This article was downloaded by: [Hodkinson, Ian D.]
On: 14 January 2009
Access details: Access Details: [subscription number 907784658]
Publisher Taylor & Francis
Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,
37-41 Mortimer Street, London W1T 3JH, UK
Journal of Natural History
Publication details, including instructions for authors and subscription information:
http://www.informaworld.com/smpp/title~content=t713192031
Life cycle variation and adaptation in jumping plant lice (Insecta: Hemiptera:
Psylloidea): a global synthesis
Ian D. Hodkinson a
a
School of Biological and Earth Sciences, Liverpool John Moores University, Liverpool, UK
Online Publication Date: 01 January 2009
To cite this Article Hodkinson, Ian D.(2009)'Life cycle variation and adaptation in jumping plant lice (Insecta: Hemiptera: Psylloidea): a
global synthesis',Journal of Natural History,43:1,65 — 179
To link to this Article: DOI: 10.1080/00222930802354167
URL: http://dx.doi.org/10.1080/00222930802354167
PLEASE SCROLL DOWN FOR ARTICLE
Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf
This article may be used for research, teaching and private study purposes. Any substantial or
systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or
distribution in any form to anyone is expressly forbidden.
The publisher does not give any warranty express or implied or make any representation that the contents
will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses
should be independently verified with primary sources. The publisher shall not be liable for any loss,
actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly
or indirectly in connection with or arising out of the use of this material.
Journal of Natural History
Vol. 43, Nos. 1–2, January 2009, 65–179
Life cycle variation and adaptation in jumping plant lice (Insecta:
Hemiptera: Psylloidea): a global synthesis
Ian D. Hodkinson*
School of Biological and Earth Sciences, Liverpool John Moores University, Liverpool, UK
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
(Received 1 February 2008; final version received 20 July 2008)
This paper integrates the scattered information on the life histories of the jumping
plant lice or psyllids, examining those aspects of their biology that contribute to
successful life cycle completion. Variation in life history parameters is reviewed
across the world’s psyllids and the relative importance of phylogeny and
environment, including host-plant growth strategy, in determining life history
strategies is assessed. Elements of life cycles considered include: development rate
and voltinism, response to high temperature and drought, cold-hardiness and
overwintering strategy, seasonal polymorphism, diapause, metabolism, host-plant
selection and range, phenological and other adaptations to host plants, disease
transmission and host amelioration, dispersal, reproduction and mate finding.
Life history parameters are analyzed for 342 species. While a phylogenetic signal
can be identified within the data, the main drivers for life history adaptation are
environmental temperatures and water availability, acting directly on the psyllids
or mediated through their host plants.
Keywords: psyllid; life-history; phylogeny; temperature; water
Introduction
Jumping plant lice, or psyllids (Psylloidea), comprise a group of around 3000 species
of small plant-sap-feeding insects allied to the aphids and whiteflies. They occur
throughout nearly all the world’s major climatic regions where suitable host plants
are found. This paper attempts to integrate the wealth of scattered information on
the life histories of the psyllids and to examine those aspects of their biology that
contribute to successful life cycle completion. A significant proportion of this
information is contained in relatively old or obscure sources, or is to be found in
current literature that escapes electronic abstracting and is thus in increasing danger
of remaining unrecognized. The broad variations that occur within psyllid life cycles
are documented and the modifications of biology and mechanisms by which psyllids
adapt to exploit a diverse range of host plants growing under varying environmental
conditions are examined. An attempt is made throughout to identify common
themes and patterns. Table 1 draws together, in taxonomic sequence, the basic data
and reference sources that are avilable for the life histories of the world’s Psylloidea
species and this forms the foundation for the subsequent analyses.
The great majority of psyllid species are narrowly host-specific and are
predominantly
associated
with
perennial
dicotyledenous
angiosperms
(Klimaszewski 1973; Hodkinson and White 1981; Hodkinson 1983a, 1986a, 1988a;
Gegechkori and Loginova 1990; Hollis 2004). A few species develop on monocots
*Email: i.d.hodkinson@ljmu.ac.uk
ISSN 0022-2933 print/ISSN 1464-5262 online
# 2009 Taylor & Francis
DOI: 10.1080/00222930802354167
http://www.informaworld.com
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
66 I.D. Hodkinson
including Livia spp. on Carex and Juncus, Bactericera tremblayi (Wagner) and B.
kratochvili (Vondracek) on Allium spp. (Alliaceae), and a few species on Palmaceae.
These include an undescribed species on Bactris gasipaes in Colombia, five Hawaiian
Megatrioza on Pritchardia, an undescribed Australian paurocephaline species on
Livistona and an undescribed Indian ‘‘Psylla’’ species on Areca catechu
(Klimaszewski 1973; Pava et al.1983; Uchida and Beardsley 1988; Hodkinson and
Bird 2000; Mondal et al. 2003; Hollis 2004). Gymnosperms are similarly poorly
exploited as host plants, with just two species of Ehrendorferiana Burckhardt
breeding on Austrocedrus and Fitzroya (Cupressaceae) in Chile and Trioza colorata
Ferris and Klyver and T. dacrydii Tuthill exploiting Dacrydium (Podocarpaceae) in
New Zealand (Tuthill 1952; Burckhardt 2005a). Conifers, however, as discussed
later, play a highly significant role as overwintering shelter plants for adults from a
wide range of psyllid species.
The psyllid life cycle typically comprises of an egg stage, five larval instars and a
sexually reproducing adult stage, with males and females usually showing only
moderate deviation from a 1:1 sex ratio at emergence. Parthenogenetic reproduction,
in which only females are found in the population, is rare but probably occurs in
some populations of Cacopsylla rara (Tuthill), Glycaspis operta (Moore), Glycaspis
atkinsoni Moore and Cacopsylla myrtilli (Wagner) (Hodkinson 1983b; Moore 1983;
Hodkinson and Bird 2006a). The latter species appears as populations of
parthenogenetic females throughout most of its circumboreal range but males
become common at higher altitudes above tree line (Hodkinson and Bird 2006a).
Within the equitable warm and wet conditions of lowland tropical evergreen
rainforests psyllid life cycles tend to be continuous, with multiple generations per
year. This probably typifies the environment under which psyllids originally
radiated. This continuous multivoltine life cycle – appropriate to a climatically
benign environment – has, however, subsequently undergone considerable modification. Psyllids have adapted to exploit a range of host plants that have themselves,
over evolutionary time, diversified their physiognomy, physiology and phenology as
they adapted to varying environmental conditions within widely different major
climatic zones. Such evolution is driven by two overriding variables, temperature and
precipitation, which vary in response to site latitude, altitude and continentality.
Host plants have also undergone concomitant chemical evolution and adaptation
during which their unique chemistry, in particular their complement of characteristic
secondary compounds, has evolved. This unique chemistry is thought to form the
basis for host selection and host fidelity in the psyllids.
In the only detailed study of psyllid plant coevolution, involving host-specific
legume feeding psyllids on the Canary Islands, .60% of host associations resulted
from phylogenetically conserved host switching among related legumes: strict
cospeciation was only evident among more recent psyllid–host associations (Percy
2003a, 2003b; Percy et al. 2004). Thus, while precise evolutionary tracking of host
plants may not always have occurred, related groups of psyllids tend usually to be
typically associated with related host-plant taxa and individual psyllid species, with a
few notable exceptions, display a high degree of host specificity (Hodkinson 1974,
1986b; van Klinken 2000). The exceptions are usually north temperate multivoltine
pest species such as Bactericera nigricornis, B. trigonica Hodkinson and B. cockerelli
feeding variously on a range of host-plant genera within the Solanaceae,
Umbelliferae and Cruciferae (Pletsch 1947; Wallis 1955; Hodkinson 1981). Some
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
67
species that were initially thought to be polyphagous, such as the univoltine Trioza
rotundata in Europe (on Stellaria (Caryophyllaceae), Saxifraga (Saxifragaceae) and
Cardamine (Cruciferae), (Conci and Tamanini 1987, 1991)) have subsequently been
shown to be complexes of species, each with a narrow host range (Burckhardt and
Lauterer 2002).
Many psyllids form galls on their host plant. These may include simple
irregular distortions of the leaf or shoot, through leaf pit galls or roll leaf galls to
complex enclosed gall structures on leaves, shoots, flowers rootlets and stems.
Detailed information on gall formation and the morphological adaptations for
living in different types of galls is reviewed by Hodkinson (1984) and Burckhardt
(2005b). This study avoids detailed description of gall formation and focuses,
where appropriate, on the adaptive significance of galls within individual psyllid
life cycles.
The paper initially considers the important constraints, adaptations and
modifications of psyllid biology that have shaped their life cycles. It analyzes how
these life cycle elements and adaptations have been assembled and combined to
produce the wide range of life histories we observe among the world’s psyllid fauna
today. The relative significance of phylogeny and environment are assessed as
predictors of psyllid life histories.
Important elements of psyllid life histories
Development rate and voltinism
Ambient temperature is a major determinant of egg and larval development rates in
non-diapausing psyllids and, as such, governs the potential number of generations
per annum (voltinism). The slowest development rate recorded is for Strophingia
ericae on Calluna at high elevation, which, including two periods of winter diapause,
takes 2 years to complete one generation, (Hodkinson 1973b). By contrast,
multivoltine tropical/subtropical species such as Heteropsylla cubana, Diaphorina
citri, Trioza erytreae and Trioza magnicauda, with free-running life cycles, may
complete between 8 and 16 generations per annum (Table 1). Development rates and
resultant generation times may, however, differ between seasons, depending on
varying ambient temperatures as in Phytolyma fusca on Milicia, Ctenarytaina
spatulata on Eucalyptus and Diaphorina citri on Citrus (Ledoux 1955; Bigornia and
Obana 1974; Shahid and Khan 1976; Perez Otero et al. 2005). Many temperate
species are univoltine, with a relatively short development period compressed within
and synchronized with the early growing season of the host plant (Table 1). Arctic
species, including many Cacopsylla, Psylla and Bactericera species, despite their
adaptations to harsher climates, almost invariably show a similar trend to
univoltinism (Table 1) (Hodkinson et al. 1979; Hodkinson and Bird in press).
Among multivoltine temperate species such as Trioza urticae on Urtica, various
Cacopsylla spp. on Pyrus and Agonoscena cisti on Pistacia the number of generations
per annum rarely exceeds six, and is usually no more than than three to four (Onillon
1969; An et al. 1996; Lauterer 1998; Souliotis and Tsourgianni 2000). Some widely
distributed species, such as Trioza cinnamomi on Cinnamomum are univoltine in the
cooler parts of their range but become multivoltine elsewhere (Miyatake 1969;
Rajapakse and Kulasekera 1982). Strophingia ericae, cited previously, becomes unirather than semivoltine at lower warmer altitudes (Hodkinson 1973b).
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Lerp former
Feeding site
Gall
Voltinism
Overwintering elsewhere
Hodkinson (1973a,
1973b), Parkinson and
Whittaker (1975),
Lauterer (1976),
Whittaker (1985), Miles
et al. (1997, 1998),
Hodkinson et al. (1999),
Butterfield et al. (2001)
Rapisarda (1990a),
Hodkinson et al. (1999)
Overwintering on host
References
Overwintering stage
Host plant(s)
Plant type
Psyllidae
Strophingiinae
Species
Climate zone
Higher taxon
Strophingia ericae
(Curtis)
TeM
Pe/C
L
S
1 or
0.5
S
Calluna
S. cinereae
Hodkinson
TeM/M
Pe/C
L
S
1
S
Erica
68 I.D. Hodkinson
Table 1. Life history characteristics of the world Psylloidea. The psyllid classification follows White and Hodkinson (1985) updated to include recent
changes. Where subfamilies comprise of a single tribe the subfamily name is given. Note: abbreviations: climatic zone: TrM, tropical moist; TrD, tropical
dry; TrS, tropical seasonal; M, Mediterranean; TeM, temperate moist; TeD, temperate dry; B, boreal; plant functional type of Raunkiær, based on
overwintering strategy: P, Phanaerophyte (tall trees and shrubs, overwintering buds above soil surface); C, Chamaephyte (low growing or prostrate dwarf
shrubs, overwintering buds ,25 cm above soil surface); H, Hemicryptophyte (overwintering bud at soil surface); G, Geophyte (overwintering bud below
soil surface); He, Helophyte (marsh plants); T, Therophyte (overwintering as seed); Par, parasitic on other plants; d, deciduous; e, evergreen; s,
semideciduous; overwintering stage: E, egg; L, larva; A, adult; overwintering site: on the host plant: T, trunk/stem; S, shoot; L, leaf; R, roots; B, buds;
elsewhere: C, conifers/evergreen shrubs; LL, leaf litter; voltinism: M, multiple generations per year, number not fully determined, otherwise number of
generations per annum stated; feeding site: S, shoot apex; L, expanded leaf; F, flower; St, stem; R, roots; B, buds; gall type: D, general distortion of leaf
and/or shoot; P, pit gall on leaf; R, roll leaf gall; Ro, root gall; Lf, leaf-fold gall; F, flower gall; El, enclosed leaf gall; Es, enclosed stem gall; Eb, enclosed
bud gall; lerp formation indicated by (X).
H
A
S
1
S
D
Carex
Ossiannilsson (1992)
TeM
H
A
S
1+
S
D
Juncus
L. maculipennis
(Fitch)
TeM
H
A
C
1?
S
D
Juncus
L. mediterranea
Loginova
M/TeD
H
A
C
1
S
Verrier (1929), HeslopHarrison (1949b),
Schmidt (1966),
Gegechkori (1984)
McAtee (1915), Weiss and
West (1922), HeslopHarrison (1949b)
Gegechkori (1984)
Phytolyma fusca
Alibert
TrS
Ps
ELA
S
M
S/L
El
Milicia
P. lata (Walker)
TrS
Ps
ELA
S
M
S/L
El
Milicia
Gyropsylla ilicis
(Ashmead)
G. spegazziniana
(Lizer)
TrS
Pe
A
S
1
L
R
Ilex
TeM
Pe
E
S
1
L
R
Ilex
Juncus
Vosseler (1906), White
(1966, 1967), Ledoux
(1955)
White (1966, 1967),
Cobbinah (1986)
Mead (1983)
Brèthes (1921), Leite and
Zanol (2001)
69
TeM
Journal of Natural History
Gall
Lerp former
Feeding site
Gyropsyllini
Voltinism
Aphalarinae
Phytolymini
Overwintering elsewhere
Livia crefeldensis
Mink
L. junci (Schrank)
References
Overwintering on host
Liviinae
Host plant(s)
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
A
C
1?
L
TeM
H
A
C/L 2+
L
TeM
H
A
C
1
TeM
TeM
He
H
A
A
C
C
1
?
TeM
H
A
C/L 1
S
Rumex
A. freji Burckhardt TeM
and Lauterer
H
A
C
S
Polygonum
TeM
H
A
C/L 1
S
Polygonum
A. longicaudata
Schaefer
P
Lerp former
H
1–2
Gall
Lauterer (1991),
Ossiannilsson (1992)
Lauterer (1979), Conci
et al. (1993)
Ossiannilsson (1992)
Lauterer (1982)
Feeding site
Polygonum
Voltinism
Ossiannilsson (1992)
Overwintering elsewhere
Stellaria
TeM
Aphalara affinis
(Zetterstedt)
A. avicularis
Ossiannilsson
A. borealis HeslopHarrison
A. calthae (L.)
A. crispicola
Ossiannilsson
A. exilis (Weber
and Mohr)
Overwintering on host
References
Overwintering stage
Host plant(s)
Plant type
Aphalarini
Species
Climate zone
Higher taxon
Polygonum
S/L/F
L
P
Caltha
Rumex
Lauterer (1976),
Gegechkori (1984),
Ossiannilsson (1992)
Gegechkori (1984),
Lauterer (1991),
Ossiannilsson (1992),
Conci et al. (1993) (all
as A. polygoni),
Burckhardt and
Lauterer (1997)
Lauterer (1976)
70 I.D. Hodkinson
Table 1. (Continued.)
Lerp former
Gall
H
A
C
1
L
Polygonum
TeD
H
A
?
1
?
Rumex
TeM
H
A
C
2
L
Rumex
TeM
H
L
R?
1
S
Achillea
TeM
H
L
S
1
S
Senecio
Lauterer and Burckhardt
(2004)
TeM
TeM
H
H
L
L
R?
S
1
1
S
S
Artemisia
Senecio
TeD
TeM
H
H
L
L
S
S
1
1
St
S
Phlox
Leontodon
Conci et al. (1993)
Lauterer and Burckhardt
(2004)
Wheeler (1994)
Lauterer and Burckhardt
(2004)
Lauterer (1982),
Gegechkori (1984),
Ossiannilsson (1992)
(all as A. rumicicola)
Burckhardt and
Lauterer (1997)
Conci et al. (1993)
71
TeM
Journal of Natural History
Feeding site
Gegechkori (1984), Conci
et al. (1993)
Gegechkori (1984)
Voltinism
References
Overwintering elsewhere
Craspedolepta
bulgarica
(Klimaszewski)
C. campestris
Lauterer and
Burckhardt
C. conspersa (Löw)
C. crispata Lauterer
and Burckhardt
C. eas (McAtee)
C. flavipennis
(Foerster)
Overwintering on host
A. maculipennis
Löw
A. nigrimaculosa
Gegechkori
A. polygoni
Foerster
Host plant(s)
Overwintering stage
Aphalarini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
TeM
H
L
R?
1
S
Artemisia
C. nebulosa
(Zetterstedt)
TeM/B
H
L
R
1
S
Chamerion
C. nervosa
(Foerster)
TeM
H
L
R
1
L
Achillea
C. omissa Wagner TeM
C. sonchi (Foerster) TeM
H
H
L
L
R
S
1?
1?
S
S
Artemisia
Leontodon
B
H
L
R?
1
S
Chamerion
TeM
H
L
R
1
R/S
Ro
Lerp former
Feeding site
Gall
Voltinism
C. malachitica
(Dahlbom)
C. schwarzi
(Ashmead)
C subpunctata
(Foerster)
Overwintering elsewhere
Conci et al. (1993),
Hodkinson
(unpublished)
Lal (1934), Sampo (1975),
Lauterer (1993a), Bird
and Hodkinson (1999,
2005), Hodkinson and
Bird (2006b)
Lauterer (1991),
Hodkinson
(unpublished)
Lauterer (1991)
Lauterer and Burckhardt
(2004)
Hodkinson and Bird
(1998, unpublished)
Lauterer and Baudys
(1968), Bird and
Hodkinson (1999,
2005), Hodkinson and
Bird (2006b)
Overwintering on host
References
Overwintering stage
Host plant(s)
Plant type
Aphalarini
Species
Climate zone
Higher taxon
Chamerion
72 I.D. Hodkinson
Table 1. (Continued.)
Lerp former
L
S
1
S
Solidago
Rhodochlanis
bicolor (Scott)
M
Th
EL
S
1
S
R. salsolae
(Lethierry)
M
Pe
EL
S
1
S
Salicornia,
Suaeda
Salsola,
Petrosimonia
Suaeda
Conci et al. (1993)
Agonoscena
cisti (Puton)
M
Pe
A
5–6
L
Pistacia
A. pistaceae
Burckhardt
and Lauterer
TeD
Pe
A
T
2–5
L
Pistacia
A. succincta (De
Geer)
M
Pe
L
L
3
L
Ruta
Lauterer et al. (1998),
Souliotis and
Tsourgianni (2000)
Tokmakoglu (1973),
Mohammed and Sheet
(1989) (as targionii),
Souliotis and
Tsourgianni (2000),
Mehrnejad (2002),
Mehrnejad and
Copland (2005, 2006a,
2006b)
Heeger (1856), Douglas
(1878), Boselli (1930),
Ramirez Gomez (1960)
73
TeM/TeD H
Journal of Natural History
C. veaziei (Patch)
L
Gall
Feeding site
Journet (1984),
Hodkinson
(unpublished)
Conci et al. (1993)
Voltinism
Overwintering elsewhere
References
Overwintering on host
Rhinocolinae
Rhinocolini
Host plant(s)
Overwintering stage
Aphalarini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Pe
A
S
M
S
Pistacia
TeM
Pd
E
S
1
S
Acer
M
Pd
EN
S
1
S
Acer
Davatchi (1958), Conci
et al. (1993)
Löw (1880), Gegechkori
(1984), Lauterer (1991),
Rapisarda and Belcari
(1999)
Conci et al. (1993)
TeD
Pe
L
S
1
S
Calligonum
Loginova (1970, 1976)
TeD
Pe
L
S
1
S
Calligonum
Loginova (1970, 1976)
TeD
Pe
L
S
1
S
Calligonum
Loginova (1970, 1976)
TeD
Pe
L
S
1
S
Calligonum
Loginova (1970, 1976)
TeD
Pe
L
S
1
S
Calligonum
Loginova (1970, 1976)
TeD
Pe
L
S
1
S
Calligonum
Loginova (1970, 1976)
TeD
Pe
L
S
.1
S
Calligonum
Loginova (1970)
74 I.D. Hodkinson
M
Es
Lerp former
Feeding site
Gall
Voltinism
Overwintering elsewhere
R. fusca
Burckhardt
Acaerus calligoni
(Baeva)
A. deminutus
(Loginova)
A. luridus
(Loginova)
A. memoratus
(Loginova)
A. tumidulus
(Loginova)
A. turkistanika
(Löw)
Pachypsylloides
aemulus
Loginova
References
Overwintering on host
Pachypsylloidini
A. targionii
Lichtenstein
Rhinocola aceris
(Foerster)
Host plant(s)
Overwintering stage
Rhinocolini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Gall
L
S
.1
S
Es
Calligonum
Loginova (1970)
TeD
Pe
L
S
.1
S
Es
Calligonum
Loginova (1970)
TeD
Pe
L
S
.1
S
Es
Calligonum
Loginova (1970)
TeD
Pe
L
S
.1
S
Es
Calligonum
Loginova (1970)
TeD
Pe
L
S
.1
S
Es
Calligonum
Loginova (1970)
M/TeD
Pd
A
2?
L
R
Populus
Pe
ELA
L
C
S/L
T
Morus
Loginova and Parfentiev
(1958), Gegechkori
(1984), Lauterer
(1993b), Conci et al.
(1993)
Hsieh and Chen (1977)
Pe
Pe
L
A
L
7
1
L
L
P
P
Kydia
Lindera
Mathur (1935, 1975)
Miyatake (1970)
C
Journal of Natural History
Feeding site
Pe
C
Lerp former
Voltinism
TeD
75
TrM
Paurocephala
psylloptera
Crawford
P. russellae Mathur TrM
TeM
Togepsylla
matsumurana
Kuwayama
Overwintering elsewhere
Togepsyllinae
References
Overwintering on host
Paurocephalinae
P. argutus
Loginova
P. errator
Loginova
P. patulus
Loginova
P. pompatus
Loginova
P. reverendus
Loginova
Camarotoscena
speciosa (Flor)
Host plant(s)
Overwintering stage
Pachypsylloidini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Lerp former
Feeding site
Gall
Voltinism
Overwintering elsewhere
Overwintering on host
References
Overwintering stage
Euphyllurinae
Diclidophlebiini Diclidophlebia
eastopi
Vondracek
D. harrisoni
Ossisanya
D. longitarsata
(Brown and
Hodkinson)
D. lucens
(Burckhardt
et al.)
D. nebulosa
(Brown and
Hodkinson)
D. tuxtlaensis
(Conconi)
D. xuani
Messi et al.
Host plant(s)
Plant type
Species
Climate zone
Higher taxon
TrM
Ps
ELA
S
C
S
Triplochiton
Kudler (1968), Osisanya
(1974a, 1974b)
TrM
Ps
ELA
S
C
S
Triplochiton
Osisanya (1974a, 1974b)
TrM
Pe
ELA
S
C
S
Miconia
Brown and Hodkinson
(1988, unpublished)
TrM
Pe
ELA
S
C
S
Miconia
Burckhardt et al. (2005)
TrM
Pe
ELA
L
C
L
Luehea
Brown and Hodkinson
(1988, unpublished)
TrM
Pe
ELA
S
C
S
Conostegia
Conconi (1973)
TrM
Pe
ELA
C
L
Ricinodendron
Aléné et al. (2005a, 2005b)
76 I.D. Hodkinson
Table 1. (Continued.)
A
S
E. pakistanica
Loginova
E. phillyreae
Foerster
TrD
Pe
A
S
M
Pe
A
S
S
Olea
1+
S
Olea
Silvestri (1934), Ramirez
Gomez (1958),
Rapisarda (1990a),
Conci et al. (1993), Del
Bene et al. (1997),
Arambourg and
Chermiti (1986),
Tzanakakis (2003,
2006)
Thakur et al. (1989)
1
S
Olea, Phillyrea, Loureiro Ferreira (1946),
Osmanthus
Ramirez Gomez (1958),
Prophetou and
Tzanakis (1977,1986),
Stavraki (1980),
Lauterer et al. (1986),
Rapisarda (1991),
Conci et al. (1993),
Prophetou (1993,1997),
Del Bene et al. (1997),
Tzanakakis (2003,
2006)
77
3+
Journal of Natural History
Pe
References
Lerp former
Overwintering on host
M
Host plant(s)
Gall
Overwintering stage
Euphyllura
olivina (Costa)
Feeding site
Plant type
Euphyllurini
Voltinism
Species
Overwintering elsewhere
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Ligustrum
Arbutus
Ferris and Hyatt (1923)
Fox Wilson (1924),
Azevedo and Figo
(1979), Bertaux et al.
(1996), Malausa and
Giradet (1997),
Rapisarda (1998),
Hodkinson (1999),
Olivares (2000), Purvis
et al. (2002)
Hodkinson (2007 and
unpublished)
A
S
Ps
A
Pe
EL
S
?
S/St
Pe
ELA
S
up
to 8
S
Eucalyptus
Pe
ELA
S
M
S
Eucalyptus
C/L 1
Lerp former
Feeding site
S
Mustafa (1984, 1989a,
1989b), Mustafa and
Najar (1985)
Konovalova (1976)
Gall
Voltinism
Olea
Pe
TrS/TeM
Overwintering elsewhere
S
Overwintering on host
2–3
M
TeM
Ligustrinia
herculeana
Loginova
Neophyllura arbuti TeM/M
(Schwarz)
Ctenarytainini Ctenarytaina
TrS/M/
eucalypti (Ferris
TeM
and Klyver)
C. peregrina
Hodkinson
References
Overwintering stage
E. straminea
Loginova
Host plant(s)
Plant type
Euphyllurini
Species
Climate zone
Higher taxon
X
78 I.D. Hodkinson
Table 1. (Continued.)
Lerp former
Pe
ELA
S
M
S
Eucalyptus
C. thysanura
(Ferris and
Klyver)
TeM
Pe
ELA
S
3
S
Boronia
Diaphorina citri
Kuwayama
TrS
Pe
ELA
S
8–16
S
Citrus
Hussain and Nath (1927),
Atwal (1962), Mangat
(1966), Catling (1970),
Atwal et al. (1970),
Pande (1971), Bigornia
and Obana (1974),
Shahid and Khan
(1976), Mead (1977),
Lakra et al. (1983), Tsai
and Liu (2000), Liu and
Tsai (2000), Nakata
(2006)
Journal of Natural History
Feeding site
Gall
Voltinism
Overwintering elsewhere
Mansilla et al. (2004),
Costanzi et al. (2003),
Perez Otero et al. (2005)
Mensah and Madden
(1992a, 1992b, 1993a,
1993b, 1994)
Overwintering on host
References
TrS/M
Ctenarytainini C. spatulata
Taylor
Diaphorininae
Diaphorinini
Host plant(s)
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
79
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Feeding site
S
Lerp former
Voltinism
9
Gall
Overwintering elsewhere
Overwintering on host
L
References
Murraya
Beeson (1941), Mathur
(1975)
Boselli (1960) (as putoni),
Rapisarda (1990a)
Lal (1934), Gegechkori
(1984)
Lauterer (1982), Conci
and Tamanini (1990)
Heslop-Harrison (1942),
Loginova (1954),
Nguyen (1970b), Conci
and Tamanini (1990)
Lal (1934), Loginva
(1954), Ramirez Gomez
(1956), Conci and
Tamanini (1990)
Loginova (1968), Conci
and Tamanini (1990)
Rapisarda (1998)
Conci and Tamanini
(1990)
Overwintering stage
Psyllopseini
D. communis
Mathur
D. lycii Loginova
Host plant(s)
Plant type
Diaphorinini
Species
Climate zone
Higher taxon
TrS
Pe
A
M/TeD
Pe
ELA
S
5
Pd
E
S
1–2
L
R
Fraxinus
Lycium
Psyllopsis
TeM
discrepans (Flor)
P. distinguenda
TeM
Edwards
P. fraxini (L.)
TeM
Pd
E
S
2
L
R
Fraxinus
Pd
E
S
1–2
L
R
Fraxinus
TeM
Pd
E
S
1–2
L
Pd
E
S
2
L
Pd
Pd
E
E
S
S
?
2
L
L
P. fraxinicola
(Foerster)
P. machinosus
M/TeD
Loginova
P. meliphila (Löw) M/TeD
P. narzykulovi
TeD
Baeva
Fraxinus
?
Fraxinus
Fraxinus
Fraxinus
80 I.D. Hodkinson
Table 1. (Continued.)
Pd
E
S
2
L
L
Fraxinus
Pd
E
S
2
L
L
Fraxinus
Pe
ELA
L
?
L
El
Baccharis
Ps
ELA
S
M
S
Acacia
Ps
ELA
S
M
S
Acacia
Ps
ELA
S
M
S
Acacia
Hoffman et al. (1975),
Webb (1977), Webb
and Moran (1978)
Journal of Natural History
Gall
Lerp former
Feeding site
Conci and Tamanini
(1990)
Conci and Tamanini
(1990)
Espirito-Santo and
Wilson Fernandez
(1998, 2002)
Conci et al. (1993),
Rapisarda
(1985,1993a),
Rapisarda and Belcari
(1999)
Palmer and Witt (2006)
Voltinism
A. melanocephala
Burckhardt and
Mifsud
A. russellae Webb TrD
and Moran
Overwintering elsewhere
Acizziinae
References
Overwintering on host
Aphalaroidinae
P. repens
TeD
Loginova
P. securicola
TeD
Loginova
TrS
Baccharopelma
baccharidis
(Burckhardt)
TrD/M
Acizzia
acaciaebaileyanae
(Froggatt)
Host plant(s)
Overwintering stage
Psyllopseini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
81
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Overwintering on host
References
Acizziinae
A. uncatoides
(Ferris and
Klyver)
TrD/M
Ps
ELA
S
M
S
Acacia,
Albizzia
S
Morus
Heslop-Harrison (1949a)
(as Neopsyllia indica),
Munro (1965), Koehler
et al. (1966),
Madubunyi (1967),
Madubunyi and
Koehler (1974), Leeper
and Beardsley (1976),
Arzone and Vidano
(1985), Rapisarda
(1993a), Rapisarda and
Belcari (1999)
Chon (1964), Kuwayama
(1971), Waku and Endo
(1987), Arai (1991,
1993)
Anomoneurinae
Anomoneura mori
Schwarz
TeD
Pd
A
S
1
Lerp former
Overwintering stage
Host plant(s)
Gall
Plant type
Feeding site
Climate zone
Voltinism
Species
Overwintering elsewhere
Higher taxon
82 I.D. Hodkinson
Table 1. (Continued.)
Ps
ELA
S
M (8– S
10)
Mimosa
Moxon (1984), Chazeau
(1987), Oka and
Bahgiawati (1988),
Singh (1988), Yasuda
and Tsurumachi (1988),
Takara et al. (1990),
Rauf et al. (1990), Patil
et al. (1994), Austin
et al. (1996), Ogol and
Spence (1997), Geiger
and Gutierrez (2000)
Willson and Garcia (1992)
H. spinulosa
Muddiman et al.
H. texana
Crawford
Arytaina genistae
(Latreille)
A. africana
Heslop-Harrison
TrS
Ps
ELA
S
,8
S
TrS
Ps
ELA
S
C
S
Prosopis
Donnelly (2002)
TeM/M
Pe
A
S
2
S
Cytisus
Watmough (1968a, 1968b)
M
Pe
E
S
2
S
Adenocarpus
Rapisarda (1988) (as
maculata), Rapisarda
(1990a), Conci et al.
(1993)
83
Leucaena
Journal of Natural History
TrD
Lerp former
Overwintering on host
Heteropsylla
cubana
Crawford
Gall
Overwintering stage
References
Feeding site
Plant type
Ciriacreminae
Arytaininae
Host plant(s)
Voltinism
Species
Overwintering elsewhere
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Lerp former
Feeding site
Gall
Voltinism
Overwintering elsewhere
Overwintering on host
Arytainilla
barbagalloi
Rapisarda
A. cytisi (Puton)
References
Overwintering stage
Arytaininae
Host plant(s)
Plant type
Species
Climate zone
Higher taxon
M
Pe
E
S
1
S
Genista
Rapisarda (1989c), Conci
et al. (1993)
M
Pe
E
ST
1
S
Genista
Calicotome
A. spartiicola
(Šulc)
M
Pe
E
S
1
S
Cytisus
A. spartiophila
(Foerster)
M
Pe
E
S
1
S
Cytisus
Cyamophila
astragalicola
Gegechkori
C. caraganae
(Loginova)
C. caucasica
(Baeva)
C. coluteae Baeva
C. dicora Loginova
C. glycyrrhizae
(Becker)
TeD
C
A
?
1
S
Astragalus
Rapisarda (1988, 1990a,
1990b), Conci et al.
(1993)
Conci and Tamanini
(1985a), Conci et al.
(1993)
Heslop-Harrison (1951),
Watmough (1968a,
1968b)
Gegechkori (1984)
TeD
H
A
?
1
S
Caragana
Gegechkori (1984)
TeD
H
A
?
1
S
Glycyrrhiza
Gegechkori (1984)
TeD
TeD
TeD/M
C
C
H/C
A
A
A
?
S
?
2
1
2–3
S
S
S
Colutea
Astragalus
Glycyrrhiza
Gegechkori (1984)
Naeem and Behdad (1988)
Gegechkori (1984)
84 I.D. Hodkinson
Table 1. (Continued.)
Lerp former
A
?
2
S
Medicago
Gegechkori (1984)
TeD
H/C
A
?
1
S
Hedysarum
Gegechkori (1984)
TeM
H
A
1
S
Anthyllis
M
Pe
EL
S
1
S
Genista
Conci and Tamanini
(1986a, 1989b), Conci
et al. (1993)
Conci et al. (1993)
TeM
Pe
E/L?
S
1
S
M
M
Pe
Pe
EL
L
S
S
1
2
S
S
Chamaecytisu, Conci et al. (1993)
Lemboptropis
Genista
Conci et al. (1993)
Genista
Rapisarda (1988, 1990b),
Conci et al. (1993)
TeM
Pe
AE
S
1
S
Genista
Conci et al. (1993)
M
Pe
E/L
S
1
S
Retama
TeM/M
Pe
A/E
S
1
S
Spartium
Rapisarda (1991, 1992),
Conci et al. (1993)
Rapisarda (1988, 1992),
Conci et al. (1993)
85
H
Journal of Natural History
TeD/M
C
Gall
Feeding site
L. horvathi (Scott)
L. magna
Hodkinson
and Hollis
L. pyrenaea
(Mink)
L. retamae
(Puton)
L. spectabilis
(Flor)
Voltinism
Livilla bimaculata
Hodkinson and
Hollis
L. cognata (Löw)
Overwintering elsewhere
C. medicaginis
(Andrianova)
C. megrelica
(Gegechkori)
C. prohaskai
(Priesner)
References
Overwintering on host
Arytaininae
Host plant(s)
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Lerp former
Feeding site
Gall
Voltinism
Overwintering elsewhere
Psyllinae
Overwintering on host
L. variegata
(Löw)
L. vicina (Löw)
References
Overwintering stage
Arytaininae
Host plant(s)
Plant type
Species
Climate zone
Higher taxon
TeM/M
Pe
E/L?
S
1
S
Laburnum
Conci et al. (1993)
TeM/M
Pe
A
S
1
S
Cytisus
Conci and Tamanini
(1988), Conci et al.
(1993)
Conci et al. (1993)
L. vittipennella
(Reuter)
Psylla alni (L.)
TeM/M
Pe
L
S
1
S
Genista
TeM
Pd
E
S
1
S
Alnus
P. alpina Foerster
P. betulae (L.)
TeM
TeM
Pd
Pd
E
E
S
S
1
1
S
S
Alnus
Betula
P. betulaenanae
Ossianilsson
B
Pd
E
S
1
S
Betula
Lal (1934), Lauterer
(1976), Gegechkori
(1984), Ossiannilsson
(1992)
Conci et al. (1993)
Gegechkori and
Djibladzne (1976),
Gegechkori (1984),
Ossiannilsson (1992)
Ossiannilsson (1992),
Hodkinson and Bird (In
press)
86 I.D. Hodkinson
Table 1. (Continued.)
Lerp former
B
Pd
E
S
1
S
Alnus
M
Pd
E
S
1
S
Alnus
P. floccosa Patch
P. fusca
(Zetterstedt)
TeM
TeM
Pd
Pd
E
E
S
S
1
1
S
S
Alnus
Alnus
P. negundinis
Mally
P. trimaculata
Crawford
Baeopelma
colorata (Löw)
B. foersteri (Flor)
TeM
Pd
E
S
1
S
Acer
Hodkinson and Bird (In
press)
Chiara et al. (1990), Conci
et al. (1993), Rapisarda
and Belcari (1999)
Patch (1909)
Lauterer (1998),
Ossiannilsson (1992),
Conci et al. (1993)
Mally (1894)
TeM
Pd
E
S
1
S
Prunus
Osborn (1922)
M
Pd
E
S
1
S
Ostrya
TeM
Pd
E
S
1
S
Alnus
Rapisarda (1990b), Conci
et al. (1993)
Lal (1934), Lauterer
(1976), Gegechkori
(1984), Ossiannilsson
(1992), Conci et al.
