1959.
1969.
SYNOPSIS OF THE SPECIES OF AGROMYZID LEAF MINERS DESCRIBED

from north AMERICA. U.S. Natl. Mus. Proc. 108 (3407): 347-445.
CEUTHORHYNCHUS (HADROPLONTUS) TRIMACULATUS (P.) AND ceuthorRHYNCHIDIUS HORRIDUS (PANZ ), TWO WEEVILS OF POTENTIAL VALUE
FOR THE BIOLOGICAL CONTROL OF THISTLES IN THE GENUS CARDUUS
L. U.S. Dept. Agr., ARS, Ent. Res. Div., Albany, Calif., Biol. Con
trol of Weeds Invest. Mimeo. Rpt. 1969, 17 pp.
Fuller, M. J.
1969.
the genus carduus in Nebraska. Nebr. Univ. Studies N.S. 39:

1-57.
Furrer, J. D., and McCarty, M. K.
1966.
MUSK THISTLE - ITS APPEARANCE, SPREAD AND CONTROL. Nebr. Agr.
Expt. Sta. Bui. EC 66-160:1-6.
Ganiev, S. G.
1975.
CONTENT OF ECDYSONES IN SOME PLANTS OF THE TRIBE CYNAREAE OF
THE FAMILY ASTERACEAE (COMPOSITAE). Rastit. Resur. 11 (1):
94-96.
Ghani, M. A.
1963.
EUBLEMMA PARVA HB., AN ENEMY OF XANTHIUM SHIPPED TO
AUSTRALIA. Commonwealth Inst. Biol. Control Tech. Bui. 3:
27-29.
Goeden, R. D.
1974.
COMPARATIVE SURVEY OF THE PHYTOPHAGOUS INSECT FAUNAS OF
ITALIAN THISTLE, CARDUUS PYCNOCEPHALUS, IN SOUTHERN CALIFOR
NIA AND SOUTHERN EUROPE RELATIVE TO BIOLOGICAL WEED
control. Environ. Ent. 3 (3): 464-474.
1976.
THE PALEARCTIC INSECT FAUNA OF MILK THISTLE, SILYBUM MARIANUM,

AS A SOURCE OF BIOLOGICAL CONTROL AGENTS FOR
Andres,CALIFORNIA.
L. A., Freeman,
Environ.
T. E., Ent.
and others.
5 (2): 345-353.
1974.
PRESENT STATUS OF PROJECTS ON THE BIOLOGICAL CONTROL OF WEEDS

WITH INSECTS AND PLANT PATHOGENS IN THE UNITED STATES AND
and Louda,
Canada.
S. M. Weed Sci. 22: 490-495.
1976.
BIOTIC INTERFERENCE WITH INSECTS IMPORTED FOR WEED
and Ricker,
control.
D. W. Ann. Rev. Ent. 21:325-342.
1974.
IMPORTED SEED WEEVILS ATTACK ITALIAN AND MILK THISTLES IN

SOUTHERN CALIFORNIA. Calif. Agr. 28: 8-9.
and Ricker, D. W.
1977.
ESTABLISHMENT OF RHINOCYLLUS CONICUS ON MILK THISTLE IN
SOUTHERN CALIFORNIA. Weed Sci. 25 (3): 288-292.
Grandi.G.
1952.
introduzione allo studio dell entomologia aggiunte e correzione. Bologna Univ. Ent. 1st. Bol. 29: 263-306.
Hacker, J.
1925.
(note.) U.S. Dept. Agr. Coop. Econ. Insect Rpt. 25 (30): 625.
89
Hannemann, H. J.
1964.
KLEINSCHMETTERLINGE ODER MICROLEPIDOPTERA. II. DIE WICKLER S.
L. (COCYLIDAE UND C ARPOSENIDAE). DIE ZUNSLERARTIGEN
(PYRAloidea). In Dahl et al., Die Tierwelt Deutschlands, v. 50, 401
PP.
Harris, P.
1964.
HOST SPECIFICITY OF ALTICA CARDUORUM GUER. (COL.
chrysomelidae). Canad. J. Zool. 42: 857-862.
1971.
biological control of weeds. Environ. Let. 2 (2): 75-88.
1973a.
WEED VULNERABILITY TO DAMAGE BY BIOLOGICAL CONTROL

agents. Commonwealth Inst. Biol. Control Misc. Pub. No. 6, pp.
29-39.
1973b.
THE SELECTION OF EFFECTIVE AGENTS FOR THE BIOLOGICAL CONTROL

of weeds. Canad. Ent. 105: 1495-1503.
1975.
general approach to biocontrol of weeds in Canada. Quebec

