ARTICLE IN PRESS

Basic and Applied Ecology 8 (2007) 97—116

www.elsevier.de/baae

REVIEW

Host plant location by Chrysomelidae

Patricia Fernandez, Monika Hilker

Freie Universität Berlin, Angewandte Zoologie/Ökologie der Tiere, Haderslebener Str. 9, 12163 Berlin, Germany

Received 30 December 2005; accepted 27 May 2006

KEYWORDS Summary
Chrysomelidae;
Chrysomelidae are a taxon with an enormous plethora of highly specialised
Leaf beetles;
herbivorous species. The present study provides an overview of the knowledge
Host search;
available so far on cues guiding chrysomelids to locate a host plant. Host location
Host plant speciali-
behaviour will be addressed from different trophic perspectives. Cues from
zation;
undamaged host plants are distinguished from cues released from damaged ones.
Adaptive evolution;
The role of host plant cues is considered with respect to the surrounding vegetation
Plant volatiles;
and to the presence of competitors. Only very little is known on how enemies
Plant damage;
searching for prey influence a chrysomelid’s choice for a plant. Finally, also
Phytopathogens
the impact of phytopathogens on host location in Chrysomelidae is addressed. The
pheno- and genotypic plasticity of chrysomelid host location behaviour might have
facilitated pioneering new host plants. For a better understanding of adaptive
evolution, we conclude that future studies on host plant location by chrysomelids
need to address in addition to ecological and behaviourals aspects more intensively
also the molecular questions of adaptation.
& 2006 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

Zusammenfassung
Chrysomelidae (Blattkäfer) bilden ein Taxon mit einer enormen Fülle an hoch
spezialisierten herbivoren Arten. Diese Studie bietet eine Zusammenstellung des
verfügbaren Wissens über Signale, die Chrysomeliden zur Wirtpflanzensuche nutzen.
Die Wirtsuche wird dabei auf verschiedenen trophischen Ebenen analysiert. Signale
ausgehend von unbeschädigten Pflanzen werden unterschieden von Signalen
beschädigter Pflanzen. Die Rolle von pflanzlichen Signalen wird auch im Kontext
der umgebenden Vegetation und der Gegenwart von Konkurrenten betrachtet. Es ist
nur wenig darüber bekannt, wie die Gegenwart von Fraßfeinden die Wirtspflanzen-
suche der Blattkäfer beeinflusst. Auch der Einfluss von Phytopathogenen auf die
Wirtssuche von Chrysomeliden wird skizziert. Die große phäno- und genotypische
Plastizität des Wirtssuchverhaltens von Blattkäfern könnte das Erobern neuer
Wirtspflanzenarten erleichtert haben. Die bisher vorwiegend verhaltensbiologischen
und ökologischen Studien zeigen, dass es für ein besseres Verständnis der adaptiven
Evolution der Chrysomelidae in Zukunft von besonderer Bedeutung sein wird, auch

Corresponding author. Tel.: +49 30 8385 3918; fax: +49 30 8385 3897.
E-mail address: hilker@zedat.fu-berlin.de (M. Hilker).

1439-1791/$ - see front matter & 2006 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.
doi:10.1016/j.baae.2006.05.001
 ARTICLE IN PRESS
98 P. Fernandez, M. Hilker

auf molekularer Ebene die Anpassung von Blattkäfern an ihre Wirtspflanzen noch
intensiver zu untersuchen.
& 2006 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

Contents

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Cues from uncolonised host plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
The plant’s environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
The plant’s plasticity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
The beetle’s plasticity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Perception of host plant cues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Detoxification of noxious plant secondary components . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Cues from colonised host plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Cues from colonised host plants emitted by the herbivores . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Cues from colonised host plants emitted by the damaged foliage . . . . . . . . . . . . . . . . . . . . . . . 106
Cues from colonised host plants emitted by the predators. . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Cues from phytopathogen-infected plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
Future challenges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110