(1993)
Journal of Natural History
Feeding site
Gall
Voltinism
Overwintering elsewhere
P. borealis
(Horvath)
P. cordata
Tamanini
References
Overwintering on host
Psyllinae
Host plant(s)
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
87
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
TeM/TeD Pe
Spanioneura
fonscolombei
(Foerster)
S. caucasica
TeD
Pe
Loginova
Cacopsylla sensu stricto
Cacopsylla mali
TeM
Pd
(Schmidberger)
Lerp former
S
Buxus
Lal (1934), Wilcke (1941),
Nguyen
(1965,1968,1969),
Sampo (1975),
Malenovsky (1999)
Ramirez Gomez (1956),
Conci et al. (1993)
(literature disagrees)
Gegechkori and
Djibladzne (1976)
Gall
EL
References
Feeding site
Pe
Host plant(s)
Voltinism
TeM/M
Overwintering elsewhere
Overwintering on host
Asphagidella buxi
(L.)
Overwintering stage
Psyllinae
Plant type
Species
Climate zone
Higher taxon
1
L
D
EA?
S
1 or
M?
L
Buxus
E
S
1
?
Buxus
E
S
1
S
Malus
C. peregrina
(Foerster)
TeM
Pd
E
S
1
S
Crataegus
C. sorbi (L.)
TeM
Pd
E
S
1
S
Sorbus
Brittain (1922a, 1922b,
1923a, 1923b), Speyer
(1929), Przybylski
(1970), Jonsson (1983),
Gegechkori (1984),
Lauterer (1999)
Missonnier (1956), Sutton
(1983, 1984),
Gegechkori (1984)
Conci et al. (1993)
88 I.D. Hodkinson
Table 1. (Continued.)
Cacopsylla (Hepatopsylla) on Salix
C. ambigua
TeM
Pd
(Foerster)
C. brunneipennis
TeM/B
Pd
(Edwards)
S
Ulmus
Gegechkori (1984),
Lauterer (1999),
Ossiannilsson (1992),
Conci et al. (1993)
E
S
1–2
S
Salix
Lal (1934), Lauterer
(1976, 1999)
Gegechkori (1984), Hill
and Hodkinson (1995),
Lauterer (1999), Hill
et al. (1998)
Gegechkori (1984)
A
L
1
F/S
Salix
TeD
Pd
A
?
1
?
Salix
B
Pd
A
C
1
F/S
Salix
TeD
Pd
A
C
1
?
Salix
B
P/Cd
A
L
1
F/S
Salix
TeM
Pd
E
1
S
Salix
S
Ossiannilsson (1992),
Lauterer (1999)
Gegechkori (1984)
Hodkinson (1997),
Hodkinson and Bird (In
press)
Lauterer (1999)
89
C. intermedia
(Löw)
1
Journal of Natural History
C. compar
(Loginova)
C. elegantula
(Zetterstedt)
C. fraterna
(Gegechkori)
C. groenlandica
(Šulc)
Lerp former
S
Gall
E
References
Feeding site
Pd
Host plant(s)
Voltinism
TeM
Overwintering elsewhere
C. ulmi (Foerster)
Overwintering on host
Psyllinae
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Pd
A
C
1
F/S
Salix
C. macleani
(Hodkinson)
C. memor
(Loginova)
C. moscovita
(Andrianova)
B
Pd
A
L
1
F/S
Salix
Conci and Tamanini
(1989a), Conci et al.
(1993)
Hodkinson et al. (1979)
TeD
Pd
A
?
1
?
Salix
Gegechkori (1984)
TeM
Cd
A
L
1
F/S
Salix
C. nigrita
(Zetterstedt)
B
Pd
A
C
1
F/S
Salix
C. palmeni (Löw)
B
Pd/Cd A
L
1
F/S
Salix
C. parvipennis
(Löw)
C. phlebophyllae
(Hodkinson)
C. propinqua
(Schaefer)
TeM
Pd/Cd A
L
1
F/S
Salix
Gegechkori (1984),
Lauterer (1993c, 1999),
Ossiannilsson (1992),
Hill and Hodkinson
(1996)
Gegechkori (1984),
Lauterer (1999),
Ossiannilsson (1992)
Hodkinson et al. (1979),
Ossiannilsson (1992),
Hill and Hodkinson
(1995), Hill et al. (1998)
Ossiannilsson (1992)
B
C
A
L
1
F/S
Salix
Hodkinson et al. (1979)
B
Pd
A
L
1
F/S
Salix
Ossiannilsson (1992), Hill
and Hodkinson (1995),
Hill et al. (1998)
90 I.D. Hodkinson
Lerp former
Feeding site
Gall
Voltinism
References
Overwintering elsewhere
C. iteophila (Löw) TeM
Overwintering on host
Psyllinae
Host plant(s)
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Pd
A
C
C. saliceti
(Foerster)
C. zaecevi (Šulc)
TeM
Pd
A
B
Pd/Cd A
F/S
Salix
C/L 1
F/S
Salix
L
1
F/S
Salix
Lauterer (1999),
Ossiannilsson (1992),
Conci et al. (1993)
Gegechkori (1984), Conci
et al. (1993)
Hodkinson et al. (1979),
Ossiannilsson (1992)
4–7
S
Pyrus
1
S
Sorbus
T
Pd
A
TeM/TeD Pd
E
T
1
S
Hippophae
B
M
B
A
A
E
?
T
S
1
M
1
S
S
S
Ledum
Pyrus
Vaccinium
Pd
Pd
Cd
C
Lauterer (1979),
Gegechkori (1984) (as
C. vasiljevi)
Lauterer (1976, 1999),
Ossiannilson (1992)
Lauterer (1982, 1993a,
1999), Gegechkori
(1984), Conci et al.
(1993)
Ossiannilsson (1992)
Conci et al. (1993)
Lauterer (1999),
Ossiannilsson (1992)
91
C. ledi (Flor)
C. notata (Flor)
C. myrtilli
(Wagner)
1
Journal of Natural History
TeM
References
Lerp former
Overwintering elsewhere
TeM
C. corcontum
(Šulc)
C. hippophaes
(Foerster)
Host plant(s)
Gall
Overwintering stage
C. pulchra
(Zetterstedt)
A
Feeding site
Plant type
Psyllinae
Other Cacopsylla (Hepatopsylla)
C. bidens (Šulc)
TeM
Pd
Voltinism
Species
Overwintering on host
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
S
Host plant(s)
References
Pyrus
Wille (1950),
Bonnemaison and
Missonnier (1955a,
1955b, 1956), Nguyen
(1964, 1967a, 1967b
1970a, 1971, 1972a,
1972b, 1973, 1975),
Wojnarowska
et al. (1960), Nucifora
(1969), Lazarev (1979),
Deronzier (1981, 1984),
Deronzier and Atger
(1980), Atger (1982),
Gegechkori (1984),
Rieux and d’Arcier
(1990), Lyoussoufi et al.
(1988, 1992, 1994),
Stratopoulou and
Kapatos (1995a,
1995b), Souliotis and
Broumas (1998),
Kapatos and
Stratopoulou (1996,
1999), Schaub et al.
(2005)
92 I.D. Hodkinson
2–8
Lerp former
T
Gall
A
Feeding site
Pd
Voltinism
TeM
Overwintering elsewhere
C. pyri (L.)
Overwintering on host
Psyllinae
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
3–5
S
Lerp former
T
Gall
A
Feeding site
Pd
Voltinism
TeM
Overwintering elsewhere
C. pyricola
(Foerster)
Overwintering on host
Psyllinae
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Pyrus
Slingerland (1892), Ross
(1919), Schaefer (1949),
Siddiqui (1949), Wilde
(1962, 1965), Wilde and
Watson (1963), Wong
and Masden (1967),
Rasmy and MacPhee
(1970), Burts (1970),
Oldfield (1970), Radjabi
and Behechti (1975),
McMullen and Jong
(1972, 1976, 1977), Fye
(1983), Mustafa and
Hodgson (1984),
Savinelli and Tetrault
(1984), Krysan (1990),
Krysan and Higbee
(1990), Horton et al.
(1990a, 1990b), Horton,
Higbee, et al. (1994),
Horton et al. (1998), An
et al. (1996)
93
References
Journal of Natural History
Host plant(s)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Overwintering on host
Pd
E
S
TeM
Pd
E
S
C. visci (Curtis)
TeM
Par on E
Pd
C. zetterstedti
(Thomson)
TeM/TeD Pd
C. alaterni
(Foerster)
C. albipes (Flor)
A
M
Pe
ELA
TeM
Pd
A
References
1
S
Rhododendron
Conci et al. (1993)
1
S
Viburnum
S
2–3
S
Viscum,
Loranthus
T
1
S
Hippophae
Gegechkori and
Djibladzne (1976),
Gegechkori (1984),
Lauterer (1999)
Bin (1970), Lauterer
(1999), Hansen and
Hodkinson (2006)
Lauterer (1982, 1993a,
1999), Ossiannilsson
(1992), Conci et al.
(1993)
C
1
S
Crataegus
S
Rhamnus
C
up to
5
1
?
Sorbus
S
Lerp former
Overwintering stage
TeM
Pd
Host plant(s)
Gall
Plant type
C. rhododendri
(Puton)
C. viburni (Löw)
Cacopsylla (Thamnopsylla)
C. affinis (Löw)
TeM
Feeding site
Climate zone
Psyllinae
E
Voltinism
Species
Overwintering elsewhere
Higher taxon
Lauterer (1982, 1999),
Sutton (1984) (as
subferruginea)
Rapisarda (1989a, 1990a),
Conci et al. (1993)
Gegechkori (1984), Conci
et al. (1993)
94 I.D. Hodkinson
Table 1. (Continued.)
Overwintering elsewhere
Voltinism
Feeding site
Psyllinae
C. breviantennata
(Flor)
TeM
Pd
A
?
1–2
S
Sorbus
Gegechkori (1984),
Lauterer (1993c, 1999),
Conci et al. (1993)
Gegechkori (1984)
C. cotoneasteris
(Loginova)
C. crataegi
(Schrank)
TeD
Pd
A
?
1
?
TeM
Pd
A
C
1
S
Crataegus
M
S
Rhamnus
Ramirez Gomez (1956),
Nguyen (1963),
Gegechkori (1984),
Ossiannilsson (1992),
Lauterer (1999)
Rapisarda (1989a)
C. euxina
(Loginova)
C. fasciata
(Horvath)
C. incerta (Baeva)
C. limbata
(Meyer-Dur)
M
Pd
ELA
TeD
Pd
A
C
2
S
Spiraea
Gegechkori (1984)
TeD
TeM
Pd
Pd
A
A
C
C
1
1
S
S
Rhamnus
Rhamnus
Gegechkori (1984)
Conci and Tamanini
(1982, 1988), Conci
et al. (1993)
S
Journal of Natural History
Overwintering stage
References
Lerp former
Plant type
Host plant(s)
Gall
Species
Overwintering on host
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
95
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
A
C. myrthi (Puton)
M
Pe
ELA
C. picta (Foerster) TeM
Pd
C. pruni (Scopoli)
TeM
C. pulchella (Löw) M
Lal (1934), Domenichini
(1967), Lazarev (1972),
Sutton (1983),
Gegechkori (1984),
Lauterer (1999), Conci
et al. (1993), Tedeschi
et al. (2002)
Conci et al. (1993)
C
1
S
Crataegus,
Malus
S
Myrthus
A
C
up to
5
1
S
Malus
Pd
A
C
1
S
Prunus
Pd
A
C
1
S
Cercis
S
Lerp former
Pd
Gall
Overwintering stage
TeM
Feeding site
Plant type
C. melanoneura
(Foerster)
References
Voltinism
Climate zone
Psyllinae
Host plant(s)
Overwintering elsewhere
Species
Overwintering on host
Higher taxon
Harisanov (1966b) (as
costalis), Lauterer
(1999)
Harisanov (1966a),
Gegechkori (1984),
Ossiannilsson (1992),
Lauterer (1999), Conci
et al. (1993), Labonne
and Lichou (2004)
Burckhardt (1999), Conci
et al. (1993), Rapisarda
and Belcari (1999)
96 I.D. Hodkinson
Table 1. (Continued.)
A
C
C. rhamnicola
(Foerster)
TeM
Pd
A
C
Pd
A
C
Pd
E
S
Pyrus
1
S
Rhamnus
1
?
Ribes
Brocher (1926),
Wojnarowska (1962),
Lazarev (1975),
Lauterer (1999),
Ossiannilsson (1992)
Gegechkori (1984),
Lauterer (1999),
Ossiannilsson (1992)
Gegechkori (1984)
S
1
S
Betula
Gegechkori (1984),
Ossiannilsson (1992),
Hodkinson
(unpublished)
E
S
2
L
Diospyros
Ashmead (1881)
ELA
S
C
S
Indigofera
ELA
S
M up
to 11
S
Bauhenia
Grove and Ghosh (1914),
Maxwell-Lefroy (1913),
Mathur (1975)
Mathur (1935, 1975)
R
97
Ps
1
Journal of Natural History
Pd
TrS
References
Lerp former
Overwintering elsewhere
TeM/M
Psylla nr. simlae
Crawford
Host plant(s)
Gall
Overwintering stage
C. pyrisuga
(Foerster)
Other Miscellaneous ‘Psylla’ spp.
Psylla diospyri
TrS
Pd
Ashmead
Psylla isitis
TrS
Pe
Buckton
Feeding site
Plant type
Psyllinae
C. steinbergi
TeD
(Loginova)
Cacopsylla (Chamaepsylla)
C. hartigii (Flor)
TW
Voltinism
Species
Overwintering on host
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Voltinism
Feeding site
Gall
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Overwintering on host
E
T
1
L
Lf
Pe
ELA
L
M
L
Pachypsyllinae
E. ostreoides
Crawford
E. vittatus
Crawford
Celtisapsis japonica
(Miyatake)
C. usabai Miyatake
Pachypsylla
celtidisgemma
Riley
P. celtidisinternis
Mally
TrS
Pe
ELA
L
M
L
TrS
Pe
E
S
5
S
TeM
Pd
E
S
2
S
TeM
TeM
Pd
Pd
E
L
S
B
1
1
S
B/L
TeM
Pd
A
?
1
P. celtidismamma
(Fletcher)
TeM
Pd
A
T
1
Lerp former
Overwintering stage
Pd
Spondyliaspididae
Euphalerinae
Euphalerus hiuri
TeM
Miyatake
E. nidifex Schwarz TrS
Overwintering elsewhere
Plant type
Species
Climate zone
Higher taxon
X
El
Host plant(s)
References
Caesalpinia
Miyatake (1973)
Piscidia
Mead (1967), Russell
(1971)
Ferreira et al. (1990)
Lonchocarpus
Cassia
X
Celtis
X
B/El
Celtis
Celtis
B
B
Celtis
L
El
Celtis
Beeson (1941), Mathur
(1935)
Miyatake (1968b, 1980,
1994)
Miyatake (1980, 1994)
Riley (1890), Weiss
(1921), Walton (1960)
Weiss (1921), Walton
(1944), Smith and
Taylor (1953)
Riley (1890), Smith and
Taylor (1953), Heard
and Buchanan (1998)
98 I.D. Hodkinson
Table 1. (Continued.)
Pd
A
T
1
L
El
Celtis
Pd
L
L
1
L
El
Celtis
Pd
A
T
1
L
Pe
ELA
L
C
L
Pe
ELA
L
2–3
L
X
Eucalyptus
X
Celtis
TrS
Pe
ELA
L
3
L
X
Eucalyptus
C. fiscella Taylor
C. maniformis
Taylor
Creiis costatus
Froggatt
TrS
TrS
Pe
Pe
ELA
ELA
L
L
5
4
L
L
X
X
Eucalyptus
Eucalyptus
TrS
Pe
ELA
L
2+
L
X
Eucalyptus
Clark and Dallwitz (1975)
99
C. densitexta
Taylor
Purcell et al. (1997),
Wineriter et al. (2003)
Clark (1962, 1963a,
1963b), Clark and
Dallwitz (1975),
Morgan (1984), Collett
(2001)
White (1968, 1970b,
1970c, 1973), Morgan
(1984), Collett (2001)
Campbell (1992)
Campbell (1992)
Melaleuca
Journal of Natural History
Gall
Lerp former
Feeding site
Riley (1890), Smith and
Taylor (1953)
Riley (1890), Smith and
Taylor (1953)
Riemann (1958)
Voltinism
Overwintering elsewhere
P. celtidisvesicula TeM
Riley
P. venusta
TeM
(Osten-Sacken)
Tetragonocephala TeM
flava Crawford
Spondyliaspidinae Boreioglycaspis
TrS
melaleucae Moore
Cardiaspina
TrS
albitextura Taylor
References
Overwintering on host
Pachypsyllinae
Host plant(s)
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Pe
ELA
L
2+
L
X
Eucalyptus
Moore (1961)
TrS
Pe
ELA
L
2+
L
X
Eucalyptus
G. fuscovena
Moore
G. prepta Moore
Spondyliapspis
occidentalis
Solomon
TrS
Pe
ELA
L
M
L
X
Eucalyptus
Clark and Dallwitz
(1975), Morgan (1984),
Brennan, Hrusa et al.
(2001), Brennan and
Weinbaum (2001a,
2001b, 2001c, 2001d)
Morgan (1984)
TrS
TrS
Pe
Pe
ELA
ELA
L
L
M
M
L
L
X
X
Eucalyptus
Eucalyptus
Clark and Dallwitz (1975)
Solomon (1936)
TrM
Pe
L
S
1
S
Mangifera
Mathur (1935, 1946),
Mani (1948), Singh M
(1959), Singh S (1954,
1960), Prasad (1957),
Chaterjee and Sebastian
(1965), Singh and Misra
(1978), Monobrullah
et al. (1998)
Es
Lerp former
Feeding site
TrS
Apsylla cistellata
(Buckton)
Gall
Voltinism
Spondyliaspidinae Glycaspis baileyi
Moore
G. brimlecombei
Moore
Calophyidae
Apsyllinae
Overwintering elsewhere
Overwintering on host
References
Overwintering stage
Host plant(s)
Plant type
Species
Climate zone
Higher taxon
100 I.D. Hodkinson
Table 1. (Continued.)
Pe
ELA
L
M
L
R
Cedrela
Pintera (1982)
TrM
Pe
ELA
L
M
L
R
Cedrela
Pintera (1982)
TeM
Pd
A
1
S
Phellodendron
TrS
Pe
ELA
L
M
L
P
Tetragastris
M/TeD
Pd
S
1
L
P/R
Cotinus
TeM
Pd
E or
A
L
T
1
L
Konovalova (1963),
Miyatake (1992)
Iglesias (1983), Brown and
Hodkinson (1988,
unpublished)
Gegechkori (1984), Conci
et al. (1996)
Weiss and Nicolay (1918)
TrS/M
TeM
TeM
Pe
Pe
Pe
ELA
A
L
L
S
S
M
1
2
L
L
L
P
TrS
Pe
L
L
3
L
Rhus
Downer et al. (1988)
Miyatake (1992)
Wheeler and Rawlins
(1993)
El
Garuga
Mathur (1935, 1946),
Raman (1987)
Phacopteronidae
Phacopteron
lentiginosum
(Buckton)
101
D
Schinus
Picrasma
Rhus
Journal of Natural History
Gall
TrM
C
Lerp former
Feeding site
C. nigripennis
Riley
C. schini Tuthill
C. shinjii Sasaki
C. triozomima
Schwarz
Voltinism
Calophyinae
Overwintering elsewhere
Mastigimas ernstii
(Schwarz)
Mastigimas
schwarzi
(Tuthill)
Calophya nigra
Kuwayama
C. nigrilineata
Brown and
Hodkinson
C. rhois (Löw)
References
Overwintering on host
Mastigimatinae
Host plant(s)
Overwintering stage
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
E
S
1
L
Ficus
Triozamia lamborni TrM
(Newstead)
Pe
ELA
S
M
S
Antiaris
Boselli (1929a), Ramirez
Gomez (1956),
Gegechkori (1984) (and
as viridis), Rapisarda
(1989b) (as viridis),
Conci et al. (1996b),
Tuncer (2002), Gencer
et al. (2007)
Akanbi (1980)
TeM
Pd
E
L
1–2
L
Firmiana
Ding et al. (1987)
TrM
Pe
ELA
S
M
S/F
Theobroma
TrM
Pe
ELA
S
M
S
Durio
Entwistle (1972),
Kaufmann (1973),
Igboekwe (1983),
Igboekwe and Adenuga
(1983), Messi (1983a,
1983b)
Gadug and Hussein
(1987)
Allocarsidara
malayensis
(Crawford)
Lerp former
Feeding site
Pd
Carsidara limbata
(Enderlein)
Mesohomotoma
tessmannii
(Aulmann)
Gall
Voltinism
M
Homotoma ficus
(L.)
Overwintering elsewhere
Overwintering on host
Carsidaridae
Carsidarinae
References
Overwintering stage
Triozaminae
Host plant(s)
Plant type
Homotomidae
Homotominae
Species
Climate zone
Higher taxon
102 I.D. Hodkinson
Table 1. (Continued.)
C
2
L
Comarum
Lauterer (1982)
TeM
Pd
A
C
2
L
Salix
B. atkasookensis
B
(Hodkinson)
B brassicae
TeM
(Vasil’ev)
B. bohemica (Šulc) TeM
Pd
A
L
1
L
Salix
Gegechkori (1984),
Lauterer (1976),
Ossiannilsson (1992),
Conci et al. (1996)
Hodkinson et al. (1979)
H
A
1
?
Brassica
Gegechkori (1984)
H
A
C
1?
L
Geum
TeM
H
A
C
1
L
Ranunculus
Gegechkori (1984),
Ossiannilsson (1992),
Conci et al. (1996)
Conci and Tamanini
(1991), Conci et al.
(1996)
Journal of Natural History
Feeding site
A
B. bucegica
(Dobreanu and
Manolache)
Lerp former
Voltinism
Hel
?
Gall
Overwintering elsewhere
References
TeM
Bactericera
acutipennis
(Zetterstedt)
B. albiventris
(Foerster)
Overwintering on host
Host plant(s)
Overwintering stage
Triozidae
Triozini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
103
Gall
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Feeding site
Lerp former
Voltinism
Compere (1916), Essig
(1917), Lehrman (1930),
Knowlton and Janes
(1931), Knowlton
(1933, 1934), Knowlton
and Thomas (1934),
Davis (1937), Janes and
Davis (1937), List
(1939a, 1939b) Swenk
and Tate (1940), Pletsch
(1947), Wallis (1946,
1955), Liu and Trumble
(2004, 2005, 2006,
2007), Liu et al. (2006)
Conci et al. (1996),
Mifsud (1997)
Gegechkori (1984),
Ossiannilsson (1992),
Conci et al. (1996)
Bin (1972), Gegechkori
(1984), Ossiannilsson
(1992), Conci et al.
(1996)
Overwintering elsewhere
Solanaceae
Overwintering on host
References
Overwintering stage
Host plant(s)
Plant type
Triozini
Species
Climate zone
Higher taxon
B. cockerelli (Šulc) TeD
C
ELA
L
L/C 3+
L
P/D
B. crithmi (Löw)
M/TeM
H
ELA
L
2+
L
B. curvatinervis
(Foerster)
TeM
Pd
A
C
1?
L
Crithmum,
Ferula
Salix
B. femoralis
(Foerster)
TeM
H
A
C
1–2
L
Alchemilla
104 I.D. Hodkinson
Table 1. (Continued.)
TeM
H
A
TeM
H
A
B. perrissii Puton
B. reuteri (Šulc)
TeM
TeM
H
H
A
A
B. salicivora
(Reuter)
TeM
Pd
A
B. modesta
(Foerster)
B. nigricornis
(Foerster)
1
L
Geum
Conci et al. (1996)
2–3
S
Allium
L
1–3?
L
C
2
L
Sanguisorba,
Poterium
Solanum
Lauterer (1965), Conci
and Tamanini (1991),
Conci et al. (1996)
Lauterer (1991), Conci
et al. (1996)
Heinz and Profft (1939),
Ossiannilsson (1943),
Biase (1983),
Gegechkori (1984),
Lauterer (1991), Conci
et al. (1996)
Lauterer (1982)
Lauterer (1963),
Ossiannilsson (1992)
Gegechkori (1984),
Ossiannilsson (1992),
Conci et al. (1996a,
1996b)
S
?
D
L/C 1
?
2?
S
L
Artemisia
Potentilla
C
L
Salix
?
105
C
Journal of Natural History
LA
Lerp former
G
Gall
TeM
Feeding site
A
References
Voltinism
H
Host plant(s)
Overwintering elsewhere
TeM
B. harrisoni
(Wagner)
B. kratochvili
Vondracek
Overwintering on host
Overwintering stage
Triozini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Lerp former
Feeding site
TeM
Pd
A
C
2?
L
B. tremblayi
(Wagner)
M/TeM
G
A
C
7–10
S
D
Allium
B. trigonica
Hodkinson
TeM
H
A
C
2–3
L
D
Daucus
Phylloplecta
tripunctata
(Fitch)
TeM
Pd/C
A
C
1
S
D
Rubus
P. trisignata
(Löw)
TeM
Pd/C
A
C
1
L
Pe
L
1
L
L
Gall
Voltinism
Gegechkori (1984),
Ossiannilsson (1992),
Conci et al. (1996a,
1996b)
Tremblay (1958, 1961 (as
nigricornis), 1965a,
1965b), Annunziata and
Clemente (1980), Conci
et al. (1996)
Biase (1983), Lauterer
(1993a), Conci et al.
(1996)
Sirrine (1895), Smith
(1911), Felt (1906),
Petersen (1923), Mead
(1966a), Stuart (1991)
Conci and Tamanini
(1984b, 1986b), Conci
et al. (1996)
Mathur (1935), Beeson
(1941)
Overwintering elsewhere
Salix
B. striola (Flor)
Egeirotrioza ceardi TrS
(Bergevin)
Overwintering on host
References
Overwintering stage
Host plant(s)
Plant type
Triozini
Species
Climate zone
Higher taxon
Rubus
El
Populus
106 I.D. Hodkinson
Table 1. (Continued.)
Pe
L
S
1
S
Es
Populus
Mathur (1935, 1975)
E
Pe
L
S
1
St
Es
Populus
Pedata (1998)
TeM
Pe
A
L/S
C
1
L
R
Eleagnus
Miyatake (1978)
TeM
Pe
A
L/S
C
1
L
R
Eleagnus
Miyatake (1978)
TeM
Pe
A
L/S
C
1
L
R
Eleagnus
Miyatake (1978)
TeD
H
A
?
1
?
TeM
H
L
S
1
S
Bupleurum
Lauterer (1979, 1991)
TeM
H
L
S
2
S
Bupleurum
TeM
Pd
E
S
1
L
Lauterer (1965, 1979,
1991)
Sampo (1975), Lauterer
(1982), Ossiannilsson
(1992), McLean (1993,
1994, 1998), Conci et al.
(1996)
Gegechkori (1984)
R
Rhamnus
107
Gall
TrS
Journal of Natural History
Feeding site
Lerp former
Voltinism
Overwintering elsewhere
References
Overwintering on host
E. bifurcata
(Mathur)
E. populi
(Horvath)
Epitrioza
marginata
Miyatake
E. mizuhonica
Kuwayama
E. yasumatsui
Miyatake
Eryngiofaga
babugani
Loginova
E. hungarica
(Klimaszewski)
E. lautereri
Loginova
Trichochermes
walkeri
(Foerster)
Host plant(s)
Overwintering stage
Triozini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Gall
L
M
L
P
Trichilia
Brown and Hodkinson
(1988, unpublished)
Pe
L
L
1
L
El
Psidium
Butignol and Pedrosa
(2003)
H
A
C
1?
L
Achillea
C
?
L
Cirsium
1–4
L
Gegechkori (1984),
Ossiannilsson (1992),
Conci et al. (1996)
Gegechkori and
Djibladzne (1976),
Gegechkori (1984),
Lauterer (1991),
Ossiannilsson (1992)
Essig (1917), Weiss (1917),
Lizer (1918), Weiss and
Dickerson (1921),
Borelli (1920), Miles
(1928), Sampo (1977),
Conci and Tamanini
(1985b), Ramirez
Gomez (1958), de
Meirleire (1971),
Gegechkori (1984),
Conci et al. (1996)
H
A
T. alacris Flor
M
Pe
A
S
R
Laurus
108 I.D. Hodkinson
Feeding site
ELA
TeM
Lerp former
Voltinism
Pe
T. agrophila Löw
Overwintering elsewhere
References
Overwintering on host
TrS
Leuronota
trichiliae Brown
and Hodkinson
Neotrioza tavearesi TrS
Crawford
Trioza sensu lato
T. abdominalis
TeM
Flor
Host plant(s)
Overwintering stage
Triozini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
A
T. apicula
Rapisarda
T. binotata Conci
and Tamanini
T. camphorae
Sasaki
M
Pd/Pe
A
TeM
Pd
A
TeM
Pe
L
L
1
S
D
1?
L
P?
S
1
L
L
1
L
S
C
References
Anthriscus,
Angelica
Ossiannilsson (1992),
Conci et al. (1996) (as
pallida)
Lundblad (1929), Bey
(1931) (both as
viridula), Balachowsky
and Mesnil (1936),
Laska (1964,1974),
Rygg (1977),
Gegechkori (1984),
Ramert and Nehlin
(1989), Ossiannilsson
(1992), Ellis and
Hardman (1992), Conci
et al. (1996),
Kristoffersen and
Anderbrandt (2007)
Rapisarda (1993b)
Daucus
Hippophae
P
Camphora
Conci and Tamanini
(1984c)
Sasaki (1910), Sorin
(1959a)
109
H
1
Host plant(s)
Journal of Natural History
TeM
C
Lerp former
T. apicalis
Foerster
Gall
A
Feeding site
H
Voltinism
TeM
Overwintering elsewhere
T. anthrisci
Burckhardt
Overwintering on host
Overwintering stage
Triozini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Voltinism
Feeding site
Gall
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Overwintering elsewhere
T. centranthi
(Vallot)
TeM
H/T
A or S
(ELA)
C
1 to M L
D
T. cerastii (L.)
T. chenopodii
Reuter
TeM
TeM
C
H/T
A
A
C
1
2–5
S
L
D
D
T. chrysanthemi
Löw
TeM
H
A
C
1
L
P
T. cinnamomi
Boselli
TeM
Pe
ELA
T. cirsii Löw
TeM
H
A
T. diospyri
(Ashmead)
TrS
Pd
E
S
L
1 to M L
C
S
1
L
2+
L
El
R
Lerp former
Overwintering stage
Overwintering on host
Plant type
Triozini
Species
Climate zone
Higher taxon
Host plant(s)
References
André (1878),
Ossiannilsson (1992),
Conci et al. (1996)
Cerastium
Conci et al. (1996)
Chenopodium, Lauterer (1982), Baloch
and Ghaffar (1984),
Atriplex,
Ossiannilsson (1992),
Halimione
Conci et al. (1996)
Chrysanthemum Conci and Tamanini
(1991), Conci et al.
(1996)
Cinnamomum
Miyatake (1969),
Rajapakse and
Kulasekera (1982)
Cirsium
Conci and Tamanini
(1990)
Diospyros
Mead (1966b)
Centranthus,
Valerianella
110 I.D. Hodkinson
Table 1. (Continued.)
M up
to 8
L
Lerp former
L
Gall
ELA
Feeding site
Ps
Voltinism
TrS
Overwintering elsewhere
Overwintering on host
T. erytreae (Del
Guercio)
Overwintering stage
Triozini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Citrus
Moran and Blowers
(1967), Moran (1968a,
1968b), Catling and
Annecke (1968),
Catling (1969a, 1969b,
1970, 1971), van
Vuuren and Moll
(1984), van den Berg
and Villiers (1987),
Samways (1987), van
den Berg and Deacon
(1988), van den Berg
(1990), van den Berg,
Anderson, et al. (1991),
van den Berg, Deacon
and Steenekamp (1991),
van den Berg, Deacon
and Thomas (1991a,
1991b), Messi and
Tamesse (1999),
Tamesse and Messi
(2004)
111
References
Journal of Natural History
Host plant(s)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
L
T. flavipennis
Foerster
T. fletcheri minor
Crawford
TeM
H
A
TrS
Pe
ELA
T. galii
TeM
H
A
T. hirsuta
(Crawford)
TrS
Pe
E
S
T. ilicina (De
Stefani Perez)
M
Pe
L
T. jambolanae
Crawford
TrS
Pe
ELA
Lerp former
ELA
Gall
Pe
Feeding site
Overwintering on host
TrS
Voltinism
Overwintering stage
T. eugeniae
Froggatt
Overwintering elsewhere
Plant type
Triozini
Species
Climate zone
Higher taxon
Host plant(s)
References
Morgan (1984), Downer
et al. (1991), Mead
(1994), Dahlsten et al.
(1995), Young (2003)
Löw (1880), Conci et al.
(1996)
Mathur (1935), Beeson
(1941), Mani (1948),
Das et al. (1988)
Boselli (1929b),
Burckhardt and
Lauterer (2006)
Mathur (1935, 1975),
Beeson (1941), Mani
(1948), Dhiman and
Singh (2003, 2004)
Conci and Tamanini
(1985c), Rapisarda and
Belcari (1999), Conci
et al. (1996)
Raman (1991)
M
L
(3–5+)
P
Syzygium
1
L
P
Aegopodium
C
L
El
Terminalia
S/L
D/R
Galium,
Aperula
2
L
R
Terminalia
L
1
L
P
Quercus
L
6–8
L
El
Syzygium
C
L
C/LL 1+?
112 I.D. Hodkinson
Table 1. (Continued.)
A
T/S
1
L
El
T. laserpitii
Burckhardt and
Lauterer
T. machilicola
Miyatake
T. magnisetosa
Loginova
T. malloticola
(Crawford)
TeM
H
A
1
L
TeM
Pe
L
L
1
L
TeM
Pd
A
?
2
L
TrS
Pe
L
L
2–3
L
Pe
ELA
L
9–11
L
L
1
L
P
C
1?
L
P
?
1
L
T. magnicauda
TrM
Crawford
T. magnoliae
TrS
(Ashmead)
T. munda Foerster TeM
Pe
L
H
A
TeM
H
A
T. nana
Gegechkori
C
?
References
Rhamnus
Machilus
Rapisarda (1989a), Conci
et al. (1996)
Burckhardt and Lauterer
(1982), Conci et al.
(1996)
Miyatake (1968a)
Rhamnus
Gegechkori (1984)
Mallotus
Mathur (1935, 1975),
Beeson (1941), Mani
(1948)
Chang et al. (1995)
Laserpitium
P
El
Diospyros
Magnolia,
Persea
Knautia,
Succisa,
Scabiosa
Valeriana
Mead (1963), Leege (2006)
Gegechkori (1984),
Ossiannilsson (1992),
Conci et al. (1996)
Gegechkori (1984)
113
Gall
Pe
Host plant(s)
Journal of Natural History
Feeding site
M
Lerp former
Voltinism
Overwintering on host
T. kiefferi Giard
Overwintering elsewhere
Overwintering stage
Triozini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
TrS
Pe
L
TrS
Pe
L
TeM
TeM
H
H
A
A
T. remota Foerster TeM
Pd
TeM
T. rhamni
(Schrank)
T. rotundata Flor TeM
(sensu
Burckhardt and
Lauterer)
Lerp former
?
Gall
A
Eleagnus
Lauterer and Janicek
(1990), Lauterer (1993a)
Vaishampayan and
Bahadur (1980)
Mathur (1935, 1975)
Feeding site
Overwintering on host
Pd
References
Voltinism
Overwintering stage
TeM
T. neglecta
Loginova
T. obsoleta
Buckton
T. pitiformis
Mathur
T. proxima Flor
T. rapisardai Conci
and Tamanini
Host plant(s)
Overwintering elsewhere
Plant type
Triozini
Species
Climate zone
Higher taxon
?
2
L
L
3
L
El
Diospyros
L
4
L
P
Mallotus
C
C
1
1
L
L
P
Hieracium
Laserpitium
A
C
1
L
P
Quercus
Pd
A
C
2
L
P
Rhamnus
H
A
C
1
L/St
P
Cardamine
Conci et al. (1996)
Conci and Tamanini
(1984d, 1988), Conci
et al. (1996)
Sorin (1959b), Gegechkori
(1984), Lauterer (1991),
Conci et al. (1996)
Löw (1877), Gegechkori
(1984), Lauterer (1991),
Conci et al. (1996)
Gegechkori (1984), Conci
and Tamanini (1987,
1991), Conci et al.
(1996), Burckhardt and
Lauterer (2002)
114 I.D. Hodkinson
Table 1. (Continued.)
Feeding site
Gall
A
C
1
F
F
T. saxifragae Löw TeM
H/G
A
C
1–2
L
T. schrankii Flor
T. scottii Löw
TeM
TeM
H
Pd/G
A
A
C
C
1
1
L
L
T. senecionis
(Scopoli)
T. soniae
Rapisarda
T. tabebuiae
Santana and
Burckhardt
T. tripteridis
Burckhardt et al.
TeM
H
A
C
1
L
M
Pd
A
C
1
TrR
Pe
L
TeM
H
A
L
C
Host plant(s)
References
Rumex
Sampo (1975),
Gegechkori (1984),
Conci et al. (1996)
Conci and Tamanini
(1986c), Lauterer
(1993a), Conci et al.