Soc. Protect. Plants, St. Hyacinthe, Quebec, J. 1975, pp. 135-141.
and Zwolfer, H.
1968.
screening of phytophagous insects for biological control of
and Zwolfer,
weeds.H.Canad. Ent. 100 (3): 295-303.
1971.
Hasan, S.
1971.
1974.
CARDUUS ACANTHOIDES L., WELTED THISTLE, AND C. NUTANS L., NOD

DING thistle. In Biological Control Programmes Against Insects
and Weeds in Canada, 1959-1968. Commonwealth Inst. Biol.
Control Tech. Commun. 4, pp. 76-79.
host specialization in chondrilla fungi. Commonwealth Inst.
Biol. Control Tech. Commun. 4, pp. 135-139.
RECENT ADVANCES IN THE USE OF PLANT PATHOGENS AS BIOCONTROL

AGENTS OF WEEDS. Pest Articles News Sum. 20 (4): 437-443.
Hawkes, R. B., Andres, L. A., and Dunn, P. H.
1972.
seed weevil released to control milk thistle. Calif. Agr. 26
(12): 14.
Heikertinger, F.
1954.
chrysomelidae, blattkafer. In Blunch, H., ed., Handbuch der
Pflanzenkrankheiten, v. 5, pt. 2, ser. 2, pp. 270-379. Berlin u.
Hamburg.
Henderson, M., and Anderson, J. G.
1966.
common weeds in south Africa. Repub. So. Afr. Bot. Survey
Mem. 37, p. 330.
Hensley.M. S.
1973.
TAXONOMY AND DISTRIBUTION OF CIRSIUM THISTLES IN ROCKINGHAM
county, virginia. Va. J. Sci. 24 (3): 140.
Hering.E. M.
1938.
NEUE PALEARKTISCHE UND exotische bohrfliegen. Deut. Ent.
Ztschr. 1938:397-417.
90
Hodgson, J. M., and Rees, N. E.

1976.
DISPERSAL OF RHINOCYLLUS CONICUS FOR BIOCONTROL OF MUSK THIS
TLE. Weed Sci. 24 (1): 59-62.
HUFFAKER, C. B.
1956.
PRINCIPLES OF BIOLOGICAL CONTROL OF WEEDS. 10th Internatl.
Cong. Ent. Proc. 4: 533-542.
Jensen, L. A.
1970.
musk thistle and scotch thistle. Utah State Univ. Ext. Serv.
Weeders Digest 6: 1-2.
Jessep.C. T.
1975.
introduction of a weevil for biological control of nodding
thistle. New Zeal. Weed Pest Control Conf. Proc. 28, pp.
205-206.
Johnson, M. F.
1974.
CYNAREAE (ASTERACEAE) IN VIRGINIA: CIRSIUM, CARDUUS,
onopordum. Va. J. Sci. 25: 152-160.
Karny.M.
1963.
haltica carduorum guer., (col. chrysomelidae), attacking cir
SIUM arvense. Commonwealth Inst. Biol. Control. Tech. Bui. 3:
99-110.
Kazmi.S. M. A.
1963.
REVISION DER GATTUNG CARDUUS. teil I. Sonderdruck aus den
Mitt, der Bot. Staatssamml. Munchen. 5: 139-198.
1964.
Kok, L. T.
1974.
1975a.
revision der gattung carduus. teil ii. Sonderdruck aus den

Mitt, der Bot. Staatssamml. Munchen. 5: 279-550.
EFFICACY OF SPRING RELEASES IN COLONIZATION OF RHINOCYLLUS CON
ICUS FOR THE biocontrol of thistles. Environ. Ent. 3 (3):
429-430.
host specificity studies on ceuthorrhynchidius horridus
(PANZER) (COLEOPTERA: CURCULIONIDAE) FOR THE BIOCONTROL OF MUSK
and plumeless thistle. Weed Res. 15: 21-25.
1975b.
SURVIVAL OF AESTIVATING ADULT RHINOCYLLUS CONICUS FROELICH AT

DIFFERENT TEMPERATURES AND PHOTOPHASES. N.Y. Ent. Soc. J. 83:
251.
and Surles, W. W.
1975.
SUCCESSFUL BIOCONTROL OF MUSK THISTLE BY AN INTRODUCED WEEVIL,
Ward, R.RHINOCYLLUS
H., and Grills,
CONICUS.
C. C. Environ. Ent. 4 (6): 1025-1027.
1975.
BIOLOGICAL STUDIES OF CEUTHORRHYNCHIDIUS HORRIDUS (PANZER), AN

