BioRisk 4( | }: 267-292 (20 l 0) Apeer-rev ‘iewed open-access journa 1]

cee ae oe BioRis

www.pensoftonline.net/biorisk

Leaf and Seed Beetles
(Coleoptera, Chrysomelidae)
Chapter 8.3

Ron Beenen', Alain Roques?

| Universiteit van Amsterdam, Zoologisch Museum Amsterdam, Plantage Middenlaan 64, 1018 DH, Amster-
dam, The Netherlands 2. INRA, UR633 Zoologie Forestiére, 2163 Av. Pomme de pin, 45075 Orléans, France

Corresponding author: Ron Beenen (r.beenen@wxs.nl)

Academic editor: David Roy | Received 4 February 2010 | Accepted 22 May 2010 | Published 6 July 2010

Citation: Beenen R, Roques A (2010) Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3. In: Roques A et
al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 267-292. doi: 10.3897/biorisk.4.52

Abstract

The inventory of the leaf and seed beetles alien to Europe revealed a total of 25 species of which 14 seed
beetles (bruchids) and 11 leaf beetles mostly belonging to the subfamilies Alticinae and Chrysomelinae. At
present, aliens account for 9.4% of the total fauna of seed beetles in Europe whereas this percentage is less
than 1% for leaf beetles. Whilst seed beetles dominated the introductions in Europe until 1950, there has
been an exponential increase in the rate of arrival of leaf beetles since then. New leaf beetles arrived at an
average rate of 0.6 species per year during the period 2000-2009. Most alien species originated from Asia
but this pattern is mainly due to seed beetles of which a half are of Asian origin whereas leaf beetles pre-
dominantly originated from North America (36.4%). Unlike other insect groups, a large number of alien
species have colonized most of Europe. All but one species have been introduced accidentally with either
the trade of beans or as contaminants of vegetal crops or stowaway. Most aliens presently concentrate in
man-made habitats but little affect natural habitats (<6%). Highly negative economic impacts have been

recorded on stored pulses of legumes and crops but very little is known about possible ecological impact.

Keywords
Coleoptera, Chrysomelidae, Bruchidae, seed beetle, leaf beetle, bioinvasion, alien, Europe, translocation,

introduction

Copyright R. Beenen,A. Roques. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

268 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

8.3.1 Introduction

The family Chrysomelidae is one of the largest Coleopteran families, including ca. 37
000 described species in the world and perhaps the same number as yet undescribed
(Jolivet and Verma 2002). Bruchidae, or seed beetles, is a relatively small family. King-
solver (2004), referring to the most recent world catalogue, mentions 1,346 valid
bruchid species. Although there are good arguments to treat Bruchidae as a subfamily
of Chrysomelidae and raise some leaf beetle subfamilies to family rank (Reid 1995),
this is still not common practice among leaf beetle researchers. We treat Bruchidae and
Chrysomelidae in this contribution as families, merely for practical reasons. According
to Fauna Europaea, the fauna presently observed in Europe includes 1532 leaf beetles
and 145 seed beetles.

Except for important agricultural pests such as the Colorado potato beetle, Lepti-
notarsa decemlineata, and more recently, the western corn rootworm, Diabrotica vir-
gifera virgifera, little was known about introductions of alien leaf beetles until Beenen
(2006) revealed that 126 species have been translocated at least once from one con-
tinent to another. More information on alien seed beetles has been available in the
literature mainly because of their potential impact on stored products (Southgate
1979). In the present work, we will show that 25 non-native species of leaf and seed
beetles of which one is of unknown origin (cryptogenic) have already established in
Europe (Table 8.3.1). Thus, aliens still represent only a very small proportion (1.5%)
of the total fauna of leaf and seed beetles in Europe. By comparison, approximately
71 alien leaf beetle species have been recorded from North America (Beenen 2006,
Beenen, unpubl.).

Within Europe, changes in the distribution of native leaf beetles have also been
noticed which can be partly associated either to human activity or to natural trends
such as delayed post-glacial expansion and global warming. For example, the recent
northwards expansion of a flea beetle, Longitarsus dorsalis, seems to result from both
the introduction of a rapidly expanding invasive plant originating from South Af-
rica, Senecio inaequidens DC., on which L. dorsalis thrives (Beenen 1992), and from
increasing temperatures during the past years. However, the role of human activity
is often difficult to ascertain in such observed range expansions of native species.
We will essentially consider the species alien to Europe, a summary of the species
alien im Europe (Table 8.3.2) and will present their characteristics at the end of the
chapter.

8.3.2 Taxonomy

A total of 25 alien species of which 14 seed beetles and 11 leaf beetles have been
recorded as established in Europe (Table 8.3.1). Thus, bruchids represent more than
a half (56.0%) of the alien species whereas they account for only 8.1% of the native
fauna of seed and leaf beetles (Figure 8.3.1). This arrival of alien seed beetles has

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 269

% species % species
30 20 10 0 0 20 40 60

Bruchidae
cae Alticinae
Chrysomelinae
Galerucinae
Cassidinae
Hispinae Alien species
Cryptocephalinae
Clytrinae
Donaciinae
Eumolpinae 4
Criocerinae 4
Zeugophorinae ~—
Orsodacninae 4
Lamprosomatidae +
Synetinae

Native species
259

o co co oOo fo Co S&S &

—iE
°o

Figure 8.3.1. Comparison of the relative importance of the subfamilies of Chrysomelidae and Bruchidae
in the alien and native entomofauna in Europe. Subfamilies are presented in a decreasing order based on

the number of alien species. The number right to the bar indicates the number of species per family.

significantly modified the composition of the total fauna of seed beetles observed
in Europe, where aliens account for 9.4% at present. The pattern is rather different
for Chrysomelidae. Although this family includes 13 subfamilies in Europe the alien
entomofauna is only distributed among five of these subfamilies. Large differences
are observed in the contribution of each subfamily without any apparent correlation
to its numerical importance in the native fauna. The recent arrival in France of an
alien palm hispine beetle, Pistosia dactylifera (Drescher and Martinez 2005), largely
modified the composition of the Hispinae subfamily which includes only three native
species (Fauna Europaea 2009). However, aliens represent much less so for the two
major subfamilies of leaf beetles, Alticinae flea beetles (four species- 0.7% of the total)
and Chrysomelinae (four species- 1.3% of the total). Other alien species include one
skeletonizing leaf beetle (Galerucinae) and one tortoise leaf beetle (Cassidinae). The
same subfamily pattern is observed for translocations of leaf beetles at world level
but Beenen (2006) also noticed other species belonging to Hispinae (e.g. Brontispa
palm leaf beetles) and Criocerinae. It is noticeable that representatives from some
important subfamilies such as Cryptocephalinae and Donaciinae have never been
introduced, or never established at least.

Leaf beetles and seed beetles largely differ in biological traits that may be involved
in the relative success of seed beetle invaders compared to other groups. Seed beetles
have several ways of egg-laying. Most species deposit their eggs on mature pods of
legumes (Fabaceae), the eggs being cemented to the pod or dropped in a self- made

270 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

hole in the pod wall. Other species lay eggs on mature seeds that are still attached to
the inside of a partly opened pod. A third group of species oviposit on mature seeds
that have fallen to the ground from a fully dehisced pod. However, some species such
as Acantoscelides obtectus use different life history strategies. Early in the season in this
species, oviposition occurs on green pods of Phaseolus, while later in the season, the
eggs are deposited on mature seeds that have fallen to the ground. These biological
features make A. obtectus fully capable of completing cycle after cycle on naked seeds in
storage (Kingsolver 2004). The larvae of seed beetles entirely develop within the seeds
until pupation and their presence cannot be recognized before adult emergence, unless
the seed is X-rayed.

In contrast, leaf beetles show a large variety of reproductive traits. Many Galeruci-
nae (e.g., Diabrotica species) and Alticinae larvae (e.g., Epitrix species) develop in or at
the roots of plants and adults feed from leaves of a specific host plant or a wide variety
of plant species. Other Chrysomelidae feed both as larva and adult externally on leaves
of their host plants. Although practically no plant species is free of leaf beetles, most
leaf beetles need fresh plant products in all or at least in the adult stage. Stored dry
plant products are not suitable for leaf beetles to complete their life cycle.