(1996)
Conci et al. (1996)
Sampo (1975),
Gegechkori (1984),
Conci et al. (1996)
Gegechkori (1984), Conci
et al. (1996)
Rapisarda (1993b), Conci
et al. (1996)
De Queiroz Santana and
Burckhardt (2001)
Saxifraga
R
Astrania
Berberis
L
P
Senecio,
Adenostyles
Quercus
M
L
R
Tabebuia
1
L
F
Valeriana
115
Burckhardt et al. (1991),
Conci and Tamanini
(1991), Conci et al.
(1996)
Journal of Natural History
Voltinism
H
Lerp former
Overwintering elsewhere
TeM
T. rumicis Löw
Overwintering on host
Overwintering stage
Triozini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
TeD
H
E
T. viridula
(Zetterstedt)
T. vitreoradiata
(Maskell)
Pauropsylla
beesoni Laing
P. depressa
Crawford
TeM
H
A
TeM
Pe
ELA
TrS
Pe
TrS
TrS
P. longispiculata
Mathur
C
1–4
L
Urtica
1
L
Valeriana
?
L
Cirsium
L
2–5
L
Pittosporum
L
L
2
L
El
Litsaea
Pe
L
L
1–2
L
El
Ficus
Pe
L
L
1
L
El
Buchanania
S
C
Lerp former
T. valerianae
Gegechkori
Gall
A
Lal (1934), Zhangeri
(1954), Onillon (1969),
Davis (1973), Sampo
(1975), Gegechkori
(1984), Conci et al.
(1996)
Gegechkori and
Djibladzne (1976),
Gegechkori (1984)
Gegechkori (1984),
Ossiannilsson (1992)
Carter (1949)
Feeding site
H
References
Voltinism
TeM
Host plant(s)
Overwintering elsewhere
T. urticae (L.)
Overwintering on host
Overwintering stage
Pauropsyllini
Plant type
Triozini
Species
Climate zone
Higher taxon
Mathur (1935), Beeson
(1941), Mani (1948)
Mathur (1935), Beeson
(1941), Mani (1948),
Abbas (1967), Negi and
Bisht (1989)
Thenmozhi and
Kandasamy (1992)
116 I.D. Hodkinson
Table 1. (Continued.)
Gall
Pe
L
L
3
L
El
Ficus
Mathur (1935, 1975)
M
Pe
ELA
L
M
L
P
Ficus
Pe
ELA
L
M
L
El
Ficus
Awadallah and Swailem
(1971)
Hill (1982)
Journal of Natural History
Feeding site
Lerp former
Voltinism
TrS
P. udei Rübsaamen TrS
Overwintering elsewhere
References
Overwintering on host
P. purpurescens
Mathur
P. trichaeta Petty
Host plant(s)
Overwintering stage
Pauropsyllini
Species
Plant type
Higher taxon
Climate zone
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 1. (Continued.)
117
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
118 I.D. Hodkinson
The relationship between the development rate of non-diapausing larvae and
temperature is asymptotic for species such as Agonoscena pistaciae, Diaphorina citri,
Cacopsylla pyri and Cacopsylla pyricola within the temperature range 5–35uC
(McMullen and Jong 1977; Kapatos and Stratopoulou 1999; Liu and Tsai 2000;
Nakata 2006; Mehrnejad and Copland 2006b). Development commences at the
lower temperature threshold for development, rises approximately linearly to a
maximum as temperature increases, but then falls back at higher temperatures,
presumably in response to increasing thermal stress. By contrast, other species such
as Psyllopsis fraxini, Mesohomotoma tessmanni and Heteropsylla cubana appear to
show a linear response, although the maximum experimental temperatures tested
(30–32uC) were lower than in the previous examples and thus probably less stressful
(Nguyen 1970a; Messi 1983b; Patil et al. 1994; Geiger and Gutierrez 2000). A similar
difference in response occurs between the development rate of eggs (linear) and
larvae (asymptotic) of C. pyri over an identical temperature range, suggesting a
divergence in thermal sensitivity of their respective development rate at higher
temperatures (Kapatos and Stratopoulou 1999).
Temperature-specific development rates may also differ significantly among
instars, although there appears to be little consistency in the trend. For example, in
Ctenarytaina thysanura and Trioza urticae, at a given temperature, speed of
development is highest in the early instars and progressively slows in the later instars
(Onillon1969; Mensah and Madden 1993b). By contrast, development rates in other
species such as Heteropsylla cubana, Mesohomotoma tessmani, Trioza magnicauda,
Trioza erytreae, Psyllopsis fraxini, Cacopsylla pyricola and Diaphorina citri, appear
highest in the intemediate instars (2–4) (Moran and Blowers 1967; Nguyen 1970b;
An et al.1996; McMullen and Jong 1977; Messi 1983b; Patil et al. 1994; Chang et al.
1995; Geiger and Guttierez 2000; Tsai and Liu 2000; Liu and Tsai 2000).
The lower temperature threshold for larval development is comparatively low
relative to ambient temperatures in tropical/subtropical species such as Heteropsylla
cubana (9.6uC), Trioza erytreae (8.6–9.2uC) and Diaphorina citri (10.9–11.7uC) but
significantly higher than in temperate species such as Strophingia ericae (c. 3uC),
Trioza urticae (,6uC) and Cacopsylla pyricola (2–3uC) (Blowers and Moran 1968;
Hodkinson et al. 1999; Kapatos and Stratopoulou 1999; Liu and Tsai 2000; Geiger
and Gutierrez 2000).
Field development times from hatching egg to emerging adult in nondiapausing tropical/subtropical species including Allocarsidara malayensis,
Mesohomotoma tessmani, Diclidophlebia xuani, Heteropsylla cubana, Trioza
erytreae and Trioza magnicauda typically range between 9.5–23 days (Blowers
and Moran 1968; Messi 1983b; Gadug and Hussein 1987; Patil et al. 1994; Chang
et al. 1995; Tsai and Liu 2000). However, some less typical tropical species, such as
Euphalerus clitoriae on Clitoria may take up to 34 days to complete development
(Junior et al. 2005). By contrast, development times of equivalent warm to cool
temperate species, including Euphyllura olivina, Anomoneura mori, Asphagidella
buxi, Cacopsylla melanoneura, C. pyricola and C. ambigua, typically span around
22–44 days (Lal 1934; Loureiro Ferriera 1946; Kuwayama 1971). Development,
however, can be significantly slower at temperatures just above the developmental
threshold, extending to 190 days in T. urticae at 6uC, 56 days for Psyllopsis fraxini
at 15uC and 47 days for Cacopsylla pyricola at 10uC (Onillon 1969; Nguyen 1970b;
McMullen and Jong 1977).
Journal of Natural History
119
Response to high temperature and drought
Psyllids in tropical/subtropical and desert ecosystems are often exposed to
potentially lethal high temperatures, particularly when such temperatures are
coupled with low humidity to produce a high Saturation Deficit Index (SDI). High
temperatures/low humidity are known to influence strongly life cycle completion in
several psyllid species, where it results in reduced fecundity, increased mortality and
slower rates of development at temperatures above the optimum for the species in
question. It may also limit the distribution of the psyllid within the broader range of
its potential host plant.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Effect on mortality, reproduction and longevity
The link between high temperature, SDI and mortality is particularly welldocumented in Trioza erytreae on Rutaceae, for which models have been produced
to predict population densities from these climatic variables by defining particularly
lethal SDI values (Moran and Blowers 1967; Catling and Annecke 1968; Catling
1969a, 1969b; van Vuuren and Moll 1984; Samways 1987; van den Berg, Anderson
et al. 1991; Tamesse and Messi 2004). Similar links between high temperature and
population crashes have been observed in Heteropsylla cubana (.36uC) and
Glycaspis baileyi on Eucalyptus, notwithstanding that larvae of the latter species
secrete a protective covering or lerp (Moore 1961; Yasuda and Tsurumachi 1988;
Geiger and Gutierrez 2000).
The relationship between SDI and mortality at a given standard high
temperature, however, is not linear. At a low SDI Acizzia russellae on Acacia died
from thermal shock associated with low evaporative cooling of host leaves. Survival
rose to an optimum at a moderate SDI as evaporative cooling became more effective
but then declined as the SDI increased and desiccation became significant (Hoffman
et al. 1975). Trioza hirsuta on Terminalia showed similar optimal survival within the
range 70–90% relative humidity (Dhiman and Singh 2003).
Among temperate species such as Cacopsylla pyricola, egg output per female and
longevity are reduced at temperatures (,35uC) that exceed the optimum of 26.7uC.
Similar suppression of egg production at high summer temperature is found in
Cacopylla pyri (Stratopoulou and Kapatos 1995b; Souliotis and Broumas 1998).
Even in tropical species, such as Diaphorina citri, the optima for oviposition and
development are 25–28uC, with larvae failing to develop at 33uC (Liu and Tsai 2000).
Gall-forming species, despite living within humidity-buffered galls, are not
immune from a high SDI, with species such as Euphalerus ostreoides on
Lonchocarpus suffering high mortality among young larvae before gall formation
(Ferreira et al. 1990). However, some species in cool temperate environments such as
Craspedolepta nebulosa and C. subpunctata appear physiologically capable of
withstanding at least short-term exposure, as older larvae, to high temperature
(40uC), provided high humidity is maintained (Bird and Hodkinson 1999).
Often the precise choice of oviposition site on a plant determines whether or not
a psyllid egg survives a low SDI. Those of Cacopsylla pyricola, for example are less
susceptible to desiccation when laid along the mid vein rather than on the leaf lamina
(Horton 1990a). This is probably linked to differences in the relative availability of
water within the leaf tissue for absorption through the basal pedicel of the psyllid
egg, which is inserted into the plant tissue (White 1968; Conci 2000).
120 I.D. Hodkinson
Effects on distribution
Temperature and humidity may thus affect both the absolute distribution and the
relative breeding success of a psyllid species across its range. Megatrioza species on
Pritchardia in the Hawaiian Islands are confined to altitudes above 425m, where
temperatures are lower and humidity is higher (Uchida and Beardsley 1988). The
range of Trioza erytreae in South Africa is limited by high temperature/SDI and even
within this restricted range populations tend to increase with altitude (Catling 1969b;
Green and Catling 1971; Human and Bedford 1985).
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Positive impact of drought
Not all effects of low water availability and high temperature are negative for
psyllids. Periods of atypically low winter rainfall, for example, produce water stress
in Eucalyptus fasciculosa that is strongly correlated with subsequent population
outbreaks of the psyllid Cardiaspina densitexta (White 1969, 1971). Stress in this
instance leads to increased mobilization of soluble nitrogen within the leaves, which
in turn enhances their suitability for larval development (White 1969, 1971). The
effect is similar to that observed in Cacopsylla pyricola when its host plant is
fertilized with nitrogen (Pfeiffer and Burts 1983, 1984; Daugherty et al. 2007).
Seasonal polymorphisms
Morphological differences
Several species of psyllid display environmentally determined seasonal polymorphisms as part of their life cycle. This has, in the past, resulted in the seasonal forms
being described as distinct species. Such polymorphisms are often closely linked to
diapause and have major implications for life history completion, involving key
differences in performance, life history parameters and dispersal characteristics in
the species concerned. These implications are discussed in detail later in the context
of the factors such as day length and temperature that determine onset and breaking
of diapause. Seasonal polymorphisms occur in the adults of several multivoltine
species within a number of distantly related taxa, including Agonoscena pistaceae on
Pistacia, Cacopyslla pyricola, and C. pyri on Pyrus, Celtisaspis japonica on Celtis,
Bactericera acutipennis on Comarum and Trioza chenopodii on various
Chenopodiaceae (Bonnemaison and Missonnier 1955a; Wong and Madsen 1967;
Oldfield 1970; Nguyen 1972a, 1985; Miyatake 1980; Lauterer 1982; Mustafa and
Hodgson 1984; Rieux and d’Arcier 1990; An et al. 1996; Mehrnejad 2002; Mehrnejad
and Copland 2005). Polymorhism may simply involve marked seasonal differences in
overall size (Acizzia uncatoides and A. acaciaebaileyani on Acacia) or colour (Acizzia
spp. and Bactericera perrissii on Artemisia) among generations (Koehler et al. 1966;
Lauterer 1982; Rapisarda 1993a). More frequently it additionally involves
differences in such things as the relative size, shape and venation of the forewing,
the presence or intensity of forewing colour patterns, the distribution and density of
surface spinules in the forewing cells, minor differences in the shape of the
terminalia, dark or light body colouration, and the relative length of the antenna and
its component segments. In Cacopsylla pyricola, C. pyri and A. pistaceae the morphs
usually consist of a smaller lighter coloured spring form with little or less-intense
wing colour pattern and an autumn form that is larger and darker, with a distinctive
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
121
darker wing colouration or pattern. Morph determination is, however, not absolute
and intermediate generations often display transitional characteristics between the
extremes, or a few autumn forms may be produced even in summer generations
(Mustafa and Hodgson 1984; Nguyen 1985; Rieux and d’Arcier 1990; Mehrnejad
and Copland 2005). There is evidence for C. pyri that different features of the
polymorphism, such as wing pattern or body colour are controlled to different
extents by particular temperature and photoperiod exposures acting during the
larval stages (Nguyen 1972a). In contrast to the aforementioned species, Trioza
chenopodii unusually has its darker autumn form characterized by shorter and
broader wings (Lauterer 1982). The darker wing-patterned autumn morph of
Pachypsylla japonica additionally shows strong sexual dimorphism in wing pattern.
The adaptive significance of this is obscure, but such sexual pattern dimorphism also
occurs sporadically in both other temperate (Livilla pogii on Genista) and tropical
(Euphalerus fossiconis, host unknown) species (Conci and Tamanini 1984a; Brown
and Hodkinson 1988). Other species, such as Crastina loginovae on Tamarix, show
similar strong sexual dimorphism in general body colouration (Conci and Tamanini
1983).
Seasonal colour change in long-lived adults
Long-lived adults of many temperate univoltine species undergo marked colour
changes throughout the season. This may have adaptive significance through
camouflage (Sutton 1983). Changes can occur gradually over several months and are
often associated with a reproductive diapause. In genera such as Cacopsylla and
Psylla, for example, the general body colouration changes gradually from pale
colours such as green or yellow to deep red, brown and black. Adults of Cacopsylla
peregrina, for instance, are bright green on emergence in spring, matching the colour
of the leaves of their host plant, Crataegus monogyna, on which they are initially
found. As the season progresses sexually maturing adults move from the leaves on to
the darker stems as a prelude to oviposition and this is accompanied by a change in
body colouration to a more cryptic brown and red (Sutton 1983).
Consequences of size differences
Temperature-induced size polymorphism resulting from varying developmental rates
has strong implications for life history completion at the edge of psyllid species’
ranges. For example, Craspedolepta nebulosa and C. subpunctata are univoltine
species feeding on Epilobium angustifolium. Both are widely distributed in the
temperate northern hemisphere. C. nebulosa shows developmental flexibility by
reducing its body size and increasing its developmental temperature reaction norm as
the day degrees available for larval development decrease along an altitudinal
transect. C. subpunctata shows no such flexibility and is thus restricted to lower
altitudes, thereby occupying a smaller portion of the host-plant range than C.
nebulosa (Bird and Hodkinson 2005; Hodkinson and Bird 2006b). Atypically high
temperature may also affect psyllid size. Both egg length and wing length of
Heteropsylla cubana on Leucaena in Thailand decreased with rising temperature over
the range 20–30uC and this was thought to be a partial explanation for a population
crash during unseasonably hot weather (Geiger and Gutierrez 2000).
122 I.D. Hodkinson
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Induction of seasonal morphs
The production of different seasonal morphs is primarly controlled by day length,
with temperature often playing a secondary role. For example C. pyricola eggs reared
experimentally under short days (LD 12:12 h) produce autumn/winter-form adults;
those reared under long day length (LD 16:8 or 18:6 h) produce summer forms
(Mustafa and Hodgson 1984). The precise day length inducing the winter form
varies, however, between studies and localities, with winter forms being produced at
day lengths between 11 and 14 h in California and British Columbia respectively
(Wong and Madsen 1967; Oldfield 1970; McMullen and Jong 1976). Day length acts
on the early larval stages, but larvae become progressively less susceptible to a
sudden switch in day length as they develop, becoming insensitive by the fourth or
fifth instar (Mustafa and Hodgson 1984; An et al. 1996). The process at the relatively
high experimental temperatures used appears relatively insensitive to temperature
(An et al. 1996). Seasonal polymorphism in C. pyri and Agonoscena pistaceae
appears to be under similar control, with short day length (LD 12:12), but coupled
with low temperature (15uC) acting on instars 1–3 to produce the autumn/winter
form and long day length linked to higher temperatures (25uC) resulting in spring/
summer forms (Bonnemaison and Missonnier 1955a, 1955b; Nguyen 1972a;
Mehmejad and Copland 2005).
Significance of diapause
For tropical multivoltine psyllids species living in non-seasonal environments, such
as Diaphorina citri in the Philippines, the host plant Citrus spp. remains suitable for
psyllid development throughout the year and life cycle progression is usually
uninterrupted (Bigornia and Obana 1974). However, for species living in seasonal
environments, close phenological synchrony of development with that of the host
plant is a prerequisite for successful life cycle completion. Diapause provides the
timing and synchronization mechanism through which psyllids are able to survive
unfavourable periods, such as extended periods of cold or drought, when their host
plant becomes unfavourable for development. Diapause, which involves a slowing
down or cessation of development, may occur in one or more of the egg, larval and
adult stages, depending on species and circumstances. It is usually controlled by
environmental cues, such as photoperiod and temperature, which signal changes in
the favourability of the external environment that will, when mediated through the
host plant, affect psyllid development. The mechanisms that instigate and control
adult diapause are well understood for a few economically important multivoltine
psyllids such as Cacopsylla pyricola and C. pyri but remain unknown for the vast
majority of species. Interestingly, diapausing adult C. pyricola are more insecticide
tolerant than non-dipausing adults and this has implications for population contol of
economically important species (Unruh and Krysan 1994).
Developmental diapause in eggs and larvae
Egg diapause occurs most frequently in univoltine species associated with deciduous
trees and shrubs in which eggs laid on exposed shoots and branches one year
overwinter, and hatch the following year. Examples include Psyllopsis fraxini on
Fraxinus, Cacopsylla peregrina on Crataegus and Psylla alni on Alnus (Lal 1934;
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
123
Nguyen 1970b; Bonnemaison 1956). Overwintering by diapausing larvae on bare
shoots is less frequent but not unknown; Calophya triozomima, for example,
overwinters at the base of the bud of its host Rhus (Wheeler and Rawlins 1993).
Larval winter diapause is more frequently found in free-living species that overwinter
on evergreen hosts such as Strophingia ericae on Calluna, Livilla magna on Genista
and Asphagidella buxi on Buxus (Nguyen 1968; Conci et al. 1993; Miles et al. 1998;
Butterfield et al. 2001). Larvae cease development in the autumn and recommence in
spring.
This period of suspended development may in some species also embrace a large
part of the previous summer. Several leaf-gall forming species, for example, Trioza
camphorae on Camphora, Trioza cinnamomi on Cinnamomum, T. machilicola on
Machilus, and T. obsoleta on Diospyros hatch in spring and develop to early-stage
larvae before entering diapause: development to adult only recommences the
following spring (Sorin 1959a; Miyatake 1968a, 1969; Vaishampayan and Bahadur
1980). A similar extended larval diapause is found in several Craspedolepta species
including C. nebulosa and C. subpunctata on Epilobium, but in these latter species
there is a larval migration onto the roots or the overwintering shoots of their
herbaceous perennial host (Bird and Hodkinson 1999, 2005). In these examples
diapause serves to retard the production of adults and synchronize the life cycle of
the psyllid with the optimum spring period for oviposition on the host. Summer
larval diapause as in Trioza saxifragae on Saxifraga and species of Acaerus on
Calligonum probably facilitates summer survival in particularly dry habitats
(Loginova 1970, 1976; Lauterer 1993a). By contrast, diapause in T. remota and T.
soniae on Quercus leaves and Trioza kiefferi on Rhamnus, which delay adult
emergence until autumn, probably corresponds with a period when mature host
leaves have become unsuitable for development. Subsequent leaf senescence in
autumn then releases the soluble amino acids required for development through to
overwintering adult (Conci et al. 1996).
Reproductive diapause in adults
Adult reproductive diapause, in which egg development by females is postponed,
again achieves similar ends in different species. In several univoltine species of
Cacopsylla sensu stricto, such as C. mali on Malus, C. sorbi on Sorbus and C.
peregrina on Crataegus, and Psylla species such as P. betulaenanae on Betula and P.
borealis on Alnus, adults emerge in late spring but egg development and oviposition
on stems and branches is delayed until autumn (Brittain 1922a; Sutton 1983;
Hodkinson and Bird in press). A possible advantage of postponed oviposition is that
eggs are not exposed to predation or desiccation throughout the summer, although
this must be offset against the probability of reduced female survival throughout this
period.
Many psyllid species overwinter as adults, either on their host or on shelter
plants (see later), and reproductive diapause ensures that eggs are not matured and
laid until the following spring when the host becomes suitable for larval
development. Thus, species that move onto shelter plants during winter undergo
an extended diapause that delays sexual maturation of eggs until the flight to and the
return from the winter host are completed. In multivoltine species such as Cacopsylla
pyricola, Cacopsylla pyri and Bactericera nigricornis it is the autumn generation that
124 I.D. Hodkinson
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
undergoes this reproductive diapause (Nguyen and Ledoux 1973; Nguyen 1975;
Mustafa and Hodgson 1984; Krysan 1990; Krysan and Higbee 1990; Lyoussoufi
et al. 1994; Horton et al. 1998). Many adult overwintering psyllids, including, for
example, several Cacopsylla species on Salix, Livia junci on Juncus, Aphalara species
on Polygonaceae, Euphyllura phillyreae Foerster on Olea and Togepsylla matsumurana on Lindera are, however, univoltine (Heslop-Harrison 1949b; Miyatake 1970;
Lauterer 1976, 1979; Prophetou and Tzanakakis 1977 (as olivina); Hill and
Hodkinson 1996; Tzanakakis 2003; Del Bene et al. 1997). They emerge in the
previous spring or early summer, necessitating an even longer period of reproductive
diapause to ensure host synchrony the subsequent year. Diapause may also play a
role in survival during summer dry periods. Diaphorina lycii on Sardinia has up to
five generations per annum, but breeding on the host Lycium is concentrated into the
wetter spring and autumn and adults undergo reproductive diapause during the
intervening summer (Rapisarda 1990a).
Control of diapause
Egg and larval diapause
Little is known about the control of diapause in psyllid eggs and larvae. Among the
few species studied, the univoltine Asphagidella buxi on Buxus overwinters in
southern France as a first instar larva beneath the chorion of the egg from which it
has emerged in autumn. September–December represents a period of true diapause
initiated by cues unknown (Nguyen 1968). This is followed by a reactivation phase
that leads up to the moult to second instar in March. Prolonged exposure to
temperatures between 0–10uC for 10–30 days is necessary to break diapause but once
broken the development rate during the reactivation phase is positively correlated
with temperature (Nguyen 1968). Developmental inhibition, acting at different
stages of larval development of Strophingia ericae on Calluna at high or low
altitudes, controls whether the species undergoes a one or two year life cycle. At both
altitudes eggs hatch over an extended summer period. In annual lowland
populations, which overwinter predominantly in instar 3, long days (LD18:6 h)
retard development through instars 1–3, and development through to adult is
delayed until the following spring. Instars 4 and 5 respond positively to elevated
temperature and long days, from mid-winter onwards. In biennial upland
populations development is slower at lower temperatures and larvae overwinter
predominantly in instars 1 and 2. Development continues the following year but is
inhibited by short autumn day length (LD 12:12 h) in instar 5, ensuring synchrony
within the population and adult emergence in spring of the following year (Miles
et al. 1998; Butterfield et al. 2001).
Adult diapause
In seasonally polymorphic multivoltine species, such as C. pyricola, C. pyri and A.
pistaciae, the factors such as photoperiod and temperature that induce the autumn
morphological forms are the ones that simultaneously initiate ovarian diapause in
overwintering female adults (Wong and Madsen 1967; Oldfield 1970; McMullen and
Jong 1976; Mustafa and Hodgson 1984; Mehmejad and Copland 2005). Less is
known about the factors inducing ovarian diapause in spring emerging adults,
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
125
although photoperiod, temperature and possibly host-plant condition are again
likely to be involved. In Euphyllura straminea on Olea a mean temperature of .20uC
rather than photoperiod or host-plant quality is thought to induce summer diapause
(Mustafa and Najar 1985). For overwintering adult female psyllids ovarian diapause
is usually broken at some point during the winter to be followed by a period of
quiescence at low temperature when ovarian development continues to be depressed
but once spring temperatures rise then egg development takes place. Adult longevity
during this late-winter period may, however, decline as temperatures rise (Hill and
Hodkinson 1996).
For species with an extended long ovarian diapause commencing in the previous
spring, such as Euphyllura phillyreae, then a succession of summer, winter and spring
conditions are necessary to terminate diapause (Prophetou and Tzanakakis 1986;
Tzanakakis 2003). A combination of increasing day length coupled with rising
temperature during winter, however, is still necessary to break ovarian diapause in
the univoltine Cacopsylla moscovita overwintering on Salix (Hill and Hodkinson
1996).
For multivoltine species such as C. pyricola and C. pyri the effectiveness of long
photoperiod in breaking female diapause diminishes as the winter progresses and the
importance of rising temperature increases (Nguyen 1964, 1967a, 1967b, 1968, 1975;
Fields and Zwick 1975; McMullen and Jong 1976; Horton et al. 1998). However, in
the latter species exposure to temperatures above 25uC breaks diapause irrespective
of photoperiod. The temperature required to terminate ovarian diapause generally
differs both within and among species and habitats, depending on the characteristic
temperatures that the species normally experience and the relative length of the
winter period.
By contrast with the reproductively suppressed females, newly emerged males of
the autumn diapausing form of C. pyricola have active sperm in the testes and
seminal vesicles, but rates of insemination are depressed at short photoperiod (LD
10:14). Initially, an exposure for 10 days at long photoperiod (LD 16:8) is required to
release sexual activity but the exposure time required decreases as the winter
progresses (Krysan 1990; Krysan and Higbee 1990). Repeated mating and
insemination of females on the overwintering host is evidenced by the presence of
multiple spermatophores (mean 5.3 to 16.5 per female), with each spematophore
representing one copulation (Burts and Fischer 1967; Krysan 1990; Krysan and
Higbee 1990). Psyllids collected from conifers on warm days during late winter show
similar promiscuous tendencies suggesting that, for psyllids, the overwintering
period is far from being a quiescent and relatively unimportant phase of the life
cycle. Mating, however, does not always lead to insemination (Van den Berg,
Deacon and Thomas 1991a).
Mating, before or immediately after adult diapauses, usually ensures that egg
development and maturation are completed ahead of the host plant becoming
suitable for oviposition, provided the temperature is sufficiently high. This applies to
both summer diapausing species such as Euphyllura straminea and winter diapausing
species such as Cacopsylla moscovita, Cacopsylla pyricola and Euphyllura phillyreae
(Prophetou and Tzanakakis 1977, 1986; Mustafa and Najar 1985; Lyoussoufi et al.
1994; Hill and Hodkinson 1996; Horton et al. 1998). Mature eggs of C. moscovita
first developed in the field, for example, 6 weeks before the Salix catkins on which
they were to be laid (Hill and Hodkinson 1996).
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
126 I.D. Hodkinson
The larger size of overwintering morphs of multivoltine psyllid species has
implications for both reproductive performance and dispersal ability of individuals.
There are often major differences in the main parameters of reproduction between
summer and post-diapause winter morphs of the same species, including the length
of the pre-reproductive and oviposition periods, the mean number of eggs produced
per female and adult longevity. For example, optimum mean fecundity in
Agonoscena pistaceae varied between 893 and 1087 eggs per female in summer and
winter morphs respectively (Mehmejad and Copland 2005). The difference is even
more marked in pear psyllids with corresponding figures for C. pyricola of 212
(summer) and 486 (winter) in Canada and 387 and 486 in South Korea (McMullen
and Jong 1977; Butt and Stuart 1986; An et al. 1996). C. pyri exhibits similar
variation (342 and 471) (Nguyen 1970a; Kapatos and Stratopoulou 1996).
The optimum temperature for maximum fecundity may also shift between
generations to match the prevailing ambient temperature. In laboratory experiments,
egg output per female of C. pyricola was optimal at 15.6uC in the winter form but
maximal at between 21.1 and 26.7uC in the summer form (McMullen and Jong
1977). The pre-reproductive period is generally shortest in summer forms, but
longevity is greatest in winter forms. The net result of these variations is that egg
output tends to be maximized at the start of the host-plant growing season. There
may also be behavioural differences between the morphs, with summer forms
showing a strong ovipositional preference for leaves but winter forms preferring
dormant bud-bearing stems (Butt and Stuart 1986).
Development of cold-hardiness
Overwintering psyllids in temperate, montane and boreal habitats are frequently
exposed, often over long periods, to sub-zero temperatures that may potentially
damage or ultimately freeze the body tissues. Freeze tolerance is unknown among
psyllids and survival depends on the ability of individual species, whether in the egg,
larval or adult stage, to resist freezing by lowering the supercooling point (SCP) of
their body tissues. Adult and larval stages may mitigate the effects of low air
temperature to some extent by seeking out overwintering sites beneath a protective
snow blanket or, in the case of adults, on evergreen trees such as conifers (Bird and
Hodkinson 1999).
Eggs
Overwintering eggs, however, are often exposed on tree branches to the full rigour of
winter. Those of Cacopsylla mali in Norway, for example, display a mean SCP that
varies between 228.0uC and 238.8uC, depending on whether or not eggs have been
acclimated at sub-zero (25uC) temperature (Skanland and Sömme 1981). This allows
eggs to avoid freezing, even at the very low winter temperatures recorded. The lowered
SCP is achieved partly by the synthesis of cryoprotectants such as glycerol within the
egg, with highest concentrations found in midwinter (Skanland and Sömme 1981).
Larvae
Recorded mean SCPs for overwintering instars include Strophingia ericae
(range5221.6 to 227.3uC), S. cinereae (223.6 to 223.7uC), Craspedolepta nebulosa
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
127
(221.6 to 223.5uC) and C. subpunctata (221.8 to 223.5uC) (Cannon 1983; Bird
and Hodkinson 1999; Hodkinson et al. 1999). SCP was slightly lower in upland
populations of S. ericae than lowland populations, although there was no
difference between the mean SCP of the S. ericae and S. cinereae where they
occurred together at the same site, despite the latter species being more typically
Mediterranean than its congener. Similarly there was little variation in SCP of
populations of C. nebulosa in lowland UK and Tromsø, northern Norway. SCPs of
all species studied to date, despite their varying evolutionary and geographical
origins, fall within a narrow range, suggesting that larvae may posses attributes
that predispose them to surviving cold. Sap feeding, in particular, ensures that icenucleating food particles are absent from the gut. There is, however, evidence for
all the aforementioned species and for Asphagidella buxi that the SCP should only
be taken to indicate the lower limit of cold-hardiness (Nguyen 1969; Bird and
Hodkinson 1999; Hodkinson et al. 1999). In both long and short time survival
experiments at sub-zero temperatures, mortality accrues above the SCP as
temperatures fall: mortality is related to the period of exposure as well as to
temperature per se. Furthermore, prior acclimation at high sub-zero temperatures
(25 or 210uC) enhances survival of fifth-instar A. buxi larvae at 215uC compared
with controls at 215uC. Survival at low temperatures in species such as A. buxi
may differ among overwintering instars, with later instars performing better.
However, in the aformentioned Strophingia and Craspedolepta species, such
differences were not apparent (Bird and Hodkinson 1999).
Adults
Overwintering adults of A. buxi and Cacopsylla melanoneura are significantly less
cold-hardy than eggs or larvae, as previously discussed. The SCP of C. melanoneura,
for example, varied between 26.8 and 214.7uC when acclimated at up to 11 days at
6uC compared with 211.1 to 215uC when acclimated at 27uC (Nguyen 1969,
Jackson et al. 1990). SCP values for adults of the overwintering form of Cacopsylla
pyricola (218 to 222uC), however, were comparable with larval values cited earlier
and Trioza apicalis adults show high survival (71–87%) when exposed to 218uC for
7 days (Rygg 1977; Horton et al. 1996; Lee et al. 1999). Freeze susceptibility,
however, was markedly increased (SCP changed from 215uC to 22–15uC) when C.
pyricola was placed in contact with surface moisture or bacteria such as
Pseudomonas syringi, which act as ice-nucleating agents (Horton et al.1996; Lee
et al. 1999).
One of the mechanisms by which freeze susceptibility can be lessened is for
overwintering psyllids to reduce freezable body water content before the onset of
winter, as occurs in adult C. melanoneura and Euphyllura straminea (Mustafa 1989b;
Jackson et al. 1990). Psyllids that become active during winter and begin fluid
feeding during temporary warm spells thus run the risk of lowering their resistance to
cold. Even in non-feeding adults, metabolic production of water from stored fat
reserves may again potentially increase cold susceptibility (Jackson et al. 1990; Hill
and Hodkinson 1996).
Psyllids have thus evolved a variety of mechanisms to prevent winter mortality in
cold environments that appear largely independent of phylogeny.
128 I.D. Hodkinson
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Metabolic adaptations
In passing through their life cycles, psyllids incur the metabolic cost of respiration:
energy expended in maintaining body tissue becomes unavailable for growth,
development and reproduction. It is thus advantageous to minimize basal
metabolism during periods of seasonal inactivity or growth cessation, especially
when this is linked to diapause. Metabolic energy expenditure for individual species,
measured as oxygen uptake, is influenced significantly by ambient temperature, body
size and sex (Krawczyk and Migula 1979; Migula et al. 1980).
Adult male psyllids generally tend to be smaller and more active than females
and, within any given species, have a higher metabolic rate per unit body weight.
Similarly across adults of species representing the larger psyllid families Psyllidae
and Triozidae there is a negative relationship between log respiration and log body
mass, indicating that smaller species tend to respire less ‘‘efficiently’’ than larger
species.
There are, however outliers to this general pattern: Rhinocola aceris and Livia
junci (Psyllidae) have much lower rates of metabolism than might be predicted from
their body size, possibly indicating their more sedentary nature (Migula et al. 1980).
When measured over the range 15–25uC an array of species belonging to Aphalara,
Craspedolepta, Rhinocola, Livia, Psyllopsis, Arytaina, Psylla, Cacopsylla, Trioza,
Trichochermes and Bactericera, genera with differing degrees of phylogenetic
relatedness and contrasting life history patterns, all showed significantly increasing
respiration with temperature. The rate of increase, however, varied significantly
among species, resulting in considerable differences in metabolism among species at
a given ambient temperature.
While the species sample size is small, and the temperature range examined
rather high, there is clear evidence for differences in metabolic rate among individual
species with different life history adaptations (Migula et al. 1980). For example,
widely distributed multivoltine species such as Bactericera nigricornis, Trioza urticae
and Cacopsylla pyri tend to have higher metabolic rates than their univoltine
congeners (Migula et al. 1980). Similarly, Aphalara species tend to have higher
metabolism than the related Craspedolepta, which may reflect differences in their
speed of development following overwintering as adults and larvae respectively.
Among several species overwintering on conifers, such as Aphalara exilis and
Bactericera nigricornis, metabolism is higher by between 20–38% during spring
reproductive activity than in autumn before overwintering (Migula et al. 1980).
Phenological synchrony with host-plant growth and host quality
Under both temperate and tropical conditions a high degree of phenological
synchrony between psyllid and host-plant growth is required for successful life cycle
completion. Among temperate psyllids that overwinter as eggs, such as many
Cacopsylla, Psylla and Psyllopsis species, egg hatch is generally timed to coincide
with bud burst, although the precise mechanisms maintaining this synchrony are
poorly understood (Nguyen 1970b; Lal 1934). In Cacopsylla mali and C. peregrina,
for example, on Malus and Crataegus respectively, larvae hatch within a few days of
the first buds breaking and move on to the newly developing plant tissues, especially
the flower clusters (Przybylski 1970; Jonsson 1983; Sutton 1984; Lal 1934). This
synchrony is generally strictly maintained among different sites and years
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
129
(Przybylski 1970). The importance of such close synchrony for the psyllids is
emphasized in Crataegus by the rapid decline in the quality of their food resource,
with soluble nitrogen concentrations of shoots and leaf clusters falling from around
0.5 mg N dry weight21 at bud burst to less than 0.1 mg N dry weight21 1 month later
(Sutton 1984). Timing of egg hatch in Homotoma ficus on Ficus appears to determine
subsequent larval abundance (Gençer et al. 2007). Cacopsylla ambigua on Salix, in
contrast to the aformentioned species, is unusual in that eggs hatch well ahead of
bud burst and the larvae remain quiescent beneath the bud scales (Lauterer 1999).
Non-gall-forming univoltine psyllids that overwinter as adults and which already
contain fully developed eggs by early spring, such as Cacopsylla moscovita mentioned
earlier, have the advantage of ovipositing directly onto newly emerging foliage as
soon as it appears. This is a more precise procedure with less natural wastage than
one involving small newly emerged larvae seeking out actively growing tissues. Eggs
laid, however, may then take further time to hatch. Cacopsylla affinis and C.
melanoneura larvae, for example, emerge 7–14 days after C. peregrina on the same
host (Sutton 1984). This direct spring oviposition strategy is employed by a diversity
of species including Gyropsylla ilicis on Ilex, several Aphalara species on
Polygonaceae and is probably best exemplified by Cacopsylla species feeding on
willow (Salix) (Mead 1983; Hodkinson et al. 1979; Hill and Hodkinson 1995; Hill
et al. 1998; Hodkinson 1997).