Introduction by different factors, among them plant parameters

Chrysomelidae form a taxon of enormous diver-
sity with nearly 40,000 species, almost all of which
are phytophagous (Futuyma, 2004; Jolivet & Haw-
keswood, 1995). The host plants range from
bryophytes over tree ferns to gymno- and angio-
sperms (Jolivet & Hawkeswood, 1995). Some
species are specialised to feed upon roots, others
use above-ground plant material and feed upon
flowers and leaves. Many chrysomelids are feeding
mono- and oligophagously on specific plant taxa,
others – especially among the Eumolpinae, Crypto-
cephalinae, and Clytrinae – are able to use a wide
range of very different plants. Out of 139 chry-
somelid genera in North America, 46.8% use one
genus of host plant, and 46.7% use between 2 and 5
genera (Mitchell, 1981). However, even when
having specialised on a narrow range of plant
species, no evolutionary dead end has been
entered. Instead, even highly specialised chry-
somelids are able to switch to new host plant
species, if, e.g., predation pressure on the ances-
tral host plant becomes too strong (Gross, Fa-
touros, & Hilker, 2004; Gross, Fatouros, Neuvonen,
& Hilker, 2004; Termonia, Hsiao, Pasteels, &
Milinkovitch, 2001).
The factors driving host plant specialisation of
insects have been discussed intensively from
different perspectives (e.g., Bernays & Graham,
1988; Thompson, 1999). The composition of the
herbivore community on host plants is influenced
such as plant primary and secondary metabolites,
as well as by plant architecture, distribution and
availability (Ehrlich & Raven, 1964; Strong, Lawton,
& Southwood, 1984). The success of a herbivorous
insect using a particular plant depends on the
herbivore’s ability to locate this plant, to use its
nutrients, and to cope with its defensive devices.
These abilities are dependant upon the pheno- and
genotypic plasticity of the herbivore (Bernays,
2001; Chapman, 2003; Futuyma, 2000; Thompson,
1996) and of the plant (Gardner & Agrawal, 2002).
A particular herbivore individual feeding upon a
plant is usually not feeding alone, but faces
conspecific and heterospecific competitors as well
as phytopathogens altering the plant quality, and
thus influencing the herbivore’s success on the
plant (Denno, McClure, & Ott, 1995; Rostás, Simon,
& Hilker, 2003). With respect to the third trophic
level, predators and parasitoids may strongly affect
the use of suitable plants by herbivores (Bernays &
Graham, 1988; Price et al., 1980; Vet & Dicke,
1992).
Thus, ecological bottom-up and top-down factors
as well as geno- and phenotypic factors of the
herbivore influence its association with a host
plant. The huge diversity of chrysomelids specia-
lised on such a wide range of host plants offers an
ideal tool to study the mechanisms of adaptive
evolution. Unlike the Lepidoptera, which form
another important taxon of herbivorous insects
that has been studied intensively with respect to
 ARTICLE IN PRESS
Host plant location
host plant interactions, larvae and adults of
Chrysomelidae generally use the same ecological
niche. Like numerous herbivorous hemimetabolous
insects, also chrysomelids need to choose a host
plant suitable for egg deposition, larvae, and
adults. In several species, the host plant even gives
shelter for the pupal stage. When choosing a host
plant that serves for the entire development of the
offspring and nutrition of the adult stage, it might
be beneficial to take into account the change of
host plant quality in the course of time and season,
as well as alteration of host plant quality due to
previous oviposition and feeding activity (Awmack
& Leather, 2002; Hilker & Meiners, 2006).
The demands on a herbivorous insect for success-
ful colonization of a suitable host plant are
immense. Refined perceptive abilities are neces-
sary to locate a plant and enormous integrative
abilities to recognize the suitability of a plant.
When having started feeding upon a plant, reliable
digestive abilities are necessary to gain sufficient
nutrients from the plant and to detoxify possible
noxious secondary plant components. De Jong and
Nielsen (2002) stated that ‘‘adaptive evolution can
only be fully understood when the genetic basis of
adaptations is unravelled’’. Even though general
adaptation such as, e.g., detoxification of plant
toxins by P450-monooxygenases (Scott & Wen,
2001) or glutathione-S-transferases (Ranson &
Hemingway, 2005) have been studied in several
insect species, so far little knowledge is available
on the genetics of specific adaptations of herbivor-
ous insects to host plants. This is true also for the
huge taxon of Chrysomelidae. A few and highly
interesting molecular studies are available on the
digestive adaptation of chrysomelids to plant toxins
(see below), but very little is known about the
genes influencing host plant search. Several studies
have considered the questions how chrysomelids
locate their host plants, which cues stimulate them
to feed upon a plant, and which factors might force
them to leave a plant (Jermy, 1994; Matsuda, 1988;
Mitchell, 1994). Many of these studies focus on
specific chrysomelid taxa or specific factors by
which the insects orient to a host plant and
‘‘decide’’ to accept it.
Our aim is (1) to outline the high diversity and
plasticity of cues and factors that chrysomelids
face when searching for a host plant, and, by
providing such an overview, (2) to elucidate the
most urgent future research questions we should
address in this field. The review uses literature of a
time period ranging from 1926, when McIndoo did
first olfactometer experiments with the Colorado
potato leaf beetle, to 2006, when recent studies on
induction of plant defence by chrysomelid egg
99
deposition have been presented (Hilker & Meiners,
2006). We will address questions such as, e.g.,
which cues might guide a chrysomelid beetle during
host plant search? Which factors influence its
searching behaviour? We will differentiate in the
following between cues and factors from non-
colonised (Table 1) and already herbivore-colonised
plant individuals (Table 2) on the one hand, and
phytopathogen-infected plants (Table 3) on the
other hand. When encountering already colonised
plants, chrysomelids need to be able to respond
selectively to the most beneficial cues among a
plethora of information, i.e., herbivore-induced
plant cues, cues released by con- and heterospe-
cific herbivores, cues released by predators or
parasitoids. A colonised host plant might indicate
on the one hand a suitable plant individual (not
only for feeding, but also for finding mates), and on
the other hand a place where intra- and inter-
specific competition might lower performance. A
colonised host plant might also indicate a higher
danger of being detected by enemies, or a place
where species interact through shared natural
enemies. In addition to herbivores and carnivores
present on a host plant, phytopathogens are well
known to change the attractiveness of a host plant.
Thus, also the response of chrysomelids to diseased
plants will be addressed.
Cues from uncolonised host plants
The search pattern of a chrysomelid beetle for its
host plant needs to show a high flexibility and to
take into account the enormous variation of: (a)
the environment in which the plant is growing; (b)
the geno- and phenotypic plasticity of the plant
itself; and (c) also the beetle’s plasticity and
motivation. Moreover, host plant cues are usually
not detected via a single perception ‘‘channel’’,
but by visual, olfactory, and contact cues, thus
requiring highly integrative abilities of the chry-
somelid. When having encountered a plant, the
beetles need to be able to efficiently digest the
plant material and to cope with plant toxins. These
aspects of host plant search and its acceptance will
be addressed below with respect to plants not yet
being colonised by a herbivore (compare Table 1).
The plant’s environment
The preferences of Chrysomelidae for host plants
were shown to change according to abiotic con-
ditions. Feeding preferences of Colorado potato
leaf beetles (CPB) for Solanum tuberosum and
 ARTICLE IN PRESS
100 P. Fernandez, M. Hilker

Table 1. Cues/Factors from undamaged host plants affecting host location in leaf beetles; reference: exemplifying
the impact of the respective cue/factor (F ¼ field study; L ¼ laboratory study; studies of a ¼ adults, l ¼ larvae)

Cue/factor Chrysomelid taxon Field, Lab Parameter studied Reference

adult, larvae

Soil texture Diabrotica spp. F; l Population density Beckler et al. (2004)

Elevation F; l Population density French et al. (2004)
Vegetation diversity Leptinotarsa L; a Attraction () Thiery and Visser (1986)
decemlineata
Vegetation density Galeruca tanaceti F; a Oviposition (+) Meiners and Obermaier (2004)
Trirhabda canadensis F; a Host colonisation (+) Morrow et al. (1989)
Plant genotype Altica spp. F, L; a Feeding Pettis et al. (2004)
Chrysophtharta F, L; a Oviposition, feeding Rapley et al. (2004)
agricola
Chrysomela scripta F, L; a Feeding Lin et al. (1998)
Phyllotreta spp. L; a Feeding Nielsen et al. (2001)
Phratora vulgatissima L; l Development Glynn et al. (2004)
Plant phenotype Agelastica alni F, L; a, l Feeding, oviposition Dolch and Tscharntke (2000)
(changed by
attacked
neighboured plant)
Plant size Leptinotarsa L; a Attraction (+) Visser (1976)
decemlineata
L; a Attraction (+) Bolter et al. (1997)
F; a Attraction (+) Hoy et al. (2000)
Altica subplicata F; a Abundance(+), feeding Bach (1993a)
(+)
F; a Colonisation (+) Bach (1993b)
Cassida canaliculata F; a Oviposition (+) Heisswolf et al. (2005)
F; l Survival (+) Heisswolf et al. (2005)
Leaf age Chrysophtarta agricola F; a Feeding (+) Nahrung and Allen (2003)
F, L; a Oviposition () Nahrung and Allen (2003)
Several willow leaf L; a Feeding () Ikonen (2002)
beetles
L; a Feeding () Miyamoto and Nakamura
(2004)
F, L; a, l Feeding, oviposition King et al. (1998)
Larval development
Galerucella F, L; a Oviposition () Tanaka and Nakasuji (2002)
nipponensis
Plant visual cues Leptinotarsa L; a Attraction (+) Zehnder and Speese (1987)
decemlineata
L; a Attraction (+) Jermy et al. (1988)
L; a Attraction (+) Szentesi et al. (2002)
Phyllotreta striolata L; a Attraction (+) Yang et al. (2003)
Oreina cacaliae L; a Attraction (+) Kalberer et al. (2001)
Plant volatiles Leptinotarsa L; a Attraction (+) Visser and Thiery (1986)
(undamaged plant) decemlineata L; a Attraction (+) Thiery and Visser (1995)
Agelastica alni L; a Attraction (+) Park et al. (2004)
Trirhabda canadiensis L; a Attraction (+) Puttick et al. (1988)
Diabroticites spp. L, F; a Attraction (+) Metcalf and Metcalf (1992)
F; a Attraction (+) Naranjo (1994)
L; l Attraction (+) Hibbard and Bjostad (1988)
Leaf morphology Willow leaf beetles F; a, l Feeding Rowell-Rahier (1984a)
L; a Feeding Rowell-Rahier (1984b)
L; a, l Abundance Soetens et al. (1991)
 ARTICLE IN PRESS
Host plant location 101