8.3.3 Temporal trends

Chrysomelids probably began to be introduced thousands of years ago. It is likely
that leaf beetles associated with crops have taken the same route as herbs associated
with cereals which are supposed to have entered Europe from the Near East (Pinhasi
et al. 2005). Beenen (2006) argued that the combination of Buglossoides arvensis (L.)
Johnston and Longitarsus fuscoaeneus Redtenbacher 1849 might have taken the route
from southwest Asia where they spread with agriculture to large parts of the temper-
ate parts of the Northern hemisphere. Thus, a number of species which are at present
considered as native may indeed be originally alien. Bruchidae must have infested
pulses grown by man since the dawn of agriculture. Southgate (1979) also mentioned
infestations of lentils from the Egyptian Ptolemaic period (305 BC — 30 BC). Rela-
tively little is known of these ancient introductions. More recent ones are much better
documented as in the case of the potato Colorado beetle (Leptinotarsa decemlineata)
(see factsheet 14.10).

From a global point of view, new records of alien species in Europe were relatively
important during the 2Ӣ half of the 19" century, due to seed beetle species. The most
important being Acanthoscelides obtectus, Callosobruchus chinensis and C. maculatus.
However, these species may have been introduced well before their first record. Since
ca. 1900, the rate of seed and leaf beetle introductions severely decreased until 1975
when it began to increase again with globalization, essentially through the arrival of
leaf beetles. The last seven years since 2000 corresponded to an acceleration of intro-
ductions, with an average of 0.8 new species of Chrysomelidae per year, again mostly

leaf beetles (Figure 8.3.2)

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 271

Mean number of new alien species
recorded per year during the period
0 0.2 04 06 08 |

|800- 1849
1850-1899
1900-1924
1925-1949
1950-1974
1975-1999
2000-2009 -

je Seed beetles
|__| Leaf beetles

Time period
Figure 8.3.2. Temporal changes in the mean number of new records per year of seed and leaf beetle spe-

cies alien to Europe from 1800 to 2009. The number right to the bar indicates the total number of seed

and leaf beetle species recorded per time period.

8.3.4 Biogeographic patterns

Asia supplied the major proportion of the alien seed and leaf beetles that have estab-
lished in Europe (Figure 8.3.3). However, this pattern is mainly due to seed beetles of
which a half are of Asian origin whereas leaf beetles predominantly originated from
North America (36.4%). No seed and leaf beetle species of Australasian origin have yet
established in Europe.

Alien species are not evenly distributed in Europe, and leaf and seed beetles do not
show the same pattern of expansion. Half of the alien seed beetles have colonized more
than ten countries with four of them present in more than 50 countries and the main
islands of Europe. In contrast, 63.6% of the alien leaf beetles have not yet spread out of
the country where they have been initially introduced. Only two species, Leptinotarsa
decemlineata and Diabrotica virgifera, are presently encountered in 38 and 20 countries
respectively (EPPO 2009, Gédéllo University 2004, Grapputo et al. 2005, Purdue
University 2008) (see maps in the spreadsheets 8 and 10). Owing to climate change, L.
decemlineata may extend its range to Finland (Valosaari et al. 2008).

Alien seed and leaf beetles appear to be concentrated in southern Europe with 18
species observed in mainland Italy and more than 10 species in continental France

2I2 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

Cryptogenic
VV. Africa
14.3%

C&S America
21.4% PES nee

North America ~~ i‘

7.1%
50.0%
C& S America
7 Ay a
___ Asia
27.3%

North America
36.4%

Figure 8.3.3. Comparative origin of seed and leaf beetle species alien to Europe

and mainland Greece. Central Europe usually hosts less than 10 species except Czech
Republic (11 species), whereas aliens have been little recorded in Northern Europe

(Figure 8.3.4).

8.3.5 Main pathways and vectors to Europe

All alien species of seed and leaf beetle except one (i.e., 95.7%) have been introduced
accidentally to Europe. Unlike North America and South Africa, where a number of
alien species were released for biological control of weeds (Beenen 2006), only the rag-
weed leaf beetle, Zygogramma suturalis, has been intentionally introduced from North
America for the biological control of common ragweed, Ambrosia artemisifolia L., since
1978 in Russia (Reznik et al. 2004) and several countries of southeastern Europe, and
subsequently established in the wild especially in the Caucasus (Kovalev 2004). A flea
beetle native of Continental Europe, Altica carduorum (Guérin- Méneville), has also
been introduced in Britain and Wales in 1969-1970 to control creeping thistles, Cir-
sium arvense (L.) Scop. but none apparently established (Baker et al. 1972, Cox 2007).
Although it is difficult to ascertain the exact pathway of introduction for most of the

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 273

Number of alien species

Figure 8.3.4. Colonization of continental European countries and main European islands by seed and

leaf beetle species alien to Europe.

other species introduced accidentally, the general behaviour of chrysomelids suggests
that most introductions are related to trade of plants and stored products, although
some may have arrived as stowaways in all forms of packaging and transport, or even
as wind-borne organisms.

The world trade of beans for agricultural purposes is probably responsible for the
nowadays wide distribution in Europe of most alien species of seed beetles, such as
Acanthoscelides obtectus, Bruchus species Callosobruchus species and Zabrotes subfasciatus
(Figure 8.3.8) which develop in legume seeds of the subfamily Papilionoideae (Phaseo-
lus, Lathyrus, Pisum, Vicia) (Béhme 2001, Kingsolver 2004). However, the arrival of
other seed beetles of the genera Bruchidius, Caryedon, Megabruchidius and Mimosestes
seems to be more related to the trade in legume tree seeds of Mimosoideae (Albizzia,
Acacia) and Caesalpinoideae (Cassia, Cercis, Tamarindus) used as ornamentals in parks
and gardens. Megabruchidius tonkineus was at first suspected to have been introduced
from Vietnam to Germany with white beans (Wendt 1980) but it was later found to

274 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

be associated with pods of honey locust trees, Gleditsia triacanthos L. (Papilionoideae),
and not capable of complete development in beans (Guillemaud et al. 2010). Similarly,
Acanthoscelides pallidipennis was probably introduced with seeds of false indigo bush
(Amorpha fructicosa L., Papilionoideae) (Tuda et al. 2006) and Bruchidius siliquastri
with these of redbuds (Cercis; Caesalpinoideae) from China (Kergoat et al. 2007).
Seeds imported for ornamental purposes may also serve as the vector of seed beetles.
Specularius impressithorax (Pic) sustained several generations indoors in the Nether-
lands after having been introduced from South Africa along with seeds of Erythrina
(Papilionoideae) used for decoration, but did not eventually establish (Heetman and
Beenen 2008) (Figure 8.3.7).

Most alien leaf beetles are associated with vegetable crops (Solanaceae, Brassicace-
ae, Gramineae including maize). With both larvae and adults feeding on foliage, these
species probably entered Europe as plant contaminants (eggs, larvae) or crop contami-
nants (adults). The Colorado potato beetle has frequently been intercepted with potato
plants and tubers, but also in all forms of packaging and transport. For example, it
usually arrived to Great Britain with commercial freight among vegetable crops such
as lettuce, Lactuca sativa L., or on ships, aircraft or private cars traveling from the con-
tinent (Cox 2007). Indeed, fresh vegetables grown on land harbouring overwintering
beetles are common means of beetle transport in international trade (Bartlett 1980).
The African tortoise beetle Aspidimorpha fabricii (= A. cincta Fabricius) was believed
to be imported in Italy as a contaminant of bananas in the late 1950s but it became
a problem in cultures of Beta vulgaris L. (Zangheri 1960). A hispine palm leaf beetle,
Pistosia dactyliferae was also probably introduced as a contaminant of palms imported
for ornamental purposes (Drescher and Martinez 2005).

The means of introduction appears different when larvae are root-feeding as in
Diabrotica and Epitrix species. Unless soil infested with larvae has been imported with
host plants, which is usually prohibited, these species probably travel as stowaways.
The western corn rootworm, Diabrotica virgifera virgifera, proved to have been trans-
located from North America to Europe at least three times in aircraft laden with goods
and materials, but probably not with maize plants (Ciosi et al. 2008, Miller et al.
2005). The outbreaks in Northwestern Italy and Central Europe probably resulted
from introductions of individuals originating in northern USA (Delaware) (Guille-
maud et al. 2010).

However, another pest species related to tobacco, Epitrix hirtipennis, is assumed to
have arrived in Europe as aerial plankton with easterly trade winds blowing from the
New World to Europe (Déberl 1994b). Similarly, Jolivet (2001) reported the translo-
cation of the Sweet potato flea beetle, Chaetocnema confinis Crotch, from the USA to
several tropical destinations by hurricanes. Adults of Colorado potato beetle are also
assumed to be capable of migrating across the Channel although this beetle does not
fly strongly (Cox 2007) or from Russia (the St Petersburg region) to Finland (Grap-
puto et al. 2005).