Many of these latter species develop on female Salix catkins and life cycle
completion within a narrow phenological window is vital. Willow catkins are of
short persistence, developing early in the year before drying out and dehiscing once
the seed has developed. Catkin ‘‘life’’ for six species of willow in northern Alaska
varied between 37–43 days, depending on species. The associated psyllids, Cacopsylla
palmeni and C. phlebophylla developed from egg to adult within 36–41 days, an
exceedingly tight phenological schedule (Hodkinson et al. 1979). Development rates
of psyllid and host are, nevertheless, independently temperature-dependent. Later
studies of C. palmeni, C. propinqua and C. brunneipennis along altitudinal transects in
Norway showed that life cycle completion was determined by the available thermal
budget and its differential effect on psyllid and host development rates (Hill et al.
1995). Each psyllid had a wide distribution along the transect but failed to complete
its life cycle at a characteristic upper altitudinal limit because host growth became
too slow to support development or the psyllid developed too slowly to exploit the
phenological window available. The upper limit for C. palmeni was, however,
significantly higher than that for C. brunnepennis (Hill et al. 1995). Soluble nitrogen
concentrations within catkins declined slightly with increasing altitude but the
decline over time during catkin development at a given altitude was far steeper,
implying a rapidly narrowing time window for psyllid development as catkins aged
(Hill et al. 1998; Hodkinson et al. 2001).
MacLean (1983) extended these ideas to propose a simple temperature-driven
phenological model of psyllid and host-plant development on a wider geographical
scale. The latitudinal distribution of nearly all Alaskan psyllid species is more
restricted than that of their host plants (MacLean and Hodkinson 1980). The model
demonstrates how a psyllid’s northern limit might be set by the failure of its host
plant to develop and grow sufficiently quickly to support life cycle completion within
one season. The southern limit, by contrast is determined by the plant developing too
quickly to permit psyllid development through to maturity. Furthermore, psyllids
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
130 I.D. Hodkinson
may, because of phenological contstraints, exploit different host plants in separate
parts of their range or even exploit different tissues on the same host species. The
Greenland willow psyllid Cacopsylla groenlandica, in the relatively benign climate of
southern Greenland, reproduces on four different Salix species, developing on both
catkins and growing shoot tips (Hodkinson 1997). Further northwards the thermal
budget available for development decreases and the psyllid life cycle can be
completed only on the catkins of one species, Salix glauca, despite other species often
being present.
Among temperate species that overwinter as larvae on evergreen plants, such as
Strophingia ericae on Calluna and Asphagidella buxi on Buxus, phenological
synchrony is probably less important: once diapause is broken: the psyllids simply
recommence development as the host plant resumes growth in the spring
(Hodkinson 1973b; Nguyen 1968). Consequently, by contrast with the aforementioned Salix-feeding species, S. ericae is less dependent on precise host synchrony
and occupies the full altitudinal range of Calluna (Hodkinson et al. 1999).
Several psyllids, particularly Craspedolepta species, feed and overwinter as larvae
on perennial herbaceous plant species that die back each year and pass the winter
with the perennating buds usually present as small rosettes at the soil surface
(Hemicryptophytes) or on tubers (Geophytes). These plants, frequently associated
with xerophytic conditions, often grow rapidly and flower early in the year,
presenting a time-limited opportunity for psyllid exploitation. Overwintering sites of
the associated psyllid larvae are usually at or below the soil surface on the rosette
buds (Craspedolepta nervosa), on fine roots (C. nebulosa and C. subpunctata) or at
the base of old woody stems (C. eas) (Wheeler 1994; Bird and Hodkinson 2005). In
spring late instar larvae migrate to the rapidly growing shoot and quickly complete
development. Larvae of the next generation then migrate back down to the
overwintering site before entering a long larval diapause, usually before midsummer. Close phenological synchrony is thus maintained and long exposure to dry
summer conditions avoided.
Many species of psyllid in tropical and subtropical regions do not undergo an
extended diapause and reproduction is continuous. However, many of their host
species do not produce new shoots and leaves continuously but put out flushes of
new growth in response to variations in ambient temperature and moisture
availability. Flushing cycles may be irregular and non-synchronous within species
and may vary over short geographical distances. Even within individual host plants
there may be marked differences in leaf quality between sun and shade leaves.
Diclidophlebia xuani on Ricinodendron, for example, tends to attain higher
population density on unshaded leaf shoots whereas psyllid galls on Persea tend
to be more numerous on shade leaves (Aléné et al. 2006; Leege 2006). This again
presents synchrony problems in psyllids that need to seek out suitable tissues on
flushing trees on which to complete their life cycle.
The relationship between the flushing cycle of tree and shrub species and the
breeding success of their associated psyllid species has been widely documented.
Good examples include Diclidophlebia harrisoni on Triplochiton, two Phytolyma
species on Milicia, Diaphorina citri and Trioza erytreae on Citrus, D. lycii on Lycium,
Acizzia uncatoides on Albizzia, several Cardiaspina and Glycaspis species on
Eucalyptus, Mesohomotoma tessmanni on Theobroma, Protyora and Diclidophlebia
species on Argyrodendron, and Heteropsylla cubana on Leucaena (Moore 1961; Clark
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
131
1962; White 1967; Catling 1969a; Entwistle 1972; Osisanya 1974a, 1974b; Bigornia
and Obana 1974; Clark and Dallwitz 1975; Leeper and Beardsley 1976; Lakra et al.
1983; Cobbinah 1986; van den Berg and Villiers 1987; Rapisarda 1990a; Basset
1991). Similar examples have also been observed in milder temperate regions
including Ctenarytaina eucalypti on planted Eucalyptus in Europe and Trioza
vitreoradiata on Pittosporum in New Zealand (Carter 1949; Purvis et al. 2002).
In both the aforementioned set of species and those non-tropical multivoltine
species with a winter diapause and several summer generations, such as Cacopsylla
pyricola, successful host-plant usage depends on females correctly discerning the
physiological state and condition of the host-plant tissue at the time of oviposition
(White 1970a; Nguyen 1972b; Moran and Buchan 1975; Butt and Stuart 1986; Stuart
et al. 1989; Horton 1990a, b; Horton and Krysan 1990, 1991; van den Berg,
Anderson, et al. 1991; Mensah and Madden 1992a; Puterka et al. 1993; Luft and
Paine 1997). Selection must favour young growing tissues rather than older mature
leaves and shoots. For certain pest species, such as Cacopsylla pyricola on Pyrus, for
which detailed information is available on the temperature dependence of diapause
termination, pre-reproductive period, oviposition period and development rates,
phenological models can be used to predict psyllid population growth characteristic
and densities for the following summer period (Schaub et al. 2005). This can help
determine when subsequent population control measures should be applied.
Many species of psyllid persist within enclosed or roll-leaf galls on the mature
leaves of their host throughout much of the year. Even in these species, however, the
timing of oviposition and subsequent gall initiation, to correspond with the flushing
cycle of new growth, is important (Raman 1994, 2003). Gall induction usually
involves the active modification of growth in young rapidly growing plant tissue and
plants are thus most susceptible to galling at time of flushing (Kumar et al. 1981).
Subsequent growth of plant tissue through to maturity would normally correspond
with a period of declining quality for psyllid feeding that is overcome by the
metabolic changes induced within the leaf as a result of psyllid feeding that allow
continued development within the gall.
The close correspondence between new tissue growth and gall initiation, and
increasing gall mortality on maturing leaves, has been observed in many psyllids
representing several different families, including Pachypsylla species on Celtis,
Phacopteron lentiginosum on Garuga, Schedotrioza and Glycaspis species on
Eucalyptus, Trioza simplifica on Terminalia, Trioza gigantea on Vaccinium,
Calophya spp and Tainarys sordida on Schinus and Baccharopelma baccaridis on
Baccaris (Smith and Taylor 1953; Walton 1960; Kandasamy 1980; Kandasamy and
Krishnan 1981; Taylor 1987; Espirito-Santo and Wilson Fernandez 1998, 2002; Saiz
and Nunez 2000). In many of these species adult emergence from the gall is timed to
correspond with a predictable seasonal flush of new growth and often involves a
larval or adult diapause designed to maintain host-plant synchrony.
A similar situation pertains in several leaf pit-gall forming species of Trioza
including T. camphorae, T. cinnamomi, T. machilicola, T. obsoleta and T. ilicina in
which oviposition occurs on new flush spring growth and larval diapause
corresponds with summer/winter leaf maturity (Sorin 1959a; Miyatake 1968a,1969;
Vaishampayan and Bahadur 1980; Rapisarda and Belcari 1999). Synchrony and
survival may be further enhanced in species such as Pachypsylla venusta on Celtis
and Trioza tabebuiae on Tabebuia in which the presence of galls prevents galled
132 I.D. Hodkinson
leaves being shed during normal autumn abscission and thus remaining on the host
tree (Smith and Taylor 1953; De Queiroz Santana and Burckhardt 2001). However,
contrary to this trend, Celtis laevigata displays early abscission of leaves with
Pachypsylla galls and this may serve as a plant defence mechanism (Stromgren and
Lanciani 2001). Even in tropical tree species, including Milicia excelsa, staggered loss
of leaves by adult trees allows small populations of associated psyllids (e.g.
Phytolyma fusca) to survive as galls through the dry season when the majority of
trees are leafless (White 1967).
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Development on different hosts
The relative success of a psyllid species in completing its life history may vary
significantly among different potential host species or even among different
provenances within the same host species. Breeding success is determined by the
initial attractiveness of a particular host, the extent to which oviposition occurs and
the survival of these eggs through to adult emergence.
In large-scale host-plant trials psyllids are often found to oviposit on a much
wider range of plant species than those on which they can successfully complete
development (Baloch and Ghaffar 1984). Prosopidopsylla flava, for example,
oviposited on 57 of 58 host species (Leguminosae and Rosaceae) tested but
developed successfully on just four species of Prosopis (Leguminosae) (van Klinken
2000). Similarly, Boreioglycaspis melaleucae laid eggs on 27 out of 43 species of
Myrtaceae but developed successfully on just two or three species of Melaleuca
(Purcell et al. 1997; Wineriter et al. 2003).
Different species of plant within a given host range often vary in their
susceptibility to the associated psyllid species, ranging across a spectrum from highly
susceptible to near resistant. Examples include Glycaspis brimblecombei on
Eucalyptus spp, Ctenarytaina thysanura on Boronia spp Cacopsylla pyricola on
Pyrus and Heteropsylla cubana on Leucaena spp (Williams et al. 1963; Westigard
et al. 1970; Mensah and Madden 1991; Brennan, Hrusa, et al. 2001; Mullen and
Shelton 2003; Pasqualini et al. 2006; Center et al. 2007).
Similar variation in susceptibility also occurs across provenances, cultivars and
varieties within single species of host plant, as in Phytolyma lata on Milicia excelsa,
Acizzia melanocephala on Acacia nilotica, Cacopsylla pyricola and C. pyri on Pyrus
communis, Heteropsylla cubana on Leucaena leucocephala and Bactericera cockerelli
on Lycopersicon (Harris 1973; Chang and Philogene 1976; Butt et al. 1989; Cobbinah
and Wagner 1995; Berrada et al. 1995; Baldassari et al. 1996; Puterka 1997; Mullen
and Shelton 2003; Finlay-Doney and Walter 2005; Liu and Trumble 2006;
Pasqualini et al. 2006; Palmer and Witt 2006). Such variation forms the basis for
selective breeding for host-plant resistance against pest psyllid species or identifying
varieties of invasive weed species susceptible to biological control (e.g. Nguyen and
Messi 1973; Lahiri and Biswas 1980; Palmer and Witt 2006; Center et al. 2006).
Sometimes the within-species variation in plant susceptibility may be as great or
greater than the between-species variation. Asphagidella buxi, for instance, breeds
naturally on Buxus sempervirens var arborescens but not on var rotundifolia, yet it
breeds successfully on B. macrophylla (Nguyen 1965). Comparable apparent
anomalies are found in the host range of Heteropsylla cubana (Mullen and Shelton
2003). Differences in susceptibility may even occur among plants of the same
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
133
provenance. The most striking example occurs in species of Myrtaceae that display
heteroblasty, strong morphological differentiation between juvenile and mature
foliage related to tree age (Brennan, Weinbaum, et al. 2001). Ctenarytaina eucalypti,
for example, oviposits and develops on juvenile shoots of Eucalyptus globulus
whereas C. spatulata develops primarily on the mature foliage (Brennan and
Weinbaum 2001a, 2001b, 2001c).
Variation in psyllid development success among host species and cultivars can
usually be explained by differences in the initial attractiveness of the foliage,
differential oviposition rates, larval survival rates and larval development period.
Diaphorina citri, when tested using four host Citrus species, developed most
successfully on C. paradisi as a result of higher fecundity, faster development time
in the final instar and higher larval survival (Tsai and Liu 2000; Nava et al.
2007). Heteropsylla cubana developed more successfully on Leucaena leucocephala
than on L. collinsii in which slower colonization resulted in 46–63% fewer eggs
being laid, a 67% reduction in larval survival and the production of smaller,
probably less fecund adults (Lapis and Borden 1993a, 1993b). The host preference
hierarchy in Bactericera cockerelli on Lycopersicon is similarly based on rates of
oviposition, development and survival, although these parameters may differ
between native and invasive populations of the psyllid (Liu and Trumble 2004,
2005, 2006, 2007). There are, however, apparently anomalous examples in which a host
plant that is most attractive for oviposition is not the most suitable for larval
development. Trioza erytreae, for example, oviposits preferentially on Citrus limon but
the development period is shorter and the adult size attained is greater on indigenous
Rutaceae such as Vepris and Clausena (Moran 1968a, 1968b).
The precise mechanisms that determine the preference hierarchy of host plants
appear to vary among psyllid species. In Cacopsylla pyricola, when offered three host
plant species in the laboratory, host acceptance for oviposition appeared determined
by interactions among plant species, female egg load and the time for which the
psyllid was deprived of a suitable host (Horton and Krysan 1991). Cues received
during probing and settling released oviposition but egg laying ceased earlier on
lower hierarchy species, suggesting that further cues received during oviposition were
also involved in prolonging egg laying.
Other factors implicated in establishing preferences include host species
phenology (Euphyllura phillyreae), amount of glaucous wax on the leaf surface
(Glycaspis brimblecombei), physical hardness of the terminal shoot (Ctenarytaina
thysanura), leaf colour (Mesohomotoma tessmanni and G. brimblecombei), the
presence of attractive chemicals such as caryophyllene (Heteropsylla cubana) and low
concentrations of repellent chemicals such as phenolics (Cacopsylla pyricola),
terpenoids (Boreioglycaspsis melaleucae) or glucosinolates (undescribed ‘‘Aphalara’’
sp.) (Moran and Brown 1973; Louda and Rodman 1983; Messi 1983a; Ullman and
McLean 1988a; Mensah and Madden 1991; Luft and Paine 1998; Luft et al. 2001;
Brennan and Weinbaum 2001a, 2001c, 2001d; Finlay-Doney and Walter 2005;
Wheeler and Ordung 2005; Prophetou 1997). In Bactericera cockerelli on
Lysopersicon jumping and leaf avoidance behaviour was greatest on the most
resistant cultivars, suggesting active repellence and not just an antixenosis response
(Liu and Trumble 2004). A single gene (Mi-1.2) from wild tomato, Solanum
peruvianum, confers resistance to B. cockerelli in some commercial tomato varieties
(Casteel et al. 2007).
134 I.D. Hodkinson
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Host-plant amelioration, disease transmission and endosymbionts
Host plants, particularly when leaves are mature, provide a low quality source of
soluble nutrients, especially available nitrogen in the form of amino acids, for sapfeeding psyllids. This frequently results, as noted earlier, in lower rates of
reproduction, slower development and reduced longevity on mature versus young
or senescing foliage (Nguyen 1972b). Psyllids, however, often display mechanisms
through which they enhance, ameliorate or partly circumvent the condition of their
mature host plant for larval growth and development (White 1970b).
Feeding involves the injection of saliva and its associated enzymes, such as
amylase, into the host, most frequently into the phloem and its associated tissues or
into leaf mesophyll (Pussard 1939; Williams and Benson 1966). This may or may not
result in wider salivary translocation within the plant. Species of Cardiaspina,
Glycaspis, Creiis and Lasiopsylla on Eucalyptus blakeleyi and E. melliodora induce
localized symptoms of varying severity in phloem tissues that resemble premature
senescence (Woodburn and Lewis 1973). In mesophyll-feeding species such as
Cardiaspina retator on Eucalyptus camaldulensis feeding similarly produces cell
degeneration that resembles senescence and involves the mobilization of lipids,
amino acids and soluble proteins (Crawford and Wilkens 1996).
Some psyllids, including known pest species such as Trioza apicalis (on carrot)
and Bactericera cockerelli (on potato and tomato), induce a wider systemic
phytotoxaemia within their host, resulting in severe growth distortion, cellular
necrosis and yellowing of leaves (Richards and Blood 1933; Eyer and Crawford
1933; Eyer 1937; Sanford 1952; Laska 1964; Markkula and Laurema 1971), which
again resemble senescence, with the associated mobilization of soluble nitrogen and
increasing the availability of nutrients to the psyllid (Laurema 1989). This may, as in
T. apicalis, be accompanied by an increase in leaf monoterpenes concentrations and
result in reduced root growth (Nissinen et al. 2005, 2007). It may also be associated
with the transmission of plant diseases, particularly mycoplasmas (see next section).
Gall formation, which is largely outside the scope of this review, similarly brings
about improvements in host tissue quality for feeding psyllid larvae through the
creation of metabolic sinks within the plant tissue (e.g.Raman 1987; Rajadurai
et al.1990; Mani and Raman 1994; Yang et al. 2006).
Enhanced amelioration may occur when psyllids feed in groups rather than
singly. Cacopsylla pyri on Pyrus communis and Cardiaspina densitexta on Eucalyptus
fasciculosa, for example, showed higher reproduction, greater longevity or enhanced
survival with increasing feeding group density up to an optimum (White 1970b;
Nguyen 1971). Galls of Pachypsylla celtidismamma on Celtis similarly grew larger
when more than one gall was present per leaf (Heard and Buchanan 1998). However,
feeding by some species at high densities, such as Boreioglycaspis melaleucae on
Melaleuca may promote increased leaf abscision and a decline in host quality
(Morath et al. 2006). Similarly, feeding-induced changes in the concentrations of leaf
nutrients, chlorophyll, minerals and phenolics may lead to an ultimate reduction in
food quality for Cacopsylla species on Pyrus (Scutareanu and Loxdale 2006).
Significance of disease transmission
Several psyllids are known vectors of plant diseases and as such are regarded as
noxious pests. However, psyllids often show close association with these pathogens.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
135
When viewed from the psyllids’ perspective, the association may prove highly
beneficial by bringing about pathogen-induced changes in the host plant that makes
it more acceptable or more nutritious for psyllid development (Weintraub and
Beanland 2006). Pathogens may, for example, induce physiological changes
resembling the premature senescence noted previously or produce reduced levels
of defensive chemicals. However, our knowledge of these psyllid–pathogen
relationships is confined to just a few crop plants but similar pathogens occur
widely in wild hosts where the insect–vector relationships remain to be established
(Weintraub and Beanland 2006).
The main plant diseases associated with and transmitted by psyllids (Table 2) are
viruses, and bacteria within three main groups, the liberibacters, the phytoplasmas
(previously known as mycoplasma-type organisms) and fireblight. Several of the
causative agents are taxonomically poorly defined, being identified solely from their
RNA, and are included in the Candidatus category of the bacterial classification. The
disease organisms are initially ingested during psyllid feeding and are then later reinjected back into other plants with the psyllid saliva. Phytoplasmas and
liberibacters in particular are restricted to the phloem sieve tubes and circulate with
the plant sap, making them ideal candidates for transmission by psyllids. Within the
insect they cross the gut wall, multiply in the haemolymph and migrate into the
salivary glands ready for onward transmission in the saliva (Hibino et al. 1971; Chen
et al. 1973; Cousin and Boudon-Padieu 2002; Hung et al. 2004; Weintraub and
Beanland 2006).
Both larvae and adults appear capable of transmitting phytoplasmas (Carraro,
Loi, et al. 1998; Tedeschi and Alma 2004). Some evidence exists for transovariole
transfer of these bacteria between female psyllids and their offspring in psyllids such
as for Phytoplasma prunorum in Cacopsylla pruni (Tedeschi et al. 2006). However,
Liberibacter asiaticum in Diaphorina citri and Phytoplasma mali in Cacopylla
melanoneura, by contrast, do not appear to be vertically transmitted between
generations (Hung et al. 2004; Tedeschi et al. 2006). Interestingly, infection of potato
by a virus provides cross-protection against psyllid yellows phytoplasma transmitted
by Bactericera cockerelli (Staples 1968).
Psyllids may also benefit from a general weakening of the plant caused by sooty
moulds growing on the larval excreta or honeydew deposited on the leaf or shoot
surface. Examples include Cacopsylla pyricola on Pyrus and Ctenarytaina thysanura
on Boronia (Savinelli and Tetrault 1984; Mensah and Madden 1992b).
Significance of endosymbionts
Endosymbiotic bacteria also play a more direct role in the nutrition of psyllids,
which in common with aphids, whiteflies and pseudococcids, support such bacteria
within, or associated with, specialized cells (bacteriocytes) that aggregate to form a
bacteriome within the insect’s body cavity (Tarsia in Curia 1934; Chang and
Musgrave 1969; Waku and Endo 1987; Fukatsu and Nikoh 1998; Thao et al. 2000a).
Phloem sap, on which many psyllids feed, is rich in sugars but poor in amino acids
and it is thought that the endosymbionts synthesize essential amino acids and
vitamins such as riboflavin that then become available to the psyllid (Thao et al.
2000a; Thao et al. 2001).
Organism
Bacteria
Candidatus status
Liberibacter asiaticus
(in Asia and Florida)
Liberibacter africanus
(in Africa)
Liberibacter americanus
(in S. America)
Phytoplasma
Phytoplasma prunorum
Phytoplasma mali
Disease
Citrus Huanglongbing
(HLB)5Greening Disease
Citrus Huanglongbing
(HLB)5Greening Disease
Citrus Huanglongbing
(HLB)5Greening Disease
Peach Yellow Leaf Roll (PYLR)
European Stone Fruit Yellows
(ESFY)5Apricot Chlorotic
Leafroll
Apple Proliferation (AP)
Psyllid vector
Reference
Trioza erytreae
Halbert and Manjunath (2004), Davis et al. (2005),
Das et al. (2007)
Van den Berg et al. (1987), Anon. (1988)
Diaphorina citri
Teixeira et al. (2005)
Cacopsylla pyricola
Purcell and Suslow (1984), Blomquist and Kirkpatrick
(2002)
Carraro, Osler, et al. (1998), Jarausch et al. (2001),
Carraro et al. (2004), Labonne and Lichou (2004),
Delic et al. (2005)
Frisinghelli et al. (2000), Jarausch et al. (2003),
Tedeschi et al. (2002), Tedeschi and Alma (2004)
Jensen et al. (1964), Ullman and MacLean (1988b),
Davies et al. (1992), Giunchedi et al. (1994),
Carraro et al. (1998), Ben Khalifa et al. (2007)
Liu et al. (2007)
Diaphorina citri
Cacopsylla pruni
Cacopsylla picta
C. melanoneura
Cacopsylla pyricola
C. pyri
Phytoplasma pyri (in
Europe and N.America)
Pear Decline (PD)
Phytoplasma (in Taiwan)
Pear Decline (PDTW)
Phytoplasma
Carrot Stolbur
Cacopsylla qianli
C. chinensis
Bactericera trigonica
Wissadula Proliferation (WP)
Paracarsidara dugesii (Löw)
Dabek (1983) (as concolor)
Fireblight of orchard trees
Cacopsylla pyricola
Psyllids generally
Wilde et al. (1971), Hildebrand et al. (2000)
Potato Rugose Stunting Virus
Russelliana solanicola
Tenorio et al. (2003)
Zebra chip disease
Bactericera cockerelli
Munyaneza et al. (2007)
Font et al. (1999), Weintraub and Beanland (2006)
Léclant et al. (1974) (misidentification as B.
nigricornis?)
‘Rickettsia type organism’
Family
Enterobacteriaceae
Erwinia amylovora
Virus
(SB26/29)
Undetermined
136 I.D. Hodkinson
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 2. List of plant diseases transmitted by psyllid vectors.
Journal of Natural History
137
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
The psyllid endosymbionts fall into two main groups, primary (P) and secondary
(S). The P endosymbionts, found within the bacteriocytes, are genetically similar
throughout the psyllids, suggesting that they have colonized the psyllids just once,
and then co-evolved with their hosts (Thao et al. 2000b). They are defined as a single
taxon Candidatus Carsonella ruddii (Thao et al. 2001; Spaulding and von Dohlen
2001). The S endosymbionts, by contrast, are present in cells associated with the
bacteriocytes and appear to be multiply derived, consisting of several distinct groups
within the Eubacteriaceae (Spaulding and von Dohlen 1998; Thao et al. 2001;
Fukatsu and Nikoh 1998). Their function in psyllid nutrition is less clear than for the
P endosymbionts (Thao et al. 2001). Secondary endosymbiont infection levels may
vary greatly among populations, as in Glycaspis brimblecombei, where infection
appears more associated with levels of parasitism than with nutrition (Hansen et al.
2007).
Dispersal
Effective dispersal is a key element in the life history of psyllids irrespective of the
habitat within which they are found. In insects that are capable of flying only limited
distances under their own power, it serves several important purposes. In particular,
it enables a species to track the changing spatial distribution of its host plant and/or
the temporal availability of the food resource that it relies on for breeding success. It
allows psyllids to move between different host-plant species and to exploit non-hostplant species as overwintering sites and it permits species to escape the effects of
strong intraspecific competition and natural enemies. Among economically
important species, such as Trioza erytreae, it expedites rapid colonization of
cultivated Citrus hosts (metapopulation sinks) by psyllids originating on indigenous
host plants within the surrounding area (metapopulation sources) (van den Berg,
Deacon and Steenekamp 1991).
Dispersal distance
There is strong evidence to suggest that, as a group, the psyllids are highly effective
dispersers over both short and long distances, although in almost all cases dispersal
is wind assisted. Dispersing psyllids belonging to the genera Cardiaspina,
Ctenarytaina, Eucalyptolyma, Psylla sensu lato and Bactericera have been taken in
drogue nets towed behind light aircraft in Australia, the Galapagos Islands and the
USA or in kite mounted nets in the Canary Islands (Glick 1939; White 1970a, 1973;
Ashmole and Ashmole 1988; Peck 1994). Psyllids form a major component of the
insect flotsom found on the surface of the sea at sites around the UK coastline and
off the west coast of the USA (Cheng and Birch 1978; Hardy and Cheng 1986).
Species belonging to several genera including Aphalara, Craspedolepta, Livia,
Cacopsylla, Euphalerus sensu lato, Acizzia, Bactericera and Trioza are a common
component of aerial deposition on high altitude snowfields in California and
Tenerife (Papp and Johnson 1979) or early successional volcanic areas in the Azores
(Ashmole et al. 1996). Thirty-seven species have been recorded as vagrants in yellow
water traps in northern Italy and several, including Cacopsylla melanoneura, C.
affinis, Bactericera albiventris and Trioza urticae, overwinter on Pinus in northern
England at a distance of around 13km from the nearest host plant (Hodkinson 1972,
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
138 I.D. Hodkinson
1983c). Trioza apicalis similarly moves up to 1km on to its overwintering shelter
plants (Kristoffersen and Anderbrant 2007).
Tropical forest species may also move considerable distances. The coastal
mangrove-feeding Limbopsylla lagunculariae has been taken inland in central
Panama, many kilometres from the coast (Brown and Hodkinson 1988).
Populations of Boreioglycaspis melaleucae, newly introduced for the biological
control of Melaeuca, spread at a rate of up to 10km per year. Further testimony to
the rapid dispersal powers of psyllids is the time (,10 years) in which Heteropsylla
cubana spread from an origin in Central America to colonize virgin plantings of its
forage legume host Leucaenca leucocephala in the Pacific, Asia, Australia and Africa
(Hodkinson 1988b).
Experimental studies suggest that in species such as Arytainilla spartiophila,
Acizzia russelli and Trioza erytreae females disperse further than males, as evidenced
by an increase in the female: male sex ratio with increasing distance, 90–1500m
depending on species, from the source (Dempster 1968; Webb 1977; van den Berg
and Deacon 1988). However, this would be an ineffectual strategy in species with a
post-dispersal ovarian diapause. For psyllids living on short herbaceous plants, such
as Trioza urticae, the effective dispersal boundary layer, within which most directed
dispersal movements take place, is probably less than 1m (Omole 1980). It is the
individuals that stray above this height that are more likely to be wind dispersed.
Adaptive significance of dispersal
Multivoltine psyllids often show differences in dispersal behaviour among
generations. Summer and autumn emerging adults of Cardiaspina densitexta in
Australia, for example tend to show what White (1970c) calls ‘‘concentrative’’
behaviour. Adults blown out of a given Eucalyptus tree usually fly back into the
same tree, and appear to neglect adjacent trees with significantly lower populations.
This results in some trees supporting high psyllid densities while other nearby trees
have low-density populations. By contrast, long-distance ‘‘dispersive’’ behaviour is a
characteristic feature of the spring generation that has developed at shorter day
length and lower temperatures. This parallels certain multivoltine north temperate
species, such as Cacopsylla pyricola in which the winter morph, with its relatively
longer wings produced under short day length, shows significantly greater dispersive
behaviour than the spring or summer generation (Hodgson and Mustafa 1984;
Horton, Burts, et al. 1994). The actual duration of the flight activity period is similar
in the two morphs but flight frequency is much greater in the former (Horton and
Lewis 1996). The actual rate of dispersal of these winter forms out of pear orchards
tends to be correlated with the rate of leaf fall, lower temperature and density (Fye
1983; Horton, Burts, et al. 1994).
Univoltine species moving onto overwintering shelter plants, by comparison,
exhibit both an autumn and a spring period of peak dispersal as they move to and
from their winter host. Where significant dispersal occurs in summer generations, as
in Arytainilla spartiophila, Acizzia russellae and Trioza erytreae it is usually
associated with interspecific competition arising from high populations and declining
host-plant favourability or increasing pressure from natural enemies (Dempster
1968; Webb 1977; van den Berg, Anderson, et al. 1991). Psyllids disperse to new,
more favourable plants.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
139
Dispersal opens up the opportunity for host-plant alternation in multivoltine
psyllids but this behaviour, so typical of many aphid species, is rare in psyllids, with
just two known examples. Bactericera crithmi on Malta undergoes a winter
generation on Ferula during the period when its normal host Crithmum is dormant
(Mifsud 1997). It moves back to Crithmum in spring. Similarly, the vector of psyllid
yellows disease of potato, Bactericera cockerelli overwinters as source populations
on Lycium and other wild Solanaceae in the warmer southern USA. In spring there is
a general northwards wind-assisted dispersal of adults to establish sink populations
breeding on potato (Solanum) in regions far to the north (Knowlton 1933; Knowlton
and Thomas 1934; Swenk and Tate 1940; Wallis 1946, 1955).
Dispersal allows psyllids to track spatial and temporal changes in the availability
of host-plant tissues suitable for growth and development. Many psyllids require
flushes of young, rapidly growing leaf tissues on which to breed. As these leaves
mature they become unsuitable for psyllid development and the psyllid must seek out
new breeding sites. This is most acute where individuals of particular tree species drop
leaves completely but asynchronously. In some Australian species, such as Cardiaspina
densitexta and C. albitextura, this can result in progressive waves of psyllid outbreak
and decline spreading across the landscape as psyllids track the flushing pattern of
their host Eucalyptus (Morgan 1984). This precludes the need for diapausing stages.
The problem may be particularly acute for tropical rainforest psyllids where individual
host plants are usually sparsely distributed within a highly species diverse tree
community and often flush asynchronously. A high level of dispersive behaviour by
psyllids is necessary constantly to track the spatial and temporal availability of their
food resource (Brown and Hodkinson 1988; Hodkinson and Casson 2000).
Even in temperate regions individual host plants may vary markedly in
suitability between successive years. Many species of Rosaceae, for example, exhibit
biennial or irregular patterns of flowering by individual plants that may affect their
suitability as psyllid hosts (Sutton 1984). It is unsurprising, therefore, that highly
dispersive psyllids like Cacopsylla melanoneura and C. affinis are associated with one
such rosaceous plant species, namely Crataegus (Sutton 1984).
Overwintering on shelter plants
Many temperate species of psyllids are known to disperse to, and overwinter as
adults on, evergreen shelter plants before moving back onto their true host in the
spring (Reuter 1909; McAtee 1915; Hodkinson 1972; Hågvar and Hågvar 1975;
Kristoffersen and Anderbrandt 2007) (Table 1). Such hosts are usually conifers,
primarily species of Pinus, Picea, Abies, Taxus, Tsuga, Cupressus and Juniperus but
may also include thorny evergreen shrubs such as Ulex. In Trioza apicalis there
appears to be a distinct order of preference, with Picea supporting higher
populations than Pinus or Juniperus (Kristoffersen and Anderbrandt 2007). The
period spent on shelter plants, which usually matches periods when the host is
dormant or unfavourable for psyllid development, is normally accompanied by an
ovarian diapause. This adaptive overwintering strategy is found in almost all
temperate psyllid families but is most frequent among species of Aphalara, Livia,
Cacopsylla, Bactericera, Phylloplecta and Trioza. It is, however, recorded more
sporadically in a wider range of genera including Pachypsylla, Calophya,
Camarotoscena, Togepsylla, Ligustrinia, Cyamophila, Livilla and Epitrioza (Table 1).
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
140 I.D. Hodkinson
Psyllids on conifers are easily caught and observed whereas those overwintering
in leaf litter or grass tussocks are much less obvious. Careful studies on some species
such as Bactericera perrisi, Aphalara avicularis, A. exilis and A. longicaudata show
that individuals overwinter both on conifers and in litter, perhaps raising the
question for other species of what actual proportion of the overwintering population
is on conifers (Lauterer 1976, 1982, 1991). Furthermore, several species of Aphalara,
Cacopsylla and Trioza, known to overwinter on conifers, can also be overwintered
successfully in grass tussocks maintained in pots (Heslop-Harrison 1937). These
individuals tend to be much lighter coloured than those spending winter on conifers
(Heslop-Harrison, 1937).
One question that has not been fully answered is whether overwintering psyllids
feed on conifers or other shelter plants. Experiments in our laboratory using Pinus
shoots labelled with C14 and P32 repeatedly failed to provide definitive evidence for
winter feeding by Cacopsylla melanoneura, although the maintenance of body
condition and levels of hydration suggest that some feeding must take place (Jackson
et al. 1990). There is, however, some evidence that overwintering Cacopsylla pyricola
may feed on transitory hosts such as Prunus persica (Ullman and McLean 1988b).
Mate finding and aggregation on host plants
Small dispersive insects, such as psyllids, are faced with the problem of finding
suitable mates, either on their breeding or overwintering host plants. Species are
frequently found as highly-aggregated, mixed-sex colonies on the tissues of their
host. Several are known to emit species- and gender-specific stridulation calls or to
make drumming sound or vibrations with their tarsi on leaf surfaces (Campbell 1964;
Ossiannilsson 1950; Heslop-Harrison 1961; KL Taylor 1985; Carver 1987;
Tishechkin 1989, 2005, 2007; Percy et al. 2006), which are thought to aid mate
location and aggregation. There is also some evidence for chemical mechanisms
leading to aggregation and mate finding. Male Cacopsylla pyricola are attracted to
volatile chemicals emanating from pear shoots with receptive post-diapause females
present or from shoots that have recently supported populations of such females.
The precise nature of the chemical stimulus is unknown and it remains to be
determined whether the chemicals involved originate from the psyllid, the host plant,
or a combination of both (Horton et al. 2007; Horton and Landolt 2007). However,
in some other species, such as Cardiaspina albitextura, host-plant tissues previously
occupied by psyllids appear less favourable for oviposition than formerly
unfrequented sites (Clark 1962, 1963b).
Variation in fecundity among species
It might be predicted that the fecundity of psyllid species is related to their type of
life cycle, with species having larvae living in protective galls or lerps producing
fewer eggs than those living on exposed growing tips. There is, however, surprisingly
little pattern in the fecundity of psyllid species (Table 3). Experimentally measured
fecundity differs widely, even among species within the same genus or family. Where
repeated measures have been made on the same species in different localities or at
different times, such as in Trioza erytreae, Heteropsylla cubana or Diaphorina citri,
mean fecundity may differ by a factor of two or more (Table 3). Furthermore, there
is little to suggest major differences in fecundity related to taxonomic position.
Journal of Natural History
141
Table 3. Fecundity of selected psyllid species on preferred host, illustrating variation in
potential reproductive output across the group. Numbers given are mean and maximum egg
production per female under non-limited experimental conditions.