Table 1. (continued )

Cue/factor Chrysomelid taxon Field, Lab Parameter studied Reference

adult, larvae

Phyllotreta cruciferae L; a Feeding Palaniswamy and Bodnyark

(1994)
Cerotoma trifurcata F, L; a Feeding Lam and Pedigo (2001)
Plant nutrients Diverse chrysomelids L; a, l Feeding Matsuda (1988)
L; F; a Feeding (+) Hunt et al. (1994)
L; a Feeding (+) Ikonen (2002)
Plant secondary Flea beetles L; a, l Feeding Nielsen (1978, 1988)
chemistry
Diabroticites L, a Feeding Metcalf et al. (1980)
L; a Feeding Abe et al. (2000)
Willow leaf beetles F, a, l Feeding Rowell-Rahier (1984a)
L, a Feeding Rowell-Rahier (1984b)
L; a, l Abundance, oviposition Rank (1992)
F; a Feeding Rank (1992)
L; a Feeding Kohlemainen et al. (1995)
L; a Feeding Ikonen (2002)
Ophraella communa L; a Feeding Tamura et al. (2004)

A positive or negative correlation between the plant cue/factor and the parameter studied is indicated by (+) and (), respectively. If
no (+) or () label is given, the cues/factors had varying effects.

Table 2. Cues/factors from damaged or colonised host plants affecting host location in leaf beetles; reference:
exemplifying the impact of the respective cue/factor (F ¼ field study; L ¼ laboratory study; studies of a ¼ adults,
l ¼ larvae)

Cue/factor Chrysomelid taxon Field/lab Parameter studied Reference

adult/
larvae

Sex pheromones Diabroticites F, L, a Attraction (+) Chuman et al. (1987)

L; a Attraction (+) Guss et al. (1984)
Longitarsus jacobaea F, L; a Attraction (+) Zhang and McEvoy (1994)
Leptinotarsa decemlineata L; a Attraction (+) Levinson et al. (1979)
L; a Male movement, mating Jermy and Butt (1991)
Aggregation Leptinotarsa decemlineata L; a Attraction (+) Dickens et al. (2002)
pheromones
L Miscellaneous Oliver et al. (2002)
Oulema melanopus L; a Electrophysiology Cossé et al. (2002)
F, L; a Attraction (+) Rao et al. (2003)
Phyllotreta cruciferae F, L; a Attraction (+) Peng and Weiss (1992)
F; a Attraction (+) Peng et al. (1999)
Acalymma vittatum F; a Attraction (+) Smyth and Hoffmann
(2003)
Chrysomelid Altica carduorum L; a Aggregation (+) Wan and Harris (1996)
faecal cues
Leptinotarsa decemlineata L; a Oviposition () Szentesi (1981)
Chrysomelid Several Chrysomelinae L; a Feeding () Raupp et al. (1986)
defensive
secretion
(repellent)
L; a Feeding (), Hilker (1989)
oviposition ()
F; a Dispersal (), Schindek and Hilker (1996)
oviposition ()
 ARTICLE IN PRESS
102 P. Fernandez, M. Hilker

Table 2. (continued )

Cue/factor Chrysomelid taxon Field/lab Parameter studied Reference

adult/
larvae

Plant volatiles Leptinotarsa decemlineata L; a Attraction (+) Bolter et al. (1997)

(induced by L; a Attraction (+) Schütz et al. (1997)
damage)
L; a Attraction (+) Landolt et al. (1999)
Oreina cacaliae L; a Attraction (+) Kalberer et al. (2001)
Phyllotreta spp. F; a Attraction (+) Vincent and Stewart,
(1984)
F; a Attraction (+) Pivnick et al. (1992)
L; a Feeding () Palaniswamy and Lamb
(1993)
Plant volatiles Xanthogaleruca luteola L; a Attraction Meiners et al. (2005)
(induced by egg Feeding (+)
deposition)
Plant volatiles Epitrix hirtipennis F; a Abundance () Thaler (1999)
(induced by
jasmonate)
Presence of Diabrotica undecimpunctata L; a Feeding () Williams et al. (2001)
enemies howardi
Acalymma vittatum L, F; a Feeding () Williams and Wise (2003)
Emigration (+)

A positive or negative correlation between the plant cue/factor and the parameter studied is indicated by (+) and (), respectively. If
no (+) or () label is given, the cues/factors had varying effects.

Table 3. Phytopathogen infection of a host plant affecting host location in leaf beetles (F ¼ field study;
L ¼ laboratory study; studies of a ¼ adults, l ¼ larvae)

Phytopathogen Chrysomelid taxon Field/lab Observed behaviour Reference

taxon adult/larvae

Uromyces rumici Gastrophysa viridula F, L; a Oviposition () Hatcher et al. (1994)

Hatcher (1995)
Puccina carduorum Cassida rubiginosa L; a Oviposition (–) Kok et al. (1996)
Feeding (–)
Phoma destructiva Cassida rubiginosa L; a Oviposition (–) Kruess (2002)
Feeding (–)
L; l Development time (–), Kruess (2002)
weight (–),
mortality (–)
Alternaria brassicae Phaedon cochleariae L; a Oviposition (–) Rostás and Hilker (2002)
feeding (–)
L; l Feeding (+) Rostás and Hilker (2002)
Melampsora allii- Plagiodera versicolora L; a Oviposition (–) Simon and Hilker (2005)
fragilis feeding (–)
L; l Development time (), Simon and Hilker (2003)
weight (–),
mortality (–)
Melampsora epitea Phratora vulgatissima L; a Feeding () Peacock et al. (2003)
Erysiphe Diabroticites F; a, l Population density () Moran and Schultz (1998)
cichoracearum
Cladosporium Diabrotica L; l Feeding (+) Moran (1998)
cucumerinum undecimpunctata