The collection and trade of orchids for greenhouses has also resulted in the arrival
of several species which caused severe damage without persisting such as a flea beetle,

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 O75

Acrocrypta purpurea Baly, a species from Southeast Asia which was accidentally intro-
duced with plant collections into a greenhouse of Leiden University in the Netherlands
(Doberl 1994a). Likewise, larvae of a criocerine species, the yellow orchid beetle Lema
pectoralis Baly, were imported to the Netherlands with an orchid collected in 1988
in Thailand (Beenen, unpubl.). Originating of the Peninsula Malaysia and Singapore
(Mohamedsaid 2004), L. pectoralis is a major pest (orchid lema’) of orchid cultures,
particularly Vanda and Dendrobium, in the Philippines (de la Cruz 2003).

Pathways within Europe are a source of particular concern because of the waiver of
formerly routine phytosanitary inspections on goods transported within the European
Union. Thus, alien species once introduced into one European country along with
alien plants or seeds, can freely move to other European countries. Spread may com-
bine long-distance, human-mediated dispersal and natural dispersal by adult flight, as
it is the case for Leptinotarsa decemlineata (Grapputo et al. 2005). Another significant
example is the present northwards expansion of a species alien in Europe, Chrysolina
americana. This leaf beetle originates from the Mediterranean Basin where it is as-
sociated to Rosmarinus and Lavendula. Because both plants are popular garden plants
throughout Europe, C. americana has been translocated outside its native range along
with its host plants, e.g. to the Netherlands along with potted Lavendula plants im-
ported from Italy (Beenen, unpubl.). Once introduced, this species, which has good
flight capacities, disperses naturally by flight.

8.3.6 Most invaded ecosystems and habitats

All alien Chrysomelidae are phytophagous. As expected from the numerical impor-
tance of Bruchidae within aliens, seeds constitute the most important larval feeding
niche (56.0%), far more important than leaves (24.0%) and roots (20.0%). Almost
all these species are only present in man-made habitats which represent 94.1% of the
colonized habitats, essentially agricultural lands, parks and gardens, glasshouses, and
warehouses for seed beetles (Figure 8.3.5). Natural and semi-natural habitats have been
very little colonized yet.

In addition to these strong habitat trends, about 40% of the alien chrysomelid
species remain strictly related to their original, alien plants. This is especially true for
leaf beetles, where only Epitrix hirtipennis out of the 11 alien species has been observed
to shift onto native Solanaceae in Italy (Beenen 2006). In contrast, most alien seed
beetles found outdoors have already switched to seeds of native plants, for example
Bruchidius siliquastri on the native redbud, Cercis siliquastrum, in France (Kergoat et
al. 2007), and Acanthoscelides obtectus and Callosobruchus chinensis on wild legumes
(Tuda et al. 2001). Under outdoor conditions, a strict dependency to the original alien
host was only observed for two Megabruchidius species, M. tonkineus and M. dorsalis,
associated with seeds of honey locust tree, Gleditsia triacanthos, in parks and gardens.
However, a number of seed beetle species still confined to greenhouses and warehouses
only develop on alien hosts of tropical origin, such as Caryedon serratus associated with

276 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

Percentage of alien leaf and
seed beetles living in the habitat

0 20 40 60

Coastal areas- B~ 0
Inland surface waters- C
Mires and bogs- D
Grasslands-E~ 1 NI Seed beetles
Heathlands- F [i 1 | Leaf beetles
Woodlands and Forests- G
Inland without vegetation- H
Agricultural lands- | (i 17
Parks and gardens- 12 (7
Buildings, houses- | [i ‘0

Greenhouses- ]100~ 0

Habitats
Figure 8.3 5. Main European habitats colonized by the established alien species of Chrysomelidae and

Bruchidae. The number over each bar indicates the absolute number of alien species recorded per habitat.

Note that a species may have colonized several habitats.

groundnuts (Arachis hypogaea L.), tamarind (Tamarindus indica L.) and other seeds of
alien Caesalpinioideae (Kingsolver 2004). Such species still cannot establish outdoors
because none of their alien hosts can survive in the wild at the present time.

8.3.7 Ecological and economic impact

Threats due to alien chrysomelid species were first pointed out by Linnaeus in a lecture
in 1752, referring to his observation of asparagus plants (Asparagus officinalis L.) that
were heavily infested in the vicinity of Hamburg by C7rioceris asparagi, a species intro-
duced from Russia at this time (Aurivillius 1909).

Alien chrysomelid species are better known for their economic impact than for
their ecological impact. Indeed, possible ecological impacts on native flora and fauna
are very little documented. Positive impact can be appreciated for only one alien spe-
cies, Zygogramma suturalis, a strict monophagous species deliberately introduced to
Europe for the control of the invasive ragweed (cf above).

Negative economic impacts have been recorded in seven of the alien seed bee-
tle species which may severely affect stored pulses of economically-important legumes
(Acanthoscelides obtectus, A. pallidipennis, Bruchus pisorum, B. rufimanus, Callosobruchus
chinensis, C. maculatus, C. phaseoli, and Zabrotes subfasciatus; see (Borowiec 1987, Hoff-
mann et al. 1962)). Most of them are capable of re-infesting stored legumes until the

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 DF,

food reserves are exhausted. In leaf beetles, large economic impacts have been shown for
the Colorado potato beetle, L. decemlineata, affecting potato crops (see factsheet 14.10)
and the western corn rootworm, D. virgifera virgifera affecting maize roots and foliage
(see factsheet 14.8). However, It must be stressed that economic damage has only been
seen on maize in Serbia, and in some bordering areas in Croatia, Hungary, Romania,
and small areas in Bosnia-Herzegovina and Bulgaria (EPPO 2009). In the United King-
dom, yield losses to be expected from the arrival and spread of D. virgifera virgifera have
been estimated to range from 0.9 to 4.1 million € over 20 years in absence of obligatory
campaign to prevent spread of western corn rootworm but the costs of such a campaign
could also range from 3.7 to 10.5 million € (Central Science Laboratory 2007). Epitrix
hirtipennis may also impact tobacco crops (Sannino et al. 1984, Sannino et al. 1985) as
well as E. cucumeris these of potato and tomato (Borges and Serrano 1989), and Phae-
don brassicae the cabbage crops (Limonta and Colombo 2004). Alien foliage-feeding
chrysomelids may also act as vectors for plant diseases, for example D. virgifera which
transmits several cowpea virus strains in North America (Lammers 2006). However,
little is yet known in this field (Jolivet and Verma 2002). Besides such economic dam-
age, aesthetic impacts are recorded on ornamental plants, such as these of the leaf beetle
Pistosia dactylifera on palm trees in southern France (Drescher and Martinez 2005).

8.3.8 Expected trends

Introduction of alien chrysomelids is still an ongoing process, especially through the
trade of ornamentals via garden centers. For example, an alien species of the genus
Luperomorpha was recently imported to Europe. L. xanthodera, originating from
China, was first found in Great Britain feeding in flowers of several plant species in
garden centers (Johnson and Booth 2004). Later it was observed in Switzerland (FE
Kohler, personal communication), Germany (Déberl and Sprick 2009) and the Neth-
erlands (Beenen et al. 2009), and also in garden centers, especially on rose flowers
(Figure 8.3.6). Other alien specimens of Luperomorpha observed in Italy (Conti and
Raspi 2007) and France (Doguet 2008) were first identified as L. nigripennis, from In-
dia and Nepal, but finally identified as L. xanthodera (Doberl and Sprick 2009). Plants
cultivated in the Mediterranean area, then transported without severe pest control and
sold in Central, Western and Northern Europe also constitute a serious threat for the
expansion of species alien im Europe. The risks associated to this pathway were esti-
mated for Norway (Staverlokk and Saethre 2007).

Species originating from subtropical and tropical regions have also been translocated
such as Aspidimorpha nigropunctata (Klug) from tropical Africa to The Netherlands and
Macrima pallida (Laboissiére) from the Himalayan region to Cyprus. These introduc-
tions usually have not led to establishment (Beenen 2006). However, they do indicate a
potential risk, especially in the context of global warming which may facilitate establish-
ments of such species in the near future. The arrival in southern Europe of additional
species associated with ornamental palms such as the hispine leaf beetle, Brontispa longis-

278 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

Figure 8.3.6. Adult of alien flea beetle, Luperomorpha xanthodera (Credit: Urs Rindlisbacher- Foto:

www.insektenwelt.ch)

Figure 8.3.7. Adult of alien seed beetle, Specularius impressithorax; a- dorsal view; b- lateral view (credit:
C. van Achterberg; photo taken using Olympus stereomicroscope SZX12 with AnalySIS Extended Focal
Imaging software).