Family
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Psyllidae
Species
Aphalara
polygoni
Paurocephala
psylloptera
Diclidophlebia
eastopi
Diclidophlebia
harrisoni
Diclidophlebia
xuani
Ctenarytaina
thysanura
Agonoscena
pistaceae
Gyropsylla
spegazziniana
Diaphorina citri
Diaphorina citri
Diaphorina citri
Diaphorina citri
Diaphorina citri
Cacopsylla
melanoneura
Cacopsylla
pyricola
Cacopsylla
pyricola
Cacopsylla
pyricola
Cacopsylla
pyricola
Cacopsylla pyri
Cacopsylla pyri
Cacopsylla pyri
Cacopsylla pyri
Acizzia
uncatoides
Acizzia
uncatoides
Arytaina genistae
Arytainilla
spartiophila
Heteropsylla
cubana
Maximum
Mean
c. 300
640{
Reference
Lauterer (1982)
(as rumicicola)
Hsieh and Chen (1977)
502
Osisanya (1974a)
131
Osisanya (1974a)
532–758
86–92
Aléné et al. (2005a)
180
108*
Mensah and Madden
(1992a, 1993b)
Mehrnejad and Copland
(2005)
Leite and Zanol (2001)
807
630{
748
858
266
210–300
84
Hussain and Nath (1927)
Liu and Tsai (2000)
Tsai and Liu (2000)
Mangat (1966)
Pande (1971)
Domenichini (1967)
664
Burts and Fischer (1967)
445
McMullen and Jong (1977)
665
Rasmy and MacPhee (1970)
893–1087
700
520
116
387–486
An et al. (1996)
47–406
Kapatos and Stratopoulou
(1996)
Lyoussoufi et al. (1988)
Nguyen (1970a)
Nguyen (1973)
Koehler et al. (1966)
2527
471
588
463{
986
962
354
435
93
Madubuny and Koehler
(1974)
Watmough (1968a)
Watmough (1968a)
758
Patil et al. (1994)
142 I.D. Hodkinson
Table 3. (Continued.)
Family
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Spondyliaspididae
Calophyidae
Carsidaridae
Triozidae
Species
Maximum
Mean
Heteropsylla
cubana
Heteropsylla
texana
‘Psylla’ isitis
Euphalerus
clitoriae
Cardiaspina
albitextura
Cardiaspina
albitextura
Boreioglycaspis
melaleucae
Apsylla cistellata
Apsylla cistellata
Carsidara limbata
Allocarsidara
malayensis
Mesohomotoma
tessmanni
Phylloplecta
tripunctata
Trioza erytreae
Trioza erytreae
857
394
Trioza erytreae
Trioza eugeniae
Trioza hirsuta
Trioza
magnicauda
Bactericera
cockerelli
Bactericera
cockerelli
Bactericera
tremblayi
Schedotrioza
multitudinea
Trichochermes
walkeri
Neotrioza
taveresi
Pauropsylla
depressa
100
Reference
Takara et al. (1990)
Donnelly (2002)
828
479{
1148
Grove and Gosh (1914)
Junior et al. (2005)
220
124*
Clark (1962, 1963b)
290
45
Morgan and Taylor (1988)
78
Purcell et al. (1997)
141–150
141
1701
50
61
48{
202
94–164
560
327
787
Monobrullah et al. (1998)
Prasad (1957)
Ding et al. (1987)
Gadug and Hussein (1987)
Igboekwe and Adenuga
(1983)
Petersen (1923)
331
180
827
198
99
692
Moran and Blowers (1967)
Van den Berg, Deacon and
Thomas (1991a)
Van den Berg (1990)
Young (2003)
Dhiman and Singh (2004)
Chang et al. (1995)
1176
439
Pletsch (1947)
1300
318
Knowlton and Janes (1931)
803
431
Tremblay (1965b)
487
GS Taylor (1985, 1987)
201
McLean (1998)
219
Butignol and Pedrosa (2003)
279
.150
Abbas (1967)
Note: occasionally, where means were not calculated in the original paper, they are calculated
as (minimum+maximum)/2 (indicated by*); alternatively, some means (indicated by{) are
calculated directly from raw data given in the original paper; a range of values indicates
recorded differences in means among seasons.
Journal of Natural History
143
Typically mean fecundity per female ranges from 40–50 to over 1000, with a majority
of species lying within the range 200–800. Lowest values (40–50) occur in some but
not all Spondyliaspididae and Carsidaridae but highest values are often found in pest
species of Psyllidae and Triozidae such as Agonoscena pistaciae, Trioza erytreae and
Bactericera cockerelli in which some individual females produce up to 1300 eggs.
Oviposition usually occurs over an extended period with females often maturing
successive batches of eggs (An et al. 1996; Dhiman and Singh 2004). In species such
as Cacopsylla pyricola, Trioza erytreae and Agonoscena pistaciae, repeated mating is
necessary for a female to produce a full egg complement (Burts and Fischer 1967;
van den Berg, Deacon and Thomas 1991a; Mehrnejad 1998; Mehrnejad and
Copland 2006a), emphasizing the importance of continuous mate finding for
successful life cycle completion.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Factors influencing fecundity
Several factors influence fecundity, including temperature, day-length and season,
host condition and crowding. Fecundity, as noted earlier, tends to decline above and
below an optimum temperature, as in Acizzia uncatoides, Agonoscena pistaciae,
Cacopsylla pyri and Bactericera cockerelli (List 1939a; Madubunyi and Koehler
1974; Nguyen 1970a; Mehrnejad and Copland 2006a). In multivoltine species such as
Agonoscena pistaciae, Diclidophlebia xuani, Cacopsylla pyri, Cacopsylla pyricola egg
output per female also varies significantly between seasons, with day length as well as
temperature often an important determining factor (Nguyen 1970a; Kapatos and
Stratapoulou 1996; Mehrnejad and Copland 2005; Aléné et al. 2005b). Moderate
crowding initially enhances fecundity in species such as Trichochermes walkeri,
Cardiaspina albitextura, Cacopsylla pyri and Arytaina genistae but increasing density
beyond the optimum leads to declining fecundity (Watmough 1968a, 1968b; Clark
1963a; Nguyen 1971, 1973; McLean 1998). The presence of eggs also acts as a
deterrent to oviposition in Trioza eugeniae (Luft and Paine 1997).
The previous discussion of life history parameters shows that psyllid species exhibit
considerable variation in their life history characteristics and their adaptive response
to their environment. It is now appropriate to examine the distribution of
characteristics across the Psylloidea and to identify how the life cycle parameters
are combined within the life histories of species both within and among higher taxa.
Analysis of psyllid life history characteristics
Table 1 shows the detailed life history characteristics of 342 psyllid species culled
from the literature. Species are arranged in descending taxonomic sequence by
family, tribe and genus. The key references from which data are drawn are listed. In
the large genus Cacopsylla the Salix-feeding species with similar life histories are
separated from the other species, which themselves are split into subgenera.
Information is provided for each species on the major climatic zone within which it is
found, the functional growth form of its host plant, the overwintering stage(s), the
overwintering site, either on or off the host, voltinism, feeding site on the host and
whether or not the larva forms galls of a particular type or lerps on the host plant. A
full explanation of the various life history categories and their abbreviations as used
in this table are listed in the note of Table 1.
144 I.D. Hodkinson
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Methods of coding and analysis
The categorical life history data in Table 1 were numerically coded as the basis for a
full analysis of the dataset, with each category of each characteristic given a separate
numerical code. Voltinism presented a slight problem of coding and species were
coded as semivoltine, univoltine or multivoltine, depending on the maximum life
history duration. Taxonomic status of the psyllids was coded at the family, tribe and
genus level. The basic objective of the analyses was to test whether there were
recognisable and consistent patterns in the data linking particular life history traits
with specific psyllid groups.
Three separate multivariate analyses were employed to explore the structure of
the dataset using the MINITAB 14 statistical package, namely Multiple
Correspondence Analysis (MCA), Linear Discriminant Analysis (LDA), and
Cluster Analysis (CA) of both the descriptive variables and of the species.
MCA attempted to measure the extent to which different characters correspond
with each other across the dataset and whether particular sets of characteristics
correspond to particular taxonomic groupings within the psyllids. Analyses were
initially conducted using life history characters alone to explore relationships among
these variables and then repeated with psyllid groupings added. These analyses used
the full dataset for all species.
LDA tests whether suggested groupings of species are justified on the basis of the
measured life history variables. This analysis was conducted three times using genus,
tribe and family as the suggested species grouping. The output displays how many
species are correctly or incorrectly allocated objectively to the proposed grouping. A
high level of correct prediction indicates that life history characteristics tend to be
relatively uniform within the group and are good predictors of taxonomic position.
A low level suggests that life history characteristics are highly variable within
proposed groups and thus poor predictors. This may, however, indicate greater
adaptive flexibility as species have evolved differing life cycles to exploit varying
opportunities within different environments. In conducting these analyses it is
necessary to remove monobasic genera and tribes from the dataset, where necessary,
as a single taxon cannot form a group.
CA measured similarity among taxa or among variables within the main dataset
using Euclidean distance: clustering was by average linkage.
Results of analyses
CA of variables (Figure 1) indicates two major groupings of characters and two
outliers. The first grouping links voltinism and overwintering stage, not unexpectedly, to climate. The second links feeding site and overwintering site to plant
functional type. The two outlying characters are gall type and lerp formation. Their
separation from other characters appears to lie in the fact that lerp formation is
largely concentrated within one family, the Spondyliaspididae, while gall instigation/
type is spread broadly but rather haphazardly across taxa.
Overall, the level of correspondence among characters in MCA (Table 4) was
low, with cumulative correspondence across species along the first five axes totalling
only 26%. Introducing psyllid tribe as an additional character actually reduced
correspondence further (19%). Gall and lerp formation were again the main outlying
characters.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
145
Figure 1. Dendrogram illustrating the level of correspondence among life history parameters
measured across psyllid species. Note: similarity is measured by Euclidean distance; clustering
is by average linkage.
The total percentage of species allocated to the correct tribe by LCA was
generally around a weighted mean of 47% (Table 5) but this hid wider variation
among taxa, with one large group the Triozini in particular, dragging the total down
but with other smaller tribes, such as Acizziini, Gyropsyllini, Mastigimatini and
Phytolymini, showing good predictability. However, the correlation between group
size and predictability (%) was non-significant (r50.01, p.0.05). Despite wide
variability among taxa, predictability increased stepwise from the genus (n5302,
40% correct) to the family level (n5322, 50% correct). This suggests that
differentiation and thus discrimination among taxa, related to life history traits,
increases with taxonomic level.
Discussion and conclusions
Within the Psylloidea there is considerable variation in the body form of the larvae
related to phylogeny and one might predict that particular physiognomies are best
Table 4. Results of multiple correspondence analysis across species showing percentage of
cumulative correspondence explained by the first five axes.
Cumulative Variation (%)
Axis
Character set
1
2
3
4
5
Excluding tribe
Including tribe
7
5
13
9
17
12
22
16
26
19
Note: analyses are presented both with and without the inclusion of psyllid tribe as a
character.
146 I.D. Hodkinson
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Table 5. Percentage of species allocated to their correct tribe using linear discriminant analysis
based on life history characteristics. n is the number of species on which the percentage is
based.
Acizzini
Aphalarini
Arytainini
Calophyini
Ciriacremini
Ctenarytainini
Diaphorinini
Diclidophlebiini
Euphalerini
Euphyllurini
Gyropsyllini
Liviini
Mastigimatini
Mesohomotomini
Pachypsyllini
Pachypsylloidini
Paurocephalini
Pauropsyllini
Phytolymini
Psyllini
Psyllopseini
Rhinocolini
Spondyliaspidini
Strophingiini
Triozini
Total
n
% correct
3
27
25
7
3
4
3
7
4
6
2
4
2
2
8
11
3
6
2
72
9
6
11
2
86
315
100
67
16
43
0
25
33
100
0
17
100
75
100
50
13
100
66
33
100
68
89
33
91
100
16
47
adapted to exploiting plants in different ways (Loginova 1982; White and
Hodkinson 1985). Thus, for example, larvae of many Triozidae and Calophyidae
are strongly flattened and rounded and one might expect them to be best adapted for
a sedentary existence, living on the surface of leaves or within open pit galls on the
leaf surface. By contrast, larvae of many Psyllidae and Spondyliaspididae are less
flattened and more robust, with relatively longer legs. They appear best adapted for
a free living existence on expanding shoots or developing within larger enclosed leaf
galls. While many species conform to this expected typology others do not. Thus,
many Triozidae and Calophyidae live on growing shoots or within enclosed galls
while several Psyllidae form pit galls and many Spondyliaspididae live on leaf
surfaces. It is against this phylogenetic history that interpretations and conclusions
regarding psyllid life history adaptations can now be made.
Several important conclusions can be drawn from the analyses of the life history
data. Firstly, and perhaps most significantly, the linkage between phylogenetic
grouping and life history characteristics, while discernible, is not of overriding
significance in determining the type of life history a psyllid undergoes. The
phylogenetic signal, as revealed by MCA is marginally stronger at the family level
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
147
than at the generic level but at best only explains about a quarter of the total
variation observed. Similarly, LDA on average, using life history characteristics as
predictors, only allocates species to the correct genus, tribe or family group with at
best 50% mean accuracy. This again suggests high within group variation in life
history characteristics, even at ascending taxonomic levels. However, within these
average figures there do appear to be individual taxa that display a higher level of
predictability, such as Craspedolepta, Aphalara and Psyllopsis, but this is
compensated for by others such as the large genus Trioza that show low
predictability. Nevetheless, these general findings accord with Danks (2006, p. 9)
who reviewed published data on life history traits across the Insecta. He concluded
that: ‘‘…phylogenetic history of a group or species does determine the core structure
of seasonal responses… but that perhaps more striking is the large number of traits
linked to habitat or its seasonal components that have evolved many times
independently.’’ Among such traits he lists diapause, cold-hardiness, reproductive
pattern, paedogenesis, gall formation etc. (see also Danks (2002, 2005, 2007)). He
concludes (2006, p. 9) that: ‘‘…how the different responses are integrated to provide
coherent, seasonally relevant development trajectories can be understood only by
reference to ecological demands.’’
Extending these conclusions to the psyllids we can observe the manner in which
environmental and host-plant factors overlay the phylogenetic signal to produce the
wide variation observed in psyllid life history traits observed today. Thus, the CA for
life history variables recognizes two major groupings of linked parameters. First,
voltinism and overwintering stage is linked to climatic environment. Second, larval
feeding site and overwintering site is linked to plant functional type. Gall formation,
by contrast appears to have evolved independently on several occasions across
disparate psyllid taxa. Lerp formation is more tightly constrained within the family
Spondyliaspididae. It should be noted, however, that plant functional type is itself
largely a plant response to climate.
It now becomes possible to suggest how environmental and ecological
constraints have led to the observed diversification of psyllid life histories and to
draw together the various threads into a coherent exposition of psyllid life history
adaptation on a global stage. Global temperature and moisture gradients and the
adaptive biology of host plants provide the backdrop against which such psyllid
adaptations have evolved.
Psyllid species living within tropical moist habitats, as typified by lowland
tropical rainforest, probably suffer the least constraint on their development. They
are usually associated with evergreen phanerophytes and chamaephytes, undergo
continuous reproduction and are thus typically multivoltine. However, even in these
benign habitats, host tree species frequently display cycles of flushing of suitable
tissues for larval development, with individual trees at different stages of the flushing
cycle at any one time. In such high diversity forests, a continuous life cycle demands
continuous and effective population dispersal to seek out the sparsely distributed
trees of the host species in an appropriate phenological state for reproduction to
continue (Hodkinson and Casson 2000; Brown and Hodkinson 1988). It may also
select for small and highly dispersive species (Hodkinson and Casson 2000).
In tropical habitats with increasing seasonality of rainfall an increasing
proportion of host tree and shrub species are deciduous, with some host species
being leafless for several months of the year and some tree species showing various
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
148 I.D. Hodkinson
levels of deciduousness between individuals. Under these circumstances adult and/or
egg diapause allows associated psyllids, independent of taxonomic provenance, to
arrest their life cycle and therby align development with the phenology of their host,
but at the expense of a reduced number of generations per year. As tropical habitats
become even more seasonally dry then deciduousness among hosts becomes the
norm and seasonal diapause responses assume even greater significance for psyllid
survival. Here again successful dispersal over a wide area may be necessary to
relocate hosts following periods of inactivity.
As one moves from the wet tropics into the moist temperate regions or at higher
altitudes within the tropics a number of developments and trends in psyllid life cycles
are observed. There is a general reduction in the temperature threshold for
development, development rates are often slower, voltinism is reduced and life
histories become strongly seasonal, with developmental or reproductive diapause
becoming increasingly important for maintaining developmental synchrony with the
host plant. Some species display morphologically distinct seasonal forms. Winter
survival mechanisms, involving increased levels of cold-hardiness, assume increasing
significance. There is also an increase in the availablity and use of herbaceous hostplant species falling within the hemicryptophyte and geophyte functional categories.
Typically the growing tips of these plants overwinter at or below the soil surface,
produce a flush of growth each year and then die back in the autumn. This
necessitates a psyllid overwintering strategy that involves movement onto the winter
bud or root or hibernation away from the host plant and movement back onto the
new foliage in spring.
Psyllids living on temperate phanerophytes and chamaephytes have alternative
life history possibilities. Many psyllid species on deciduous phanerophytes overwinter
as diapausing eggs on the buds or apical shoots where they are exposed to the full
rigours of winter. Hatching is timed to coincide with spring bud burst. Alternatively,
adults may overwinter either on the host or on shelter plants. In both cases a
reproductive diapause is necessary to delay oviposition until the spring growth of the
plant commences. Movement onto and from winter shelter plants necessitates the
development of two phases of pre-reproductive dispersal in autumn and spring.
On evergreen phanerophyte/chamaephyte species many psyllid species overwinter
as diapausing larvae on the green shoots or leaves. On such plants the requirement
for precise phenological synchrony with plant growth is less demanding as larval
growth simply recommences as shoots or leaves resume growth in the spring.
Development time can, as a consequence, be potentially extended beyond one year.
Overwintering as mixed populations of eggs, larvae and adults also becomes feasible
on such plants.
Within temperate regions, where rainfall becomes strongly seasonal, as in areas
of Mediterranean climate, periods of psyllid development and reproduction often
become compressed into the short period of the year, such as early spring, when
temperatures are sufficiently high but not too hot and when rainfall is adequate to
stimulate the flushing of new plant growth. Such compression of life histories, with
long periods of spent inactive as diapausing eggs, larvae or adults, becomes even
more pronounced in psyllid species associated with host plants growing in steppe and
desert environments.
As one moves north from the temperate to the cold boreal regions psyllid faunas
become much less diverse and individual species are usually associated with
Journal of Natural History
149
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
prominent host taxa such as Salix, Alnus and Betula on which they typically undergo
rapid annual life cycles during a short summer growing period. These woody host
plants almost invariably exhibit a low chamaephyte growth form: other herbaceous
hemicryptophytes and geophytes appear less commonly as hosts. Overwintering is
either as eggs on growing shoots or as adults that overwinter in leaf litter around the
base of the plants, often covered by a protective snow layer.
It is reasonable to conclude on the previous evidence that the two prime
environmental variables, temperate and moisture availability, acting within an
ecological context, are either directly, or mediated through the physiognomy and
ecological adaptations of host plants, the major pressures acting on the evolution of
psyllid life histories. Together they have frequently resulted in similar strongly
convergent life histories across taxonomically disparate sets of psyllid species and
divergent life histories among related species. Danks’s (2006) conclusions regarding
the comparatively low importance of group phylogenetic history in determining life
history parameters are well supported by the psyllid data.
References
Abbas SR. 1967. Biology of Pauropsylla depressa Crawford (Homoptera: Psyllidae).
Cecidologica Indica 2:43–52.
Akanbi MO. 1980. Preliminary notes on Triozamia lamborni: a potentially dangerous pest of
Antiaris africana. Entomol Mon Mag. 116:113–115.
Aléné DC, Messi J, Quilici S. 2005a. Contribution a la connaissance de la faune d’arthropodes
associée a Ricinodendron heudelotii Baill. (Euphorbiaceae) au Cameroun. Fruits 60:121–132.
Aléné DC, Messi J, Quilici S. 2005b. Quelques aspects de la biologie de Diclidophlebia xuani
Messi et al. (Hemiptera: Psyllidae), ravageur de Ricinodendron heudelotii Baill.
(Euphorbiaceae) au Cameroun. Fruits 60:279–287.
Aléné DC, Messi J, Quilici S. 2006. Influence of shade on the sensitivity of seedlings of
Ricinodendron heudelotii (Baill.) to attacks of Diclidophlebia xuani Messi et al. in the
natural environment in Cameroon. Fruits 6:273–280.
An JH, Yiem MS, Kim DS. 1996. Effects of photoperiod and temperature on formation and
fecundity of two seasonal forms of Psylla pyricola (Homoptera: Psyllidae). Korean J
Appl Entomol. 35:205–208.
André E. 1878. Mémoire pour servir a l’histoire de la la Trioza centranthi Vallot. Annal Soc
Entomol Fra. 8:77–86.
Annunziata F di, Clemente N. 1980. La psilla della cipollo (Trioza tremblayi Wagn.). Inf
Agrar. 36:10159–10161.
Anon. 1988. Citrus greening bacterium and its vectors Diaphorina citri and Trioza erytreae
(Homoptera Psyllidae). Bull OEPP 18:497–508.
Arai Y. 1991. On the habitat of the adult of the mulberry sucker in winter Anomoneura mori
Schwarz (Homoptera Psyllidae). J Seric Sci Jpn. 60:390–393.
Arai Y. 1993. Observation on the mating behavior of the mulberry sucker, Anomoneura mori
(Homoptera: Psyllidae). J Seric Sci Jpn. 62:297–302.
Arambourg Y, Chermiti B. 1986. Psyllidae. Traite d’entomologie oleicole. Madrid (Spain):
Conseil Oleicole International.
Arzone A, Vidano C. 1985. Il fitomiza Psylla uncatoides su mimosa in Liguria. Inf Fitopatol.
35:31–34.
Ashmead WH. 1881. On the Aphididae of Florida with descriptions of new species (Family
Psyllidae). Can Entomol. 13:220–225.
Ashmole NP, Ashmole MJ. 1988. Insect dispersal on Tenerife, Canary Islands: high altitude
fallout and seaward drift. Arctic Alpine Res. 20:1–12.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
150 I.D. Hodkinson
Ashmole NP, Ormomi P, Ashmole MJ, Martin JL. 1996. The invertebrate fauna of early
successional volcanic habitats in the Azores. Bol Mus Municipal Funchal. 48:5–39.
Atger P. 1982. Le psylle du poirier. Paris (France): CentreTechnical Interprofessionnel des
Fruits et Legumes.
Atwal AS. 1962. Insect pests of citrus in the Punjab. Vol 1. Biology and control of citrus psylla
Diaphorina citri Kuwayama (Hemi: Psyllidae). Punjab Hortic J. 2:104–108.
Atwal AS, Chaudhary JP, Ramzan M. 1970. Studies on the development and field population
of citrus psylla Diaphorina citri. J Res Punjab Agric Univ. 7:333–338.
Austin MT, Williams MJ, Hammond AC, Frank JH, Chambliss CG. 1996. Psyllid population
dynamics and plant resistance of Leucaena selections in Florida. Trop Grasslands.
30:223–228.
Awadallah KT, Swailem SM. 1971. On the bionomics of the sycamore-fig psyllid Pauropsylla
trichaeta. Bull Soc Entomol Egypt. 55:193–199.
Azevedo F, Figo ML. 1979. Ctenarytaina eucalypti Mask (Homoptera, Psyllidae). Boletino
Servicio de Plagas Forestales 5:41–46.
Balachowsky A, Mesnil L. 1936. Les insectes nuisibles aux plantes cultivées. Vol. 2. Paris
(France): Mery.
Baldassari N, Baronio P, Rocchetta G, Salvaterra I. 1996. Research on the development of
Cacopsylla pyri (L.) (Hemiptera Psyllidae) on different mutants and selections of pear.
Boll Ist Entomol ‘‘Guido Grandi’’ Univ Stud Bologna. 50:201–213.
Baloch GM, Ghaffar A. 1984. Natural enemies of Chenopodium spp in Pakistan with notes on
Trioza chenopodii, a promising biocontrol agent. Entomophaga 29:409–414.
Basset Y. 1991. The taxonomic composition of the arthropod fauna associated with an
Australian rainforest tree. Aust J Zool. 39:171–190.
Beeson CFC. 1941. The ecology and control of the forest insects of India and the neighbouring
countries. Dehra Dun (India): Vasant Press.
Ben Khalifa M, Marrakchi M, Fakhfakh H. 2007. Candidatus Phytoplasma pyri infections in
pear orchards in Tunisia. J Plant Pathol. 89:269–272.
Berrada S, Nguyen TX, Lemoine J, Vanpoucke J, Fournier D. 1995. Thirteen pear species and
cultivars evaluated for resistance to Cacopsylla pyri (Homoptera: Psyllidae). Environ
Entomol. 1995:1604–1607.
Bertaux F, Phalip M, Martinez M, Schumacher J-C. 1996. Le psylle de l’eucalyptus, nouveau
ravageur des eucalyptus en France. Phytoma –Defense Veg. 487:48–50.
Bey NS. 1931. Untersuchungen uber Mohrenblattsauger. Z Angew Entomol. 18:175–188.
Biase LMD. 1983. La Trioza (Bactericera) nigricornis Forst e la Trioza trigonica Hod
(Homoptera Psylloidea) nel fucino. Difesa Piante. 2:103–110.
Bigornia AE, Obana SP. 1974. Studies on the population dynamics of Diaphorina citri
(Kuway.) and the flushing rhythm of citrus. Philipp J Plant Ind. 39:37–52.
Bin F. 1970. Lygus viscicola Put. (Miridae) e Psylla visci Curt. (Psyllidae) rincoti del vischio
nuovi per la fauna italiana. Boll Zool Agrar Bachic. 10:133–143.
Bin F. 1972. Presenza in Italia della Trioza femoralis Foerster e notize su due calcidoidei suoi
parassiti. Entomologica, Bari 8:45–53.
Bird JM, Hodkinson ID. 1999. Species at the edge of their range: the significance of the thermal
environment for the distribution of congeneric Craspedolepta species (Sternorrhyncha:
Psylloidea) living on Chamerion angustifolium (Onagraceae). Eur J Entomol. 96:103–109.
Bird JM, Hodkinson ID. 2005. What limits the altitudinal distribution of Craspedolepta
species (Sternorrhyncha: Psylloidea) on fireweed? Ecol Entomol. 30:510–520.
Blomquist CL, Kirkpatrick BC. 2002. Identification of phytoplasma taxa and insect vectors of
peach yellow leaf roll disease in California. Plant Dis. 86:759–763.
Blowers JR, Moran VC. 1968. Notes on the female reproduction system of the South African
citrus psylla, Trioza erytreae (Del Guercio) (Homoptera: Psyllidae). J Entomol Soc South
Afr. 36:71–81.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
151
Bonnemaison L, Missonnier J. 1955a. Influence du photoperiodisme sur le déterminisme des
formes estivales ou hivernales et de la diapause chez Psylla pyri (Homopteres). Comptes
Rendus Acad Sci, Paris. 240:1277–1279.
Bonnemaison L, Missonnier J. 1955b. Recherches sur le determinisme des formes estivales ou
hivernales et de la diapause chez le psylle du poirier (Psylla pyri L.). Ann Epiphyt C.
4:457–528.
Bonnemaison L, Missonnier J. 1956. Le psylle du poirier (Psylla pyri L) morphologie et
biologie, methodes de lutte. Ann Epiphyt. 2:263–331.
Borelli N. 1920. Contributo alla conoscenza della vita nelle galle dell’alloro. Boll Soc Entomol
Ital. 51:3–37.
Boselli FB. 1929a. Studii sugli psyllidi I. Contributio alla conoscenza della psylle del fico
(Homotoma ficus L.). Boll Lab Zool Gen Agrar Fac Agrar Portici 21:218–251.
Boselli FB. 1929b. Studii sugli psyllidi IV. Biologia e svillupo della Spanioza gallii
aspinovelutina (Šulc). Boll Lab Zool Gen Agrar Fac Agrar Portici 23:13–27.
Boselli FB. 1930. Studii sugli psyllidi (Homoptera: Psyllidae), VIII. Biologia e sviluppo della
Rhinocola succincta Heeger. Boll Lab Zool Gen Agrar Fac Agrar Portici 24:211–222.
Boselli FB. 1960. Studii sugli psyllidi (Homoptera: Psyllidae o Chermidae) XI. Biologia e
sviluppo di Diaphorina putoni Loew specie nuova par la fauna italiana. Ann Sper Agri,
Roma 14:149–156.
Brennan EB, Hrusa GF, Weinbaum SA, Levison W. 2001. Resistance of Eucalyptus species to
Glycaspis brimblecombei (Homoptera: Psyllidae) in the San Francisco Bay area. Pan-Pac
Entomol. 77:249–253.
Brennan EB, Weinbaum SA. 2001a. Effect of epicuticular wax on adhesion of psyllids to
glaucous juvenile and glossy adult leaves of Eucalyptus globulus Labillardiere. Aust J
Entomol 40:270–277.
Brennan EB, Weinbaum SA. 2001b. Performance of adult psyllids in no-choice experiments on juvenile and adult leaves of Eucalyptus globulus. Entomol Exp Appl. 100:
179–185.
Brennan EB, Weinbaum SA. 2001c. Psyllid responses to colored sticky traps and the colors of
juvenile and adult leaves of the heteroblastic host plant Eucalyptus globulus. Environ
Entomol. 30:365–370.
Brennan EB, Weinbaum SA. 2001d. Stylet penetration and survival of three psyllid species on
adult leaves and ‘‘waxy’’ and ‘‘de-waxed’’ juvenile leaves of Eucalyptus globulus. Entomol
Exp Appl. 100:355–363.
Brennan EB, Weinbaum SA, Rosenheim JA, Karban R. 2001. Heteroblasty in Eucalyptus
globulus (Myricales: Myricaceae) affects ovipositional and settling preferences of
Ctenarytaina eucalypti and C. spatulata (Homoptera: Psyllidae). Environ Entomol.
30:1144–1149.
Brèthes J. 1921. Un nuevo Psyllidae de la Republica Argentina (Gyropsylla ilicicola). Rev
Facult Agron, Univ Nacion la Plata. 14:82–89.
Brittain WH. 1922a. The adult habits of the apple sucker (Psylla mali Schmidberger). Sci Agri.
3:58–64.
Brittain WH. 1922b. The apple sucker (Psylla mali). J Econ Entomol. 15:96–101.
Brittain WH. 1923a. The European apple sucker. Bull Nova Scotia Dep Agri. 10:1–69.
Brittain WH. 1923b. Injuries, life history and control of the apple sucker (Psylla mali
Schmidberger). Sci Agri 3:176–188.
Brocher F. 1926. Observations biologiques sur Psylla pyrisuga (Hemipt.). Ann Soc Entomol
Fr. 95:183–188.
Brown RG, Hodkinson ID. 1988. Taxonomy and ecology of the jumping plant-lice of Panama
(Homoptera Psylloidea). Entomonograph. Vol. 9. New York: EJ Brill.
Burckhardt D. 1999. Cacopsylla pulchella (Löw) a plant-louse species on Cercis siliquastrum
from Basel (Hemiptera, Psylloidea). Mitt Entomol Ges Basel 49:71–76.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
152 I.D. Hodkinson
Burckhardt D. 2005a. Ehrendorferiana, a new genus of Neotropical jumping plant lice
(Insecta: Hemiptera: Psylloidea) associated with conifers (Cupressaceae). Org Divers
Evol. 5:317–319.
Burckhardt D. 2005b. Biology, ecology, and evolution of gall-inducing psyllids (Hemiptera:
Psylloidea). In: Raman A, Schaefer CW, Withers TM, editors. Biology, ecology, and
evolution of gall-inducing arthropods. Enfield (NH): Science Publishers Inc. p. 143–158.
Burckhardt D, Conci C, Lauterer P, Tamanini L. 1991. Taxonomy and biology of Trioza
tripteridis on Valeriana spp. (Homoptera: Psylloidea). Boll Soc Entomol Ital. 122:165–172.
Burckhardt D, Hanson P, Madrigal L. 2005. Diclidophlebia lucens, n. sp (Hemiptera:
Psyllidae) from Costa Rica, a potential control agent of Miconia calvescens
(Melastomataceae) in Hawaii. Proc Entomol Soc Wash. 107:741–749.
Burckhardt D, Lauterer P. 1982. Trioza laserpitii sp.n., a new Central European psyllid
(Homoptera, Psyllodea). Reichenbachia 20:145–153.
Burckhardt D, Lauterer P. 1997. Systematics and biology of the Aphalara exilis (Weber and
Mohr) species assemblage (Hemiptera: Psyllidae). Entomol Scand. 28:271–305.
Burckhart D, Lauterer P. 2002. Revision of the Trioza rotundata Flor complex (Hemiptera:
Psylloidea): taxonomy and biology. Mitt Schwizerischen Entomol Ges. 75:21–34.
Burckhardt D, Lauterer P. 2006. The palaeartic triozids associated with Rubiaceae
(Hemiptera, Psylloidea): a taxonomic re-evaluation of the Trioza galii Foerster complex.
Rev Suisse Zool. 113:269–286.
Burts EC. 1970. The pear psylla in central Washington. Wash Agri Exp Stn Circ. 516:1–13.
Burts EC, Fischer WR. 1967. Mating behaviour, egg production and egg fertility in the pear
psylla. J Econ Entomol. 60:1297–1300.
Butignol CA, Pedrosa MJH. 2003. Biologia de Neotrioza tavaresi Crawford, 1925 (Hemiptera,
Psyllidae), galhador da folha do aracazeiro (Psidium cattleianum). Rev Brasil Entomol.
47:1–7.
Butt BA, Stuart C. 1986. Oviposition by summer and winter forms of pear psylla Psylla
pyricola on dormant pear budwood. Environ Entomol. 15:1109–1110.
Butt BA, Stuart LC, Bell RL. 1989. Feeding, longevity, and development of pear psylla
(Homoptera, Psyllidae) nymphs on resistant and susceptible pear genotypes. J Econ
Entomol. 82:458–461.
Butterfield J, Whittaker JB, Fielding CA. 2001. Control of the flexible annual/biennial life
cycle of the heather psyllid Strophingia ericae. Physiol Entomol. 26:266–274.
Campbell KG. 1964. Sound production by Psyllidae (Hemipteta). J Entomol Soc Aust. 1:3–4.
Campbell KG. 1992. The biology and population ecology of two species of Cardiaspina
(Hemiptera: Psyllidae) in plague numbers on Eucalyptus grandis in New South Wales.
Proc Linn Soc NSW. 113:135–150.
Cannon RJC. 1983. Cold hardiness in the heather psyllid Strophingia ericae (Homoptera:
Psylloidea). Cryo Lett. 4:113–118.
Carraro L, Ferrini F, Labonne G, Ermacora P, Loi N. 2004. Seasonal infectivity of Cacopsylla
pruni, vector of European stone fruit yellows phytoplasma. Ann Appl Biol. 144:191–195.
Carraro L, Loi N, Ermacora P, Gregoris A, Osler R. 1998. Transmission of pear decline by
using naturally infected Cacopsylla pyri. Acta Horti. 472:665–668.
Carraro L, Osler R, Loi N, Ermacora P, Refatti E. 1998. Transmission of European stone fruit
yellows phytoplasmas by Cacopsylla pruni. J Plant Pathol. 80:233–239.
Carter MW. 1949. The Pittosporum chermid Powellia vitreoradiata Mask. NZ J Sci Tech. B
31:31–42.
Carver M. 1987. Distinctive motory behavior in some adult psyllids (Homoptera, Psylloidea).
J Aust Entomol Soc. 26:369–372.
Casteel CL, Walling LL, Paine TD. 2007. Effect of Mi-1.2 gene in natal host plants on
behavior and biology of the tomato psyllid Bactericerca cockerelli (Šulc) (Hemiptera:
Psyllidae). J Entomol Sci. 42:155–162.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
153
Catling HD. 1969a. The bionomics of the South African citrus psylla Trioza erytreae
(Homoptera: Psyllidae) part 1. The influence of the flushing rhythm of citrus and factors
which regulate flushing. J Entomol Soc South Afr. 32:191–208.
Catling HD. 1969b. The bionomics of the South African citrus psylla Trioza erytreae
(Homoptera: Psyllidae) part 3. The influence of extremes of weather on survival. J
Entomol Soc South Afr. 32:273–290.
Catling HD. 1970. Distribution of psyllid vectors of citrus greening disease, with notes on
biology and bionomics of Diaphorina citri. FAO Plant Prot Bull. 18:8–15.
Catling HD. 1971. The bionomics of the South African citrus psylla Trioza erytreae
(Homoptera: Psyllidae) part 5. The influence of host plant quality. J Entomol Soc South
Afr. 34:381–391.
Catling HD, Annecke DP. 1968. The ecology of the citrus psylla in the Letaba district of
Northern Transvaal. S Afr Citrus J. 410:1–7.
Center TD, Pratt PD, Tipping PW, Rayamajhi MB, Van TK, Wineriter SA, Dray FA. 2007.
Initial impacts and field validation of host range for Boreioglycaspis melaleucae Moore
(Hemiptera: Psyllidae), a biological control agent of the invasive tree Melaleuca
quinquenervia (Cav.) Blake. Environ Entomol. 36:569–576.
Center TD, Pratt PD, Tipping PW, Rayamajhi MB, Van TK, Wineriter SA, Dray FA, Purcell
M. 2006. Field colonization, population growth, and dispersal of Boreioglycaspis
melaleucae Moore, a biological control agent of the invasive tree Melaleuca quinquenervia
(Cav.) Blake. Biol Control. 39:363–374.
Chang JF, Philogene BJR. 1976. The development and behavior of the pear psylla Psylla
pyricola (Homoptera Psyllidae) on different pear rootstocks and cultivars.
Phytoprotection 57:137–149.