A positive or negative effect of the plant infection by a pathogen on the parameter studied is indicated by (+) and (), respectively.
 ARTICLE IN PRESS
Host plant location
S. dulcamara appeared to depend on the tempera-
ture (Bongers, 1970). Wind speed affected move-
ments of Trirhabda canadensis towards host plants.
At high wind speeds, host plants located upwind
were colonised, whereas at low wind speeds, also
host plants located not upwind were found (Mor-
row, Tonkyn, & Goldburg, 1989).
The landscape structure in which a host plant
grows may significantly influence its detectability
and, thereby, the degree of plant attack
(Tscharntke & Brandl, 2004). For example, the
abundance of Diabrotica virgifera virgifera and D.
barberi was found to be associated with soil texture
and elevation (Beckler, French, & Chandler, 2004;
French, Beckler, & Chandler, 2004). Also vegetation
diversity may influence host plant finding. Diverse
vegetation might lower the detectability of a
specific host plant species. Specific host plant
odour might be masked by volatiles from non-host
plants (Bruce, Wadhams, & Woodcock, 2005). For
example, mixing non-host odour with host odour
neutralised the usually positive response of the
Colorado potato beetle (CPB), Leptinotarsa decem-
lineata, to host odour (Thiery & Visser, 1986).
When searching for a host plant, the patch size
and host density can influence the herbivore0 s
choice (Bach, 1984; Meiners & Obermaier, 2004;
Schoonhoven, Jermy, & van Loon, 1998). The
vegetation structure where the host plant grows
is well known to influence the success of locating a
host plant (Bach, 1984; Hassel, 1978; Meiners &
Obermaier, 2004). The polyphagous tansy leaf
beetle Galeruca tanaceti, for example, preferred
to lay eggs in a structurally highly complex
vegetation (in terms of host density and vegetation
height and coverage), which might be a strategy to
hide from an abundant egg parasitoid (Meiners &
Obermaier, 2004). The goldenrod leaf beetle,
Trirhabda canadensis, preferred to colonise plots
with a higher density of host plants (Morrow et al.,
1989).
The plant’s plasticity
Like other herbivores, chrysomelids need to cope
with the genotypic plasticity of their host plants.
For several species, preferences for specific geno-
types are well known, like Altica spp. preferring
certain cultivars of crape myrtles, Lagerstroemia
indica L., for feeding (Pettis, Boyd, Braman, &
Pounders, 2004), or Chrysophtharta agricola pre-
ferring particular strains of Eucalyptus globulus for
oviposition (Rapley, Allen, & Potts, 2004). Feeding
preferences of the cottonwood leaf beetle, Chry-
somela scripta, varied significantly among poplar
103
clones depending on the amount of a specific
surface component, alpha-tocopheryl quinone
(Lin, Binder, & Hart, 1998). In contrast, Phyllotreta
spp. did not discriminate between wild Arabidopsis
thaliana and metabolically engineered plants con-
taining four-fold more sinalbine, suggesting that
only the presence (and not the concentration) of
this compound stimulated feeding (Nielsen, Han-
sen, Agerbick, Petersen, & Halkier, 2001). Neither
did Phratora vulgatissima show any preference for
one of the 10 genotypes generated from a cross
between Salix viminalis and S. dasyclados, even
though the genotypes differed significantly in foliar
nitrogen concentrations and phenolic substances
(Glynn, Ronnberg-Wastljung, Julkunen-Tiitto, &
Weih, 2004).
The phenotype of an uncolonised host plant can
also change with the neighbourhood to colonised
plants from the same species (see review in Bruin,
Dicke, & Sabelis, 1995; Dicke & Bruin, 2001; Karban
& Baldwin, 1997). Volatiles induced by herbivores
in one plant may enhance the production of
volatiles in undamaged, neighboured plants which
may change their attractiveness to herbivores. A
field study showed that undamaged alder trees
were less attractive to the alder leaf beetle
Agelastica alni when located in the neighbourhood
of alder trees damaged by this species (Dolch &
Tscharntke, 2000).
The size of a plant may influence the chrysome-
lid’s choice. When encountering patches of unco-
lonised host plants, the largest one might be the
most attractive one because it might release the
highest quantities of attractive odour or the most
conspicuous visual cues. The CPB was shown to be
attracted by large (460 cm height), but not by
small plants (o 25 cm height) (Bolter, Dicke, van
Loon, Visser, & Posthumus, 1997; Visser, 1976). In
field studies, larger potato plants also attracted
more CPB than smaller ones (Hoy, Vaughn, & East,
2000). Willow flea beetles, Altica subplicata,
tended to colonise and damage taller hosts (Bach,
1993a, b). Furthermore, the monophagous tortoise
beetle Cassida canaliculata preferred to oviposit on
large host plants, Salvia pratensis. These large
plants provided significantly higher nitrogen con-
tent. Larvae feeding upon leaves of large plants
developed significantly faster than those on leaves
of small plants (Heisswolf, Obermaier, & Poethke,
2005).
Intra-plant differential foliage selection by chry-
somelids has been found in several studies. No
general pattern of preference for, e.g., young
leaves is detectable so far. Different foliage may
be used for different purposes, a possible strategy
to provide intra-plant niches for different
 ARTICLE IN PRESS
104
developmental stages. The paropsine leaf beetle
Chrysophtharta agricola preferred adult eucalypt
foliage to juvenile leaves for feeding, but juvenile
foliage was preferred for oviposition (Nahrung &
Allen, 2003). Some willow leaf beetles were shown
to prefer feeding upon young host plant leaves with
the highest nitrogen content (Ikonen, 2002; Miya-
moto & Nakamura, 2004). Likewise, female adults
of Galerucella nipponensis preferred young or
middle-aged leaves to old leaves as oviposition
sites on its aquatic host plant Trapa japonica
(Tanaka & Nakasuji, 2002). The willow leaf beetle
Plagiodera versicolora preferred to oviposit in the
field on leaves near the centre of branchlets (i.e.,
older leaves), whereas in the laboratory, females
preferred young leaves to old ones (King, Crowe, &
Blackmore, 1998). These results indicate that in the
field other cues such as light or danger of predation
might have influenced the choice for oviposition
sites in P. versicolora.
The beetle’s plasticity
Host plant preference of the individual herbivor-
ous insect usually changes during its life time,
because the selection of a plant depends on the
internal state of the individual herbivore, e.g., its
age and hormonal state, hunger, or experience
(Bernays, 1995; Courtney, Chen, & Gardner, 1989;
Jermy & Szentesi, 2003; Papaj & Prokopy, 1989;
Schoonhoven et al., 1998; Szentesi & Jermy, 1990).
Diabrotica virgifera virgifera larvae were found to
be strictly monophagous, feeding only on corn
roots, while the adult beetles are polyphagous
(Chyb, Eichenseer, Hollister, Mullin, & Frazier,
1995). Age, sex, and dietary history affected the
feeding response to cucurbitacins in Acalymma
vittatum feeding on cucumber (Smyth, Tallamy,
Renwick, & Hoffmann, 2002). Orientation of adult
CPB to host odour was shown to depend on the
satiation of the beetles (Thiery & Visser, 1995). In
Galerucella lineola, preferences seemed to be
unstable because they could be modified by food
experience in the early adult stage (Ikonen, Sipura,
Miettinen, & Tahvanainen, 2003). Experience was