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 Lip

Figure 8.3.8. Adult of Mexican bean weevil, Zabrotes subfasciatus. a- dorsal view; b- lateral view (credit:
C. van Achterberg; photo taken using Olympus stereomicroscope SZX12 with AnalySIS Extended Focal
Imaging software)

sima (Gestro), already invasive in other parts of the world (Nakamura et al. 2006), is thus
probable, considering the current increase in alien pests related to palms (see Chapter X).
Finally, it is difficult to make serious predictions about the results of future transloca-
tions because the species may react differently to the new habitats and hosts when compared
with the situation in their native environment. Furthermore, translocations may enhance
evolutionary changes partly because of founder effects and genetic bottlenecks and partly
because of the triggering of evolution by new environmental factors (Whitney and Gabler
2008). Zygogramma suturalis when introduced to the Northern Caucasus for biological
control of ragweed, showed rapid evolutionary changes in flight capacity (development
of flight ability and morphological changes) within only five generations (Kovalev 2004).

References

Aurivillius C (1909) Carl von Linné als Entomolog. Verlag von Gustav Fischer.

Baca F (1994) [New member of the harmful entomofauna of Yugoslavia, Diabrotica virgifera vir-
gifera LeConte (Coleoptera, Chrysomelidae)] (in serbo-croatian). Zastita Bilja 45: 125-131.

Baker CRB, Blackman RL, Claridge MF (1972) Studies on Haltica carduorum Geurin (Cole-

potera: Chrysomelidae) an alien beetle released in Britain as a contribution to the bio-

280 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

logical control of creeping thistle, Cirsium arvense (L.) Scop. Journal of Applied Ecology
9: 819-830.

Bartlett PW (1980) Interception and eradication of Colorado beetle in England and Wales,
1958-1977. Bulletin OEPP/EPPO Bulletin 10: 481-490.

Beenen R (1992) Een bijzondere voedselplant voor een bijzondere bladkever. Natuurhistorisch
Maandblad 81: 220.

Beenen R (2006) Translocation in leaf beetles (Coleoptera: Chrysomelidae). Bonner zoologische
Beitriige 54 [2005]: 179-199.

Beenen R, unpublished records.

Beenen R, Winkleman JK (2001) Aantekeningen over Chrysomelidae in Nederland 5. Entomo-
logische Berichten, Amsterdam 61: 63-67.

Beenen RJ, Winkelman F van Nunen, Vorst O (2009) Aantekeningen over Chrysomelidae (Coleo-
ptera) in Nederland 9. Entomologische Berichten, Amsterdam 69: 9-12.

Binaghi G (1947) Un nuovo tonchio introdotto dal Brasile. Giornale di Agricoltura 57: 4.

Biondi B, Daccordi M, Regalin R, Zampetti M (1994) Coleoptera Polyphaga XV (Chrysome-
lidae, Bruchidae). In: Minelli A, Ruffo S, La Posta (Eds) Check list delle specie della fauna
italiana 60. Bologna: Calderini. 34 pp.

Bohme J (2001) Phytophage Kafer und ihre Wirtspflanzen in Mitteleuropa. Ein Kompendium.
Heroldsberg: Bioform.

Borges PAV, Serrano ARM (1989) New records of the Coleopterous fauna (Insecta, Coleopte-
ra) from the Azores. Boletim do Museu Municipal do Funchal 41: 5-24.

Borges PAV, Vieria V, Dinis F, Jarroca S (2005) A list of terrestrial fauna (Mollusca and Arthro-
poda) and flora (Nryophyta, Pteridophyta and Spermatophyta) from the Azores. Horta, Angra
do Hero: Direccao Regional do Ambiente and Unversidade dos Acores.

Borowiec, L. (1983) A survey of seed-beetles of Bulgaria (Coleoptera, Bruchidae). Polsk. Pismo
Entomol. 53: 107-127.

Borowiec L (1987) The genera of seed-beetles (Coleoptera, Bruchidae). Polskie Pismo Entomo-
logiczne 57: 3-207.

Borowiec L (1988) Fauna Polski, Tom 11, Bruchidae. Polska Akad. Nauk. Inst. Zool., Warszawa.
2257 pp!

Bouchelos CT, Chalkia CA (2003) [First record in Greece of seed-eating insect of Gazia] (in
Greek). Georgia Ktinotrofia, 3: 22-24.

Buckland PC, Skidmore P (1999). Xanthogaleruca luteola (Miller) (Chrysomelidae) in Britain.
The Coleopterist 8: 97-99.

CABI/EPPO, 2003. Leptinotarsa decemlineata, Distribution Maps of Plant Pests 139. Walling-
ford, UK: CAB International.

Central Science Laboratory (2007) Cost Benefit Analysis for Diabrotica virgifera virgifera.
http://www.defra.gov.uk/planth/phnews/diab07. pdf.

Ciosi M, Miller NJ, Kim KS, Giordano R, Estoup A, Guillemaud T (2008) Invasion of Europe
by the western corn rootworm, Diabrotica virgifera virgifera: multiple transatlantic intro-
ductions with various reductions of genetic diversity. Molecular Ecology 17: 3614-3627.

Conti B, Raspi A (2007) Prima segnalazione in Italia di Luperomorpha nigripennis Duvivier
(Coleoptera Chrysomelidae). /nformatore Fitopatologico 57: 51-52.

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 281

Cox ML (1995) Psylliodes cucullata (Mliger, 1807) (Coleoptera: Chrysomelidae: Alticinae) - a
species new to Britain. Entomologists Gazette 46: 271-276.

Cox ML (2007) Adlas of the seed and leaf-beetles of Britain and Ireland. Oxford: Information
Press. 336 pp.

Cruz RT de la (2003) Understanding “Orchid lema’. BAR digest 5, no 3. Available at [www.bar.
gov.ph/bardigest/2003/julsep03_orchidlema.asp]. Retrieved August 4 2008.

Del Bene G, Conti B (2009). Notes on the biology and ethology of Luperomorpha xanthodera,
a flea beetle recently introduced into Europe. Bulletin of Insectology 62: 61-68.

Delobel A, Delobel B (2003) Les plantes hétes des bruches (Coleoptera Bruchidae) de la faune
de France, une analyse critique. Bulletin de la Société linnéenne de Lyon 72: 199-221

Déberl M (1994a) Bemerkenswerte Alticinenfunde aus Westeuropa (Col., Chrysomelidae).
Entomologische Nachrichten und Berichte 38: 179-182.

Déberl M (1994b) Auffallige Ausbreitung einiger Alticinen-Arten in Westeuropa (Coleoptera,
Chrysomelidae, Alticinae). Verhandlungen des 14. Internationalen Symposiums fiir Entomo-
faunistiek in Mitteleuropa, Miinchen, SIEEC: 276-280.

Déberl M (2000) Beitrag zur Kenntnis der Gattung Epitrix Foudras, 1860 in der Palaarktis.
Mitt. Internat. Mitteilungen des internationalen Entomologischen Vereins Frankfurt 25: 1-23.

Déberl M, Sprick P (2009). Luperomorpha Weise, 1887 in Western Europe (Coleoptera: Chry-
somelidae: Alticinae). Entomologische Blatter 105: 51-56.

Doguet S (2008) Présence en France de Luperomorpha nigripennis Duvivier, 1892. (Col. Chry-
somelidae, Alticinae). Le Coléoptériste 11: 62-63.

Doguet S (2009) Présence en Europe de deux espéces nord-américaines d’ Epitrix (Coleoptera
Chrysomelidae Alticinae). L’Entomologiste 65 : 89-90.

Drescher J, Martinez M (2005) Le coléoptére Pistosia dactyliferae menace les palmiers du sud de
la France. PHM - Revue Horticole 468: 34-35.

EPPO, 2006. Leptinotarsa decemlineata. In: PQR database (version 4.5). Paris, France: European
and Mediterranean Plant Protection Organization. www.eppo.org.

EPPO (2009). Present situation of Diabrotica virgifera in Europe. 2008. http://www.eppo.org/
QUARANTINE/Diabrotica_virgifera/diabrotica_virgifera.htm#map-dia. [accessed March
1, 2010].

Essl F, Rabitsch W (Eds) (2002) Neobiota in Osterreich. Wien: Umweltbundesamt. 432 pp.

Fauna Europaea. http://www. faunaeur.org/.

Gobierno de Canarias (2010). Especies introducidas en Canarias. http://www.interreg-biona-
tura.com/especies/index.php?opt=verDatos#. [accessed March 1, 2010].

Gédéllo University. Spread of Western Corn Rootworm in Europe. Maps 2000-2004. http://
w3.mkk.szie.hu/dep/nvtt/wernet/wernet-2.htm.