Chang KP, Lee CS, Hsiao WF. 1995. Life cycle of Philippine ebony persimmon psyllid, Trioza
magnicauda Crawford (Homoptera: Psyllidae) and insecticide screening of their control.
Plant Prot Bull Taichung. 37:423–432.
Chang K, Musgrave A. 1969. Histochemistry and ultrastructure of the mycetome and
its ‘‘symbiotes’’ in the pear psylla, Psylla pyricola (Homoptera). Tissue Cell.
1:597–606.
Chaterjee PH, Sebastian VO. 1965. Inhibition of inflorecence in Mangifera indica Linn. by the
gall-forming psyllid Apsylla cistellata Buckt (Psyllidae- Homoptera). Control measures.
Indian For. 91:228–230.
Chazeau J. 1987. Le psylle du faux-mimosa en Asie du sud-est et dans la Pacifique: etat du
probleme et perspectives de lutte (Leucaena leucocephala (Lam de Wit) – Heteropsylla
cubana Crawford. Rev Elev Med Vet Nouv Caladonie. 9:23–27.
Chen M, Miyakawa T, Matsui C. 1973. Citrus likubin pathogens in salivary glands of
Diaphorina citri. Phytopathology 63:194–195.
Cheng L, Birch MC. 1978. Insect flotsam: an unstudied marine resource. Ecol Entomol.
3:87–97.
Chiara SR, Rapisarda C, Russo A, Zagami S. 1990. Alcuni fitofagi dell’otano napoletano in
Calabria. Inf Fitopatol. 40:25–30.
Chon DR. 1964. Experiment on the life history and the control of Anomoneura mori Schwarz
(mulberry psyllid). Seric J Korea 4:33–39.
Clark LR. 1962. The general biology of Cardiaspina albitextura (Psyllidae). Aust J Zool.
10:537–586.
Clark LR. 1963a. Factors affecting the attractiveness of foliage for oviposition by Cardiaspina
albitextura (Psyllidae). Aust J Zool. 11:20–34.
Clark LR. 1963b. The influence of population density on the number of eggs laid by females of
Cardiaspina albitextura (Psyllidae). Aust J Zool. 11:190–201.
Clark LR, Dallwitz MJ. 1975. The life system of Cardiaspina albitextura (Psyllidae) 1950–
1974. Aust J Zool. 23:523–561.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
154 I.D. Hodkinson
Cobbinah JR. 1986. Factors affecting the distribution and abundance of Phytolyma lata
(Homoptera, Psyllidae). Insect Sci Appl. 7:111–115.
Cobbinah JR, Wagner MR. 1995. Phenotypic variation in Milicia excelsa to attack by
Phytolyma lata (Psyllidae). For Ecol Manag. 75:147–153.
Collett N. 2001. Biology and control of psyllids, and the possible causes for defoliation of
Eucalyptus camaldulensis Dehnh. (red river gum) in south-eastern Australia – a review.
Aust For. 64:88–95.
Compere H. 1916. Notes on the tomato psylla. Mon Bull Calif Comm Horti. 5:189–191.
Conci C. 2000. Iconography of eggs of Italian Psylloidea (Insecta: Homoptera). Atti Accad
Roveretana Degli Agiati (Ser VII) 10B:5–32.
Conci C, Tamanini L. 1982. Psylla limbata, nuova per l’Italia, da Rhamnus alpinus
(Homoptera: Psylloidea). Atti Soc Ital Sci Nat Mus Civic Stor Nat Milan. 123:
483–494.
Conci C, Tamanini L. 1983. Crastina (Eustigmatia) loginovae n. sp. dell’Italia centrale, da
Tamarix gallica, un genere nuova per l’Europa Centro-Occidentale. Atti Soc Ital Sci Nat
Mus Civic Storia Nat Milan. 124:97–104.
Conci C, Tamanini L. 1984a. Floria (Floria) poggii n.sp., from Sardinia, host plant Genista
corsica (Homoptera: Psylloidea). Ann Mus Civic Storia Nat Genova. 85:43–49.
Conci C, Tamanini L. 1984b. Phylloplecta trisignata (Low, 1986), host plant Rubus sp., of the
complex Rubi Corylifolii. Stud Trent Sci Nat, Acta Biol. 61:249–261.
Conci C, Tamanini L. 1984c. Trioza (Hippophaetrioza n. subgen.) binotata from Alto Adige,
new for Italy (Homoptera: Psylloidea). Stud Trent Sci Nat, Acta Biol. 61:239–248.
Conci C, Tamanini L. 1984d. Trioza (Trioza) rapisardai n.sp. from Piemonte, host plant
Laserpitium siler. Atti Soc Ital Sci Nat Mus Civic Storia Nat Milan. 125:201–208.
Conci C, Tamanini L. 1985a. Arytainilla spartiicola from Puglia, new for Italy (Homoptera:
Psylloidea). Atti Accad Roveretana Degli Agiati (Ser VI) 25:123–136.
Conci C, Tamanini L. 1985b. Lauritrioza n. gen. for Trioza alacris (Homoptera: Psylloidea).
Atti Soc Ital Sci Nat Mus Civic Storia Nat Milan. 126:237–256.
Conci C, Tamanini L. 1985c. Redescription of Trioza ilicina (De Stefani Perez, 1901) comb.n.
from Quercus ilex (Homoptera: Psylloidea). Boll Lab Entomol Agrar Filippo Silvestri,
Portici. 42:33–46.
Conci C, Tamanini L. 1986a. Cyamophila prohaskai from Alto Adige and Trentino, genus and
species new for Italy. Stud Trent Sci Nat, Acta Biol. 62:59–68.
Conci C, Tamanini L. 1986b. The nymph of Phylloplecta trisignata (Löw) and new data on the
morphology and the life history of the species (Homoptera: Psylloidea: Triozidae). Boll
Ist Entomol ‘‘Guido Grandi’’ Univ Bologna Lab Entomol Agrar Filippo Silvestri,
Portici. 41:93–100.
Conci C, Tamanini L. 1986c. Trioza saxifragae in Trentino, new for Italy from Saxifraga
azoides (Homoptera: Psylloidea). Ann Mus Civici Rovereto 2:159–168.
Conci C, Tamanini L. 1987. Observations on Trioza rotundata Flor (Homoptera: Psylloidea).
Ann Mus Civici Rovereto 3:265–284.
Conci C, Tamanini L. 1988. Rare or interesting species of Italian Psylloidea 1 (Homoptera).
Atti Accad Roveretana Degli Agiati (Ser VI) 28B:47–72.
Conci C, Tamanini L. 1989a. Cacopsylla iteophila in Alto Adige and Trentino, new for Italy
(Homoptera Psylloidea). Ann Mus Civici Rovereto 5:205–218.
Conci C, Tamanini L. 1989b. Life history, nymphs and egg of Cyamophila prohaskai, host
plant Anthyllis vulneraria ssp. alpestris (Homoptera: Psylloidea). Stud Trent Sci Nat,
Acta Biol. 65:137–146.
Conci C, Tamanini L. 1990. Notes on the genus Psyllopsis (Homoptera: Psylloidea). Atti
Accad Roveretana Degli Agiati (Ser VI) 29B:57–85.
Conci C, Tamanini L. 1991. Triozidae new or interesting for Italy (Homoptera: Psylloidea).
Atti Accad Roveretana Degli Agiati (Series VI) 30B:37–60.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
155
Conci C, Rapisarda C, Tamanini L. 1993. Annotated catalogue of the Italian Psylloidea. First
Part (Insecta: Homoptera). Atti Accad Roveretana Degli Agiati (Ser VII) 2B:33–136.
Conci C, Rapisarda C, Tamanini L. 1996. Annotated catalogue of the Italian Psylloidea.
Second part (Insecta: Homoptera). Atti Accad Roveretana Degli Agiati (Ser VII)
5B:5–207.
Conconi RE de J. 1973. Biological observations on Paurocephala tuxtlaensis n.sp.
(Homoptera: Psyllidae). Folia Entomol Mexican. 25–26:33–34.
Costanzi M, Santana DLS, Terra AL, Penteado SR, Iede ET, Morey CS. 2003. Un nouveau
psylle sur les eucalyptus de la Riviere Ligure et de la Côte d’Azur. Premieres observations
de Ctenarytaina spatulata Taylor dans le Bassin Mediterranean Occidental. Phytoma –
Defense Veg. 566:48–51.
Cousin MT, Boudon-Padieu E. 2002. Phytoplasmes et phytoplasmoses: vecteurs, methodes de
lutte et themes de récherche. Cahiers Agri 11:115–126.
Crawford SA, Wilkens S. 1996. Ultrastructural aspects of damage to leaves of Eucalyptus
camaldulensis by the psyllid Cardiaspina retator. Micron 27:359–366.
Dabek AJ. 1983. Wissadula proliferation in Jamaica. Vol. 2. Psyllid transmission and
antibiotic therapy. Phytopathol Z. 107:345–361.
Dahlsten DL, Kent DM, Rowney DL, Copper WA, Young TE, Tassan RL. 1995. Parasitoid
shows potential for biocontrol of Eugenia psyllid. Calif Agri. 49:36–40.
Danks HV. 2002. The range of insect dormancy responses. Eur J Entomol. 99:127–142.
Danks HV. 2005. Key themes in the study of seasonal adaptation in insects I. Patterns of cold
hardiness. Appl Entomol Zool. 40:199–211.
Danks HV. 2006. Key themes in the study of seasonal adaptation in insects II. Life history
patterns. Appl Entomol Zool. 41:1–13.
Danks HV. 2007. The elements of seasonal adaptation in insects. Can Entomol. 139:1–44.
Das AK, Rao CN, Singh S. 2007. Presence of citrus greening (Huanglongbing) disease and its
psyllid vector in the North-Eastern region of India confirmed by PCR technique. Curr
Sci. 92:1759–1763.
Das PK, Singh RN, Brahmachari BN, Sharan SK, Sengupta K. 1988. Seasonal intensity of
infestation of the gall insect Trioza fletcheri minor Crawford on Terminalia tomentosa and
Terminalia arjuna and its control through systemic insecticides. Indian J Seric. 27:117–121.
Daugherty MP, Briggs CJ, Welter SC. 2007. Bottom-up and top-down control of pear psylla
(Cacopsylla pyricola): fertilization, plant quality, and the efficacy of the predator
Anthocoris nemoralis. Biol Control. 43:257–264.
Davatchi AG. 1958. Étude biologique de la faune entomologique des Pistacia sauvages et
cultivés. Rev Pathologie Veg Entomol Agri Fr. 37:1–166.
Davies DL, Guise CM, Clark MF, Adams AN. 1992. Parry’s disease of pears is similar to pear
decline and is associated with mycoplasma-type organisms transmitted by Cacopsylla
pyricola. Plant Pathol. 41:195–203.
Davis AC. 1937. Observations on the life history of Paratrioza cockerelli (Šulc) in southern
California. J Econ Entomol. 30:377–379.
Davis BNK. 1973. The Hemiptera and coleopterous fauna of stinging nettle in East Anglia. J
Appl Ecol. 10:213–237.
Davis RI, Gunua TG, Kame MF, Tenakanai D, Ruabete TK. 2005. Spread of citrus
huanglongbing (greening disease) following incursion into Papua New Guinea. Aust
Plant Pathol. 34:517–524.
Del Bene G, Gargani E, Landi S. 1997. Observations on the life cycle and diapause of
Euphyllura olivina (Costa) and Euphyllura phillyreae Foerster (Homoptera: Aphalaridae).
Adv Horti Sci. 11:10–16.
Delic D, Martini M, Ermacora P, Carraro L, Myrta A. 2005. First report of fruit tree
phytoplasmas and their psyllid vectors in Bosnia and Herzegovina. J Plant Pathol.
87:150–150.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
156 I.D. Hodkinson
De Meirleire H. 1971. Le Psylle du laurier sauce. Phytoma 226:38.
Dempster JP. 1968. Intra-specific competition and dispersal as exemplified by a psyllid and its
anthocorid predator. In: Southwood TRE, editor. Insect abundance. Oxford (UK):
Blackwell Scientific Publications. p. 8–17.
De Queiroz Santana DL, Burckhardt DH. 2001. A new triozid pest (Hemiptera, Psylloidea,
Triozidae) on ornamental trumpet trees (Tabebuia spp., Bignoniaceae) in Brazil. Rev
Suisse Zool. 108:541–550.
Deronzier S. 1981. Bioécologie et dynamique des populations de Psylla pyri L. (Homoptera:
Psyllidae) [doctoral thesis]. [Marseille (France)]: University of Marseille.
Deronzier S. 1984. Dynamique des populations de Psylla pyri L. en verger abandonné dans le
sud-est de France. Agronomie 4:549–556.
Deronzier S, Atger P. 1980. Elements d’étude de la dynamique des populations de
Psylla pyri L. dans la basse vallée du Rhône: periode hivernale et printaniere. Acta
Oecol. 1:247–258.
Dhiman SC, Singh S. 2003. Some ecological aspects of Trioza hirsuta Crawford (Homoptera:
Psyllidae): A pest of Terminalia tomentosa. J Exp Zool India. 6:373–376.
Dhiman SC, Singh S. 2004. Biology of Trioza hirsuta Crawford (Homoptera: Psyllidae), a pest
of Terminalia tomentosa W & A. Indian For 130:680–690.
Ding J, Du J, Zhang Y, Yu J, Wu G. 1987. Studies on Thysogyna limbata Enderlein
(Homoptera: Psyllidae) in Nanjing. J Nanjing Agri Univ. 3:47–57.
Domenichini G. 1967. La Psylla melanoneura Foerster (Homoptera) nel Nord Italia. Boll Zool
Agrar Bachic. 8:169–180.
Donnelly GP. 2002. The host range and biology of the mesquite psyllid Heteropsylla texana.
Biocontrol 47:363–371.
Douglas JW. 1878. The natural history of Psylla succincta. Entomol Mon Mag. 15:68–69.
Downer JA, Avihra P, Molinar RH, Fraser JB, Koehler CS. 1988. New psyllid pest of
California pepper tree. Calif Agri. 42:30–32.
Downer JA, Koehler CS, Paine TD. 1991. Biology and management of the the eugenia psyllid
Trioza eugeniae Froggatt. J Environ Horti. 9:137–141.
Ellis PR, Hardman JA. 1992. Pests of umbelliferous crops. In: McKinlay RG, editor.
Vegetable crop pests. Boca Raton (FL): CRC Press. p. 327–383.
Entwistle PF. 1972. Pest of Cocoa. Bristol (UK): Longman.
Espirito-Santo MM, Wilson Fernandez G. 1998. Abundance of Neopelma baccharidis
(Homoptera: Psyllidae) galls on the dioecious shrub Baccharis dracunculifolia
(Asteraceae). Environ Entomol. 27:870–876.
Espirito-Santo MM, Wilson Fernandes GW. 2002. Host plant effects on the development and
survivorship of the galling insect Neopelma baccharidis (Homoptera: Psyllidae). Aust
Ecol. 27:249–257.
Essig EO. 1917. Tomato and laurel psyllids. J Econ Entomol. 10:433–444.
Eyer JR. 1937. Physiology of psyllid yellows of potatoes. J Econ Entomol. 30:891–898.
Eyer JR, Crawford RF. 1933. Observations on the feeding habits of the potato psyllid
(Paratrioza cockerelli Šulc) and the pathological history of the psyllid yellows which it
produces. J Econ Entomol. 26:847–850.
Felt EP. 1906. Bramble flea louse. Mem NY State Mus. 1:688–689.
Ferreira SA, Wilson Fernandes G, Carvalho LG. 1990. Biology and natural history of
Euphaleurus ostreoides (Homoptera: Psyllidae) a gall former on Lonchocarpus
guilleminiaus. Rev Brasil Biol. 50:417–424.
Ferris GF, Hyatt P. 1923. The life history of Euphyllura arbuti Schwarz (Hemiptera:
Chermidae). Can Entomol. 55:88–92.
Fields GJ, Zwick RW. 1975. Elimination of ovarian diapause in pear psylla Psylla pyricola in
the laboratory. Ann Entomol Soc Am. 68:1037–1038.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
157
Finlay-Doney M, Walter GH. 2005. Discrimination among host plants (Leucaena species and
accessions) by the psyllid pest Heteropsylla cubana and implications for understanding
resistance. Agri For Entomol. 7:153–160.
Font I, Abad P, Albinana M, Espino AI, Dally EL, Davis RE, Jorda C. 1999. Amarilleos y
enrojecimiento en zanahoria: una enfermedad a diagnostico. Boll Sanidad Veg Plagas
25:405–415.
Fox Wilson G. 1923. The eucalyptus psyllid Eurhinocola eucalypti. Gardener’s Chron. 76:425.
Frisinghelli C, Delaiti L, Grando MS, Forti D, Vindimian ME. 2000. Cacopsylla costalis (Flor
1861) as a vector of apple proliferation in Trentino. J Phytopathol. 148:425–431.
Fukatsu T, Nikoh N. 1998. Two intracellular symbiotic bacteria from the mulberry psyllid
Anomoneura mori (Insecta, Homoptera). Appl Environ Microbiol. 64:3599–3606.
Fye RE. 1983. Dispersal and winter survival of the pear psylla. J Econ Entomol. 76:311–315.
Gadug S, Hussein MY. 1987. The biology of the durian carsidarid Tenaphalara malayensis
Crawf. Pertanika 10:243–246.
Gegechkori AM. 1984. Psyllids (Homoptera: Psylloidea) of the Caucasus. Tibilisi (Georgia):
Academii Nauk Gruzinskoi SSR.
Gegechkori AM, Djibladzne DS. 1976. The psyllids of Colchida. Tibilisi (Georgia): Institut
A.C. Pushkin.
Gegechkori AM, Loginova MM. 1990. The psyllids (Homoptera: Psylloidea) of the USSR.
Tibilisi (Georgia): Academii Nauk Gruzinskoi SSR.
Geiger CA, Gutierrez AP. 2000. Ecology of Heteropsylla cubana (Homoptera: Psyllidae):
psyllid damage, tree phenology, thermal relations, and parasitism in the field. Environ
Entomol. 29:76–86.
Gençer NS, Coskuncu KS, Kumral NA. 2007. The colonization preference and population
trends of larval fig psylla, Homotoma ficus L. (Hemiptera: Homotomidae). J Pest Sci.
80:1–8.
Giunchedi L, Pollini CP, Biondi S, Babini AR. 1994. PCR detection of MLOs in quick
decline-affected pear trees in Italy. Ann Appl Biol. 124:399–403.
Glick PA. 1939. The distribution of insects, spiders and mites in the air. Technical Bull US
Dep Agri. 673:1–151.
Green GC, Catling HD. 1971. Weather Induced mortality of the citrus psylla Trioza erytreae
(Homoptera Psyllidae), a vector of greening virus in some citrus-producing areas of
Southern African. Agri Meteorol. 8:305–317.
Grove AJ, Gosh CC. 1914. The life history of Psylla isitis Buckt. (Psyllopa punctipennis
Crawford). The ‘‘psylla’’ disease of indigo. Mem Dep Agri in India (Entomol.)
4:329–357.
Hågvar EB, Hågvar S. 1975. Studies on the invertebrate fauna on branches of spruce Picea
abies during winter. Nor J Entomol. 22:23–30.
Halbert SE, Manjunath KL. 2004. Asian citrus psyllids (Sternorrhyncha: Psyllidae) and
greening disease of citrus: A literature review and assessment of risk in Florida. Fl
Entomol. 87:330–353.
Hansen AK, Jeong G, Paine TD, Stouthamer R. 2007. Frequency of secondary symbiont
infection in an invasive psyllid relates to parasitism pressure on a geographic scale in
California. Appl Environ Microbiol. 73:7531–7535.
Hansen LO, Hodkinson ID. 2006. The mistletoe associated psyllid Cacopsylla visci (Curtis,
1835) (Homoptera, Psyllidae) in Norway. Nor J Entomol. 53:89–91.
Hardy AC, Cheng L. 1986. Studies in the distribution of insects by aerial currents. Vol. 3.
Insect drift over the sea. Ecol Entomol. 11:283–290.
Harisanov A. 1966a. Biological and ecological studies on the peach psyllid (Psylla pruni
Scopoli). Nauchni Trudove-Vissh Selskostopanski Institut Plovdiv 15:249–259.
Harisanov A. 1966b. Biological studies on the southern apple leaf hopper Psylla costalis Flor.
Nauchni Trudove-Vissh Selskostopanski Institut Plovdiv 15:261–270.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
158 I.D. Hodkinson
Harris MK. 1973. Host resistance to the pear psylla in a Pyrus communis6P. ussuriensis
hybrid. Environ Entomol. 2:883–887.
Heard SB, Buchanan CK. 1998. Larval performance and association within and between two
species of hackberry nipple gall insects, Pachypsylla spp. (Homoptera: Psyllidae). Am
Midland Nat. 140:351–357.
Heeger E. 1856. Beitrage zur naturgeschichte der Insecten. Sitzung Akad Wiss Wien, Math
Naturwiss. 18:33–47.
Heinze K, Profft J. 1939. Uber Psylliden und Psyllidenschaden an der Kartoffel. Arb Physiol
Angew Entomol Berlin-Dahlen 6:198–208.
Heslop-Harrison G. 1937. Observations on the biology of certain British Psyllidae.
Entomologist 70:1–4.
Heslop-Harrison G. 1942. Notes on the genus Psyllopsis Loew (Hem., Psyllidae) with special
reference to the British species found on Fraxinus. Entomol Mon Mag. 78:155–160.
Heslop-Harrison G. 1949a. A new Indo-Malayan genus and species of the family Psyllidae
(Hemiptera-Homoptera). Entomol Mon Mag. 85:161–164.
Heslop-Harrison G. 1949b. The subfamily Liviinae Loew of the homopterous family Psyllidae
II. Ann Mag Nat Hist (12th ser.) 2:241–270.
Heslop-Harrison G. 1951. The Arytainini of the subfamily Psyllinae, Hemiptera-Homoptera,
family Psyllidae. Ann Mag Nat Hist (12th ser.) 4:417–454.
Heslop-Harrison G. 1961. Sound production in the Homoptera with special reference to the
sound producing mechanisms in the Psyllidae. Ann Mag Nat Hist. 3:633–640.
Hibino H, Kaloostian G, Schneider H. 1971. Mycoplasma-like bodies in the pear psylla vector
of pear decline. Virology 43:34–40.
Hildebrand M, Dickler E, Geider K. 2000. Occurrence of Erwinia amylovora on insects in a
fire blight orchard. J Phytopathol. 148:251–256.
Hill DS. 1982. Hong Kong insects. Vol. 2. Hong Kong: The Urban Council. p. 33–34.
Hill JK, Hamer KC, Hodkinson ID. 1998. Variation in resource exploitation along an
altitudinal gradient: the willow psyllids (Cacopsylla spp.) on Salix lapponum. Ecography
21:289–296.
Hill JK, Hodkinson ID. 1995. Effects of temperature on phenological synchrony and
altitudinal distribution of jumping plant lice (Hemiptera, Psylloidea) on dwarf willow
(Salix lapponum) in Norway. Ecol Entomol. 20:237–244.
Hill JK, Hodkinson ID. 1996. Effects of photoperiod and raised winter temperatures on egg
development and timing of oviposition in the willow psyllid Cacopsylla moscovita.
Entomol Exp Appl. 78:143–147.
Hodgson CJ, Mustafa TM. 1984. The dispersal and flight activity of Psylla pyricola Foerster
in southern England. Lutte integré contre les psylles du poirier. Bull OILB/SROP, p.
330–353.
Hodkinson ID. 1972. Long range dispersal of certain species of Psyllidae in the Northern
Pennines. Entomol Mon Mag. 108:21–22.
Hodkinson I. 1973a. Population dynamics and host plant interactions of Strophingia ericae
(Curt.) (Homoptera: Psylloidea). J Anim Ecol. 42:565–583.
Hodkinson ID. 1973b. The biology of Strophingia ericae (Homoptera Psylloidea) with notes
on its primary parasite Tetrastichus actis (Hymenoptera Eulophidae). Norsk Entomol
Tidsskrift 20:237–243.
Hodkinson ID. 1974. The biology of the Psylloidea (Homoptera): a review. Bull Entomol Res.
64:325–338.
Hodkinson ID. 1981. Status and taxonomy of the Trioza nigricornis complex (Hemiptera:
Homoptera:Triozidae). Bull Entomol Res. 71:671–680.
Hodkinson ID. 1983a. Facultative parthenogenesis in Psylla myrtilli (Homoptera: Psyllidae):
the saga continues in Norway. Fauna Nor Ser B. 30:1–2.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
159
Hodkinson ID. 1983b. The psyllids (Homoptera, Psylloidea) of the Austro-Oriental, Pacific
and Hawaiian zoogeographical realms – an annotated check list. J Nat Hist. 17:341–377.
Hodkinson ID. 1983c. Psyllids (Homoptera: Psylloidea) collected in yellow water trays in
northern Italy. Entomol Gaz. 34:279–280.
Hodkinson ID. 1984. The biology and ecology of the gall-forming Psylloidea. In:
Ananthakrishnan R, editor. The biology of gall forming insects. London: Edward
Arnold. p. 59–77.
Hodkinson ID. 1986a. Co-evolution between psyllids (Homoptera: Psylloidea) and rain forest
trees: the first 120 million years. In: Chadwick AC, Sutton S, editors. Tropical rain forest:
the Leeds symposium. Leeds: Leeds Philosophical Society. p. 187–194.
Hodkinson ID. 1986b. The psyllids (Homoptera: Psylloidea) of the Oriental zoogeographical
region: an annotated checklist. J Nat Hist. 20:299–357.
Hodkinson ID. 1988a. The hitchhiker’s guide to foreign food. New Sci. 118(1611):47–51.
Hodkinson ID. 1988b. The Nearctic Psylloidea (Insecta: Homoptera): an annotated check list.
J Nat Hist. 22:1179–1244.
Hodkinson ID. 1997. Progressive restriction of host plant exploitation along a climatic
gradient: The willow psyllid Cacopsylla groenlandica in Greenland. Ecol Entomol.
22:47–54.
Hodkinson ID. 1999. Biocontrol of eucalyptus psyllid Ctenarytaina eucalypti by the
Australian parasitoid Psyllaephagus pilosus: a review of current programmes and their
success. Biocontrol News Inf. 20:129N–134N.
Hodkinson ID. 2007. A new introduced species of Ctenarytaina (Hemiptera: Psylloidea)
damaging cultivated Eucalyptus parvula (5parvifolia) in Europe. Dtsche Entomol Z.
54:27–33.
Hodkinson ID, Bird JM. 1998. Host-specific insect herbivores as sensors of climate change in
arctic and Alpine environments. Arctic Alpine Res. 30:78–83.
Hodkinson ID, Bird JM. 2000. Sedge and rush-feeding psyllids of the subfamily Liviinae
(Insecta: Hemiptera: Psylloidea): a review. Zool J Linn Soc. 128:1–49.
Hodkinson ID, Bird JM. 2006a. Facultative parthenogenesis in Cacopsylla myrtilli (Wagner)
(Hemiptera: Psylloidea) in northern Sweden. Entomol Tidskrift 127:157–160.
Hodkinson ID, Bird JM. 2006b. Flexible responses of insects to changing environmental
temperature – early season development of Craspedolepta species on fireweed. Global
Change Biol. 12:1308–1314.
Hodkinson ID, Bird JM. In press. The jumping plant lice or psyllids (Homoptera: Psylloidea)
of Greenland. In: Bocher J, editor. The insects of Greenland. Copenhagen (Denmark):
Danish Polar Centre.
Hodkinson ID, Bird JM, Hill JK, Baxter R. 2001. Host plant growth characteristics as
determinants of abundance and phenology in jumping plant-lice on downy willow. Ecol
Entomol. 26:376–387.
Hodkinson ID, Bird JM, Miles JE, Bale JS, Lennon JJ. 1999. Climatic signals in the life
histories of insects: the distribution and abundance of heather psyllids (Strophingia spp.)
in the UK. Funct Ecol. 13:83–95.
Hodkinson ID, Casson D. 2000. Patterns within patterns: abundance-size relationships within
the Hemiptera of tropical rain forest or why phylogeny matters. Oikos 88:509–514.
Hodkinson ID, Jensen TS, Maclean SFJ. 1979. The distribution, abundance and host plant
relationships of Salix feeding psyllids (Homoptera: Psylloidea) in Arctic Alaska. Ecol
Entomol. 4:119–132.
Hodkinson ID, White IM. 1981. The Neotropical Psylloidea (Homoptera, Insecta) – an
annotated check list. J Nat Hist. 15:491–523.
Hoffmann JH, Moran VC, Webb JW. 1975. Influence of host plant and saturation deficit on
temperature tolerance of a psyllid (Homoptera). Entomol Exp Appl. 18:55–67.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
160 I.D. Hodkinson
Hollis D. 2004. Australian Psylloidea. Jumping plant lice and lerp insects. Canberra
(Australia): Australian Government, Australian Biological Resource Study.
Horton DR. 1990a. Distribution and survival of eggs of summer form pear psylla
(Homoptera: Psyllidae) affected by leaf mid-vein. Environ Entomol. 19:656–661.
Horton DR. 1990b. Oviposition by overwintering morph of pear psylla (Homoptera,
Psyllidae) with information on conditioning. Environ Entomol. 19:357–361.
Horton DR, Broers DA, Hinojosa T, Lewis TM. 1998. Ovarian development in overwintering
pear psylla, Cacopsylla pyricola (Homoptera: Psyllidae): seasonality and effects of
photoperiod. Can Entomol. 130:859–867.
Horton DR, Burts EC, Unruh TR, Krysan JL, Coop LB, Croft BA. 1994. Phenology of fall
dispersal by winterform pear psylla (Homoptera, Psyllidae) in relation to leaf fall and
weather. Can Entomol. 126:111–120.
Horton DR, Guedot C, Landolt PJ. 2007. Diapause status of females affects attraction of
male pear psylla, Cacopsylla pyricola, to volatiles from female-infested pear shoots.
Entomol Exp Appl 123:185–192.
Horton DR, Higbee BS, Krysan JL. 1994. Postdiapause development and mating status of
pear psylla (Homoptera, Psyllidae) affected by pear and nonhost species. Ann Entomol
Soc Am. 87:241–249.
Horton DR, Krysan JL. 1990. Probing and oviposition-related activity of summerform pear
psylla (Homoptera, Psyllidae) on host and nonhost substrates. Environ Entomol.
19:1463–1468.
Horton DR, Krysan JL. 1991. Host acceptance behavior of pear psylla (Homoptera:
Psyllidae) affected by plant species, host deprivation, habituation and egg load. Ann
Entomol Soc Am. 84:612–627.
Horton DR, Landolt PJ. 2007. Attraction of male pear psylla, Cacopsylla pyricola, to femaleinfested pear shoots. Entomol Exp Appl. 123:177–183.
Horton DR, Lewis TM. 1996. Tethered flight activity of pear psylla, Cacopsylla pyricola:
seasonal, host and morphotypic effects. Entomol Exp Appl. 78:39–49.
Horton DR, Lewis TM, Neven LG. 1996. Reduced cold-hardiness of pear psylla (Homoptera:
Psyllidae) caused by exposure to external water and surfactants. Can Entomol.
128:825–830.
Hsieh FK, Chen S. 1977. Morphology and bionomics of the Taiwan mulberry psyllid
Paurocephala psylloptera Crawford. Plant Prot Bull. 19:37–46.
Human NB, Bedford ECG. 1985. Relationship between meteorological factors and the
incidence of citrus psylla Trioza erytreae (Del Guercio) at Rustenburg from 1978 until
1982. Subtropica 6:19–23.
Hung T-H, Hung S-C, Chen C-N, Hsu M-H, Su H-J. 2004. Detection by PCR of Candidatus
Liberibacter asiaticus, the bacterium causing citrus huanglongbing in vector psyllids:
application to the study of vector–pathogen relationships. Plant Pathol. 53:96–102.
Hussain MA, Nath D. 1927. The citrus psylla (Diaphorina citri Kuw.) (Psyllidae: Homoptera).
Mem Dep Agri India 10:5–27.
Igboekwe AD. 1983. Studies on the damage to young cocoa seedlings by the cocoa psyllid
Tyora tessmanni (Aulmann) (Homoptera, Psyllidae). Café Cacao Thé 27:67–70.
Igboekwe AD, Adenuga AO. 1983. Studies on the bionomics of cocoa psyllid Tyora tessmanni
(Aulmann) (Homoptera: Psyllidae). Rev Zool Afr. 97:896–904.
Iglesias CA. 1983. Incidencia de la fenologia foliar de Tetragastris panamensis en la
distribucion de Calophya sp. (Homoptera: Psyllidae). Conciencia 10:28–30.
Jackson CS, Hodkinson ID, Stanley P. 1990. Cold-hardiness in the hawthorn psyllid
Cacopsylla melanoneura (Foerster) (Homoptera: Psylloidea). Entomologist 109:224–230.
Janes MJ, Davis AC. 1937. Observations on the life history of Paratrioza cockerelli in
California. J Econ Entomol. 30:377–378.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
161
Jarausch B, Schwind N, Jarausch W, Krczal G, Dickler E, Seemuller E. 2003. First report of
Cacopsylla picta as a vector of apple proliferation phytoplasma in Germany. Plant Dis.
87:101.
Jarausch W, Danet JL, Labonne G, Dosba F, Broquaire JM, Saillard C, Garnier M. 2001.
Mapping the spread of apricot chlorotic leaf roll (ACLR) in southern France and
implication of Cacopsylla pruni as a vector of European stone fruit yellows (ESFY)
phytoplasmas. Plant Pathol. 50:782–790.
Jensen DD, Griggs WH, Gonzales CQ, Schneider H. 1964. Pear decline virus transmission by
pear psylla. Phytopathology 54:1346–1351.
Jonsson N. 1983. The life history of Psylla mali (Homoptera, Psyllidae) and its relationship to
the development of the apple blossom. Fauna Nor Ser B. 30:3–8.
Journet ARP. 1984. Associations of Craspedolepta species (Homoptera: Psylloidea) with
goldenrod (Solidago spp.) in Quebec, Canada. Rev Entomol Quebec. 29:22–26.
Junior M, Barros R, da Silva FR, de Vasconcelos GJN. 2005. Occurrence and biological
aspects of the Clitoria tree psyllid in Brazil. Sci Agricol. 62:281–285.
Kandasamy C. 1980. A new leaf gall by Trioza gigantea Craw. (Homoptera, Psyllidae) on
Vaccinium neilgherrense (Vacciniaceae). Curr Sci. 49:754–754.
Kandasamy C, Krishnan BH. 1981. First record of leaf galls on Litsea stocksii (Meissn) Hook
F (Lauraceae) caused by a psyllid. Curr Sci. 50:967–968.
Kapatos ET, Stratopoulou ET. 1996. Demographic study of the reproductive potential of pear
psylla, Cacopsylla pyri. Entomol Exp Appl. 80:497–502.
Kapatos ET, Stratopoulou ET. 1999. Duration times of the immature stages of Cacopsylla
pyri L. (Hom., Psyllidae), estimated under field conditions, and their relationship to
ambient temperature. J Appl Entomol. 123:555–559.
Kaufmann T. 1973. Biology and ecology of Tyora tessmanni (Homoptera: Psyllidae) with
special reference to its role as cocoa pollinator in Ghana. J Kansas Entomol Soc.
46:285–293.
Klimaszewski SM. 1973. The jumping plant lice or psyllids (Homoptera: Psyllodea) of the
Palearctic: an annotated checklist. Ann Zool Warsaw. 30:155–286.
Knowlton GF. 1933. Note on host plants of Paratrioza cockerelli. J Econ Entomol. 26:730.
Knowlton GF, Janes MJ. 1931. Studies on the biology of Paratrioza cockerelli (Šulc). Ann
Entomol Soc Am. 24:283–291.
Knowlton GF, Thomas WL. 1934. Host plants of the potato psyllid. J Econ Entomol. 27:547.
Koehler CS, Kattoulas ME, Frankie GW. 1966. Biology of Psylla uncatoides. J Econ
Entomol. 59:1097–1100.
Konovalova ZA. 1963. On the biology of Calophya nigra Kuw. Trudy Akad Nauk Sootsza
SSR, Vladivostok 17:91–95.
Konovalova ZA. 1976. Leafhoppers (Homoptera, Psylloidea) in Primorsky Kray. Trudy Biolpochvennogo Inst (New Ser.) 43:3–5.
Krawczyk A, Migula P. 1979. Metabolism in Psyllodea. Acta Physiol Polonica. 30:114.
Kristoffersen L, Anderbrant O. 2007. Carrot psyllid (Trioza apicalis) winter habitats – insights
in shelter plant preference and migratory capacity. J Appl Entomol. 131:174–178.
Krysan JL. 1990. Laboratory study of mating behavior as related to diapause in overwintering
Cacopsylla pyricola (Homoptera, Psyllidae). Environ Entomol. 19:551–557.
Krysan JL, Higbee BS. 1990. Seasonality of mating and ovarian development in overwintering
Cacopsylla pyricola (Homoptera, Psyllidae). Environ Entomol. 19:544–550.
Kudler J. 1968. Diclidophlebia eastopi (Vondracek) a psyllid harmful to Triplochiton
scleroxylon (K Schum). Technical Newsletter, Forest Products Research Institute,
Ghana, 11–14.
Kumar SU, Raman A, Kader MSA. 1981. Physiological alterations in the leaves of
Buchanania lanzan due to psyllid galls. Curr Sci. 50:963–964.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
162 I.D. Hodkinson
Kuwayama S. 1971. Observastions on the biology of the mulberry sucker. Jpn J Appl Entomol
Zool. 15:115–120.