found to change feeding preference or host plant
acceptance also in bruchid species (e.g., Szentesi &
Jermy, 1990). Such changes in feeding preference
can be explained as changes in the sensitivity of
deterrent and phagostimulant receptors, as has
been shown for other taxa in relation to age,
nutrient status, and experience (Chapman, 2003;
Schoonhoven, Simmonds, & Blaney, 1991; Sim-
monds, Blaney, & Schoonhoven 1992; Simmonds,
Schoonhoven, & Blaney, 1991).
P. Fernandez, M. Hilker
Perception of host plant cues
The role of visual cues for host plant location in
chrysomelids has most intensively been studied in
CPB. These beetles use a light compass reaction to
maintain a course when foraging. They approach
host and non-host plants by orientating to silhou-
ettes (Jermy, Szentesi, & Horvath, 1988) and colour
(Szentesi, Weber, & Jermy, 2002; Zehnder &
Speese, 1987). But also for a few other chrysome-
lids, visual cues were shown to influence orienta-
tion. Sensitivity to different wavelengths may even
change according to the time of the day, as was
shown for the flea beetle Phyllotreta striolata
(Yang, Lee, & Wu, 2003). The perception of plant
visual cues may also modify the response to host
plant volatiles, as was shown by Kalberer, Turlings,
and Rahier (2001): More adult Oreina cacaliae were
found to respond to host plant volatiles in a wind
tunnel when the beetles could see the plant instead
of only perceiving the volatiles.
Olfactory attraction to host plants has been
studied intensively in several chrysomelid species.
The first demonstration that volatile emissions by
plants attract a chrysomelid species was performed
by McIndoo (1926) who showed the attractiveness
of volatiles emitted by potato plants to CPB adults,
which was later confirmed in more detail by several
other studies (e.g., Thiery & Visser, 1995; Visser &
Thiery, 1986). Since then numerous lab and field
studies have been conducted on the attractiveness
of host plant volatiles to chrysomelids. For exam-
ple, the alder leaf beetle Agelastica coeruleae was
shown to clearly distinguish between leaf odour
from nine different Betulaceae species (Park, Lee,
Shin, Kim, & Ahn, 2004). Also the goldenrod leaf
beetle Trirhabda canadensis moved upwind towards
host odours in an olfactometer (Puttick, Morrow, &
Lequesne, 1988). In the field these beetles move
towards the highest concentration of host plants
(Morrow et al., 1989). Several adult Diabroticites
have been shown to be attracted by volatiles
released from blossoms of Curcurbitaceae (Metcalf
& Metcalf, 1992). Diabrotica species have also been
shown to display oriented movement towards their
host when forced to fly at a distance of 1 m from
the edge of the crop (Naranjo, 1994). Furthermore,
CO2 was found to be an attractant for larvae of the
western corn rootworm Diabrotica virgifera virgi-
fera. The attraction was enhanced when corn
volatiles were added to the CO2 source (Hibbard
& Bjostad, 1988).
Once on the host plant, several traits perceived
by contact such as morphological leaf character-
istics and the concentration of chemical constitu-
ents (both nutrients and secondary metabolites)
 ARTICLE IN PRESS
Host plant location
are well known to affect host plant choice in
Chrysomelidae (Chapman, 2003; Jermy, 1994;
Matsuda, 1988; Mitchell, 1994; Morris, Grevstad,
& Herzig, 1996; Schoonhoven et al., 1998). Leaf
trichomes, for example, were shown to influence
host selection in a wide range of chrysomelid
species (e.g., Lam & Pedigo, 2001; Palaniswamy &
Bodnaryk, 1994; Rowell-Rahier, 1984a, b; Soetens,
Rowell-Rahier, & Pasteels, 1991).
The role of plant nutrients has mainly been
studied with respect to performance of chrysome-
lids, but several former studies reviewed by
Matsuda (1988) also addressed the role of primary
plant metabolites such as sugars, amino acids,
vitamins, and lipids as feeding stimulants. For
example, some studies showed that preference of
Chrysomelidae for specific leaves was due to
nutritive traits such as nitrogen (Hunt, Liptay, &
Drury, 1994; Ikonen, 2002).
Several classes of secondary plant metabolites
have intensively been studied as feeding stimulants
for Chrysomelidae, especially glucosinolates of
Brassicaceae in flea beetles (e.g., Nielsen, 1978,
1988; Renwick, 2002), cucurbitacins of Cucurbita-
ceae in Diabroticites (Abe, Matsuda, & Tamaki,
2000; Metcalf, 1994; Metcalf, Metcalf, & Rhodes,
1980), phenolglycosides of Salicaceae in willow leaf
beetles (e.g., Ikonen, 2002; Kohlemainen, Julku-
nen-Tiitto, Roininen, & Tahvanainen, 1995; Pas-
teels, Braekman, Daloze, & Ottinger, 1982; Rank,
1992; Rowell-Rahier, 1984a, b; Topp, Kulfan, Zach,
& Nicoloni, 2002), or ragweed triterpenoid and
caffeic acid derivatives in an Ophraella species
(Tamura et al., 2004).
Detoxification of noxious plant secondary
components
Other secondary plant metabolites, especially
pyrrolizidine alkaloids (PA) from Asteraceae and
Boraginaceae, have been investigated with parti-
cular respect to the physiological detoxification
mechanisms by chrysomelids. Different strategies
of PA detoxification evolved in the Chrysomelidae.
Neotropical Chrysomelinae (Platyphora) take up
the toxic plant tertiary PA and shift them to
defensive glands where they are stored unchanged.
The efficient transport of the toxins from the gut
into the glands obviously prevents detrimental
effects of the toxins (Hartmann, Theuring, Witte,
& Pasteels, 2001; Pasteels, Theuring, Windsor, &
Hartmann, 2003). Alpine Chrysomelinae (Oreina)
convert toxic plant tertiary PA into non-toxic
glucosides, while non-toxic plant PA-N-oxides are
sequestered in the haemolymph and defensive
105
glands (Hartmann, Theuring, Schmidt, Rahier, &
Pasteels, 1999; Rowell-Rahier, Witte, Ehmke, Hart-
mann, & Pasteels, 1991). The N-oxides are trans-
ferred through the gut without previous reduction,
most probably by specific membrane carriers
(Narberhaus et al., 2004). In contrast to Oreina,
flea beetles (Longitarsus) do not conjugate tertiary
PA with glucosides, but are instead able to oxidise
these toxins, so that also in these species PA-N-
oxides are accumulating in the haemolymph and
defensive glands (Dobler, Haberer, Witte, & Hart-
mann, 2000; Narberhaus et al., 2004). Several
other plant toxins are known to be sequestered by
chrysomelids and stored in the haemolymph or
defensive glands (Pasteels, Rowell-Rahier, Braek-
man, & Daloze, 1994). However, the mechanisms of
transfer of the plant toxins through the gut into
glands of the leaf beetles have most intensively
been studied for PA.
Cues from colonised host plants
Both cues released by the herbivores already
colonizing the plant and plant cues induced by
herbivore feeding or by oviposition can influence
the choice for a host plant (compare Table 2).
Cues from colonised host plants emitted by
the herbivores
Conspecifics already colonizing a plant may
release pheromones attracting more conspecifics.
There is ample evidence for the use of pheromones
in Chrysomelidae (literature overview: Metcalf,
1994; Morris et al., 1996; Müller & Hilker, 2004).
One of the first chrysomelid species, in which
female-produced sex pheromones were detected,
was the banded cucumber beetle, Diabrotica
balteata (Cuthbert & Reid, 1964). Later, the female
sex pheromones of 11 further Diabrotica spp. have
been identified (Chuman et al., 1987; Guss, Sonnet,
Carney, Branson, & Tumlinson, 1984). These pher-
omones are aliphatic ketones as well as propano-
