Apeer-reviewed open-access journa 1] 

BioRisk 4(1): 315-406 (2010) 2 . 
doi: 10.3897/biorisk.4.6 | RESEARCH ARTICLE B | O R IS 

www.pensoftonline.net/biorisk 

Coleoptera families other than Cerambycidae, 
Curculionidae sensu lato, 

Chrysomelidae sensu lato and Coccinelidae 
Chapter 8.5 

Olivier Denux', Pierre Zagatti* 

| INRA, UR633 Zoologie Forestiére, 2163 Av. Pomme de pin, 45075 Orléans, France, 45075 Orléans Cedex 
2 INRA — Centre de recherche de Versailles, Unité PISC, Route de Saint-Cyr, 78026 Versailles Cedex, France 

Corresponding authors: Olivier Denux (olivier.denux@orleans.inra.fr), Pierre Zagatti (pierre.zagatti@ver- 

sailles.inra.fr) 

Academic editor: David Roy | Received 4 February 2010 | Accepted 23 May 2010 | Published 6 July 2010 

Citation: Denux O, Zagatti P (2010) Coleoptera families other than Cerambycidae, Curculionidae sensu lato, Chryso- 
melidae sensu lato and Coccinelidae. Chapter 8.5. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 
4(1): 315-406. doi: 10.3897/biorisk.4.61 

Abstract 

Here we consider 274 alien Coleoptera species belonging to 41 of the 137 beetle families in Europe (Cer- 
ambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae are treated separately else- 
where). Among the families we consider as having invaded the European fauna, Acanthocnemidae and 
Ptilodactylidae represent new arrivals. Many species-rich families have surprisingly few aliens, whereas some 
relatively minor families such as Dermestidae, Nitidulidae and Anobiidae have a relatively high representa- 
tion of alien species. Since the start of the 19th century, the number of coleopteran aliens introduced into 
Europe has continued to increase. Alien species colonizing Europe derive from a wide range of geographic 
regions as well as ecozones, but the most important source area is Asia. The countries with the largest number 
of alien species established are France, Germany and Italy. The majority have been introduced accidentally 
via international transport mechanisms. The most important route for importation is stored products and 
crops, followed by transport of wood, then horticultural and ornamental plants. Most alien species in these 
families are found within anthropogenic habitats in Europe. The introduction of invasive alien beetles in 
these families has had significant economic impacts, particularly as pests of stored foodstuffs, as well as seri- 
ous ecological impacts. For example, the buprestid species Agrilus planipennis, recently recorded in Russia, 

is an important potential economic threat which may also impact the biodiversity associated with ash trees. 
Keywords 

Europe, beetles, Dermestidae, Nitidulidae, Anobiidae, alien species, invasive species, stored products, pests 

Copyright O. Denux, P. Zagatti. 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. 


316 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

8.5.1. Introduction 

Introductions of alien species in Europe started in ancient times (Genovesi and Shine 
2003), but this phenomenon has grown rapidly during the two last centuries. This is 
considered largely to be a consequence of the globalization of trade (Smith et al. 2007). 
Among these introductions, Coleoptera dominate the alien terrestrial invertebrates in Eu- 
rope, where the fauna consists of over 27,000 species in 137 families (Fauna Europaea 
Web Service). In addition to the alien species observed in the families Cerambycidae, Cur- 
culionidae (sensu lato), Chrysomelidae (sensu lato) and Coccinelidae, which were treated 
in the preceding chapters, 274 other beetles of exotic or cryptogenic origin have been 
established to date in Europe (Table 8.5.1). These alien species belong to 41 different 
families. Additionally, 237 species are considered to have been introduced through human 
activity from one region of Europe to another (Table 8.5.2). However, the cause of such 
movements are often difficult to ascertain, particularly where the original range is poorly 
known. Thus, the analyses detailed below will mostly consider the species alien to Europe. 