Labonne G, Lichou J. 2004. Data on the life cycle of Cacopsylla pruni vector of European
stone fruit yellows (ESFY) phytoplasma, in France. In: Llacer G, editor. Proceedings of
the XIXth international symposium on virus and virus-like diseases of temperate fruit
crops: fruit tree diseases. Leuven (Belgium): International Society for Horticultural
Science. p. 465–470.
Lahiri AR, Biswas S. 1980. Observations on the relative intensity of infection on three species
of cultivated citrus plants by Psylla murrayi Mathur (Homoptera: Psyllidae) at Shillong,
Meeghalaya. Bull Zool Surv India. 2:123–127.
Lakra RK, Singh Z, Kharub WS. 1983. Population dynamics of citrus psylla Diaphorina citri
Kuwayama in Haryana. Indian J Entomol. 45:301–310.
Lal KB. 1934. The biology of the Scottish Psyllidae. Trans R Entomol Soc Lond. 82:363–385.
Lapis EB, Borden JH. 1993a. Components of resistance of Leucaena collinsii (Leguminosae) to
Heteropsylla cubana (Homoptera: Psyllidae). Environ Entomol. 22:319–325.
Lapis EB, Borden JH. 1993b. Olfactory discrimination by Heteropsylla cubana (Homoptera:
Psyllidae) between susceptible and resistant species of Leucaena (Leguminosae). J Chem
Ecol. 19:83–90.
Laska P. 1964. Zusammenfassung: Beitrag zu Bionomie und Bekampfung der Trioza apicalis
Forst. (Triozidae, Homoptera). Zool Listy. 13:327–332.
Laska P. 1974. Study of Trioza apicalis (Triozidae: Homoptera). Prirodoved Prace Ustavu
Ceskoslovenske Akad Ved v Brne 8:1–44.
Laurema S. 1989. Free amino acids in the psyllid Trioza apicalis (Homopt., Triozidae) and in
carrot leaves. Ann Agri Fenn. 28:113–120.
Lauterer P. 1963. A contribution to the knowledge of the psyllid fauna of Czechoslovakia. Cas
Morav Mus, Brne. 48:145–156.
Lauterer P. 1965. A contribution to the knowledge of the psyllid fauna of Czechoslovalia II.
Cas Morav Mus, Brne. 50:171–190.
Lauterer P. 1976. Psyllids of wetland nature reserves of the German Democratic Republic with
notes on their biology, taxonomy and zoogeography. Faun Abh Dresden. 6:111–122.
Lauterer P. 1979. New and interesting records of psyllids from Czechoslovakia (Homoptera:
Psylloidea). Cas Morav Mus, Brne. 64:93–102.
Lauterer P. 1982. New data on the occurence, bionomics and taxonomy of some
Czechoslovakian Psylloidea (Homoptera). Cas Morav Mus, Brne. 67:133–162.
Lauterer P. 1991. Psyllids (Homoptera: Psylloidea) of the limestone cliff zone of the Pavlovske
Vrchy Hills (Czeckoslovakia). Cas Morav Mus, Brne. 76:241–263.
Lauterer P. 1993a. Notes on the bionomics and occurrence of some psyllids (Homoptera:
Psylloidea) in Czechoslovakia and the Balkan Peninsula. Cas Morav Mus, Brne.
77(1992):147–156.
Lauterer P. 1993b. Psyllids (Homoptera: Psylloidea) from the area flooded by the Nove
Milyny reservoir system and its environs in southern Moravia. Cas Morav Mus, Brne.
78:165–200.
Lauterer P. 1993c. Three faunistic novelties from the order Homoptera in the Czech and
Slovak Republics (Auchenorrhyncha and Psylloidea). Cas Morav Mus, Brne.
78:213–214.
Lauterer P. 1998. Results of the investigations on Hemiptera in Moravia, made by the
Moravian Museum. Cas Morav Mus, Brne. 83:99–126.
Lauterer P. 1999. Results of the investigations on Hemiptera in Moravia, made by the
Moravian Museum (Psylloidea 2). Cas Morav Mus, Brne. 84:71–151.
Lauterer P, Baudys E. 1968. Description of a new gall on Chamaenerion angustifolium (L.)
Scop. produced by the larva of Craspedolepta subpunctata (Foer.) with notes on the
bionomics of this psyllid. Cas Morav Mus, Brne. 53:243–248.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
163
Lauterer P, Broumas T, Drosopoulos S, Souliotis C, Tsourgianni A. 1998. Species of the genus
Agonoscena (Homoptera, Psyllidae), pests on Pistacia and first record of A. pistaciae in
Greece. Ann Inst Phytophathol Benaki. 18:123–128.
Lauterer P, Burckhardt D. 2004. The west palaearctic species of the Craspedolepta
flavipennis (Foerster) complex (Hemiptera: Psylloidea). Mitt Schweiz Entomol Ges.
77:251–275.
Lauterer P, Janicek R. 1990. Trioza neglecta Loginova, a new species for the fauna of Hungary
and Bulgaria (Homoptera: Psylloidea). Folia Entomol Hung. 51:163–165.
Lauterer P, Prophetou DA, Tzanakakis ME. 1986. Occurrence of Euphyllura phillyreae
Foerster (Homoptera: Aphalaridae) on olives of the Greek mainland. Ann Entomol Soc
Am. 79:7–10.
Lazarev MA. 1972. The crimean apple sucker Psylia melanoneura taurica new-form
(Homoptera: Psylloidea). Entomol Obozr. 51:23–26.
Lazarev MA. 1975. New data on the biology of the pear sucker Psylla pyrisuga (Homoptera:
Psylloidea) in the Crimea. Entomol Obozr. 54:758–759.
Lazarev MA. 1979. New data on the biology of the common pear psylla Psylla pyri
(Homoptera, Psyllidae) in the Crimea. Entomol Obozr. 58:53–56.
Léclant F, Marchoux G, Giannotti J. 1974. Mise en evidence du rôle vecteur du psylle Trioza
nigricornis Forst (Insecte, Homoptere) dans la transmission d’une maladie a proliferation
de Daucus carota L. Comptes Rendus Acad Sci, Paris Ser D. 278:57–59.
Ledoux A. 1955. Obsevations sur la biologie de Phytolyma lata Scott varieté fusca, Psylloidea,
gallicole sur l’Iroko (Clorophora excelsa). Ann Sci Nat, Zool (11th Ser.). 291–310.
Lee RE, Litzgus JD, Mugnano JA, Lee MR, Horton DR, Dunley J. 1999. Evaluation of icenucleating microorganisms for reducing the supercooling capacity and cold-hardiness of
Cacopsylla pyricola (Hemiptera: Psyllidae). Can Entomol. 131:715–723.
Leege LM. 2006. The relationship between psyllid leaf galls and redbay (Persea borbonia)
fitness traits in sun and shade. Plant Ecol. 184:203–212.
Leeper JR, Beardsley JWJ. 1976. The biological control of Psylla uncatoides (Homoptera:
Psyllidae) on Hawaii. Proc Hawaiian Entomol Soc. 22:307–321.
Lehrman RS. 1930. Some observations on the life history of the tomato psyllid (Paratrioza
cockerelli Šulc) (Homoptera). J NY Entomol Soc. 38:307–312.
Leite MSP, Zanol KMR. 2001. Biologia e morfologia de Gyropsylla spegazziniana (Lizer y
Trelles) (Hemiptera, Psyllidae). Acta Biol Parana. 30:19–34.
List GM. 1939a. The effect of temperature upon egg deposition, egg hatch and nymphal
development of Paratrioza cockerelli (Šulc). J Econ Entomol. 32:30–36.
List GM. 1939b. The potato and tomato psyllid and its control on tomato. Bull Colo Agri Exp
Stn. 454:1–33.
Liu D, Johnson L, Trumble JT. 2006. Differential responses to feeding by the tomato/potato
psyllid between two tomato cultivars and their implications in establishment of injury
levels and potential of damaged plant recovery. Insect Sci. 13:195–204.
Liu D, Trumble JT. 2004. Tomato psyllid behavioral responses to tomato plant lines and
interactions of plant lines with insecticides. J Econ Entomol. 97:1078–1085.
Liu D, Trumble JT. 2005. Interactions of plant resistance and insecticides on the development
and survival of Bactericerca cockerelli (Šulc) (Homoptera: Psyllidae). Crop Prot.
24:111–117.
Liu D, Trumble JT. 2006. Ovipositional preferences, damage thresholds, and detection of the
tomato-potato psyllid Bactericera cockerelli (Homoptera: Psyllidae) on selected tomato
accessions. Bull Entomol Res. 96:197–204.
Liu D, Trumble JT. 2007. Comparative fitness of invasive and native populations of the
potato psyllid (Bactericera cockerelli). Entomol Exp Appl. 123:35–42.
Liu HS, Chen CC, Lin CP. 2007. Detection and identification of the phytoplasma associated
with pear decline in Taiwan. European J Plant Pathol. 117:281–291.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
164 I.D. Hodkinson
Liu YH, Tsai JH. 2000. Effects of temperature on biology and life table parameters of the
Asian citrus psyllid, Diaphorina citri Kuwayama (Homoptera: Psyllidae). Ann Appl Biol.
137:201–206.
Lizer C. 1918. Sobre la presencia en Argentina de un psilido exotico (Trioza alacris F.). Ann
Zool Apl. 5:16–21.
Loginova MM. 1954. On the biology of leafhoppers of the genus Psyllopsis Löw in the
Stalingrad region. Trudy Zool Inst Acad Nauk SSR, Leningrad. 38:88–104.
Loginova MM. 1968. New data on the fauna and biology of the Caucasian Psylloidea
(Homoptera). Trudy Vses Entomol Obschchestva. 52:275–328.
Loginova MM. 1970. New psyllids (Homoptera: Psylloidea) from Soviet Central Asia.
Entomol Obozr. 49:370–385.
Loginova MM. 1976. Psyllids of the Tribe Pachypsylloidini. Zool Zhurnal. 55:612–614.
Loginova MM. 1982. Structure and morpho-ecological types of the psyllid nymphs
(Homoptera: Psylloidea). Trudy Zool Inst Acad Nauk SSR, Leningrad. 105:20–52.
Loginova MM, Parfentiev VJ. 1958. Leafhoppers (Homoptera: Psylloidea) injurious to
Populus diversifolia and P.pruinosa in the vicinity of Lake Bakhash, Kazakhstan.
Entomol Obozr. 38:88–104.
Louda SM, Rodman JE. 1983. Ecological patterns in the glucosinolate content of a native
mustard, Cardamine cordifolia, in the Rocky Mountains. J Chem Ecol. 9:397–422.
Loureiro Ferreira J. 1946. Algumas observacoes biologicas e biometricas efectuadas em
Euphyllura olivina Costa. Agron Lusit, Lisbon. 7:63–95.
Löw F. 1877. Zur Biologie un Charakteristik der Psylloden nebst Beschreibung. Verh ZoolBot Ges Wien. 26:187–216.
Löw F. 1880. Mitteilungen uber Psylloden. Verh Zool-Bot Ges Wien. 29:549–598.
Luft PA, Paine TD. 1997. Behavioral cues associated with oviposition by Trioza eugeniae.
Entomol Exp Appl. 84:293–299.
Luft PA, Paine TD. 1998. Potential biotic factors influencing settling of Trioza eugeniae
nymphs (Homoptera: Triozidae) on young foliage. Environ Entomol. 27:1425–1430.
Luft PA, Paine TD, Redak RA. 2001. Limiting the potential for intraspecific competition:
regulation of Trioza eugeniae oviposition on unexpanded leaf tissue. Ecol Entomol.
26:395–403.
Lundblad O. 1929. Morotbladloppan Trioza viridula Zett. dess biologi och upptradande som
skadedjur i Sverige. Medd Cent Lantsbruksentomologiska Avd. 350(55):1–45.
Lyoussoufi A, Armand E, Rieux RDAFF. 1992. Population dynamics of pear psylla Psylla
pyri (L.) (Homoptera: Psyllidae) and beneficials in a sprayed orchard of south-eastern
France. Acta Phytopathol Entomol Hung. 27:413–417.
Lyoussoufi A, Gadenne C, Rieux R, Faivredarcier F. 1994. Diapause development of the pear
psylla, Cacopsylla pyri (L.) under natural conditions. Entomol Exp Appl. 70:193–199.
Lyoussoufi A, Rieux R, Faivre d’Arcier F. 1988. Evolution du potentiel de ponte et de l’efectif
des oeufs du pylle du poirier Psylla pyri (L.) au cours de la periode hivernale et
printaniere dans la basse vallée du Rhône. J Appl Entomol. 106:97–107.
MacLean SF. 1983. Life cycles and the distribution of psyllids (Homoptera) in arctic and
subarctic Alaska. Oikos 40:445–551.
MacLean SF, Hodkinson ID. 1980. Distribution of psyllids (Homoptera: Psylloidea) in Arctic
and subarctic Alaska. Arctic Alpine Res. 12:369–376.
Madubunyi LC. 1967. Ecological investigations on the albizzia psyllid Psylla uncatoides
(Ferris and Klyver)(Homoptera: Psylloidea) [masters thesis]. [Berkeley (CA)]: University
of California.
Madubunyi LC, Koehler CS. 1974. Development, survival and capacity for increase of
albizzia psyllid at various constant temperatures. Environ Entomol. 3:1013–1016.
Malenovsky I. 1999. Contribution a la faunistiqe des psylles d’Alsace (Hemiptera: Psylloidea).
Bull Soc Entomol Mulhouse., (April–June):17–34.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
165
Mally CW. 1894. Psyllidae found at Ames, Iowa. Proc Iowa Acad Sci. 2:152–171.
Malausa JC, Giradet N. 1997. Lutte biologique contre le psylle de l’eucalyptus. Phytoma –
Defense Veg. 498:49–51.
Mangat BS. 1966. Biology and controlof citrus psylla Diaphorina citri Kuw. Plant Prot Bull, N
Delhi. 18:18–20.
Mani MS. 1948. Cecidozoa and zoocecidia from India. J R Asiatic Soc Bengal. 14:27–195.
Mani T, Raman A. 1994. Biochemical changes in relation to growth in two leaf gall systems
induced by Trioza jambolanae and Microceropsylla longispiculata (Homoptera:
Psylloidea). Phytophaga 6:59–64.
Mansilla JP, Pérez R, Del Estal P, Blond A. 2004. Deteccı́on en España de Ctenarytaina
spatulata Taylor sobre Eucalyptus globulus Labill. Bol Sanid Veg, Plagas. 30:57–63.
Markkula M, Laurema S. 1971. Phytotoxemia caused by Trioza apicalis (Homoptera,
Triozidae) on carrot. Ann Agri Fenn. 10:181–184.
Mathur RN. 1935. On the biology of the Psyllidae (Homopt.). Indian For Rec (New Ser.)
1:35–71.
Mathur RN. 1946. On the immature stages of some Psyllidae. Indian J Entomol. 8:224–236.
Mathur RN. 1975. Psyllidae of the Indian Subcontinent. New Delhi (India): Indian Council of
Agriculural Research.
Maxwell-Lefroy H. 1913. The Psylla disease of indigo in Behar. Agri J India. 8:1–26.
McAtee WL. 1915. Psyllidae wintering on conifers about Washington DC. Science 41:940.
McLean IFG. 1993. The host plant association and life history of Trichochermes walkeri
(Forster) (Psylloidea: Triozidae). B J Entomol Nat Hist. 6:13–16.
McLean IFG. 1994. The population dynamics of a gall forming psyllid. In: Leather SR, Watt
AD, Mills NJ, Walters KFA, editors. Individuals, populations and patterns in ecology.
Andover (UK): Intercept. p. 97–107.
McLean IFG. 1998. The population ecology of Trichochermes walkeri. In: Dempster JP,
McLean IFG, editors. Insect populations in theory and practice. London: Klewer
Academic Publishers. p. 341–366.
McMullen RD, Jong C. 1972. Influence of temperature and host vigor on fecundity of the pear
psylla (Homoptera, Psyllidae). Can Entomol. 104:1209–1212.
McMullen RD, Jong C. 1976. Factors affecting induction and termination of diapause in pear
psylla (Hemiptera, Homoptera, Psyllidae). Can Entomol. 108:1001–1006.
McMullen RD, Jong C. 1977. Effect of temperature on developmental rate and fecundity of
the pear psylla Psylla pyricola (Homoptera, Psyllidae). Can Entomol. 109:165–170.
Mead FW. 1963. A psyllid, Trioza magnoliae (Ashmead) (Homoptera: Psyllidae). Fl Dep Agri,
Div Plant Ind, Entomol Circ. 15:1–2.
Mead FW. 1966a. The blackberry psyllid Trioza tripunctata (Fitch). Fl Dep Agri, Div Plant
Ind, Entomol Circ. 52:1–2.
Mead FW. 1966b. Persimmon psyllid Trioza diospyri (Ashmead) (Homoptera: Psyllidae). Fl
Dep Agri, Div Plant Ind, Entomol Circ. 50:1–2.
Mead FW. 1967. A nest-making psyllid Euphalerus nidifex in Florida. Homoptera: Psyllidae
Fl Dep Agri, Div Plant Ind, Entomol Circ. 67:1–2.
Mead FW. 1977. The Asiatic citrus psylla Diaphorina citri Kuwayama (Homoptera: Psyllidae).
Fl Dep Agri, Div Plant Ind, Entomol Circ. 180:1–4.
Mead FW. 1983. Yaupon psyllid Gyropsylla ilicis (Ashmead) (Homoptera: Psyllidae). Fl Dep
Agri, Div Plant Ind, Entomol Circ. 247:1–2.
Mead FW. 1994. Eugenia psyllid, Trioza eugeniae Froggatt (Homopter: Psyllidae). Fl Dep
Agri, Div Plant Ind, Entomol Circ. 367:1–3.
Mehrnejad MR. 1998. Evaluation of the parasitoid Psyllaephagus pistaciae (Hymenoptera:
Ecyrtidae) as a biocontrol agent of the common pistacio psylla Agonoscena pistaciae
(Hemiptera: Psylloidea) [doctoral thesis]. [London (UK)]: University of London. 271.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
166 I.D. Hodkinson
Mehrnejad MR. 2002. Bionomics of the common pistachio psylla, Agonoscena pistaciae, in
Iran. In: Batlle I, Hormaza I, Espiau MT, editors. Proceedings of the 3rd International
Symposium on Pistachios and Almond. Leuven (Belgium): International Society
Horticultural Science. p. 535–539.
Mehrnejad MR, Copland MJW. 2005. The seasonal forms and reproductive potential of the
common pistachio psylla, Agonoscena pistaciae (Hem., Psylloidea). J Appl Entomol.
129:342–346.
Mehrnejad MR, Copland MJW. 2006a. Host-stage selection and oviposition behaviour of
Psyllaephagus pistaciae, parasitoid of the common pistachio psylla Agonoscena pistaciae.
Biol Control. 36:139–146.
Mehrnejad MR, Copland MJW. 2006b. Biological parameters of parasitoid Psyllaephagus
pistaciae and its host Agonoscena pistaciae in relation to temperature. J Entomol Res Soc.
8:1–20.
Mensah RK, Madden JL. 1991. Resistance and susceptibility of Boronia megastigma cultivars
to infestations by the psyllid Ctenarytaina thysanura. Entomol Exp Appl. 61:189–198.
Mensah RK, Madden JL. 1992a. Factors affecting Ctenarytaina thysanura oviposition on
Boronia megastigma terminal shoots. Entomol Exp Appl. 62:261–268.
Mensah RK, Madden JL. 1992b. Feeding behaviour and pest status of Ctenarytaina thysanura
Ferris and Klyver (Hemiptera: Psyllidae) on Boronia megastigma Nees in Tasmania. J
Aust Entomol Soc. 31:71–78.
Mensah RK, Madden JL. 1993a. Development and application of an integrated pest
management programme for the psyllid, Ctenarytaina thysanura on Boronia megastigma
in Tasmania. Entomol Exp Appl. 66:59–74.
Mensah RK, Madden JL. 1993b. Life history and biology of Ctenarytaina thysanura Ferris
and Klyver (Hemiptera: Psyllidae) on Boronia megastigma Nees ex Bartl. (Rutaceae) in
Tasmania. J Aust Entomol Soc. 32:327–337.
Mensah RK, Madden JL. 1994. Life table and key factor studies of Ctenarytaina thysanura
Ferris and Klyver (Hemiptera: Psyllidae) on Boronia megastigma Nees. ex Bartl.
(Rutaceae) in Tasmania. J Aust Entomol Soc. 33:353–361.
Messi J. 1983a. Facteurs influecant le comportement de ponte de Mesohomotoma tessmanni
Aulmann, psylle parasite de cacoyer. Bull Soc Entomol Fr. 88:368–374.
Messi J. 1983b. Influence of temperature and photoperiod on the embryonic and
postembryonic development of Mesohomotoma tessmanni Aulmann (Homoptera,
Psyllidae) a pest of the cacao plant. Bull Inst Fondam Afr Noire, Ser A, Sci Nat.
45:274–287.
Messi J, Tamesse JL. 1999. Fluctuations numeriques des populations de Trioza erytreae (Del
Guercio), psylle des agrumes dans un verger fruitier a Yaounde (Hemiptera:
Psylloidea:Triozidae). Ann Soc Entomol Fr. 35:238–241.
Mifsud D. 1997. The jumping plant-lice (Hemiptera: Psylloidea) of the Maltese Islands
[masters thesis]. [Msida (Malta)]: University of Malta. 84 p.
Migula P, Glowacka E, Krawczyk A. 1980. Respiratory metabolism in jumping plant lice
(Psylloidea, Homoptera). Prace Naukowe Uniwersytetu Slaskiego w Katowicach
375:185–205.
Miles HW. 1928. The bay psyllid. Northwest Nat, Arbroath. 3:8–14.
Miles JE, Bale JS, Hodkinson ID. 1997. Effects of temperature elevation on the population
dynamics of the upland heather psyllid Strophingia ericae (Curtis) (Homoptera:
Psylloidea). Global Change Biol. 3:291–297.
Miles JE, Bale JS, Hodkinson ID. 1998. Life cycle regulation in the heather psyllid
Strophinigia ericae: responses to temperature and photoperiod. Physiol Entomol.
23:376–381.
Missonnier J. 1956. Note sur la biologie du psylle de l’aubepine (PsylIa peregrina). Ann
Epiphyt 7:253–262.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
167
Miyatake Y. 1968a. A new Japanese species of Trioza from Machilus thunbergii, with
descriptions of the immature stages and notes on biology. Trans Shikoku Entomol Soc.
10:1–10.
Miyatake Y. 1968b. Pachypsylla japonica sp. nov., a remarkable lerp-forming psyllid from
Japan (Homoptera: Psyllidae). Bull Osaka Mus Nat Hist. 21:5–12.
Miyatake Y. 1969. On the life history and the immature stages of Trioza cinnamomi (Boselli),
with the redescription of the adult (Hemiptera: Psyllidae). Bull Osaka Mus Nat Hist.
22:19–30.
Miyatake Y. 1970. Some taxonomical and biological notes on Togepsylla matsumurana
Kuwayama (Hem., Psyllidae). Bull Osaka Mus Nat Hist. 23:1–10.
Miyatake Y. 1973. Notes on the genus Euphalerus of Japan with description of a new species
(Homoptera Psyllidae). Bull Osaka Mus Nat Hist. 27:23–28.
Miyatake Y. 1978. Notes on the genus Epitrioza of Japan with descriptions of two new species
(Homoptera: Psyllidae). Bull Osaka Mus Nat Hist. 31:93–111.
Miyatake Y. 1980. Notes on the genus Pachypsylla of Japan with description of Pachypsylla
usubai new species (Homoptera: Psyllidae). Bull Osaka Mus Nat Hist. 33:61–70.
Miyatake Y. 1992. A revision of the genus Calophya from Japan (Homoptera: Psylloidea).
Bull Osaka Mus Nat Hist. 46:11–23.
Miyatake Y. 1994. Further knowledge on the distribution and biology of two species of the
genus Celtisaspis (Homoptera: Psylloidea: Spondyliaspididae). Bull Osaka Mus Nat Hist.
48:27–30.
Mohammed MA, Sheet AI. 1989. Ecological study on the pistachio psyllid (Agonoscena
targionii (Licht.)) (Homoptera, Psyllidae) in Mosul region, Iraq. Arab J Plant Prot.
7:138–142.
Mondal S, Ray R, Mukhopadhyay TK. 2003. Destruction of Areca catechu by Psylla sp.
(Hemiptera: Psyllidae). Bionotes 5:93.
Monobrullah M, Singh PP, Singh R. 1998. Life-history and morphology of different stages of
mango shoot gall psyllid, Apsylla cistellata Buckton (Homoptera: Psyllidae). J Entomol
Res, N Delhi. 22:319–323.
Moore KM. 1961. Observations on some Australian forest insects 8. The biology and
occurrece of Glycaspis baileyi Moore in New South Wales. Proc Linn Soc NSW.
86:185–200.
Moore KM. 1983. New species and records of Glycaspis Taylor (Homoptera,
Spondyliaspidae) with phyletic groupings. J Aust Entomol Soc. 22:177–184.
Moran VC. 1968a. The development of the citrus psylla Trioza erytreae (Homoptera,
Psyllidae) on Citrus limon and four indigenous host plants. J Entomol Soc South Afr.
31:391–402.
Moran VC. 1968b. Preliminary observations on the choice of host plants by adults of the
citrus psylla Trioza erytreae (Homoptera: Psyllidae). J Entomol Soc South Afr.
31:403–410.
Moran VC, Blowers JR. 1967. On the biology of the South African citrus psylla, Trioza
erytreae (Del Guercio) (Homoptera: Psyllidae). J Entomol Soc S Afr. 30:96–106.
Moran VC, Brown RP. 1973. The antennae, chemoreception and probing activity of the citrus
psylla Trioza erytreae (Del Guercio) (Homoptera:Psyllidae). J Entomol Soc S Afr.
36:191–202.
Moran VC, Buchan PR. 1975. Oviposition by the citrus psylla Trioza erytreae (Homoptera,
Psyllidae) in relation to leaf hardness. Entomol Exp Appl. 18:96–104.
Morath SU, Pratt PD, Silvers CS, Center TD. 2006. Herbivory by Boreioglycaspis melaleucae
(Hemiptera: Psyllidae) accelerates foliar senescence and abscission in the invasive tree
Melaleuca quinquenervia. Environ Entomol. 35:1372–1378.
Morgan FD. 1984. Psylloidea of South Australia. Woolman DJ, editors. Handbook of the
Flora and Fauna of South Australia. Adelaide: South Australia, Illustrated. Paper.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
168 I.D. Hodkinson
Morgan FD, Taylor GS. 1988. The white lace lerp in southeastern Australia. In: Berryman
AA, editor. Dynamics of forest insect populations. patterns, causes, implications.
London: Plenum Press. p. 130–140.
Moxon JE. 1984. D.P.I. Entomology Bulletin 43, Psyllid pest of Leucaena. Harvest 12:19–22.
Mullen BF, Shelton HM. 2003. Psyllid resistance in Leucaena. Part 2. Quantification of
production losses from psyllid damage. Agrofor Syst. 58:163–171.
Munro JA. 1965. Occurrence of Psylla uncatoides on Acacia and Albizzia with notes on
control. J Econ Entomol. 58:1171–1172.
Munyaneza JE, Crosslin JM, Upton JE. 2007. Association of Bactericera cockerelli
(Homoptera: Psyllidae) with ‘‘zebra chip’’, a new potato disease in southwestern
United States and Mexico. J Econ Entomol. 100:656–663.
Mustafa TM. 1984. Factors affecting the distribution of Euphyllura olivina (Hom. Psyllidae)
on olive. Z Angew Entomol. 97:371–375.
Mustafa TM. 1989a. Bionomics of the olive psylla Eupyllura olivina Costa (Hom. Psyllidae) in
Jordan. J Biol Sci Res. 20:159–166.
Mustafa TM. 1989b. Seasonal changes in fat and water content in field populations of olive
psylla Euphyllura olivina Costa (Homoptera, Psyllidae). Insect Sci Appl. 10:181–186.
Mustafa TM, Hodgson CJ. 1984. Observation, diapause induction and termination in Psylla
pyricola in England. Lutte integré contre les psylles du poirier, Bull OILB/SROP,
127–138.
Mustafa TM, Najar YH. 1985. Contributions to the reproductive biology of olive psylla
Euphyllura olivina (Homoptera, Psyllidae). Z Angew Entomol. 100:79–83.
Naeem A, Behad E. 1988. The biology of the gas psyllid. Entomol Phytopathol A. 55:111–121.
Nakata T. 2006. Temperature-dependent development of the citrus psyllid, Diaphorina citri
(Homoptera: Psylloidea), and the predicted limit of its spread based on overwintering in
the nymphal stage in temperate regions of Japan. Appl Entomol Zool. 41:383–387.
Nava DE, Torres MLG, Rodrigues MDL, Bento JMS, Parra JRP. 2007. Biology of
Diaphorina citri (Hem., Psyllidae) on different hosts and at different temperatures. J Appl
Entomol. 131:709–715.
Negi DS, Bisht RS. 1989. On the biology of the psyllid Pauropsylla depressa Crawford
(Homoptera: Psyllidae). Uttar Pradesh J Zool. 9:253–257.
Nguyen T-X. 1963. Note préliminaire sur quelques psyllides du sud-ouest de la France. Rev
Pathol Veg Entomol Agri Fr. 42:169–176.
Nguyen T-X. 1964. Observations préliminaires concernant l’élimination de la diapause chez
Psylla pyri L. Rev Pathol Veg Entomol Agri Fr. 43:3–12.
Nguyen T-X. 1965. Observations sur la ponte preferentielle de Psylla buxi L (Homopteres
Psyllides) sur les differentes variétés de buis. Bull Soc Hist Nat Toulouse. 100:299–311.
Nguyen T-X. 1967a. Influence des facteurs externes sur l’élimination anticipée de la diapause
de Psylla pyri L. (Homoptères-Psyllidae). Comptes Rendus Acad Sci, Paris Ser D.
264:1445–1448.
Nguyen T-X. 1967b. Observations sur l’élimination de la diapause de Psylla pyri (Hom.
Psyllidae) dans les conditions naturelles. Ann Soc Entomol Fr (New Ser.) 3:151–164.
Nguyen T-X. 1968. Rôle de la temperature dans l’évolution et élimination de la diapause
larvaire de Psylla buxi (Hom. Psyllidae). Ann Soc Entomol Fr (New Ser.) 4:69–74.
Nguyen T-X. 1969. Étude de la resistance au froid et de la capacité d’acclimation de
Psylla buxi L (Homoptera: Psyllidae). Comptes Rendus Acad Sci, Paris Ser D. 268:
1410–1413.
Nguyen T-X. 1970a. Influence de la temperature et de la photopériode sur la reproduction
d’un psylle du poirier, Psylla pyri L. (Insecte: Homoptera-Psyllidae). Comptes Rendus
des Academie de Sciences, Paris Ser D. 271:2336–2338.
Nguyen T-X. 1970b. Récherches sur la morphologie et la biologie de Psyllopsis fraxini (Hom.
Psyllidae). Ann Soc Entomol Fr (New Ser.) 6:757–773.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
169
Nguyen T-X. 1971. Effet de groupement sur la reproduction et la longevité de Psylla pyri L.
(Insectes, Homoptera-Psyllidae). Comptes Rendus Acad Sci, Paris Ser D. 272:1782–1784.
Nguyen T-X. 1972a. Études de la diapause imaginale de Psylla pyri L. (Homoptera: Psyllidae)
1. Determinisme du polymorphisme saisonier des adultes. Ann Zool Ecol Anim.
4:281–309.
Nguyen T-X. 1972b. Influence de la nature des plantes hôtes sur la longevité et la fecondité de
Psylla pyri (L.) (Insecte, Homoptera-Psyllidae). Comptes Rendus Acad Sci, Paris Ser D.
274:546–548.
Nguyen T-X. 1973. Action de surpeuplement sur la reproduction de Psylla pyri (L.) (Insectes,
Homoptere, Psyllidae). Comptes Rendus Acad Sci, Paris Ser D. 276:2389–2390.
Nguyen T-X. 1975. Évolution et la termination de la diapause ovarienne de Psylla pyri
(Homoptera-Psyllidae) dans les conditions naturelles de la region Toulousaine. Bull Soc
Zool Fr. 100:241–246.
Nguyen T-X. 1985. Établissement d’une échelle morphométrique pour les Psyllidae (InsectaHomoptera); polymorphisme saisonnier de Psylla pyri L. Comptes Rendus Acad Sci,
Paris Ser D. 301:369–372.
Nguyen T-X, Ledoux A. 1973. Évolution de la diapause ovarienne des femelles de Psylla pyri
L. (Homoptera: Psyllidae) pendant la periode automne-hiver. Comptes Rendus Acad Sci,
Paris Ser D. 277:2385–2387.
Nguyen T-X, Messi J. 1973. Observations sur la ponte des femelles de Arytaina genistae elevée
sur des genets d’origine differente. Comptes Rendus Acad Sci, Paris Ser D. 276:
1691–1696.
Nissinen A, Ibrahim M, Kainulainen P, Tiilikkala K, Holopainen JK. 2005. Influence of
carrot psyllid (Trioza apicalis) feeding or exogenous limonene or methyl jasmonate
treatment on composition of carrot (Daucus carota) leaf essential oil and headspace
volatiles. J Agri Food Chem. 53:8631–8638.
Nissinen A, Vanhala P, Holopainen JK, Tiilikkala K. 2007. Short feeding period of carrot
psyllid (Trioza apicalis) females at early growth stages of carrot reduces yield and causes
leaf discolouration. Entomol Exp Appl. 125:277–283.
Nucifora A. 1969. La Psylla piri (L.) nei frutteti dell’Etna. Tecnica Agri, Catania 21:
348–361.
Ogol CKPO, Spence JR. 1997. Abundance, population dynamics and impact of the leucaena
psyllid Heteropsylla cubana Crawford in a maize-leucaena agroforestry system in Kenya.
Insect Sci Appl. 17:183–192.
Oka IN, Bahgiawati AH. 1988. Comprehensive program towards integrated control of
Leucaena psyllid, a new insect pest of Leucaenca trees in Indonesia. Indones Agri Res
Devel J. 10:23–30.
Oldfield GN. 1970. Diapause and polymorphism in California populations of Psylla pyricola
(Homoptera. Psyllidae). Ann Entomol Soc Am. 63:180–184.
Olivares TS. 2000. Ctenarytaina eucalypti (Maskell,1890): el psilido de eucalipto en Chile
(Hemiptera: Sternorryncha: Psylloidea: Spondysliaspinae). Gayana 64:239–241.
Omole MM. 1980. Attraction to colour traps and height of flight in Trioza urticae (Hemiptera,
Heteroptera, Psyllidae). Entomol Mon Mag. 116:173–175.
Onillon JC. 1969. Effect of the numerical density of Trioza urticae (Homoptera, Psyllidae) on
the ovarian activity of Tetrastichus upis (Hymenoptera, Tetrastichidae). In: Labeyrie V,
editor. Colloques internationaux de centre national de la récherche scientifique No. 189.
l’influence des stimuli externes sur la gametogenese des insectes. Paris (France): Centre
National de la Récherche Scientifique. p. 57–70.
Osborn H. 1922. Life history notes on Cranberry Lake Homoptera. Technical Publication of
the New York State College of Forestry at Syracuse 16:87–104.
Osisanya EO. 1974a. Aspects of the biology of Diclidophlebia eastopi and Diclidophlebia
harrisoni (Homoptera: Psyllidae). Bull Entomol Res. 64:9–18.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
170 I.D. Hodkinson
Osisanya EO. 1974b. Seasonal fluctuations of a natural population of Diclidophlebia eastopi
and Diclidophlebia harrisoni (Homoptera: Psyllidae). Bull Entomol Res. 64:289–296.
Ossiannilsson F. 1943. Studier over de Svenska potatifaltens insectfauna och dess betyldelse
for spridning av virussjukdomar. 1. Hemiptera, Forekmost och utbredning. Statens
Vaxtskyddsanstalt Medd. 39:1–72.
Ossiannilsson F. 1950. Sound production in psyllids (Hem. Hom.). Opusc Entomol. 11:82–84.
Ossiannilsson F. 1992. The Psylloidea (Homoptera) of Fennoscandia and Denmark. Fauna
Entomol Scandin. 26:1–347.
Palmer WA, Witt ABR. 2006. On the host range and biology of Acizzia melanocephala
(Hemiptera: Psyllidae), an insect rejected for the biological control of Acacia nilotica
subsp indica (Mimosaceae) in Australia. Afr Entomol. 14:387–390.
Pande YD. 1971. Biology of citrus psylla Diaphorina citri (Hemiptera, Homoptera, Psyllidae).
Israel J Entomol. 6:307–311.
Papp RP, Johnson JB. 1979. Origins of psyllid fallout in the central Sierra-Nevada of
California. Pan-Pac Entomol. 55:95–98.
Parkinson JD, Whittaker JB. 1975. A study of two physiological races of the heather psyllid
Strophingia ericae (Homoptera, Psylloidea). Biol J Linn Soc. 7:73–81.
Pasqualini E, Civolani S, Musacchi S, Ancarani V, Dondini L, Robert P, Baronio P. 2006.
Cacopsylla pyri behaviour on new pear selections for host resistance programs. Bull
Insectol. 59:27–37.
Patch EM. 1909. Downy psyllid of alder, Psylla floccosa, new species. Can Entomol.
41:301–303.
Patil NG, Baker PS, Pollard GV. 1994. Life history parameters of the leucaena psyllid
Heteropsylla cubana (Crawford) (Homoptera: Psyllidae) under various temperature and
relative humidity regimes. Insect Sci Appl. 15:293–299.