ates and acetates. Female sex pheromones are also
supposed to be present in the ragwort flea beetle
Longitarsus jacobaea (Zhang & McEvoy, 1994) and
the CPB (Jermy & Butt, 1991; Levinson, Levinson, &
Jen, 1979).
Also several aggregation pheromones have been
detected in chrysomelids. A male-produced aggre-
gation pheromone was recently identified in the
CPB (Dickens, Oliver, Hollister, Davis, & Klun, 2002;
Oliver, Dickens, & Glass, 2002). Males of the
cereal leaf beetle Oulema melanopus produce an
 ARTICLE IN PRESS
106
aggregation pheromone (Cossé, Bartelt, & Zilkows-
ki, 2002; Rao, Cossé, Zilkowski, & Bartelt, 2003).
Furthermore, the presence of an aggregation
pheromone produced by males was proposed for
the flea beetle Phyllotreta cruciferae (Peng &
Weiss, 1992; Peng, Bartelt, & Weiss, 1999) and for
the striped cucumber beetle Acalymma vittatum
(Smyth & Hoffmann, 2003).
Several studies indicate that chrysomelids aggre-
gate in response to faecal cues on the plant as has
been reviewed by Müller and Hilker (2004). For
example, adults of Altica carduorum aggregated on
faeces of larvae and adults that had fed on the
thistle Cirsium arvense. When other Cirsium spp.
served as hosts, faeces were not attractive (Wan &
Harris, 1996). This result suggests that attractive
plant compounds and/or their metabolites were
concentrated in the gut and excreted with faeces.
However, conspecifics on a plant do not only
release cues attracting further conspecifics. The
CPB oviposited less on sites with larval traces,
which suggests a deterrence effect of larval faeces,
but the role of other herbivore or plant semio-
chemicals cannot be excluded (Szentesi, 1981).
Several chrysomelid larvae also release exocrine
defensive secretions when disturbed by conspecific
adults (Pasteels et al., 1982). The adults are
strongly deterred by secretion of these larvae
(Hilker, 1989; Raupp, Milan, Barbosa, & Leonhardt,
1986; Schindek & Hilker, 1996).
Heterospecific Chrysomelidae have also been
shown to deter each other. Larvae of the willow
leaf beetle species Phratora vulgatissima and P.
vitellinae mutually deterred adults of the other
species by their defensive secretions (Hilker, 1989;
Hilker, unpublished data). Chrysomelids have also
been shown to deter heterospecific non-chrysome-
lid insects. The willow leaf beetle Plagiodera
versicolora was able to deter caterpillars of
Nymphalis antiopa by its exocrine defensive secre-
tion (Raupp et al., 1986). Female potato leaf
hoppers Empoasca fabae oviposited less on plants
that were infested by CPB or covered by their
faeces (Tomlin & Sears, 1992).
In several chrysomelid species, attraction of
conspecifics to an already colonised plant was
reported, but the causes are not clear. The loose-
strife leaf beetle Galerucella calmariensis was
strongly attracted by conspecifics when settling
on a plant after dispersal (Grevstad & Herzig,
1997). Females of the blue willow beetle, Phratora
vulgatissima, were shown to be much stronger
attracted to potted willow plants with conspecific
beetles and feeding damage than to control plants
with either only damage or beetles (Peacock,
Lewis, & Herrick, 2001).
P. Fernandez, M. Hilker
Cues from colonised host plants emitted by
the damaged foliage
For a wide range of plants it has been shown that
attack by herbivorous insects induces defensive
plant responses. Especially the induction of plant
volatiles in response to herbivore feeding (Agrawal,
Tuzun, & Bent, 1999; Arimura, Kost, & Boland,
2005; Dicke & van Loon, 2000; Karban & Baldwin,
1997) or egg deposition by herbivorous insects
(Hilker & Meiners, 2002, 2006) can affect the
search of herbivores for a host plant.
The odour released from damaged host plants
often differs qualitatively and quantitatively from
the one of undamaged plants (Vet & Dicke, 1992).
Since herbivore-infested plants emit volatiles in
much larger amounts than undamaged ones, it was
proposed that herbivore-infested plants are more
easily perceived from a distance (Dicke, Sabelis,
Takabayashi, Bruin, & Posthumus, 1990; Turlings,
Tumlinson, & Lewis, 1990). Indeed, several chry-
somelids have been shown to orient preferably to
feeding-damaged plants. For example, odour from
damaged potato plants was more attractive to the
CPB than odour from undamaged ones (Bolter et
al., 1997; Landolt, Tumlinson, & Alborn, 1999;
Schütz, Weissbecker, Klein, & Hummel, 1997). Also
Oreina cacaliae was preferentially attracted to
damaged host plants of Petasites paradoxus and
Adenostyles alliariae (Kalberer et al., 2001). In the
field, Phyllotreta cruciferae and P. striolata were
attracted by traps baited with allyl isothiocyanate,
one of the compounds released especially by
damaged crucifers (Pivnick, Lamb, & Reed, 1992;
Vincent & Stewart, 1984).
The herbivore’s response to induced plant vola-
tiles may significantly depend on the degree of
damage, and thus, the quantity and quality of
volatiles released by the damaged plant. Most of
the studies that considered different levels of
herbivore damage found that the strength of
induced responses usually correlates positively with
the level of damage (reviewed in Karban &
Baldwin, 1997). Behavioural assays in a locomotor
compensator showed that mechanical damage
inflicted with scissors in potato foliage resulted in
short-term (less than 15 min) attraction of CPB
(Bolter et al., 1997). On the contrary, more severe
damage with carborundum powder resulted in a
longer lasting attraction (at least 1 h). The elm leaf
beetle Xanthogaleruca luteola has been shown to
adjust its behavioural response to oviposition-
induced elm leaf volatiles with respect to the egg
density. When elm twigs were carrying a small egg
load, the oviposition-induced volatiles attracted
elm leaf beetle females. However, when elm twigs
 ARTICLE IN PRESS
Host plant location
were heavily laden with eggs, the volatiles induced
deterred X. luteola (Meiners, Hacker, Anderson, &
Hilker, 2005). Volatiles from a plant with low egg
load might indicate a suitable host plant to the elm
leaf beetle females, whereas odour from a heavily
egg-laden plant might indicate a high risk of egg
parasitisation. The eulophid egg parasitoid Oomy-
zus gallerucae is known to be attracted by elm
volatiles induced by a high egg load (Meiners &
Hilker, 2000).
Other studies show that chrysomelids are also
able to adjust their behaviour according to herbi-
vore density on the host plant. However, it is not
clear from these studies whether induced plant
volatiles or other insect-derived cues mediate
these adjustments. For example, the golden rod
leaf beetle Trirhabda virgata was observed to leave
host patches that had a high density of conspecific
adults and heavy larval damage, but no emigration
flights were detected from low-density patches
with little defoliation (Herzig, 1995). For larvae of
Diabrotica virgifera virgifera, it was suggested that
under low to moderate plant infestation levels,
little or no larval movement occurs after initial
establishment on a susceptible host. However,
under high infestation levels, significant movement
occurred from damaged to undamaged neighbour
plants (Hibbard, Higdon, Duran, Schweikert, &
Ellersieck, 2004). This behaviour might protect
them from infestation by entomopathogens, since
entomopathogenic nematodes have been shown to
be attracted to root volatiles induced by Diabrotica
feeding (Rasmann et al., 2005).