8.5.2 Diversity of alien coleopteran species 

The Coleoptera families treated here with the greatest number of species in Europe are 
Staphylinidae (rove beetles), Carabidae (ground beetles) and Tenebrionidae (darkling 
beetles) but these have proportionally few alien species (figure 8.5.1). These three fami- 
lies constitute an important component of the European ground fauna (Dajoz 2002). 
Conversely, the families with the most aliens in Europe and significant economic im- 
pact tend to be families with relatively few native species such as Dermestidae (carpet 
beetles), Nitidulidae (sap-feeding beetles) and Anobiidae (death-watch beetles). Two 
of the 41 families do not have any native species in Europe and they are new arrivals 
for the European fauna: Acanthocnemidae (little ash beetles) and Ptilodactylidae (toe- 
winged beetles). The following presentation of families follows the taxonomic classifi- 
cation of Fauna Europaea (Fauna Europaea Web Service) and of the Tree of Life Web 
Project (Maddison et al. 2007) (for Ptilodactylidae, not included in Fauna Europaea). 

ADEPHAGA 

The Carabidae, are widespread and known to colonize a great diversity of ecological 
niches (Denux et al. 2007, Holland 2002). They are typically predators (as larvae and 
adults), although some groups (e.g. Harpalinae) have evolved toward granivory (feeding 
on seeds). These life traits do not favour passive transportation by humans, and thus, 
only eight alien species have been established in Europe, accounting for approximately 
0.2% of the European carabid fauna. Among these, Somotrichus unifasciatus, Trechicus 
nigriceps and Plochionus pallens have benefited from the global trade in food products to 
become cosmopolitan, being introduced with cargos of groundnuts, rice, broad beans, 


Coleoptera families other than Cerambycidae, Curculionidae... eae 

No. species % species 
5000 4000 3000 2000 #1000 0 0 4 8 12 16 20 

perestee 
taphylinidae 
4728 Ni ulidae 
Anobiidae 
Latridiidae 
Tenebrionidae 
c Bales'ncl 
tophagidae 
"“Givamicise 
Histeridae 
Hydrophilidae 
Carabidae 
Cleridae 
Bostrichidae 
Laemophloeidae 
yctidae 

Anthicidae 4 
Aphodiidae 4 
lateridae 

Alien species 

Native species Buprestidae 

| 
Zopheridae 3 
Trogossitidae 3 
Languriidae 3 
Endomychidae 2 
2 
2 
2 
2 
2 

Mycetophagidae 
Cybocaphilicaes 
Trogidae 
ral ionidae 
Ptilodactylidae 
Dytiscidae 
Mordellidae 
Rutelidae 
Ciidae 
Phalacridae 
Corylophidae 
Erotylidae 
Clambidae 
Salpingidae 
Ripiphoridae 
Passandridae 
Acanthocnemidae 

Figure 8.5.1. Relative importance of the Coleoptera families other than Cerambycidae, Curculionidae 
sensu lato, Chrysomelidae sensu lato and Coccinelidae families in the alien and native fauna in Europe. 
Right - Relative importance of the families in the alien entomofauna. Families are presented in a decreas- 
ing order based on the number of alien species. Species alien zo Europe include cryptogenic species. The 
number over each bar indicates the number of alien species observed per family. Left - Species richness 
of the same families in the native European entomofauna. The number over each bar indicates the total 

number of species observed per family in Europe. 

cocoa, etc. (Jeannel 1942, Weidner et al. 1984). Only one species is established through- 
out Europe: Trechicus nigriceps (recorded in 30 countries). This species seems to have 
been imported from the Eastern coast of Africa several centuries ago (Jeannel 1942). 

The Dytiscidae (predaceous diving beetles) are all aquatic carnivores. Only one 
dytiscid beetle has been reported in our database (DAISIE). This large South American 
species, Megadytes costalis, has been recorded once in Great Britain, but there is no data 
on its establishment in the wild. 