Grapputo A, Boman S, Lindstrém L, Lyytinen A, Mappes J (2005) The voyage of an invasive
species across continents: genetic diversity of North American and European Colorado
potato beetle populations. Molecular Ecology 14: 4207-4219.

Gruev B (1981) A new species of Epitrix Foudras from the Azores (Insecta, Coleoptera, Chrys-
omelidae). Boletim da Sociedade Portuguesa de Entomologia 10: 1-2.

Gruev B, Déberl M (2005) General distribution of the flea beetles in the Palaearctic subregion
(Coleoptera, Chrysomelidae: Alticinae. Sofia and Moscow: Pensoft, Supplement Series Fau-
nistica No 42. 239 pp.

282 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

Gu J, Mao YQ, Wang LP, Xu JJ, Zhang Y, Du YZ (2009) [Genetic differentiation among dif-
ferent geographic populations of the cowpea weevil, Callosobruchus maculatus (Coleoptera:
Bruchidae)] (in Chinese, English abtsract). Acta Entomologica Sinica 52: 1349-1355.

Guillemaud T, Beaumont MA, Ciosi M, Cornuet JM, Estoup A (2010) Inferring introduction
routes of invasive species using approximate Bayesian computation on microsatellite data.
Heredity 104: 88-99.

Gyorgy Z (2007) To the biology of the honey locust seed beetle, Megabruchidius tonkineus (Pic,
1904) (Coleoptera: Chrysomelidae: Bruchinae). Folia Entomologica Hungarica 68 : 89-96.

Hansen M (1996) [Katalog over Danmarks biller] Catalogue of the Coleoptera of Denmark.
Entomologiske Meddelelser 64: 1-231.

Hansen V. (1964) [Fortegelse over Danmarks biller] Catalogue of the Coleoptera of Denmark.
Entomologiske Meddelelser 33: 1-507.

Heetman AJA, Beenen R (2008) Twee exotische zaadkevers ingevoerd met bonen (Coleoptera,
Bruchidae). Entomologische Berichten, 68: 187-188.

Hill M, Baker R, Broad G, Chandler PJ, Copp GH, Ellis J, Jones D, Hoyland C, Laing I,
Longshaw Moore N, Parrott D, Pearman D, Preston C, Smith RM, Waters R (2005) Audit
of non-native species in England. Peterborough, UK: Report to English Nature N°662.

Hodge PJ (1997) Bruchidius varius (Olivier) (Chrysomelidae) new to the British. Isles. The
Coleopterist 5: 65-68.

Hoffmann A (1945) Coléoptéres Bruchidae et Anthribidae. Faune de France 44. Paris: Lecheval-
lier. 184 pp.

Hoffmann J, Labeyrie V Balachowsky AS (1962) Les Bruchidae. In: Balachowsky AS (Ed) En-
tomologie appliquée a lagriculture, Tome 1, Volume 2. Paris : Masson, 434-494.

Hurst GW (1970) Can the Colorado potato beetle fly from France to England ? Entomologist’s
Monthly Magazine 105: 269-272.

Igrc J, Deloach CJ, Zlof V (1995) Release and Establishment of Zygogramma suturalis F (Cole-
optera: Chrysomelidae) in Croatia for Control of Common Ragweed (Ambrosia artemisii-
folia L.). Biological Control 5: 203-208.

Izquierdo I, Martin JL, Zurita N, Arechavaleta M (2004) Lista de especies silvestres de Canarias
(hongos, plantas y animales terrestres). Consejeria de Medio Ambiente y Ordenacién Ter-
ritorial, Gobierno de Canarias; Santa Cruz de Tenerife.

Jermy T, Szentesi A, Anton KW (2002) Megabruchidius tonkineus (Pic, 1904) (Coleoptera:
Bruchidae) first found in Hungary. Folia entomologica hungarica 63: 49-51.

Johnson C (1963) Chrysolina americana (L.) (Col. Chrysomelidae) in Britain. Entomologists
Monthly Magazine 99: 228-229.

Johnson C, Booth RG (2004) Luperomorpha xanthodera (Fairmaire): a new British flea beetle
(Chrysomelidae) on garden centre roses. The Coleopterist 13: 81-86.

Jolivet P (2001) Vers la mondialisation des Chrysomelides? L ‘Entomologiste 57: 123-141.

Jolivet PR, Verma KK (2002) Biology of Leaf Beetles. Andover, Hampshire, UK: Intercept Pub-
lishers. 332 pp.

Kergoat GJ, Delobel P, Delobel A (2007) Phylogenetic relationships of a new species of seed-
beetle infesting Cercis siliquastrum L. in China and in Europe (Coleoptera: Chrysomelidae:
Bruchinae: Bruchini). Annales de la Société Entomologique de France 43: 265-271.

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 283

Kingsolver JM (2004) Handbook of the Bruchidae of the United States and Canada (Insecta,
Coleoptera). United States Department of Agriculture: Technical Bulletin 1912. 324 pp.

Kovalev OV (2004) ‘The solitary population wave, a physical phenomenon accompanying the
introduction of a chrysomedlid. In: Jolivet P, Santiago-Blay JA, Schmitt M (Eds) New devel-
opments in the biology of Chrysomelidae. Amsterdam: SPB Academic Publishing, 591-601.

Lammers W (2006) Report of a Pest Risk Analysis. Netherlands Plant Protection Service.

Lays P (1988) Chrysolina americana (Linné) Belg. nov. sp. (Coleoptera, Chrysomelidae, Chrys-
omelinae), une espéce mediterranéenne en Belgique. Bulletin et Annales de la Société Royale.
Belge d'Entomologie 124: 29-33.

Limonta L, Colombo M (2004) Record in Italy of Phaedon brassicae Baly (Coleoptera Chrysomel-
idae Chrysomelinae). Bollettino di Zoologia Agraria e di Bachicoltura Serie II 36(3), 369-371.

Michieli G (1957) Ricerche sulla biologia della Phytodecta fornicata Bruggem. (Col. Chrysome-
lidae) e del suo parassita Meigenia floralis Fall. (Diptera Larvevoridae). Bollettino del Museo
Civico di Storia Naturale di Venezia 10: 49-86.

Migliaccio E, Zampetti MF (1989) Megabruchidius dorsalis e Acanthoscelides pallidipennis, spe-
cie nuove per la fauna italiana (Coleoptera, Bruchidae). Bollettino Associazione Romana di
Entomologia 43: 63-69.

Miller N, Estoup A, Toepfer S, Bourguet D, Lapchin L, Derridj S$, Kim KS, Reynaud P, Furlan
L, Guillemaud T (2005) Multiple transatlantic introductions of the western corn root-
worm. Science 310: 992.

Mohamedsaid MS (2004) Catalogue of the Malaysian Chrysomelidae (Insecta: Coleoptera).
Sofia & Moscow: Pensoft. 239 pp.

Nakamura S, Konishi K, Takasu K (2006) Invasion of the coconut beetle, Brontispa longissima:
Current situation and control measures in Southeast Asia. In: Food and Fertilizer Tech-
nology Center (FFTC) meeting ‘Development of Database (APASD) for Biological Invasion’,
Taichung, Taiwan September 2006, ROC. www.agnet.org/activities/sw/2006/589543823/
paper-899851121.pdf.

Oliviera R, Chatot C, Dedryver CA (2008) Détectée en Europe, une nouvelle altise. Potato
Planet: 30-34.

Pinhasi R, Fort J, Ammerman AJ (2005) Tracing the origin and spread of agriculture in Europe.
PLoS Biology 3: e410.

Purdue University. Map of expansion of Western Corn Rootworm in Europe in 2008. http://
extension.entm.purdue.edu/wer/images/jpg/2008/EUROPE08. jpg.

Ramos YR, Fernandez-Carrillo JL, Fernandez-Carrillo E (2007) Sobre la presencia del gorgojo
de las acacias, Pseudopachymerina spinipes (Erichson, 1833) en la Peninsula Ibérica (Cole-
optera: Bruchidae). Boletin la Sociedad Entomologica Aragonese 40: 511-522.

Reid CAM (1995) A cladistic analysis of subfamilial relationships in the Chrysomelidae sensu
lato (Chrysomeloidea). In: Pakaluk J, Slipinski SA (Eds) Biology, phylogeny, and classifica-
tion of Coleoptera. Papers celebrating the 80th birthday of Roy A. Crowson. Warszawa:
Muzeum i Ynstitut Zoologii PAN, 559-631.