introduced weevil for thistle control. Amer. Ent. Soc. Ann.
68 (3): 503-505.
Letendre, M., Ritchot, C, Guibord, O. C, and Leduc, C.
1976.
ESSAI D'ERADICATION DU CHARDON PENCHE, CARDUUS NUTANS L. A
L'AIDE DU CHARANCON, RHINOCYLLUS CONICUS FROEL. PhytOprotection 57 (1): 47-55.
McCarty.M. K.
1964.
new and problem weeds musk thistle. N. Cent. Weed Control
Conf. Proc. 20: 62-63.
91
and Hatting, J. L.
1975.
EFFECTS OF HERBICIDES OR MOWING ON MUSK THISTLE SEED PRODUC
TION. Weed Res. 15: 363-367.
and Scifres, C J.
1969.
life cycle studies with musk thistle. Nebr. Univ. Col. Agr. and
Scifres,Home
C. J., Econ.
and Robison,
Res. Bui.L.230,
R. pp. 1-15.
1973.
a descriptive guide for major Nebraska thistles. Nebr. Univ.

Agr. Expt. Sta. Pub. SB 493, 23 pp.
Scifres, C. J., Smith, A. L., and Horst, G. L.
1969.
GERMINATION AND EARLY SEEDLING DEVELOPMENT OF MUSK AND
plumeless thistles.' Nebr. Univ. Col. Agr. and Home Econ. Res.
Bui. 229, pp. 1-28.
Madrigal, R. V., Smith, C. R., Jr., and Plattner, R. D.
1974.
CARDUUS NIGRESCENS SEED OIL, A RICH SOURCE OF PENTACYCLIC
triterpenes. Amer. Oil Chem. Soc. J. 51 (7): 531.
Martin, A. C, Zim, H. S., and Nelson, A. L.
1961.
AMERICAN wildlife and plants. 500 pp. Dover Pub., New York.
Maw.M. G.
1976.
AN ANNOTATED LIST OF INSECTS ASSOCIATED WITH CANADA THISTLE
(CIRSIUM ARVENSE) IN CANADA. Canad. Ent. 108: 235-244.
Mellini.E.
1951a. insetti del carduus nutans l. i. lixus elongatus goeze (col.:
CURCULIONIDAE). Bologna Ent. Bol. 18: 272-292.
1951b.
INSETTI DEL CARDUUS NUTANS L. II. RHINOCYLLUS CONICUS FROEL. E

LARINUS JACEAE F. (COL.: CURCULIONIDAE). Bologna Ent. Bol. 18:
319-349.
1952.
INSETTI DEL CARDUUS NUTANS L. III. EURIBIA SOLSTITIALIS

L. Bologna Ent. Bol. 19: 97-119.
Mohr, K. H.
1966.
familie chrysomelidae. In Frende, H., Harde, K. W., Lohse, G.

A., eds., Die Kafer Mitteleuropas, v. 9, pp. 95-280. Krefeld.
Moore, R. J., and Frankton, C.
1974.
the thistles of Canada. Canada Dept. Agr. Res. Br. Monog. 10,
111pp.
and Mulligan. G. A.
1956.
NATURAL HYBRIDIZATION BETWEEN CARDUUS ACANTHOIDES AND CAR
and Mulligan,
DUUS nutans
G. A. in ONTARIO. Canad. J. Bot. 34: 71-85.
1959.
A STUDY OF NATURAL HYBRIDIZATION BETWEEN CARDUUS ACAN

THOIDES and carduus nutans. 10th Internatl. Cong. Genet., Mon
treal, 1958, Proc. 2: 193.
Morihara, D. K., and Balsbaugh, E. U., Jr.
1976.
PHYTOPHAGOUS INSECTS COLLECTED ON MUSK THISTLE, CARDUUS
nutans, IN southeastern SOUTH Dakota. Environ. Ent. 5 (4):
692-696.
Mulligan, G. A.
1965.
RECENT COLONIZATION BY HERBACEOUS PLANTS IN CANADA. In
Baker, H. G., and Stebbins, G. L., eds., The Genetics of Colonizing
Species, pp. 127-146. Academic Press, New York.
92
Mulligan, G. A., and Frankton, C.