The induced release of volatiles is not limited to
the site of damage, but can occur systemically,
which means that undamaged leaves near damaged
ones also emit attractants (Dicke et al., 1990;
Turlings & Tumlinson, 1992). When the water lily
Nuphar luteum macrophyllum was damaged by
feeding of Galerucella nymphaeae, adult beetles
consumed significantly more immature leaf tissue
(undamaged leaves that had not yet reached the
pond surface) from heavily damaged plants than
from the less damaged ones (Bolser & Hay, 1998).
Also feeding and oviposition of the elm leaf beetle
Xanthogaleruca luteola induced field elm leaves
(Ulmus minor) to emit volatiles systemically (Mei-
ners & Hilker, 2000; Wegener, Schulz, Meiners,
Hadwich, & Hilker, 2001). Beetles preferred to feed
upon systemically induced leaves compared to
uninfested ones (Meiners et al., 2005).
The volatiles from a herbivore-infested plant
represent a food source with competitors and
elevated risk of influx of predators and parasitoids.
Thus, it is difficult to predict whether herbivores
will be attracted or repelled. Even when attracted
107
by induced plant volatiles, other cues encountered
when already arrived at the damaged plant might
induce the insect to leave this plant. For example,
even though allyl isothiocyanate was attractive to
P. cruciferae, induced plants were obviously
avoided for feeding. When exposing feeding-in-
duced crucifer plants to P. cruciferae several days
after wounding, the induced plants suffered less
further feeding damage than the non-induced
control ones (Palaniswamy & Lamb, 1993). In the
field, the abundance of the flea beetle Epitrix
hirtipennis was reduced on tomato plants that had
been treated with jasmonate over 3 years (Thaler,
1999).
The attractiveness of herbivore-induced plant
volatiles to chrysomelids was shown to depend
upon the internal state of a chrysomelid beetle (see
above) as well as on the background of odour at
which specific volatiles are perceived. For exam-
ple, in CPB the amplitudes of electroantennograms
for (Z)-3-hexenyl acetate, a typical green leaf
volatile released from potato after damage, in-
creased from the day of beetle emergence over at
least 6–8 days of adulthood (Dickens, 2000a).
However, CPB was not attracted to a single green
leaf volatile (Visser & Avé, 1978). A green leaf
volatile obviously needed to be perceived in
context with other plant volatiles to become
attractive, since CPB was shown to significantly
respond to specific blends of green leaf volatiles in
behavioural tests (Dickens, 2000b). In Cassida
denticollis, the common green leaf volatile (Z)-3-
hexenol did not attract larvae, when offered singly,
but the presence of this green leaf volatile
enhanced the herbivore’s ability to differentiate
between host and non-host plants (Müller & Hilker,
2000).
Cues from colonised host plants emitted by
the predators
Volatiles from herbivore-induced plants are well
known to attract enemies of the feeding stages or
of the eggs (Hilker & Meiners, 2006; Hilker,
Rohfritsch, & Meiners, 2002; Price et al., 1980;
Vet & Dicke, 1992). Several studies are available
showing or strongly indicating that herbivores are
able to recognise the presence of enemies by
infochemicals (Dicke & Grostal, 2001). However,
there is only very little evidence that chrysomelids
possess the ability to detect enemies in time.
The cucumber leaf beetle Diabrotica undecim-
punctata howardi was able to detect the presence
of a spider. The mere presence of the spider
Rabidosa rabida caused females D. u. howardi to
 ARTICLE IN PRESS
108
spend less time on the host plant and to feed less
(Williams, Snyder, & Wise, 2001). Adults of D. u.
howardi were able to instantaneously recognise
dangerous predators versus taxonomically related
ones that are no true risk. Beetles might use both
size and shape to recognise a predator (Snyder &
Wise, 2000). When present at higher densities on
host plant leaves, spiders were detected by the
striped cucumber leaf beetle, Acalymma vittatum.
It was shown that this chrysomelid species mainly
relies on tactile cues, but also visual ones to detect
the enemy (Williams & Wise, 2003). So far, it is
unknown whether these chrysomelid species are
also able to respond to deposits released by a
spider (silk, feces), as was shown for the scarab
beetle Popillia japonicae and the coccinellid
Epilachna varivestis (Hlivko & Rypstra, 2003).
Cues from phytopathogen-infected
plants
In the last years, an increasing number of studies
is focusing on microorganisms affecting the inter-
action between plants and herbivores, adding a
new level of complexity to this system (for a
literature overview see Hatcher & Paul, 2001;
Rostás et al., 2003). A fungal infection can change
the host plant’s suitability for herbivores and vice
versa (Hatcher, 1995). Although both, beneficial
and detrimental impacts can be found, most
studies demonstrated that fungal infection of the
host plant had negative or no effects on the
herbivore (Rostás et al., 2003). Infection of plant
tissue by rust fungi is well known to affect
herbivore behaviour (compare Table 3). The leaf
beetle Gastrophysa viridula fed and oviposited less
on dock plants (Rumex obtusifolius) infected by the
rust fungus Uromyces rumicis than on healthy
plants in laboratory experiments. These results
were partly confirmed in manipulative field experi-
ments (Hatcher, 1995; Hatcher, Paul, Ayres, &
Whittacker, 1994). When giving the choice, adults
of Cassida rubiginosa consumed more of healthy
leaves of Carduus thoemeri than of tissue infected
by the rust fungus Puccina carduorum. But the
number of eggs laid did not differ between healthy
and diseased plants. On infected leaves, feeding
and oviposition were confined to pustule-free areas
(Kok, Abad, & Baudoin, 1996). In cage experiments,
adults of C. rubiginosa also fed and oviposited less
on leaves of Cirsium arvense if they were infected
by the necrotroph Phoma destructiva (Kruess,
2002). Likewise, Chinese cabbage leaves infected
by Alternaria brassicae were avoided by females of
P. Fernandez, M. Hilker
the mustard leaf beetle Phaedon cochleriae. Leaf
consumption and oviposition were higher on
healthy leaves (Rostás & Hilker, 2002). The adult
willow leaf beetle Plagiodera versicolora avoided
feeding and oviposition on leaves infected by the
rust Melampsora allii-fragilis. Leaves closely ad-
jacent to the infection site were avoided while
leaves farther away from the site of damage were
accepted as readily as healthy ones (Simon &
Hilker, 2005). In the same system, larval perfor-
mance was also detrimentally affected by rust
infection: increase of mortality, decrease of larval
weight and prolonging of developmental time
(Simon & Hilker, 2003). Likewise, Phratora spp.
avoided feeding upon rust infected willow leaves
(Peacock, Hunter, Yap, & Arnold, 2003). By asses-
sing the insect and fungus community structure in
the field, the density of Diabrotica undecimpunc-
tata howardi, D. virgifera virgifera, and Acalymma
vittatum was found to correlate negatively with
the presence of powdery mildew Erysiphe cichor-
acearum on leaves of Cucurbita cepo x texana
(Moran & Schultz, 1998).
Although it is less common, herbivores may also
prefer plant tissue infected by pathogenic fungi.
Diabrotica undecimpunctata removed more leaf
area from cucumber leaf discs if these had been
infected with the necrotrophic fungus Cladospor-
ium cucumerinum for 3 days (Moran, 1998). Like-