Pava OJ, Gonzalez OA, Castillo CE, Patino CH. 1983. Interesting phyto sanitary aspects of
the chontaduro palm Bactris gasipaes in some regions of Valle and Choco, Colombia.
Acta Agron Palmira 33:25–36.
Peck SB. 1994. Aerial dispersal of insects between and to islands in the Galapagos
Archipelago, Ecuador. Ann Entomol Soc Am. 87:218–224.
Pedata PA. 1998. Morpho-biological observations on Egeirotrioza (Astutia) populi Horvath,
new for Italy. Atti Accad Roveretana Degli Agiati (Series VII) 8B:75–86.
Percy DM. 2003a. Legume-feeding psyllids (Hemiptera, Psylloidea) of the Canary Islands and
Madeira. J Nat Hist. 37:397–461.
Percy DM. 2003b. Radiation, diversity, and host–plant interactions among island and
continental legume-feeding psyllids. Evolution 57:2540–2556.
Percy DM, Page RDM, Cronk QCB. 2004. Plant–insect interactions: Double-dating
associated insect and plant lineages reveals asynchronous radiations. Syst Biol.
53:120–127.
Percy DM, Taylor GS, Kennedy M. 2006. Psyllid communication: acoustic diversity, mate
recognition and phylogenetic signal. Invertebr Syst 20:431–445.
Perez Otero R, Mansilla Vazquez JP, Mansilla Salinero P. 2005. Distribucion y biologia de
Ctenarytaina spatulata Taylor sobre Eucalyptus globulus Labill. en la provincia de
Pontevedra. Bol Sanid Veg Plagas. 31:25–30.
Petersen A. 1923. The blackberry psyllid, Trioza tripunctata Fitch. Bull NJ Agri Exp Stn.
378:1–32.
Pfeiffer DG, Burts EC. 1983. Effects of tree fertilization on numbers and development of pear
psylla Psylla pyricola (Homoptera: Psyllidae) and on fruit damage. Environ Entomol.
12:895–901.
Pfeiffer DG, Burts EC. 1984. Effect of tree fertilization on protein and free amino-acid content
and feeding rate of pear psylla Psylla pyricola (Homoptera Psyllidae). Environ Entomol.
13:1487–1490.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
171
Pintera A. 1982. Some observations on the ecology and population dynamics of psyllas found
on cedar. Poeyana Inst Zool Acad Ciencias Cuba. 252:1–12.
Pletsch DJ. 1947. The potato psyllid Paratrioza cockerelli (Šulc). Its biology and control. Bull
Montana Agri Exp Stn. 446:1–95.
Prasad D. 1957. On the distribution, bionomics, and control of the mango shoot gall psyllid,
Apsylla cistellata Buckton. Indian J Entomol. 19:78–83.
Prophetou DA. 1993. Diapause termination and phenology of the olive psyllid, Euphyllura
phillyreae on two host plants in coastal northern Greece. Entomol Exp Appl. 67:
193–197.
Prophetou DA. 1997. Occurrence of immature stages of olive psyllid Euphyllura phillyreae
(Hom, Aphalaridae) in Phillyrea latifolia and Olea europaea in coastal northern Greece. J
Appl Entomol. 121:383–387.
Prophetou DA, Tzanakakis ME. 1977. Seasonal development and number of generations of
Euphyllura olivina (Hemiptera-Psyllidae) in Halkidiki (N Greece). Ann Entomol Soc Am.
70:707–710.
Prophetou DA, Tzanakakis ME. 1986. Diapause termination in the olive psyllid Euphyllura
phillyreae in the field and in the laboratory. Entomol Exp Appl. 40:263–272.
Przybylski Z. 1970. Studies on the synchronization of phytophenological phenomenon with
the development of Psylla mali (Psyllidae). Ekol Polska Ser A. 18:1–40.
Purcell MF, Balciunas JK, Jones P. 1997. Biology and host-range of Boreioglycaspis
melaleucae (Hemiptera: Psyllidae), potential biological control agent for Melaleuca
quinquenervia (Myrtaceae). Environ Entomol. 26:366–372.
Purcell AH, Suslow KG. 1984. Surveys of leafhoppers (Homoptera: Cicadellidae) and pear
psylla (Homoptera: Psyllidae) in pear and peach orchards and the spread of peach yellow
leaf roll disease. J Econ Entomol. 77:1489–1494.
Purvis G, Chauzat MP, Segonds-Pichon A, Dunne R. 2002. Life history and phenology of the
eucalyptus psyllid, Ctenarytaina eucalypti in Ireland. Ann Appl Biol. 141:283–292.
Pussard MR. 1939. Contribution a l’étude de la nutrition des psyllides. Presence d’une amylase
dans les glandes salivaires de quelques psyllides adultes. Comptes Rendus Soc Savantes
Paris 71:291–292.
Puterka GJ. 1997. Intraspecific variation in pear psylla (Psyllidae: Homoptera) nymphal
survival and development on resistant and susceptible pear. Environ Entomol.
26:552–558.
Puterka GJ, Bell RL, Jones SK. 1993. Ovipositional preference of pear psylla (Homoptera,
Psyllidae) for resistant and susceptible pear. J Econ Entomol. 86:1297–1302.
Radjabi G, Behechti ND. 1975. Bioecological studies and control of Psylla pyricola
(Homoptera, Psyllidae) in Esfahan. Entomol Phytopathol Appl. 39:39–53.
Rajadurai S, Mani T, Balakrishna P, Raman A. 1990. On the digestive enzymes and soluble
proteins of the nymphal salivary glands of Trioza jambolanae Crawford (Triozinae:
Psyllidae: Homoptera), the gall maker on the leaves of Syzygium cumini (L.) Skeels
(Myrtaceae). Phytophaga 3:47–53.
Rajapakse RHS, Kulasekera K. 1982. Some observations on the insect pests of cinnamon in
Sri Lanka. Entomon 7:221–223.
Raman A. 1987. On the cecidogenesis and nutritive tissues of the leaf galls of Garuga pinnata
Roxburgh (Burseraceae) induced by Phacopteron lentiginosum Buckton (Pauropsyllinae:
Psyllidae: Homoptera). Phytophaga 1:141–159.
Raman A. 1991. Cecidogenesis of leaf galls on Syzygium cumini (L.) Skeels (Myrtaceae)
induced by Trioza jambolanae Crawford (Homoptera, Psylloidea). J Nat Hist.
25:653–663.
Raman A. 1994. Adaptational integration between gall-inducing insects and their host plants.
In: Ananthakkrishnan TN, editor. Functional dynamics of phytophagous insects. New
Delhi (India): Oxford & IBH Publishing Co. p. 249–275.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
172 I.D. Hodkinson
Raman A. 2003. Cecidogenetic behavior of some gall-inducing thrips, psyllids, coccids, and
gall midges, and morphogenesis of their galls. Oriental Insects 37:359–413.
Ramert B, Nehlin G. 1989. Alternative plant protection methods in small scale cultivation.
Vaxtskyddsnotiser Suppl. 2:1–39.
Ramirez Gomez C. 1956. Los psilidos de Espana. Bol Real Soc Esp Hist Nat (Biol.)
53:151–219.
Ramirez Gomez C. 1958. Los psilidos de Espana. Bol Real Soc Esp Hist Nat (Biol.)
54:63–106.
Ramirez Gomez C. 1960. Los psilidos de Espana. Bol Real Soc Esp Hist Nat (Biol.) 57:4–87.
Rapisarda C. 1985. Sulla presenza in Italia di Acizzia acaciae baileyanae (Froggatt)
(Homoptera,Psylloidea), nuovo parassita di acacie ornimentali. Inf Fitopatol. 35:45–49.
Rapisarda C. 1988. Note tassinomiche e bio-ecologiche sulle specie di psilloidei viventi su
Genistinae in Sicilia (Homoptera). In: Proceedings of the XV Italian National Congress
of Entomology, l’Aquila. p. 497–504.
Rapisarda C. 1989a. Contributo alla conoscenza degli psilloidei viventi su Rhamnus alaternus
L. in Italia (Homoptera: Psylloidea). Phytophaga 3:21–60.
Rapisarda C. 1989b. Presenza in Sicilia della psilla verde del fico (Homotoma viridis
Klimaszewski) e descrizione della sua ninfa di ultima eta (Homoptera: Psylloidea). Bol
Soc Entomol Ital. 121:13–18.
Rapisarda C. 1989c. A new Sicilian species of Arytainilla Loginova (Homoptera: Psylloidea),
from Genista aetensis (Biv.) DC, with notes on its biology. Frustula Entomol (New Ser.)
9:23–39.
Rapisarda C. 1990a. Faunistic and ecological notes on the psyllids of Sardinia (Homoptera:
Psylloidea). Mem Soc Entomol Ital. 69:7–52.
Rapisarda C. 1990b. Morphological and biological notes on Arytainilla cytisi (Homoptera,
Psylloidea). Phytophaga 3:115–130.
Rapisarda C. 1992. Nymphal description and life-cycle of Livilla retamae (Puton) and Livilla
spectabilis (Flor) (Homoptera Psylloidea). Boll Zool Agrar Bachic. 24:195–204.
Rapisarda C. 1993a. Seasonal dimorphism in two Acacia-feeding psyllids of the grenus Acizzia
(Homoptera: Psylloidea). Phytophaga 3:131–146.
Rapisarda C. 1993b. Trioza soniae and T. apulica n. spp., on Quercus spp., from southern
Italy. Mem Soc Entomol Ital. 72:175–188.
Rapisarda C. 1998. New data on the Sicilian psyllids (Homoptera: Psylloidea). Atti Accad
Roveretana Degli Agiati (Ser VII). 8B:97–104.
Rapisarda C, Belcari A. 1999. Notes on some psyllids (Homoptera: Psylloidea) infesting urban
trees in Italy. In: Lemattre M, Lemattre P, Lemaire F, editors. International symposium
on urban tree health. Louvain (Belgium): International Society Horticultural Science. p.
155–164.
Rasmy AM, MacPhee AW. 1970. Studies on the pear psylla in Nova Scotia. Can Entomol.
102:586–591.
Rauf A, Hidayat P, Maryana N, Winasa IW. 1990. Biology and demography of Heteropsylla
cubana Crawford (Homoptera: Psyllidae). In: Napompeth NA, editor, Leucaena psyllid:
problems and management. Bangkok: Winrock Institute of Agricultural Development. p.
114–118.
Reuter OM. 1909. Charakteristik und Entwickelungeschichte der Hemipteren-Fauna
(Heteroptera, Auchenorryncha und Psyllidae) der Palaearctischen Coniferen. Acta Soc
Sci Fenn. 36:1–129.
Richards BL, Blood HL. 1933. Psyllid yellows of the potato. J Agri Res. 46:189–216.
Riemann JG. 1958. Notes on the host and biology of Tetragonocephala flava Crawford
(Homoptera: Psyllidae). J Kansas Enomol Soc. 31:256.
Rieux R, D’Arcier FF. 1990. Seasonal polymorphism of Psylla pyri L. (Homoptera, Psyllidae)
adults in natural populations. J Appl Entomol. 109:120–135.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
173
Riley CV. 1890. Hackberry psyllids. In: Packard AS, editor. Forest and shade tree insects.
United States Entomological Commission 5th Report, Washington. p. 614–622.
Ross WA. 1919. The pear psylla in Ontario. Annu Rep Entomol Soc Ont. 49:81–90.
Russell LM. 1971. Notes on Euphalerus nidifex Schwarz and related nest-making New World
psyllids. Fl Entomol. 54:3–12.
Rygg T. 1977. Biological investigations on carrot psyllid Trioza apicalis Forster (Homoptera,
Triozidae). Meldinger Nor Landbrukshogskole 56:1–20.
Saiz F, Nunez C. 2000. Cecidias de hoja y de rama de Schinus polygamus (Cav.) Cabr.
(Anacardiaceae): doble agente formador o secuencia temporal de cecidias formadas por
huespedes diferentes? Rev Chilena Entomol. 27:57–63.
Sampo A. 1975. Di alcuni psillocecidi nuovi o poci noti della valle d’Aosta. Rev Valdotaine
Hist Nat (Aosta) 29:153–174.
Sampo A. 1977. La psilla del lauro (Trioza alacris Flor). Il Floricultore, Milan 14:79–83.
Samways MJ. 1987. Weather and monitoring the abundance of the adult citrus psylla Trioza
erytreae Del Guercio (Homoptera, Triozidae). J Appl Entomol. 103:502–508.
Sanford GB. 1952. Phloem necrosis of potato tubers associated with infestation of vines by
Paratrioza cockerelli Šulc. Sci Agri. 32:433–439.
Sasaki C. 1910. On the life history of Trioza camphorae n.sp. of camphor tree and its injuries. J
Coll Agri Imperial Univ Tokyo. 2:277–285.
Savinelli CE, Tetrault RC. 1984. Analysis of pear psylla (Homoptera: Psyllidae) populations
and associated damage in a Pennsylvania pear orchard. Environ Entomol. 13:278–281.
Schaefer HA. 1949. Biologische und okologische Beobachtungen an Psylliden (Hemiptera).
Verhandlungen Naturforschenden Ges Basel 60:25–41.
Schaub L, Graf B, Butturini A. 2005. Phenological model of pear psylla Cacopsylla pyri.
Entomol Exp Appl. 117:105–111.
Schmidt E. 1966. Cycle biologique et phases cecidogenes de Livia juncorum Latr. Marcellia
33:223–235.
Scutareanu P, Loxdale HD. 2006. Ratio of nutrient and minerals to defensive compounds
indicative of plant quality and tolerance to herbivory in pear trees. J Plant Nutr.
29:629–642.
Shahid M, Khan S. 1976. Biology and control of Diaphorina citri Kuw (citrus psylla) in
Pashawar. Pakistan J Sci Res. 28:24–28.
Siddiqui ZA. 1949. Occurrence of Psyllia pyricola Forst on apple and pear trees in Kamaum.
Indian J Entomol. 8:237.
Silvestri F. 1934. Psyllidae. Compendio di Entomologia Applicata. Portici: Tipografia
Ballavista. p. 368–387.
Singh G, Misra PN. 1978. The mango shoot gall psyllid Apsylla cistellata Buckton and its
control. Pesticides 12:15–16.
Singh MP. 1959. Studies on the mango shoot gall psyllid Apsylla cistellata 1. Description of
development stages and habits. Indian J Entomol. 21:273–281.
Singh P. 1988. Heteropsylla cubana Crawford, a new pest of Leucaena in India. Indian For.
114:200–205.
Singh SM. 1954. Studies on Apsylla cistellata Buckton causing mango galls in India. J Econ
Entomol. 47:563–564.
Singh SM. 1960. Studies on the mango shoot gall in Tarai region of UP, its causes and control.
II Distribution, nature, extent, intensity of damage and bionomics of the pest. Horti Adv.
4:97–114.
Sirrine FA. 1895. The bramble flea louse. Annu Rep NY State Agri Exp Stn. 14:619–623.
Skanland HT, Sömme L. 1981. Seasonal-variation in cold-hardiness of eggs of the apple
psyllid Psylla mali (Schmidb.) in Norway. Cryo-Letters 2:87–92.
Slingerland MV. 1892. The pear tree psylla. Cornell Univ Agri Exp Stn Bull. 44:161–186.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
174 I.D. Hodkinson
Smith JB. 1911. The bramble flea louse Trioza tripunctata Fitch. Annu Rep NJ Agri Exp Stn.
32:416–418.
Smith RC, Taylor RS. 1953. The biology and control of the hackberry psyllids in Kansas. J
Kansas Enomol Soc. 26:103–115.
Solomon ME. 1936. Description and life-history of a new western Australian psyllid. J R Soc
West Aust. 22:41–48.
Sorin M. 1959a. On the life history and immature stages of Trioza camphorae Sasaki
(Psyllidae, Homoptera). Kontyu 27:244–248.
Sorin M. 1959b. On the life history and immature stages of Trioza remota Foerster
(Homoptera: Psyllidae). Kontyu 27:181–186.
Souliotis C, Broumas T. 1998. Study of the population dynamics of the Cacopsylla pyri L.
(Homoptera: Psyllidae) and its predators in Eastern Attiki. Ann Inst Phytophathol
Benaki. 18:97–109.
Souliotis C, Tsourgianni A. 2000. Population dynamics of Psyllidae on pistachio (Pistacia
vera): Bioecological data on Agonoscena pistaciae Burck. and Laut. (Homop.
Sternorrhyncha). Boll Zool Agrar Bachic. 32:49–58.
Spaulding AW, von Dohlen CD. 1998. Phylogenetic characterization and molecular evolution
of bacterial endosymbionts in psyllids (Hemiptera: Sternorrhyncha). Mol Biol Evol.
15:1506–1513.
Spaulding AW, von Dohlen CD. 2001. Psyllid endosymbionts exhibit patterns of co-speciation
with hosts and destabilizing substitutions in ribosomal RNA. Insect Mol Biol. 10:57–67.
Speyer W. 1929. Der Apfelblattsauger, Psylla mali Schmidb. Berlin (Germany): Springer.
Staples R. 1968. Cross protection between a plant virus and potato psyllid yellows. J Econ
Entomol. 61:1378–1380.
Stavraki HG. 1980. Biologie de Euphyllura sp. (Homoptera: Psyllidae) dans un oliveraie
d’attiki (Greece). Mededelingen van de Faculteit Landbouwwetenschappen, Universiteit
Gent 45:603–611.
Stratopoulou ET, Kapatos ET. 1995a. Distribution of population of immature stages of pear
psylla, Cacopsylla pyri, within the tree and development of sampling strategy. Entomol
Hell. 10:5–10.
Stratopoulou ET, Kapatos ET. 1995b. The dynamics of the adult population of pear psylla,
Cacopsylla pyri L. (Hom., Psyllidae) in the region of Magnesia (Greece). J Appl Entomol.
119:97–101.
Stromgren ES, Lanciani CA. 2001. Early abcission in hackberry leaves bearing Pachypsylla
galls (Homoptera: Psyllidae). Fl Entomol. 84:727–728.
Stuart LC. 1991. Blackberry psyllid. In: Ellis MA, Converse RH, Williams RN, Williamson B,
editors. Compendium of raspberry and blackberry diseases and insects. St. Paul (MN):
American Phytological Society Press. p. 73–74.
Stuart LC, Butt BA, Bell RL. 1989. Effect of host phenology on ovipositional preference of
winter form pear psylla (Homoptera, Psyllidae). J Entomol Soc BC. 86:34–38.
Sutton RD. 1983. Seasonal colour changes, sexual maturation and oviposition in Psylla
peregrina (Homoptera: Psylloidea). Ecol Entomol. 8:195–202.
Sutton RD. 1984. The effect of host plant flowering on the distribution and growth of
hawthorn psyllids (Homoptera, Psylloidea). J Anim Ecol. 53:37–50.
Swenk MH, Tate HD. 1940. The potato flea beetle and the potato psyllid in Nebraska. Bull
Neb Agri Exp Stn. 327:1–19.
Takara JM, Dinker RJ, Nagamine WT, Teramoto KK. 1990. Biology and reproductive rate of
leucaena psyllid, Heteropsylla cubana Crawford. Proc Hawaiian Entomol Soc. 30:23–30.
Tamesse JL, Messi J. 2004. Facteurs influencant la dynamique des populations du psylle
africain des agrumes Trioza erytreae Del Guercio (Hemiptera: Triozidae) au Cameroun.
Int J Trop Insect Sci. 24:213–227.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
175
Tarsia in Curia I. 1934. La simbiosi ereditaria in Trioza alacris Flor. Archiv Zool Ital.
2:215–235.
Taylor GS. 1985. The taxonomic status of Schedotrioza multitudinea (Psylloidea: Triozidae)
with notes on its biology. J Aust Entomol Soc. 24:305–312.
Taylor GS. 1987. The gall forming Psylloidea of Eucalyptus obliqua in the Mount Lofty
Ranges of South Australia. J Aust Entomol Soc. 26:223–228.
Taylor KL. 1985. A possible stridulatory organ in some Psylloidea (Homoptera). J Aust
Entomol Soc. 24:77–80.
Tedeschi R, Alma A. 2004. Transmission of apple proliferation phytoplasma by Cacopsylla
melanoneura (Homoptera: Psyllidae). J Econ Entomol. 97:8–13.
Tedeschi R, Bosco D, Alma A. 2002. Population dynamics of Cacopsylla melanoneura
(Homoptera: Psyllidae), a vector of apple proliferation phytoplasma in northwestern
Italy. J Econ Entomol. 95:544–551.
Tedeschi R, Ferrato V, Rossi J, Alma A. 2006. Possible phytoplasma transovarial
transmission in the psyllids Cacopsylla melanoneura and Cacopsylla pruni. Plant
Pathol. 55:18–24.
Teixeira DD, Danet JL, Eveillard S, Martins EC, Junior WCJ, Yamamoto PT, Lopes SA,
Bassanezi RB, Ayres AJ, Saillard C. 2005. Citrus huanglongbing in Sao Paulo State,
Brazil: PCR detection of the ‘‘Candidatus’’ Liberibacter species associated with the
disease. Mol Cell Probes. 19:173–179.
Tenorio J, Chuquillanqui C, Garcia A, Guillen M, Chavez R, Salazar LF. 2003.
Sintomatologia y efecto en el rendimiento de papa por el achaparramiento rugoso.
Fitopatologia 38:32–36.
Thakur JR, Sharma PC, Gupta PR. 1989. Ocurrence of olive psylla Euphyllura pakistanika
Loginova (Homoptera: Aphalaridae) on olive in India, a new record. Trop Pest Manag.
35:331.
Thao ML, Clark MA, Baumann L, Brennan EB, Moran NA, Baumann P. 2000a. Secondary
endosymbionts of psyllids have heen acquired multiple times. Curr Microbiol.
41:300–304.
Thao ML, Moran NA, Abbot P, Brennan EB, Burckhardt DH, Baumann P. 2000b.
Cospeciation of psyllids and their primary prokaryotic endosymbionts. Appl Environ
Microbiol. 66:2898–2905.
Thao ML, Clark MA, Burckhardt DH, Moran NA, Baumann P. 2001. Phylogenetic analysis
of vertically transmitted psyllid endosymbionts (Candidatus Carsonella ruddii) based on
atpAGD and rpoC: Comparisons with 16S–23S rDNA-derived phylogeny. Curr
Microbiol. 42:419–421.
Thenmozhi K, Kandasamy C. 1992. Studies of the biology of gall psyllids of India 1.
Pauropsylla longispiculata Mathur (Psyllidae, Homoptera). Agri Biol Res. 8:44–47.
Tishechkin DY. 1989. Acoustic signalization in the psyllids (Homoptera, Psyllinea) from the
Moscow Oblast, Russian SSR. Vestnik Moskovskogo Universiteta Seriya XVI Biologiya,
20–24.
Tishechkin DY. 2005. Vibratory communication in Psylloidea (Hemiptera). In: Drosopoulos
S, Claridge MM, editors. Insect sounds and communications. London: CRC Taylor &
Francis. p. 357–363.
Tishechkin DY. 2007. New data on vibratory communication in jumping plant lice of the
families Aphalaridae and Triozidae (Homoptera, Psyllinea). Entomol Rev. 87:394–400.
Tokmakoglu C. 1973. The studies on some features of biology and control methods of
Agonoscena targionii Licht. Bitki Koruma Bülteni 13:67–72.
Tremblay E. 1958. La Trioza nigricornis Foerster (Nota preliminare). Boll Lab Entomol Agrar
Filippo Silvestri, Portici. 16:160–170.
Tremblay E. 1961. La psilla e il misurino della cipolla. Laboratorio Enomologia Agrar ‘‘F.
Silvestri’’ Oss Mal Piante, Portici, Circ. 24:1–10.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
176 I.D. Hodkinson
Tremblay E. 1965a. Risultati di prove di lotta contro la psilla della cipolla (Trioza tremblayi
Wagner). Ann Fac Sci Agrar Univ Napoli. 30:15–27.
Tremblay E. 1965b. Studio morfo-biologico sulla Trioza tremblayi Wagner (Hemiptera:
Homomptera, Psyllidae). Boll Lab Entomol Agrar Filippo Silvestri, Portici. 23:37–138.
Tsai JH, Liu YH. 2000. Biology of Diaphorina citri (Homoptera: Psyllidae) on four host
plants. J Econ Entomol. 93:1721–1725.
Tuncer C. 2002. Investigations on the biology of Homotoma ficus L. (Homoptera:
Homotomidae) and the development threshold for eggs in Samsun Province. Turkiye
Entomol Dergisi 26:33–44.
Tuthill LD. 1952. On the Psyllidae of New Zealand. Pac Sci. 6:83–125.
Tzanakakis ME. 2003. Seasonal development and dormancy of insects and mites feeding on
olive: a review. Neth J Zool. 52:87–224.
Tzanakakis ME. 2006. Insects and mites feeding on olive. Distribution, importance, habits,
seasonal development and dormancy. Leiden (The Netherlands): Brill.
Uchida GK, Beardsley JWJ. 1988. Taxonomy and biology of Megatrioza palmicola group
(Homoptera, Psyllidae) in Hawaii. Proc Hawaiian Entomol Soc. 28:57–100.
Ullman DE, McLean DL. 1988a. Feeding behavior of the winter-form pear psylla Psylla
pyricola (Homoptera, Psyllidae) on reproductive and transitory host plants. Environ
Entomol. 17:675–678.
Ullman DE, McLean DL. 1988b. The probing behavior of the summer-form pear psylla.
Entomol Exp Appl. 47(2):115–126.
Unruh TR, Krysan JL. 1994. Reproductive diapause and host plants afffect insecticide
tolerance of adult pear psylla (Homoptera, Psyllidae). J Econ Entomol. 87:858–865.
Vaishampayan SM, Bahadur A. 1980. Observations on biology and bionomics of tendu gall
fly, Trioza obsoleta Buckton (Homoptera: Psyllidae), and assessment of losses to tendu
Diospyros melanoxylon Roxb. Proceedings of the 2nd Forestry Conference Dehra Dun,
India. p. 1–7.
van den Berg MA. 1990. The citrus psylla Trioza erytreae del Guercio (Hemiptera: Triozidae),
a review. Agri Ecosyst Environ. 30:171–194.
van den Berg MA, Anderson SH, Deacon VE. 1991. Population studies of the citrus psylla
Trioza erytreae: factors influencing dispersal. Phytoparasitica 19:283–290.
van den Berg MA, Deacon VE. 1988. Dispersal of the citrus psylla Trioza erytreae
(Hemiptera, Triozidae) in the absence of its host plants. Phytophylactica 20:361–368.
van den Berg MA, Deacon VE. 1989. Flight activities of the citrus psylla Trioza erytreae
(Hemiptera, Triozidae). Phytophylactica 21:391–396.
van den Berg MA, Deacon VE, Steenekamp PJ. 1991. Dispersal within and between citrus
orchards and native hosts and nymphal mortality of citrus psylla Trioza erytreae
(Hemiptera, Triozidae). Agri Ecosyst Environ. 35:297–310.
van Den Berg MA, Deacon VE, Thomas CD. 1991a. Ecology of the citrus psylla Trioza
erytreae (Hemiptera Triozidae) 3. Mating, fertility and oviposition. Phytophylactica
23:195–200.
van den Berg MA, Deacon VE, Thomas CD. 1991b. Ecology of the citrus psylla Trioza
erytreae (Hemiptera, Triozidae) 4. Settling and general behaviour of nymphs.
Phytophylactica 23:201–206.
van den Berg MA, de Villiers EA. 1987. Psyllids, aphids and whiteflies. Bull Dep Agri Water
Supply Repub S Afr. 411:43–48.
van den Berg MA, van Vuuren SP, Deacon VE. 1987. Cross-breeding and greening disease
transmission of different populations of the citrus psylla Trioza erytreae (Hemiptera:
Triozidae). Phytophylactica 19:353–354.
van Klinken RD. 2000. Host-specificity constrains evolutionary host change in the psyllid
Prosopidopsylla flava. Ecol Entomol. 25:413–422.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Journal of Natural History
177
van Vuuren SP, Moll JN. 1984. Population dynamics and greening transmission by the psyllid
Trioza erytreae (Del Guercio). In: Proceedings of the greening symposium, Citrus &
Subtropical Fruit Research Institute, Nelspruit, p. 90–94.
Verrier ML. 1929. Contribution a l’étude de la cecidie de Livia juncorum Latr. (Hem Psyllidae)
sur Juncus conglomeratus L. Bull Soc Entomol Fr. 19:77–80.
Vosseler J. 1906. Eine Psyllide als Erzeugerin von Gallen am Mwulebaum (Chlorophora
excelsa (Welw.) Benth et Hook). Z Wiss Insectenbiol. 2:276–285 and 305–316.
Waku Y, Endo Y. 1987. Ultrastructure and life cycle of the symbionts in a Homopteran Insect
Anomoneura mori Schwartz (Psyllidae). Appl Entomol Zool. 22:630–637.
Wallis RL. 1946. Seasonal occurrence of the potato psyllid in the North Platte Valley. J Econ
Entomol. 39:689–694.
Wallis RL. 1955. Ecological studies on the potato psyllid as a pest of potatoes. Tech Bull US
Dep Agri. 1107:1–24.
Walton BCJ. 1944. The biology of Pachypsylla celtis-gemma Riley (Homoptera: Psyllidae) and
the relationship of the parasite to the bud of Celtis occidentalis [doctoral thesis]. [New
York]: Fordham University.
Walton BCJ. 1960. The life cycle of the hackberry gall-former Pachypsylla celtidis-gemma
(Homoptera: Psyllidae). Ann Entomol Soc Am. 53:265–277.
Watmough RH. 1968a. Notes on the biology of Artytaina spartiophila and Artytaina genistae
(Homoptera, Psyllidae) on broom Sarothamnus scoparius. J Entomol Soc South Afr.
31:115–122.
Watmough RH. 1968b. Population studies on two species of Psyllidae (Homoptera:
Sternorhyncha) on broom Sarothamnus scoparius. J Anim Ecol. 37:283–314.
Webb JW. 1977. The life history and population dynamics of Acizzia russellae (Homoptera,
Psyllidae). J Entomol Soc South Afr. 40:37–46.
Webb JW, Moran VC. 1978. Influence of host plant on the population dynamics of Acizzia
russellae (Homoptera Psyllidae). Ecol Entomol. 3:313–321.
Weintraub PG, Beanland L. 2006. Insect vectors of phytoplasmas. Annu Rev Entomol.
51:91–111.
Weiss HB. 1917. The bay flea-louse, Trioza alacris Flor, as a new pest in New Jersey. Can
Entomol. 49:73–75.
Weiss HB. 1921. Notes on the life history of Pachypsylla celtidis gemma Riley. Can Entomol.
53:19–21.
Weiss HB, Dickerson EL. 1921. Notes on Trioza alacris Flor in New Jersey. Psyche 25:59–63.
Weiss HB, Nicolay AS. 1918. The life history and early stages of Calophya nigripennis. J Econ
Entomol. 11:467–471.
Weiss HB, West E. 1922. Notes on Livia maculipennis (Fitch) (Homoptera: Chermidae).
Psyche 29:226–229.
Westigard PH, Westwood MN, Lombard PB. 1970. Host preference and resistance of Pyrus
species to the pear psylla, Psylla pyricola Foerster. J Am Soc Horti. 95:34–36.
Wheeler AG. 1994. Craspedolepta eas: distribution, host, and habits of a phlox specialist
(Homoptera: Psylloidea: Aphalaridae). Proc Entomol Soc Wash. 96:91–97.
Wheeler AG, Rawlins JE. 1993. Calophya triozomima Schwarz, a sumac-feeding psyllid new to
the eastern United States (Homoptera: Psylloidea: Calophyidae). Proc Entomol Soc
Wash. 95:99–106.
Wheeler GS, Ordung KM. 2005. Secondary metabolite variation affects the oviposition
preference but has little effect on the performance of Boreioglycaspis metalencae: a
biological control agent of Melaleuca quinquenervia. Biol Control. 35:115–123.
White IM, Hodkinson ID. 1985. Nymphal taxonomy and systematics of the Psylloidea
(Homoptera). Bull Brit Mus (Nat Hist.) 50:153–301.
White MG. 1966. The problems of the Phytolyma gall bug in the establishment of Clorophora.
Inst Pap Commonw For Inst. 37:1–52.
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
178 I.D. Hodkinson
White MG. 1967. Research in Nigeria on the iroko gall bug (Phytolyma sp.) injury to iroko
tree (Chlorophora sp.). Niger For Inf Bull (New Ser.) 18:1–72.
White TCR. 1968. Uptake of water by eggs of Cardiaspina densitexta (Homoptera, Psyllidae)
from leaf of host plant. J Insect Physiol. 14:1669–1683.
White TCR. 1969. An index to measure weather-induced stress of trees associated with
outbreaks of psyllids in Australia. Ecology 50:905–909.
White TCR. 1970a. Airborne arthropods collected in South Australia with a drogue-net towed
by a light aircraft. Pac Insect. 12:251–259.
White TCR. 1970b. The nymphal stage of Cardiaspina densitexta (Homoptera, Psyllidae) on
leaves of Eucalyptus fasciculosa. Aust J Zool. 18:273–293.
White TCR. 1970c. Some aspects of the life history, host selection, dispersal and oviposition of
adult Cardiaspina densitexta (Homoptera, Psyllidae). Aust J Zool. 18:105–117.
White TCR. 1971. Lerp Insects (Homoptera: Psyllidae) on red gum Eucalyptus camaldulensis
in South Australia. S Aust Nat. 46:20–23.
White TCR. 1973. Aerial dispersal of adult Cardiaspina densitexta (Homoptera, Psyllidae) in
South Australia. Trans R Soc S Aust. 97:29–31.
Whittaker JB. 1985. Population cycles over a 16-year period in an upland race of Strophingia
ericae (Homoptera: Psylloidea) on Calluna vulgaris. J Anim Ecol. 54:311–322.
Wilcke J. 1941. Biologie en morphologie van Psylla buxi. Tijdschr Plantenzeikten 47:41–89.
Wilde WHA. 1962. Bionomics of the pear psylla, Psylla pyricola Foerster, in pear orchards of
the Kootenay Valley of British Columbia 1960. Can Entomol. 94:845–849.
Wilde WHA. 1965. The pear psylla, Psylla pyricola Foerster, in Ontario (Hemiptera:
Chermidae): a review. Proc Entomol Soc Ont. 95:5–10.
Wilde WHA, Watson TK. 1963. Bionomics of the pear psylla, Psylla pyricola Foerster, in the
Okanagan Valley of British Columbia. Can J Zool. 41:953–961.
Wilde WHA, Carpenter J, Liberty J, Tunnicliffe J. 1971. Psylla pyricola (Hemiptera:Psyllidae)
vector relationships with Erwinia amylovora. Can Entomol. 103:1175–1178.
Wille HP. 1950. Untersuchungen uber Psylla piri L. und andere Birnblattsaugerarten im
Wallis. Eidgenossischen Technischen Hochschule in Zurich: 1–113.
Williams MW, Benson NR. 1966. Transfer of C14 components from Psylla pyricola Foer. to
pear seedlings. J Insect Physiol. 12:251–254.
Williams MW, Batjer LP, Degman ES, Burts EC. 1963. The susceptibility of some pear species
to injury from pear psylla. Proc Am Soc Horti Sci. 82:109–113.
Willson BW, Garcia CA. 1992. Host specificity and biology of Heteropsylla spinulosa (Hom,
Psyllidae) introduced into Australia and Western-Samoa for the biological control of
Mimosa invisa. Entomophaga 37:293–299.
Wineriter SA, Buckingham GR, Frank JH. 2003. Host range of Boreioglycaspis melaleucae
Moore (Hemiptera: Psyllidae), a potential biocontrol agent of Melaleuca quinquenervia
(Cav.) S.T. Blake (Myrtaceae), under quarantine. Biol Control. 27:273–292.
Wojnarowska P. 1962. Psylla pyrisuga Foerst. Red pear psylla. Prace Naukowe Inst Ochrony
Roslin 4:153–177.
Wojnarowska P, Baranowna I, Lipowa I. 1960. Psylla pyri L. Prace Naukowe Inst Ochrony
Roslin 2:143–161.
Wong TTY, Madsen HF. 1967. Laboratory and field studies on the seasonal forms of pear
psylla in northern California. J Econ Entomol. 60:163–168.
Woodburn TL, Lewis EE. 1973. A comparative histological study of the effects of feeding by
nymphs of four psyllid species on the leaves of eucalypts. J Aust Entomol Soc.
12:134–138.
Yang MM, Liao LH, Lou MF, Chen WC, Huang SS, Tung GS, Weng YC, Shen CC. 2006.
Diversity, biology, and nutritional adaptation of psyllids and their galls in Taiwan. In:
Ozaki K, Yukawa J, Ohgushi T, Price PW, editors. Galling arthropods and their
associates: ecology and evolution. Tokyo (Japan): Springer-Verlag Tokyo. p. 33–42.
Journal of Natural History
179
Downloaded By: [Hodkinson, Ian D.] At: 10:37 14 January 2009
Yasuda K, Tsurumachi M. 1988. Seasonal prevalence of leucaena psyllid Heteropsylla cubana
Crawford (Homoptera: Psyllidae) in relation to temperature in Ishigaki Island. Proc
Assoc Plant Prot, Kyushu. 34:208–211.
Young GR. 2003. Life history, biology, host plants and natural enemies of the lilly pilly
psyllid, Trioza eugeniae Froggatt (Hemiptera: Triozidae). Aust Entomol. 30:31–38.
Zhangeri S. 1954. Nota sulla Trioza urticae L. Boll Ist Entomol Univ Bologna. 20:257–273.