1954.
THE PLUMELESS THISTLES (CARDUUS SPP.) IN CANADA. Canad. Field
and Kevan,
Nat. P.
68:31-36.
G.
1973.
COLOR, BRIGHTNESS AND OTHER FLORAL CHARACTERISTICS ATTRAC

TING INSECTS TO THE BLOSSOMS OF SOME CANADIAN WEEDS. Canad. J.
and Moore,
Bot. R.
51:J.1939-1952.
1961.
NlBLETT.S.
1957.
Parmenter, L.
1952.
NATURAL SELECTION AMONG HYBRIDS BETWEEN CARDUUS ACAN

THOSES and c. nutans in Ontario. Canad. J. Bot. 39: 269-279.
notes on leaf-mining diptera. So. London Ent. and Nat. Hist.
Soc. Proc. 1956: 151-153.
FURTHER ADDITIONS TO THE LIST OF AGROMYZIDAE (DIPT.) FOUND IN
BRITAIN. Ent. Monthly Mag. 88 (1063): 279-280.
Peschken, D. P.
1971.
CIRSIUM ARVENSE (L.) SCOP., CANADA THISTLE (COMPOSITAE). In
Biological Control Programmes Against Insects and Weeds in
Canada, 1959-1968. Commonwealth Inst. Biol. Control Tech.
Commun. 4, pp. 79-83.
and Beecher, R. W.
1973.
CEUTHORHYNCHUS LITURA (COLEOPTERA: CURCULIONIDAE): BIOLOGY
AND FIRST RELEASES FOR BIOLOGICAL CONTROL OF THE WEED CANADA
THISTLE (CIRSIUM ARVENSE) IN ONTARIO, CANADA. Canad. Ent. 105
Friesen,(12):
H. A.,
1489-1494.
Tonks, N. V., and Banham, F. L.
1971.
RELEASES OF ALTICA CARDUORUM (CHRYSOMELIDAE: COLEOPTERA)
AGAINST THE WEED CANADA THISTLE (CIRSIUM ARVENSE) IN
and Harris,
CANADA.
P. Canad. Ent. 102: 264-271.
1975.
HOST SPECIFICITY AND BIOLOGY OF UROPHORA CARDUI (DIPTERA:

TEPHRITIDAE) A BIOCONTROL AGENT FOR CANADA THISTLE. Canad.
Ent. 107(10): 1101-1110.
Ross, H., and Hedicke, H.
1927.
DIE PFLANZENGALLEN (CECIDIEN) MITTEL-UND NORDEUROPAS, IHRE ERREGER UND BIOLOGIE UND BESTIMMUNGSTABELLEN. 348 pp. G.
Fischer, Jena.
Savulescu.T.
1955.
monografia uridenalelor din republica populara
romana. 1166 pp. Acad. Repub. Pop. Romfine, Bucharest,
Romania.
1957.
USTILAGINALELE DIN REPUBLICA POPULARA ROMANA. 1168

pp. Acad. Repub. Pop. Romane, Bucharest, Romania.
Schaber, B. D., Balsbaugh, E. V., Jr., and Kantack, B. H.
1975.
BIOLOGY OF THE FLEA BEETLE, ALTICA CARDUORUM (COL.:
CHRYSOMELIDAE) ON CANADA THISTLE (CIRSIUM ARVENSE) IN SOUTH
DAKOTA. Entomophaga 20 (4): 325-335.
Schutze, K. T.
1931.
DIE BIOLOGIE DER KLEINSCHMETTERLINGE UNTER BESONDERER
BERUCKSICHTIGUNG IHRER NAHRPFLANZEN UND ERSCHEINungszeiten. 235 pp. Frankfurt.

Seguy.E.
1934.
Spencer, K. A.
1973.
93
dipteres(brachyceres). faune de france. V. 28, 744 pp.
AGROMYZIDAE (DIPTERA) OF ECONOMIC IMPORTANCE. Entomologies 9,

418 pp. The Hague.
Stuckey, R. L., and Forsyth, J. L.
1971.
DISTRIBUTION OF NATURALIZED CARDUUS NUTANS (COMPOSITAE) MAP
PED IN RELATION TO GEOLOGY IN NORTHWESTERN OHIO. Ohio. J. Sci.
71(1): 1-15.
SURLES.W. W.
1975.
NATIVE HYMENOPTERAN PARASITOIDS ATTACKING AN INTRODUCED
weevil, rhinocyllus conicus in virginia. Environ. Ent. 3 (6):
and Kok,1027-1028.
L. T.
1975.
SEQUENTIAL RELEASES OF RHINOCYLLUS CONICUS FROELICH FOR THE
BIOCONTROL OF CARDUUS THISTLES. N.Y. Ent. Soc. J. 83: 250.
and Kok, L. T.
1976.
PILOT STUDIES ON AUGMENTATION OF RHINOCYLLUS CONICUS FOR CAR
DUUS thistle control. Environ. Ent. 5 (5): 901-904.
and Kok, L. T.
1977.
OVIPOSITIONAL PREFERENCE AND SYNCHRONIZATION OF RHINOCYLLUS
CONICUS WITH CARDUUS NUTANS AND C. ACANTHOIDES. Environ. Ent.
6 (2): 222-224.
Kok, L. T., and Pienkowski, R. L.
1974.
RHINOCYLLUS CONICUS ESTABLISHMENT FOR BIOCONTROL OF THISTLES
Pienkowski,
IN VIRGINIA.
R. L., and
Weed
Kok,Sci.
L. T.
22 (1): 103.
1975.
MORTALITY OF THE IMMATURE STAGES OF RHINOCYLLUS CONICUS, A