Reznik SYa, Volkovitsh MG, Dolgovskaya MYu, Cristofaro M (2004) Introduction of the rag-
weed leaf beetle Zygogramma suturalis (Coleoptera, Chrysomelidae) into Russia for biologi-

cal control of common ragweed Ambrosia artemisiifolia (Asteraceae) as a model of biologi-

284 Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

cal invasion. In: Abstract Proceedings of Neobiota, 3" International Conference on Biological
Invasions, Bern, Switzerland, September - October 2004, 111.

Sainte-Claire Deville J (1938) Catalogue raisonné des Coléoptéres de France. L’Abeille 36:
373-467.

Sannino L, Balbiani A, Espinosa B (1984) Un nuovo fitofago devasta il tabacco nel beneventa-
no: Epithrix hirtipennis Melsch. Informatore Agrario 29: 55-57.

Sannino L, Balbiani A, Biondi M (1985) Epitrix hirtipennis (Melsheimer, 1847) (Coleoptera,
Chrysomelidae): Considerazioni tassonomiche, ecologich ed etologiche. Atti XIV Con-
gresso nazionale Italiano Entomologia, Palermo: 285-292.

Simmonds MSJ, Blaney WM, Birch ANE (1989) Legume Seeds: the Defences of Wild and
Cultivated Species of Phaseolus Against Attack by Bruchid Beetles. Annals of Botany 63:
177-184.

Southgate BJ (1979) Biology of the Bruchidae. Annual Review of Entomology 24: 449-473.

Staverlokk A, Saethre MG (2007) Stowaways in imported horticultural plants: alien and inva-
sive species — assesing their bioclimatic potential in Norway. Bioforsk Report 2, N°66. 70 pp.

Stephens JF (1839) A Manual of British Coleoptera or Beetles. London: Longman, Orme
Brown, Green & Longmans. 443 pp.

Strejéek J (1990) Brouci celedt Bruchidae, Urodontidae a Anthribidae. Praha : Academia. 88 pp.

Strejéek J (1993) Faunistic records from the Czech Republic 9. Klapalekiana 29: 169-171

Szentesi, A. (1999) Predispersal seed predation of the introduced false indigo, Amorpha fruticosa
L. in Hungary. Acta Zool. Acad. Sci. Hung. 45: 125-141.

Tomov R, Trencheva K, Trenchev G, Kenis M (2007) A review of the non-indigenous insects
of Bulgaria. Plant Science 44: 199-204.

Tuda M, Shima K, Johnson CD, Morimoto K (2001) Establishment of Acanthoscelides pal-
lidipennis (Coleoptera: Bruchidae) feeding in seeds of the introduced legume Amorpha
fruticosa, with a new record of its Eupelmus parasitoid in Japan. Applied Entomology and
Zoology 36: 269-276.

Tuda M, Ronn J, Buranapanichpan S, Wasano N, Arnqvist G (2006) Evolutionary diversifica-
tion of the bean beetle genus Callosobruchus (Coleoptera: Bruchidae): traits associated with
stored-product pest status. Molecular Ecology 15: 3541-3551.

Valosaari KR, Aikio S, Kaitala V (2008) Spatial simulation model to predict the Colorado
potato beetle invasion under different management strategies. Annales Zoologici Fennici
ASP 1A

Wendt H (1980) Erstmaliges Auftreten des Vorratsschadlings, Bruchidius tonkineus (Pic, 1904)
in der DDR. Deutsche Entomologische Zeitschrift 27: 317-318.

Wendt, H. (1981) Eine ftir Sudost-Europa neue Samenkafer-Art (Coleoptera: Bruchidae). Fo-
lia Entomologica Hungarica 42: 223-226.

Whitney KD, Gabler CA (2008) Rapid evolution in introduced species, “invasive traits’ and
recipient communities: challenges for predicting invasive potential. Diverstity and Distri-
butions 14: 569-580.

Zangheri S (1960) L'Aspidimorpha cincta F. (Coleoptera Chrysomelidae, Cassidinae) in Italia.
Bollettino di Zoologia agraria e di Bachicoltura Ser. II 3: 219-220.

285

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3

vn “XS
‘IS “AS ‘NU “SU “OU “AVW-.Ld

‘OZV-Ld “Ld “IAS-ON ‘ON
“IN “LN SIA ‘GW ‘AT ‘7
s1oopmo ‘L1 ‘TT OIS-.LI UVS-LI “LI

(4007) sounsa] ‘SI “TI “AI ‘NH “WH “DaAS-UD
[e 19 AOWO T, (Z9G6T) ‘Te poyeanyno ‘DANTUD “AYORUD “WD “AD
19 uuEWTyOH] “(CQOZ)| ue PTA MOOR Ud ‘NVO-SA “TWE-SA

PqoPC Pur jeqopeq ‘spaos ‘Sd “AA “NC “AC “ZO ‘AD SHO voroury | snoseyd I€8I
‘(S00Z) ‘Te 39 saBiog] —smoaswy ‘Ad ‘Od “Ad ‘Va “LV ‘CGV “TV| LI ‘6881 SRO} -oyd ‘Avg sng221G0 sapyaassoqquvay
S9]199q pges = sepryonig

(Y007)

yloog pure uosuyo[
‘(800Z) JoN3oq ‘(€00Z)| snoseyddjod
PAPled Pus Pde re. ynpe
‘(600Z) BUOD pu suag| ‘(earep) sloos

Pd ‘(Z00Z) tdsey pure snuuuong eaIoy | snoseyd (S881 ‘osreutre,7)
puod “}qndun ‘ususegq pur say fal IN ‘LI ‘€D ‘Ud “AC “HO] @D “€007 ‘eury)| -o14yq veapoqsunx vydaouosadnT
(8007) ‘Je 29 ULNSOLIGNI snoseyd DVGI
PIOIATO ‘(600Z) 20nS0q WNUDIOS Il Ld! Ld ‘8007 vsn| -oMyg| WV ‘auquay suppres xpd
"POLIOUIY
(S861) Gnes (pue
‘ye 19 OuTUUS ‘(F86T) aeaoeUrTOS yeqque7)
‘ye 19 OUTUURS ‘(000Z) Joy10 pure ‘VS | snoseyd (ZPQ | “aWToYspPy\)
F249 “(44661) F°q9d DUUSLOIENS I] OZV-Ld SW “LISD ‘DA} LI ‘861] weynos} -ovyg) Vv stuuadyany x1andy
JEIOVULTOS POLIOUIY
(6861) Joyo pure OZV| Yinog pue| snoseyd (IS8I
OURIIIS pur sasiog DUVIZOIINT II OZV-Ld| -Ld ‘Z861 Jenuey}| -oi4yg) VW ‘sIIep]) stawnang xtagdq
sapieaq vay -oeulDnTy
satgads
SdIUIIIFOY s}sO}] euqey SdIIJUNOD poprauy snqeig Ayrureyqns so Ayrure.j

‘O107 Arensqay [ orepdn seq *({] x1puadde 99s) SIN (A 02 JoJor suoneraciqqe yergepy ‘(| xipuodde sas) 991 ¢€ OS] 01 Jafar suoNerAsIgqe
sapoos AnuNO’) ‘saidads stuasoidAID F adoring 01 uaITy Ww ssnzeig ‘odorny 01 uate so1eds seprpauroskry) poysiqeisa oy] JO sonstiaoesIeyo pur Is] *|°E°g BIGEL

Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

286

(8€61)

IAI slepD-o2ures
‘(GPG[) UUeWIOPY
‘(OIOZ) selyeuey op
OUISIGOL) ‘(600Z)
voedoinyg eune,y ‘(€Q0Z)
Pde Ce Pee qo PGI

(S€61)

ITPIAIC VTepD-a1 ules
‘(ShGT1) uUPUIPOPY
‘(OLOZ) seteueD op
OUIIIGOL) ‘(600Z)

(poieanyns
pur pli)
MIND ‘SUaT
WMNSIT
‘snu1dnT
‘SMAMGYIDT
VIILA
‘Snjoasvg
‘suvaq Patol

VIIA UNSIT

Vn AMS

‘IS “AS ‘NU ‘SU ‘OU “AVW-.Ld
‘OZV-Ld ‘Ld “Td “TAS-ON
‘ON “IN “LW XW “CW AT
Pel hed Sel AVS Lia
‘SI “AI ‘NH “UH “DAS-UD “DOAN
1 SAD USD AOS
Ud ‘Ud ‘Td “NVO-Sd “TVa-Sa
‘Sa “Ad “Md “AC *ZO ‘AD “HD
‘Ad “Od “Ad “Vd “LV “TV ‘dV
VN “MS ‘TS “AS ‘NU