POLYPHAGA STAPHYLINIFORMIA 

The Hydrophilidae (water scavenger beetles) are another family of aquatic beetles, eas- 
ily distinguished from the Dytiscidae by the length of their maxillary palpi. One tribe, 
the Sphaeridiini, is exceptional due to its terrestrial, saprophagous and coprophagous 
habits. Many species share mammal dung with scarab beetles. Significantly, among 
eight hydrophilids reported as aliens in Europe, seven belong to the Sphaeridiini. 


318 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

The Histeridae (clown beetles) are mainly predators, specializing on sapropha- 
gous, coprophagous or necrophagous prey. Eight species have been reported in the da- 
tabase, but little is known about their life traits, except for the widespread, cryptogenic 
Carcinops pumilio, which is common everywhere in natural and anthropized habitats. 

The Ptiliidae (featherwinged beetles) are a very small family (120 species in Eu- 
rope and 180 in the world) of which 12 alien species have been recorded in Europe. 
These are very tiny beetles, including the smallest of all, with a length of just 0.5 mm, 
whilst even the largest members of the family do not exceed 2 mm. Adults and larvae 
are usually found in rotting organic material in a wide range of habitats. Their small 
size and lifestyle means that they are easily dispersed via the movements of soil. 

Staphylinidae is the most important group of Coleoptera in Europe and the second 
richest in the world (with over 46,000 species), but the number of alien species (31) in 
Europe is proportionally low, representing 0.7% of the whole of the Europeans staphyli- 
nid fauna. Many genera were not included in Fauna Europaea (Fauna Europaea Web 
Service), due to the lack of taxonomic expertise. Staphylinidae alien species found in Eu- 
rope are essentially predatory (Coiffait 1972, Paulian 1988) and mainly species associated 
with compost, humus and decomposing matter (Cho 2008, Odegaard and Tommeras 
2000, Tronquet 2006), such as Bisnius parcus, Lithocharis nigriceps and Oxytelus migra- 
tor. One predatory species, Philonthus rectangulus, has been reported from 36 countries/ 
islands. Originating from temperate East Asia, it may have expanded westward naturally. 

POLYPHAGA SCARABAEIFORMIA 

The Trogidae (hide beetles) are a small family of beetles related to the scarabs. They 
feed on mammal skins and furs, or on bird feathers, either as late arriving necrophages 
on carrion, or as commensals of vertebrates in their nests. Two species from Australasia 
have been recorded in Spain in our database. 

The Aphodiidae (dung beetle) are mainly small dung beetles, frequently included 
in the Scarabaeidae. Four species have been recorded as aliens, in one country only. 
Both Saprosites species introduced in Great Britain seem to be saproxylic beetles (An- 
gus et al. 2003). 

The Rutelidae (leaf chafers) are a family of brightly-coloured beetles, especially 
diverse in the tropics. Only one species of this family has been found in the Azores, the 
well-known Japanese beetle, Popilia japonica, which is considered as a severe pest in the 
United States, where it was introduced from Japan in 1912. 

POLYPHAGA ELATERIFORMIA 

The Clambidae (minute beetles) are very small beetles that have the capability to roll 
into a ball. One species is listed here, the Australian Clambus simsoni, a saprophagous 
species which seems to be rapidly expanding in western Europe. 


Coleoptera families other than Cerambycidae, Curculionidae... 319 

The Buprestidae (metallic wood-boring or jewel beetles) are a well-known family 
of xylophagous beetles. In most cases, the larvae develop in living wood, and a few spe- 
cies became major pests in orchards or forests. Only three buprestid species have been 
reported as aliens in the database, each observed in only one country. 

The Ptilodactylidae, the “toed-winged beetles”, are a group of elateriform Co- 
leoptera, which was formely treated as part of the Dascilloidea and included in Byr- 
thoidea (Maddison et al. 2007). Little is known of the biology of adults (Aberlenc and 
Allemand 1997). The habit of soil-leaf litter dwelling of both the adults and larvae 
facilitates their distribution with potted plants (Mann 2006). 