thistle head weevil, in virginia. Environ. Ent. 4 (3): 371-372.
Sydow, H., and Sydow, P.
1904.
monographiauredinearum. V. 1, 972 pp. Leipzig.
Tsao, C, and Wang, K.
1965.
A NEW PEST OF ZIZYPHUS JUJUBA IN THE SANCHOW AREA. Acta Ent.
Sinica 14 (4): 420-422.
Uphof.J. C. T.
1942.
ECOLOGICAL RELATIONS OF PLANTS WITH ANTS AND TERMITES. Bot.
Rev. 8 (9): 563-598.
Wapshere, A. J.
1974.
A STRATEGY FOR EVALUATING THE SAFETY OF ORGANISMS FOR
BIOLOGICAL WEED CONTROL. Ann. Appl. Biol. 77: 201-211.
Ward, R. H., Pienkowski, R. L., and Kok, L. T.
1974.
HOST SPECIFICITY OF THE FIRST INSTAR OF CEUTHORRHYNCHIDIUS HORRIDUS, A WEEVIL FOR BIOLOGICAL CONTROL OF THISTLES. J. Econ.
Ent. 67 (6): 735-737.
Williamson, G. D.
1963- SUMMARY OF PARASITE AND PREDATOR LIBERATION IN CANADA AND OF
65. insect shipments from Canada. Canad. Insect Pest Rev. 41 (7):
137-151; 42 (7): 125-140; 43 (5): 127-139.
196670.
insect liberations in Canada. Canada Dept. Agr. Liberation

Bui. 21: 1-18; 32: 1-19; 33: 1-9; 34: 1-16.
1974.
insect liberations in Canada. Canada Dept. Agr. Liberation

Bui. 37: 1-21.
94
Wilson, C L.
1969.
use of plant pathogens in weed control. Ann. Rev. Phytopath.

7:411-434.
Wilson, F.
1964.
the biological CONTROL OF weeds. Ann. Rev. Ent. 9: 225-244.
WUNDERLIN.R. P.
1969.
the genus carduus in Illinois. 111. State Acad. Sci. Trans. 62:
98-99.
ZWOLFER, H.
1963.
larinus - rhinocyllus weed projects for Canada. Common
wealth Inst. Biol. Control Progr. Rpt. 10, pp. 1-23. [Processed.]
1964.
THE HOST SPECIFICITY OF CEUTHORHYNCHUS LITURA F. (COL. CURcul.). weed projects for Canada. Commonwealth Inst. Biol.
Control Prog. Rpt. 12, pp. 1-11. [Processed.]
1965a.
PRELIMINARY list of PHYTOPHAGOUS insects ATTACKING WILD

cynareae (Compositaei species in Europe. Commonwealth Inst.
Biol. Control Tech. Bui. 6: 81-154.
1965b.
OBSERVATIONS ON THE DISTRIBUTION AND ECOLOGY OF ALTICA CARduorum guer. (col. chrysomelidae). Commonwealth Inst. Biol.
Control Tech. Bui. 5: 129-141.
1967.
THE HOST-RANGE, DISTRIBUTION AND LIFE-HISTORY OF RHINOCYLLUS

CONicus froel. Commonwealth Inst. Biol. Control Prog. Rpt. 18:
1-21.
1969.
EXPERIMENTAL FEEDING RANGES OF CHRYSOMELIDAE SPP. (COL.)

associated with cynareae host plants (COMPOSITAE). Common
wealth Inst. Biol. Control Tech. Bui. 12: 115-130.
1970.
COLLECTIONS OF ALTICA CARDUORUM GUER. FOR RELEASE AND STUDIES

in the united kingdom. Commonwealth Inst. Biol. Control Rpt.
1970, pp. 1-10. Del6mont, Switzerland. [Processed.]
1971.
COMPETITION AND COEXISTENCE IN PHYTOPHAGOUS INSECTS ATTACK

ING THE HEADS OF CARDUUS NUTANS L. In Dunn, P. H., ed., 2d Internatl. Symp. Biol. Control of Weeds Proc. Commonwealth Inst.
Biol. Control Misc. Pub. 6, pp. 74-81.
1974.
DAS TREFFPUNKT - PRINZIP ALS KOMMUNIKATIONSSTRATEGIE UND