wise, second instar larvae of P. cochleariae
consumed more from Chinese cabbage leaves
infected with A. brassicae than from healthy ones
(Rostás & Hilker, 2002).
While the effects of plant infection by phyto-
pathogens on herbivores have been addressed in a
considerable range of studies, only little is known
about how these interactions are mediated (Rostás
et al., 2003). Changes in nitrogen contents were
found in rust-infected dock plants that are avoided
by G. viridula. But these diseased plants also
contained higher concentrations of calcium oxalate
(Hatcher, 1995; Hatcher, Paul, Ayres, & Whittaker,
1995). Whether these or other cues were indeed
responsible for the reduced feeding activity on
rust-infected dock plants is unclear. In Chinese
cabbage infected by Alternaria brassicae, probably
an increased peroxidase activity caused avoidance
of diseased plants by the mustard leaf beetle
(Rostás, Bennett, & Hilker, 2002).
Future challenges
Huge knowledge is available on factors influen-
cing chrysomelid behaviour during host location.
 ARTICLE IN PRESS
Host plant location
However, only little is known on the mechanisms
how these factors affect a chrysomelid, and,
moreover, how they act in concert. Further knowl-
edge on this proximate level will endow us to
elucidate the evolution of host specialization in
Chrysomelidae. Knowledge on the receptors by
which chrysomelids perceive plant cues as well as
more information about the enzymes and transport
proteins necessary for digestion and detoxification
will show us which physiological and molecular
changes are necessary to successfully use a host
plant.
When addressing the perception of host plant
cues by chrysomelids, we need to ask how specific
are the receptors detecting host plant volatiles or
feeding stimulants. How finely tuned is the percep-
tion of plant cues and how are all the signals
integrated by the beetles? Electrophysiological
recordings of neural activity of chemosensory
sensilla exposed to combinations of components
provide some information on the integration of
signals on the neuron level, as, e.g., the study by
Hollister, Dickens, Perez, and Deahl (2001) on
galeal taste neurons in CPB showed. Even simple
structure–activity bioassays may give us first hints
on the binding domains of receptors and, thus,
provide us with information on receptor specificity,
as did the studies by Metcalf and Lampman (1991),
Chyb et al. (1995), or Kim and Mullin (1998) on
taste and olfactory receptors in Diabroticites.
Phylogenetic analyses using molecular and mor-
phological data sets may shed some light on the
monophyly of chrysomelids feeding on plants with
specific secondary metabolites. By such studies,
Gillespie, Kjer, Duckett, and Tallamy (2003) could
show that feeding upon cucurbitacins in Chrysome-
lidae has developed convergently in several taxa.
They suggest that the affinity of Old and New World
rootworm species to cucurbitacins is due to slight
taste receptor modifications (loose receptor hy-
pothesis, Tallamy, Mullin, & Frazier, 1999). Simi-
larly, feeding upon pyrrolizidine alkaloid containing
plants by Longitarsus flea beetles has evolved
several times independently (Dobler, 2001), thus
again suggesting that the evolution of the physio-
logical prerequisites of feeding on these toxic
plants may require only slight modifications of an
ancestral character. Labeyrie and Dobler (2004)
could demonstrate that a very tiny change of a
protein enabled a chrysomelid to conquer a new
host plant. They showed that two Chrysochus
species evolved the ability to feed on plants with
toxic cardenolides by substituting a single amino
acid of the binding site of the cardenolide sensitive
enzyme, thus acquiring cardenolide insensitivity.
Likewise, the ability of larvae of the flea beetle
109
Phyllotreta nemorum to survive on certain types of
Barbarea vulgaris depends on the presence of
major, dominant R-genes. Some of these genes
are located on the sex chromosomes (X and Y). The
B. vulgaris types studied cannot be successfully
attacked by beetles lacking the R-genes. A single Y-
linked R-gene has been shown to be sufficient to
convert a flea beetle genotype incapable of living
on the B. vulgaris types into a genotype that is able
to survive on these plants (Nielsen, 1997, 1999).
When considering the adaptation of leaf beetles
to so many plant toxins, more studies are needed
that address detoxification mechanisms of leaf
beetles. The ability of sequestration of plant toxins
is widespread in Chrysomelidae. The exciting
studies on the fate of plant pyrrolizidine alkaloids
in leaf beetles, as outlined above, provide deep
insight into the transport mechanisms of these
plant secondary components and their storage
(e.g., Dobler & Rowell-Rahier 1994; Hartmann et
al., 1999; Hartmann, Theuring, Witte, Schulz, &
Pasteels, 2003; Narberhaus, Theuring, Hartmann, &
Dobler, 2003; Narberhaus et al., 2004). The
stimulating studies by Termonia, Pasteels, Windsor,
& Milinkovitch (2002) on sequestration abilities of
Platyphora leaf beetles revealed that these species
may sequester both plant amyrins and pyrrolizidine
alkaloids. They suggest that this dual sequestration
could be a key mechanistic strategy to enable host
plant switches. Also the dual abilities of sequestra-
tion of toxic plant components and of de novo
synthesis of defensive devices could facilitate host
plant shifts. The elegant studies by Feld, Pasteels,
and Boland (2001) and Kuhn et al. (2004) showed
that some chrysomeline larvae possess the ability
both to transport plant glycosides to defensive
glands and to produce iridoid monoterpene de
novo.
When trying to elucidate the evolution of host
plant specialisation in a chrysomelid species, a
multitrophic view is needed taking into account
also the impact of microorganisms, predators, and
parasitoids in addition to plant characters. For
example, Gross, Podsiadlowski, and Hilker (2002)
and Gross, Fatouros, and Hilker (2004), Gross,
Fatouros, Neuvonen et al. (2004) studied the
driving forces of a shift from Salix spp. to birch in
Chrysomela lapponica. Costs and benefits of feed-
ing on either plant type were evaluated. Finally,
the results suggest that not a better nutrition on
birch, but a specialist natural enemy present on
Salix spp. might have forced this leaf beetle species
to pioneer birch as new host plant.
A straightforward combination of molecular,
physiological, (bio)chemical, and ecological studies
in the future will help to understand how
 ARTICLE IN PRESS
110
chrysomelids evolved the ability to conquer so
many different plants. As outlined here, numerous
ecological and behavioural studies are available.
Thus, more of our future studies need to use
genomics, quantitative genetic analyses, as well as
the candidate gene approach as tools to provide a
deeper insight into the ecology and evolution of
host location behaviour in chrysomelids (Anholt &
Mackay, 2004; Fitzpatrick et al., 2005; Thomas &
Klaper, 2004; Via, 1990).
Acknowledgements
We would like to thank two anonymous reviewers
as well as Klaus Hövemeyer, Göttingen, for their
very useful and valuable comments on an earlier
version of this manuscript. Many thanks are also
due to all members of our lab in Berlin, especially
to Roland Schroeder, who helped to collect the
literature. P.F. was supported by the Fundación
Antorchas, Argentina.
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