The Elateridae (click beetles) are a large family of beetles with quite diverse life 
history traits. Some species have soil-living larvae, either predators or rhizophages, with 
reported agricultural pests in the latter category. Other species are saproxylic (predators 
or saprophages), some of which are very specialized, and have high conservation value. 
‘Three species are reported as aliens here, occuring in one country each. ‘The life history 
traits of these species remain unknown. 

POLYPHAGA BOSTRICHIFORMIA 

The European Dermestidae comprise only 139 species (less than 1% of the European 
Coleoptera fauna) yet they are the largest contributor to the database, with 40 species 
reported as aliens. Many species are synanthropic and associated with animal remains, 
leathers and skins, dried meats, woollens and furs (Delobel and Tran 1993), such as 
Dermestes vorax, D. frischi, D. maculatus, D. lardarius and Anthrenus flavidus. Some 
species eat stored seeds such as Trogoderma granarium. The protraction of the number 
of larval stages and longevity in suboptimal nutritive media (Delobel and Tran 1993), 
as well as the relevance of the food product trade, explain partly how the damaging 
pests of this family have easily conquered new territories. 

The Lyctidae (true powder-post beetles) are a very small family (13 species in Eu- 
rope) closely related to the Bostrichidae. All species are wood-borers, specializing on 
hardwoods. ‘They usually attack dry wood that is less than five years old, and may be- 
come important pests of structural wood or furniture. As inhabitants of raw or manu- 
factured wood products, they are easily transported. Six species have been reported as 
aliens in Europe, but only one, Lyctus brunneus, has been established throughout the 
continent for more than 150 years. 

The Bostrichidae (horned powder-post beetles) are a small family (37 native spe- 
cies in Europe). The native species are saproxylophages, whereas the aliens are either 
wood-borers or grain-feeders (apparently, some species show both feeding habits) 
(Lesne 1901). Seven species have been reported as aliens, and have been found in many 
countries. The wood-borers may cause important damage to manufactured objects, 
but the stored-product feeders (Dinoderus spp., Rhyzopertha dominica) are the most 
economically harmful. Among these, the lesser grain borer, Rhyzopertha dominica, has 
been observed in 34 countries/islands. 


320 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

The Anobiidae have 19 alien species compared to 402 native species in Europe. 
About 11 species are associated with stored food products and include devastating 
pests such as Lasioderma sericorne which attacks a wide variety of dried products of 
animal or vegetable origin (Espanol 1992, Weidner et al. 1984). Several species attack 
soft woody matter: wood in the case of Ernobius mollis, but also books in the case of 
Nicobium castaneum, which can cause irrepairable damage. Many cryptogenic anobiid 
species are established in Europe for centuries, and may be found in many countries. 

POLYPHAGA CUCUJIFORMIA 

The Nitidulidae have 26 aliens compared with 219 native species in Europe. A third 
of these have occurred as far west as Macaronesia, but the other species have expanded 
their range in many countries of mainland Europe. As the majority of species are pollen- 
eaters, phytophagous, mycetophagous or predatory, they have a particular agronomic 
importance, damaging crops and stored food products. Among these, the 13 aliens 
species of the genus Carpophilus cause damage to dried fruits (Weidner et al. 1984). 

The Cybocephalidae are a very small family, frequently subsumed within Nitidul- 
idae. Cybocephaline beetles are well known predators of armoured scale insects (Coc- 
coidea: Diaspididae) throughout tropical, sub-tropical and temperate regions of the 
world (Kirejtshuk et al. 1997).They are minute beetles, very convex and able to roll 
into a ball, as for Clambidae. 

The Silvanidae (silvanid flat bark beetles) are a small family (34 native species in 
Europe) of flat beetles, formerly included in the Cucujidae. These insects were original- 
ly mycetophages, living under the bark of trees, but the feeding habits of many species 
have adapted to grain and fruit feeding, so that they have become synanthropic pests 
of stored products (Ratti 2007). Nine species are listed in the database, among which 
three are cryptogenic, long-established species occuring in several countries, such as the 
sawtoothed grain beetle, Oryzaephilus surinamensis. 