ISOLATIONSMECHANISMUS BEI BOHRFLIEGEN (DIPTERA:
trypetidae). Ent. Germ. 1 (1): 11-20.
_ and Eichhorn.O.
1966.
THE HOST RANGES OF CASSIDA SPP. ATTACKING CYNAREAE IN
EUROPE. Ztschr. f. Angew. Ent. 58: 384-397.
_ Frick, K. E., and Andres, L. A.
1971.
A STUDY OF THE HOST PLANT RELATIONSHIPS OF EUROPEAN MEMBERS
OF THE GENUS LARINUS (COL.: CURCULIONIDAE). Commonwealth Inst.
Biol. Control Tech. Bui. 14: 97-143.
95
_ and Harris, P.
1966.
CEUTORHYNCHUS LITURA F. (COL. CURCULIONIDAE), A POTENTIAL INSECT
FOR THE BIOLOGICAL CONTROL OF THISTLE, CIRSIUM ARVENSE L. IN
CANADA. Canad. J. Zool. 44: 23-38.
_ and Harris, P.
1971.
HOST SPECIFICITY DETERMINATION OF INSECTS FOR BIOLOGICAL CON
TROL OF WEEDS. Ann. Rev. Ent. 16: 157-178.
_ and Pattullo, W.
1970.
ZUR LEBENSWEISE UND WIRTSBINDUNG DES DISTEL-BLATTKAFERS LEMA
CYANELLA L. (PUNCTICOLLIS CURT.) (COL. CHRYSOMELIDAE). Anz. f.
Schadlingsk. Pflanzenschutz 43 (4): 53-59.
INDEX
Page
Fungi
Aecidium
Albugo
Ascochyta
Bremia
Cercosporella
Cladosporium
Epitea
Erysiphe
Fusicladium
Heteropatella
Leptosphaeria
Leveillula
Macrosporium
Oidium
Ophiobolus
Ovularia
Passalora
Pelospora
Phoma
Phragmidium
Phyllosticta
Pseudohelotium
Puccinia
83
82
82
82
82
82
84
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
44,
83-85, 86
82
82
82
82
82
83-84
85-86
82
Ramularia
Rhabdospora
Sclerotinia
Scolecotrichum
Septoria
Uredo
Ustilago
Venturia
Agallia
Agalliopsis
Agapanthia
Agonopteryx
Agromyza
Agromyzidae
Agrotis
Aliolis
Alleculidae
Altica
Anaphothrips
Anobiidae
Anthomyiidae
Anuraphis
Apanteles
Aphididae
Aphis
Aphthona
Apion
Archips
Arctiidae
Arima
Aritranis
Autographa
Brachycaudus
Brachymeria
Brachynotus
Bracon
Braconidae
Buprestidae
Byturidae
Page
18
18
26
71
79
79-81
64
59
24
28-30,
85
10
24
81
20
63
19
19
30
40-42
69
62
30
56
63
20
63
42
49,
51, 52, 56, 59
49
62
24
Insects
Acalles
Acanalonia
Acanaloniidae
Acanthiophilus
Acanthocnemus
Acertagallia
Acrididae
Acroclita
Actio.
Adelphocoris
Aelia
96
40
22
22
74
23
18
11
68
68
12
15
Calocoris
Campoplex
Capitophorus
Carduiphila
Carpocoris
Cassida
Cecidomyiidae
Cerambycidae
Ceratapion
Cercopidae
12
59
20,21
81
15
31-34,
86
72
26-27
41
21-22