‘Su ‘OU “Ld “AVW-.Ld ‘OZV
“Ld “Ld “Td “TAS-ON “ON “IN
“LW ‘OW IW “CW SAT ‘NT
ET TT DISS AVS SEY ST
“aI ‘NH “UH “DAS-UD “DOAN
WD AMOUD AD: "AD "AOD
Ud “Ud ‘Td “NVO-Sd “TVa-Sa

Ld ‘681

snoseyd

eouyy| -o1dyq

CCQT ‘UURWaYyog
snupUult{ns sngmnsg

voedoing vune, ‘(€00Z) ‘snahqav Ts ‘Sd “AA “NC “AC “ZO ‘AD SHO orerodua [| snoseyd (SLI
Pqopeq pur peqopaq| ‘seed parq ‘kd “Od “AG ‘VE LV “IV ‘CV! ZD ‘OS8I1 -eisy| -o1dyg ‘snoeuury) wnsosid sngonag

3 | snoseyd Z00Z 29°

(Z00Z) ‘Je 39 WoSIsy | — spaes sz24a7D Wil Ws ‘E007 -ovdy J zugspnbyts snipiggnsg

(1861)

Ipue/\ ‘(6661) !s931U9Ez¢
(6861) nIeduez

pure o1poriystpy

‘(8861) 29IMoIOg
‘(€E8G6T1) 2IMOIOg

SIDUIIIFOY

sours]
Joyo pue
(Ysnqosipur)
VSOIIINAL

vg qsoulyy

s}sOP]

‘peat

JeuqeH

Su ‘OU “Id AW ‘NT SLI ‘NH
WH “AC “ZO “HO “O¢ “Vd “LV

S311}UNOD popeAuy

Dd ‘0861

adoimy ur
psosas IST

vououry | snoseyd

YON | -014y4q

asuUeI
SAeN = | awIsay

snjyejs

(PZ81 ‘Ays~nypsiopy)
stuuadipiyyd sapyassogqUvoy
satgadg

Ayreueyqns 10 Ayrore,y

287

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3

(7007) “Te 39
Aural “(Z007Z) £81945

(6861) edweZ
pure o1oriystyy

(€007)
Pqoped pur eqopqd

(4007)

Je 19 AOWOT, “(CH6T)
uuewlyoH “(€00Z)
PqoPC pur jeqopqd

spoos

HESHP AL)
spoos
HEELS)
spoos s7dosody
VISUD
VIIIY
(Sunsoyul-o4
jo a]qedeo)
soumn3s2]
pesois 1410
pur snuidnT
‘SNJOASVY

If ‘A
4 ba

Tisal

OFS a a
“AMOAUD UD Ud “SA “ZO “TV

AYOAUD UD “AC *ZD AO] ZO ‘0061

(WeuiI A)
jeordon| snoseyd

snoseyd
-ovdyg

vOLyy

aesrodura | | snoseyd

UA “SHOT

L007 481044)
snaUulyuoy SNIPIGINAGVSaTA

(6E8I ‘snorye.)
SYDSAOP SnIPIYINAGVEI

(O61
‘IOIAIQ) snqvstas uopaliv’yy

(CEST TeyuaTAD)
yhoaspyd SNGINAG OSO]]P)

(Z00Z) ‘Je 39 AoW T,
‘(GPG[) UUPWIOPY
AGO0G) E42 1D
‘(600Z) eoedomy eune,y
‘(€00Z) PPC pue
PqoPd ‘(Zy61) ryseurg

(4007)

TE soe E(BEGT)
SPEAWC] aureyD-a1ures
‘(ChG[) UUEWTOT]
‘(O10Z) seHeueD ap
OUTSIGOF) “(600Z)
voedoing eune,y ‘(Z700Z)
(Spx) Yosuqey pur [ssy
‘(P661) ‘Te 39 Ipuorg

$INUDIIFOY

(Sunsayur-or
jo afqedeo)
sours]
posois Joy10
pure smjoasvg

(Sunsayur-os
jo afqedeo)

soumn33]

Petors

s}soP]

Lise

Hes

JeuqeH

OZV-Ld

Ld LI OIss LL “ard
AD AO aa sa Zo Da Ty’
vn MS

‘IS “AS ‘NU ‘SU ‘OU “AVW-.Ld
‘OZV-Ld “Ld “TAS-ON ‘ON
“IN “LW SIA CCW ‘AT ‘NT SLT
TIOIS3 LT AVS" LDL “SIT
“AI ‘NH “UH “DAS-UD “DOAN
“1S. OA: UO Ae AOD
Ud ‘Ud ‘Td “NVO-SH “TVa-Sa
‘Sd “Ad “MC “Ad “ZO ‘AD “HO

‘Ad ‘Od “AG “Vd “LV “TV “dV

S311JUNOD popeAuy

snoseyd

WA ‘SZ81 eoryy| -oydg

aesodura | | snoseyd

snjyejs

(SZLTI ‘sntoriqe.y)

SNIDINIDUL SAGINAG OSOTTV’)

(8¢ZI ‘snovuurT)
SISUIUIYI SNYINAGOSOTIV)
satgad§

Ayroueyqns 10 Ayrore,y

Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

288

DY OfIQUioyA OZV-Ld| vole | snoseyd (TZ8 1 ‘rewIN4))
(1007) arf ups OZV-Ld| ‘1002 < YON] -o44d] OD ypidstjod vg douse)
“UIEUIOT A,
UPMIP T
(F007) ‘uede{| snoseyd
OquIO]OT) puke eIUOUTTT| seadROISseIg Il LI| LI ‘000z ‘euryy| -ovdyg] WwW | PZ8T ‘Ajeg amozssusqg uopavgy
VO AS ‘IS “AS ‘NU “SY ‘OU
(4002) ‘Ld “Td “IN ‘OW “IW “CW ‘AT
‘TE 19 AOWIOT, (CQO) Te] FeaoeuR[OS ATA TOs; EE VSL
yo oinddesy ‘eaedoing Joyo pure SAH YH YD YOON WA vOLIOUYy
euney (9007) Odd | = “#So4aqna “Tvd-Sa ‘Sa “Ad “A “ZO SHO jeruac | snoseyd (PRI ‘Aes)
‘(€007) Odda/IAVO WHLUHIOS, I] ‘Ad ‘Dd “Ad ‘Va “LV “IV ‘CGV) Ud ‘7Z6T] Pur PION] -oyd) Vv pypauywuarap wsavqourda]

S9]199q Jeo] = aeuTauosAry)

8007 “PI949S

(0961) Hoysuez | stuvsyna vag na1qu, vy dsowopidsy
Say129q FEI] ISIOIIO [, — SVUTPISse7)
(suNsoyut-o4
jo a[qedeo)
(S¥61) sown] OZV-.Ld
uUPWYJOH ‘(€Q0Z)) pez0o3s soyI0 “Td “IN, ‘OIS- EL “LT AMOATD bl snoseyd (CEQ ‘uUuRWAYOg)
Vs Wi an SU Sees Sp bee scr Alea ed UWS “Ud SNVO-SH “SH “ZO “TV | Us ‘881 SRO} -oMyd SHIDLISL{GNS SA10LQUT
(002)
‘ye Jo soueYy “(G00Z) spoas
voedoing eune,y ‘(€Q0Z) DUDISIULD voluoury | snoseyd (CEB ‘uosysIy)
eryey pure sojayonog VIWIP ral -LI “LI AYOND UD UA ‘SA| SA ‘GIGI SRD -ovdy J sadiuids vursauhgovdopnasy
spoas
(eadyoryo)
AIL DIILA
(CPG) UUeWOPY] ‘SNJOISDY J aesiodura | | snoseyd (T8ZI ‘sniorqe.)
‘(9661) UesueH UIIVIY Uf LI ‘Ud “NC “Ad| Ud ‘Srol -o1ky ASE SOT ASU Af
satgads
SIIUIIIFOY s}sOP] euqey SoIIJUNOD popeauy snqvig Ayrureyqns so Ayrure.y

289

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3

(S007)
ZoUTeYy pure Joyosoiq

SUITE]

Cl

dd

UA “YOO

V

(6161
yrney]) avafylgovp visoqsig

(6007) Alsat)
enping “(010Z)

‘Te 19 pheuroyjMer)
‘(600Z) Atssoaruy)
OT]2PE (6007)
Oddi ‘(Z007Z) ‘Te #9
ISOID “(H66T) BORG

“SMDUL DAF

II

"VN MS ‘IS “SY ‘OU “Td
“IN ‘OW ‘LI ‘NH ‘YH “AD
“Ad “ZO “HO “Od “Ad “Vd “LV

Su ‘C661

voroury | snoseyd

[enusg a

sapisaq Jeay ourdstpy — aeurdsrpy

898T “IU0D2T
DAALIBAIA DAALIBAIA VINOLGVICT

(S661) ‘Je 39 18]