The Laemophloeidae (lined flat bark beetles) are a small family of flat beetles 
with 23 native species in Europe, which was formerly included in the Cucujidae. They 
are closely related to the Silvanidae, and show the same life history traits. Six species, 
belonging to the genus C7yptolestes, are reported as aliens in Europe. They have estab- 
lished successfully in many countries. 

The Phalacridae (shining flower beetles) are a small family of minute, rounded 
beetles. One North American species of Phalacrus has been recorded in the Azores, 
whose biological traits remain unknown (many species are micro-mycetophages). 

The Cryptophagidae (silken fungus beetles) are an important family of myce- 
tophagous insects with 228 native species in Europe, living in various habitats. Ten 
species have been reported as aliens in Europe, which are now established in many 
countries (the Cryptophagidae have the widest species range). The majority of these 
species (Cryptophagus spp.) are cryptogenic, feeding on fungal spores or decaying veg- 
etal material, sometimes on stored products. 


Coleoptera families other than Cerambycidae, Curculionidae... 321 

The Languriidae (lizard beetles) are a small family (12 native species in Europe) of 
phytophagous or saprophagous beetles. ‘Three alien species are considered here, with a 
rather low dispersal rate. Nevertheless, Cryptophilus integer and Pharaxonotha kirschii 
are reported as pests of stored products. 

The Erotylidae (pleasing fungus beetles) are a small family of mycetophagous bee- 
tles, with many species in saproxylic habitats. One Japanese species, Dacne picta, has 
possibly been introduced in Central Europe. 

The Cerylonidae (minute bark beetles) are a small family of saproxylic beetles. 
They just appear here because a well-known pest of stored grain, Murmidius ovalis, is 
now included in this family (formerly Murmidiidae). This is a cosmopolitan species 
probably originating from tropical Asia. 

The Endomychidae (handsome fungus beetles) are a small family of mycetopha- 
gous beetles (Shockley 2009, Shockley et al. 2009b), closely related to the Coccinel- 
lidae. Two very small species (Holoparamecus spp.) are cryptogenic and may be found 
in many countries worldwide. 

The Corylophidae (minute hooded beetles) are another small family of micro- 
mycetophagous beetles, which occur in a variety of habitats. One species, Orthoperus 
aequalis, from Australia, has now established in 10 countries within Europe. 

The Latridiidae (minute hooded beetles) are also a small family with 171 native 
species in Europe and 17 aliens which are essentially mycetophagous and associated 
with stored food products, such as Dianerella filum or Cartodere nodifer. These species 
are also plaster beetles which occupy wet places in the plastered walls of houses (Bouget 
and Vincent 2008). However, these latridiids do not appear to have an economic im- 
pact (Delobel and Tran 1993) and merely indicate bad food storage conditions. 

The Trogositidae (bark-gnawing beetles) are a small family of saproxylic insects, 
living as saprophages or predators of other insects under the bark of trees. The three 
species reported here are predators of cosmopolitan pests of stored products. 

The Cleridae (checkered beetles) are a conspicuous family of brightly coloured 
insects. Nearby all species are predators of other insects. Seven species are reported as 
aliens in the database, some of them (Necrobia spp.) established in Europe for a long 
time. These are either predators of xylophagous beetles or predators of stored product 
insects, and thus likely to be transported everywhere with their prey. We include here 
in the Cleridae the small family Thanerocleridae, which shows life traits similar to the 
typical Cleridae, with one introduced species, Thaneroclerus buqueti. 

The Acanthocnemidae, have only one alien species: Acanthocnemus nigricans 
which is attracted by forest fires (Schmitz et al. 2002). The recent worldwide expansion 
of this species is due to the commercial export of Australian wood (Kreiss et al. 2005). 

The Mycetophagidae (hairy fungus beetles) are a family of saproxylic insects, feed- 
ing on tree fungi. Two species, specialized on fungi growing on rotten vegetal material, 
are reported in the database. Typhaea stercorea is a well-known cryptogenic species, 
whereas Litargus balteatus is an American species found only recently in Europe 

The Ciidae (minute tree-fungus beetles) are another family of saproxylic insects 
feeding on tree fungi. Only one species (out of 76 occurring in Europe) is reported 


322 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

here as alien, Xylographus bostrichoides. This small insect probably originates from Asia 
and has to date been found in 19 European countries. 