Cerodontha
Cerosipha
Cetonia
Ceuthorrhynchidius
Ceutorhynchus
Chaetocnema
Chaetostomella
Chalcidoidea
Cheilosia
Chlamydatus
Chlorochroa
Chlorocytus
Chlorophanus
Chloropidae
Choreutis
Chrysomela
Chrysomelidae
Cicadellidae
Cleonus
Clepsis
Cnephasia
Cnephasiella
Cochylidae
Cochylis
Coleophora
Coleophoridae
Conotrachelus
Coreidae
Coreus
Cosmopepla
Cosmopterigidae
Crepidodera
Cryptocephalus
Curculionidae
Cynthia
Page
79
21
25
60
42-45,
86
34
74
51-52
72-73
12
15
67
45
79
71
34
28-40
18
45-46
69
69
69
69
69-70
71
71
46
14
15
15
70
34
34
40-61
62-63
Dactynotus
Delphacidae
Deltocephalus
Diabrotica
Diachus
Dictyopharidae
Dolycoris
Donus
Drosophilidae
21
22
18
34
35
22
15
46
78
Elymana
Empoasca
Endria
Endrosis
Enneamera
18
18
18
71
35
97
Ephialtes
Epiblemma
Epicauta
Epithrix
Estigmene
Euaresta
Eublemma
Eucosma
Euco9midae
Eupithecia
Eurydema
Eurygaster
Eurytoma
Euscelis
Euschistus
Euxanthoides
Euxoa
Evacanthus
Exeristes
Page
49,67
67
24
35
62
74
63-64
68
67-68
65
16
17
64
18
16
70
64
16
50
Forficula
Forficulidae
Frankliniella
: 11
11
11
Galeruca
Gelechiidae
Geocoris
Geometridae
Glyphipterygidae
Graphocephala
Grapholitha
Gryllidae
35-36
70
14
65
71
19
68
11
Hadroplontus
Haltica
Haplothrips
Harpalus
Heliothis
Herpystes
Hippodamia
Holocryptus
Homoeosoma
Hoplasoma
Hylemya
Hypera
42
28
10
30
64
68
59
56
65-66
36
81
46-47
Irbisia
Issidae
12
22
Jaapiella
72
Lachnea
36
98
Larinus
Lasioderma
Latalus
Lathridiidae
Lebia
Lema
Leptinotarsa
Leptocoris
Leptomias
Leptus
Liorhyssus
Liriomyza
Lita
Lixochelus
Lixus
Longitarsus
Lopidea
Lycanades
Lygaeidae
Lygaeus
Lygus
Page
47-52
24-25
19
62
30
36-37
37
17
52
30
17
79
70
53
52-55
37
13
64
14
14
13
Macrophthalmus
Macrosiphum
Macrosteles
Melanagromyza
Melanocoryphus
Melanoplus
Meligethes
Meloidae
Melyridae
Merista
Mesomorphus
Microctonus
Miridae
Monolepta
Mordella
Mordellidae
Mordellistena
Myclopsis
Myelois
62
21
19
79
14
11-12
23
24
22
37
22
37
12
37
24
23-24
24
65
66
Napomyza
Nemognatha
Neosteles
Netelia
Nezara
Nitidulidae
Noctuelia
Noctuidae
Nymphalidae
80
24
19
64
16
23
67
63
62
Odontotarsus
Oecanthus
Oecophoridae
Oedemera
Oedemeridae
Olethreutidae
Orellia
Ortholomus
Oscinella
Ostrinia
Otiorhynchus
Oulema
Oxythyrea
Papaipema
Paracantha
Paragonia
Paraholaspis
Paraphlepsius
Parascaptomyza
Pegomya
Pentatomidae
Peridroma
Phaenobremia
Phalacridae
Phalacrus
Phalonia
Phaloniidae
Phaonia
Philaenus
Phlaeothripidae
Phycita
Phycitodes
Phytometra
Phytomyza
Phytonomus
Piesma
Piesmatidae
Plagiognathus
Platyptilia
Platystoma
Platystomatidae
Poecilocapsus
Polyodaspis
Porphyrinic^
Potosia
Procerallus
Psammotettix
Pseudapion
Pseudocleonus
Page
17
11
71
23
23
67-68
75
14
79
67
55
37
25
64-65
75
19
64,
75, 76
19
78
81
15
65
20
23
23
70
69
44
22
10
67
65
63
80
47
17
17
13
68
73
73
13
79
63
25-26
23
19
42
55
Pseudococcidae
Psylliodes
Pterophoridae
Purpuricenus
Pyralidae
Pyrausta
Pyroderces
Pyrrhocoridae
Pyrrhocoris
Raphidia
Rhinocyllus
Page
22
37-38
68
27
65
67
70
17
17
Rhodobaenus
Rhopalidae
Rhopalosiphum
59
56-60,
86
60
17
21
Scaptomyza
Scarabaeidae
Scrobipalpa
Scutelleridae
Semium
Sitona
Sminthuridae
Sphaeroderma
:
Sphaerolophus - -
Staphylinidae
Syromastes
Syrphidae
Systena
78
25-26
70
17
14
60
10
38-39
30
22
15
72-73
39-40
Tanyderidae
Tanymecus
Tenebrionidae
Tephritidae
Tephritis
Terellia
Tetrastichus
Tettigoniidae
Thalpochares
Thripidae
Thyanta
Tingidae
Tingis
Tortricidae
Toxopterina
Trichosirocalus
Tropinota
Trypetidae
73
60
62
74-78
75
76
50,
56, 75, 76
11
64
10
17
14
14
69
21
60-61,
85
26
74
Urophora
99
Page
76-78,
86
Vanessa
62
Xyphosia
78
Plants (Carduus Species)