SIDUDIIFOY

DU OfIISIULAJLD
DISOLQUL]Z

s}soP]

JeuqeH

eoLoury | snos
UH | YH ‘S861 EHO

$311JUNOD popeAuy

snjyejsg

(GZLT ‘smiorqe.)
SUVANINS DULULDABOM
satgads

Ayruseygns 10 Ayrore,y

Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

290

spoos saiivdogs snoseyd (CEST ‘yeyuaT[Asy)
(SOQ0Z) ‘Je 39 sadr0g SNUULVYIOAVS II OZV-Ld| doing yerusunuo7 -oudyg snqpjoaaof snipiganag
sapieaq poss — sepryonig

(Aasinds uso7)) snoseyd (ZO8T

(Z00Z) X°D ‘(S66I1) XD sisuaaay vynssads ral q5| edomy jerusunuo; -ovdyg) Iasy[]) vwyynons saporydsgy
snoseyd (QSZ| ‘snoeuury)

(6861) OURIIASg pur sadi0g IEIOVISSLIG | =; ee odoingq yetusunu07) -o1fy snypgdazoskaga saporydsy
IeIIVOY snseone7)| snoseyd (€9Z] ‘yodoos)

(6861) OURIIAg pur sadi0g pur svaovioisy ‘odomgy [eusunuo7 -ovdy vautsnssal Déapopidasdoany
snoseyd (POGL ‘Jarueq

(FOG) UssueTT WNISAL) sdiy -oikyg| ‘[) syjostaasg vlapopidasz0ayy

snoseyd €/6] ‘Aonssy

(Z00Z) XO) | suusspuusoy ‘snucdgy adoing yetusunu07) -o1dyg SAP1OIVAAT]GO SNSAVIIGUOT

(9S81

deIOVUISEIOY JOYyI0 snoseyd| ‘aneyuasoy) sugwyoundisaqy

(6861) OURsIAg pure sodi0g | pur syvuzsif[o ospi0g UOTSII ULIULIIUpIy[ -oudy SNIVIIUNGILAV] SNSADIIBUOT
snoseyd (ZZ81

(6861) CURIA pur sadi0g OBVIUD] J adoinyq yeyuounuo07) -oudyg| ‘ady) amagosine snsavqisuoT
snoseyd (COST

(6861) OURIIAGg pur sadi0g WnUuvIOS I OZV-Ld| doing yerusunu0s -ouy ‘yooy) suassagnd xiugdq

snoseyd (S8Z] ‘Aosyjoor))

(6861) OURIIAg pur sadi0g IeUTUIR IS) I OZV-Ld| 2doimq yerusunu0; -ouy SISUMLOY VULIUIOIAVY’)
(SUT[YDI0A MOpesut) snoseyd QQ]

(Z00Z) X°D sssuaqvad SnAhGIVT ZI q5| edommg jerusunuo; -ouyg ‘ISIN VIVIYIULADI VIL

odoing yenuay pue| snoseyd BCR ‘aT[Aouayy

(6861) OURIIAg pur sadi0g SULA ) eee UJOYINOS “UIOISIA/\\, -oiyg) -ulionr vevgdojaduv vouty

sapisaq vay -oeUlDTy
sa1gad
SIDUDIIFIY eugey | selunos popeauy aSuvs sANeNy suIsaY Ajreseyqns so Ayrure.y

‘OL07 Arensqay [ aepdn aseq *({] xrpuadde das) SIN (A 0} JoJor suoMerAcIq
-qe ie1qey *([ xipuedde as) 99, ¢ OS] 02 JoJo suonerAcrqqe sopoo ANuNO’ ‘adorng wz uarye soieds seprpaurosdryy dy} JO soMsTIsIOVIeYO pur IST] *7'°E"Q BIqUL

291

Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3

(snseredse pyr)

snypqsoad sypuisifo
‘Vy ‘(snseredse
uapies) syvuzoiffo eIsy [enuay| snoseyd (SSLI
(S007) ‘Te 2° [EH ‘(Z00Z) X°D | syvuzoifo snévivdsy (‘I q5| ‘odomg jerusunuoe; -oudyg|] ‘snoeuury) tyuvdsv st1a9014)
Sa]120q FEI] -SeUTIODOTIZ)
VN “AS “aS ‘NU
‘ON “IN ‘CW ‘AT
‘AT ‘LI TI “aI “NH
spaas ‘aD ‘Id “Ad “IG snoseyd CEST TeyusyAsy
(0661) yarlens | vz, ‘suaT snskqwT Ad ‘ZD ‘HD “Ag | vorser uvrsurisupsyy -outy STULOILIVUBIS SNUINAT
(OLOZ) selzeuesz spoos 2191, snoseyd
IP OUIIIGO) *(CQOZ) ‘Te 19 sads0g UNSIT SNAMGYIVT I| NVOD -Sd ‘OZV-Ld OTOTwARTeY ISI -oidyg| €8Z] IsqsoH sadifns sngonag
(S661) ‘Te snoseyd
19 913] ‘(Q[OZ) SeTeUeD Op OUTATGOr) Spoos VIIA SUdT NVO-Sa odoiny usoynosg -oVdyd| 66Z1 ‘Yolyory s1uaq sngonag
VN ‘AS “AS ‘OU
‘AVW-Ld “OZV-Ld
‘ON “IN ‘AT ‘OT
‘TT “aI ‘NH “dD
(0661) 422/235 “(O10Z) seEeUR_ ap ‘Td ‘NVO-Sa “IG snoseyd
OUIOIGO) ‘(600Z) eovdomny euney spaos suaT “Ad ‘ZD ‘HD “Ag | vorser uvsuersoipoyy -ouyg 66L1 “Yor 2049 sngonag
spoas (Yysni
-QND BIS) SHUIILADUL
snusogasoquog
‘(as103) snavdosna
x9) ‘(J2AojO
SeZ-31Z) WNIpaU
‘I, §(JaA0]9 pos) snoseyd
(L661) 28PoH ‘(Z007Z) X°D suawad unyofisy adoinq yeyuounuo7) -oudy (JOIAI[Q) Saitva snipigong
spoas sasiqd’) snoseyd (CEST ‘yeyuoTfAs)
(SOQOZ) ‘Je 39 seds0g ‘StUOUQ) “vasstuaty OZV-Ld | Uorser uvsuespayy -ouky snupuIplay snipigonag
sated
,eugqep] | sarjunos popeauy oSues sAneyy suISAY Ayrureyqns so Ayrore,y

SIDUDIIFIY

Ron Beenen & Alain Roques / BioRisk 4(1): 267-292 (2010)

292,

(6661) Aowphrys pue pueppng

snug)

ZI qy adomy

snoseyd
“oud

(991 TITRW)
mjoaqny] vInsavBoyIUvy

sajjaag va] BUIZIU0IIIIYS -aVULINAIVE)
snoseyd (€Z8] ‘uueWwossnig)
(ZS6T) FPP osvoIpayy adoiny usoiseq -ouyg VIVILULOL DUMIOLUOL
JeIDVIWIL'T
ayo pure “dds
pyiuayy ‘(punoyaroy
rq) mdru vaoyyg
‘(urequeyd 10MqI1) snoseyd (SLLI
(L007) XOD| wsvjoaaun] osviur)T UOISII ULIULIIONPI/l -oudyg | ‘smoiqey) wyuvq vuroslig’)
snug],
(8861) sdk] ‘(€9G6T) UOsUYOf[ *(Z00Z)| wayvs ‘wnpuvavT snoseyd QC/] ‘snoeuuryT
xO’) ‘(TQ0Z) UeWaPyUIA\ pure UsuIeg ‘SNULADULSOM 7A IN ‘GD “ag | uorsar UvoUPIIOUpPI/ -ouyg DUVITAIULD pUurjOshLG)

(6861) OURAIag pue so810g

(GE81) susydarg (2007) XOD

SIDUDIIFIY

DIADUOUWIN

UNIVINIDU WNL
dvIOR TIT Joyo pue
DIADIJUIAY UNIT

URIUPIIDNPIy

UIOISIAN\

adoiny yeyuounuo07)

,euqep | soisjunoo popeauy aSuvs sANeN

snoseyd
“oud

UIIBIY

saqaaq Jeo] — seuTaWOSATY)
SO8I FIATO "V
"xy snasngdns snjvqdas0jday

Jareaqased — seutpeydasoid Ar)

(€9Z1 ‘Hodoos) 29747 sesavonrT
sated
Ayruseyqns 10 Aprure.y