C. acanthoides L.
2,
4, 5, 14-81, 84, 85
C. acantholepis Kar. & Kit.
84
C. albidus Bieb.
5
C. ammophilus Hoff. ex Link
83, 84
C. arabicus ssp. marmoratus (Boiss. &
Heldr.) Kazmi
85
C. armatus Boiss. & Heldr.
83
C. ballii Hook. f.
83, 84
C. candicans Waldst. & Ket.
84
C. carpetanus Boiss. & Reut.
84
C. cernua Cass.
84
C. chevallieri Barr.
84
C. chrysacanthus Ten.
49, 84
C. clavulatus Link
83
C. collinus Waldst. & Kit.
84
C. cornius Boiss. & Heldr.
83
C. crassifolius Willd.
75,
76,81
C. x crispo-tenuiflorus Merat
3
C. crispus L.
2,
3, 20-82, 84, 85
C. defloratus L.
10,
14-81,83
C. x dubius Balbis
3
C. edelbergii Rech.
15-81
C. gayanus Dur.
84
C. getulus Pomel
84
C. glaucus Ledeb.
82, 84
C. hamulosus Ehrhart
5
C. kerneri Simk.
83, 84
C. kikuyorum Fries
85
C. leiophyllus Petr.
3, 84
C. x leptocephalus Petermann
3
C. leptocladus Dur.
84
C. litigiosus Nocc. ex Balbi
27,
48, 52, 54, 56
C. macrocephalus Desf.
84
C. macrolepis Petermann
3
C. meonanthus Hoffm. & Link
84
C. micropterus ssp. broteri (Welw.)
Kazmi
82
C. x-mixtus Corb.
3
100
Page
44,
83, 84
C. nigrescens Villars
5,
48, 54
C. nivea Kar. & Kit.
84
C. nutans L.
1,
3-5, 11-81,83, 85
C. nutans ssp. leiophyllus (Petr.) J. Ar. - 3
C. nutans ssp. macrolepis (Petermann)
Kazmi
3
C. nutans ssp. nutans
3
C. onopordioides Fisch.
84
C. x orthocephalus Wallroth
3
C. x pernutanti-pycnocephalus Massal - 3
C. personatus (L.) Jacq.
31-81,
84
C. myriacanthus Salzm.
Page
2,
5, 10-81,83, 84
C. pycnocephalus var. albidus (Bieb.)
Kazmi
85
C. reuterianus Boiss.
84
C. scardicus (Gris.) Wettst.
84
C. seminudus Bieb.
83
C. x semiperegrinus Aellen
3
C. spachianus Dur.
83
C. tenuiflorus Curtis
2,
23-81, 83, 84
C. x theriottii Rouy
3
C. thoermeri (Weinm.)
1,84
C. uncinatus Bieb.
75
C. pycnocephalus L.
ft U.S. GOVERNMENT PRINTING OFFICE: 1981 O324-570
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Insects and Fungi Associated With Cardus

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Insects and Fungi Associated With Cardus

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This is a reproduction of a library book that was digitized

by Google as part of an ongoing effort to preserve the

A terminal inflorescence of musk thistle (Carduus nutans) with its introduced

By S. W. T. Batra, J. R. Coulson, P. H. Dunn, and P. E. Boldt1

ECONOMIC IMPORTANCE, CONTROL, AND

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

commun.). Plumeless thistle (C. acanthoides) is a problem in the Ap

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

portant introduced weed in South Africa (Henderson and Ander

x C. crispus (2N = 16)

More cytogenetic and ecological studies of possible Carduus

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

within 2 days of the first terminal heads in a population showing

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

PHYTOPHAGOUS INSECTS ASSOCIATED WITH

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

sects not in the Beneficial Insect Introduction Laboratory (BIIL)

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

Polytechnic Institute and State University, Blacksburg. Data from

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

N.J. - New Jersey

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

Carduus nutans (S. Dak.)

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

Odontotarsus purpureolineatus (Rossi)

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

Graphocephala coccinea Forster

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

Winter host is Robinia; polyphagous

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

Carduus pycnocephalus (S. Calif., S. Europe)

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

Plant recs.: Carduus pycnocephalus (S. Calif.)

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

Carduus nutans (S. Dak.)

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

Lasioderma baudii Schilsky

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

Agapanthia dahli Richter

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

tenuiflorus (A), C. pycnocephalus (A), Cirsium (D,

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

Grandi (1952); Heikertinger (1954); Zwblfer

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

Adults and larvae skeletonize and severely

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

(1965a, 1969); Zwblfer and Eichhorn (1966);

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

Diachus auratus (F.)

TECHNICAL BULLETIN 1616, U.S. DEPT. OF AGRICULTURE

INSECTS AND FUNGI ASSOCIATED WITH CARDUUS THISTLES

limited to Carduus, Cirsium, and Silybum; also