The Mordellidae (tumbling flower beetles) are a large family (256 native species in 
Europe) of flower-dwelling insects, with endophytic larvae. Only one species, Mordel- 
listena cattleyana, is considered as an alien in Europe. This is a neotropical insect whose 
larvae develop inside tissues of ornamental orchids (Costa Lima 1955). This behaviour 
may have enabled its importation through the horticultural trade, since it has been 
found in Germany and the Netherlands. 

The Ripiphoridae, formerly spelled Rhipiphoridae (wedge-shaped beetles), are a 
small family of strange parasitic insects. Their larvae develop in other insect orders, 
namely Hymenoptera, Orthoptera or Dictyoptera. One species, Ripidius pectinicornis, 
has sometimes been found in harbours, along with its host cockroaches (mainly Blatta 
orientalis). 

The Zopheridae (ironclad beetles) were previously included in the Colydiidae. 
This is a family of saproxylic, bark-living insects with 125 native species in Europe. The 
three species reported as aliens in Europe are probably predators of other saproxylic 
insects. They are established in one country only, or a small number of countries in the 
case of Pycnomerus inexpectus, a species found in tropical greenhouses. 

The Tenebrionidae is mainly composed of saprophagous species. Many species are 
xerophiles or thermophiles, which explains their predominance in areas with hot climate 
and their low representation in more temperate zones (Dajoz 2002). About 15 tenebrio- 
nid alien species are present in Europe (1.1% of European tenebrionid fauna). The ma- 
jority of these species are associated with spoiled or wet cereals (Weidner et al. 1984). 
They include very damaging pests, such as species of Tribolium, which enter cracks in wet 
or already damaged seeds, and Alphitobius spp., which feed on mildewed food products. 

The Salpingidae (narrow-waisted bark beetles) are a small family of saproxylic bee- 
tles with 18 native species in Europe. One species only is mentioned here, Aglenus brun- 
neus, formerly included in the Colydiidae (Zopheridae). It is a very small, blind insect, 
often found in stables or poultry houses, where it feeds on animal waste (Dajoz 1977). 

The Anthicidae (antlike flower beetles) are small beetles resembling ground bee- 
tles. Four species are considered as aliens, among 310 native species living in Europe. 
These insects typically feed on rotten vegetal material, which has been heated through 
fermentation. These life history traits probably enable a wide tolerance to cold tem- 
peratures, and some species are cosmopolitan, found everywhere in the world, from 
tropical to boreal climates, e.g. Omonadus floralis, recorded in 40 countries. 

8.5.3 Temporal trends 

Some Coleoptera species were introduced to Europe a very long time ago. Fossils of 
alien species have even been found in archeological sites, such as the blind flightless 
beetle Aglenus brunneus in Iceland (Buckland et al. 2009) and Amara aulica (alien 
but native in Europe), which arrived in the Faroe islands with the Viking settlers 


Coleoptera families other than Cerambycidae, Curculionidae... 323 

(Brandt 2006). But the first date of introduction of a new species into a country is 
often difficult to establish. A species could have been present for years without its 
presence being noticed immediately. Particularly relevant here are small or incon- 
spicuous species lacking agronomic or economic impact (e.g. Ptiliidae), and mem- 
bers of neglected or hard to identify taxonomic groups (e.g. Cryptophagidae and 
Staphylinidae). 

The precise date of the first record is available for 201 species (i.e. 73.1% of aliens). 
The first statistical data derives from the beginning of the 19th century with the intro- 
duction of the nitidulid Carpophilus hemipterus in 1800 by the historical opening of 
trade routes (Audisio 1993). Then comes the trogossitid Tenebroides mauritanicus in 
1803, and the anobiid Nicobium castaneum in 1807. The endomychid Holoparamecus 
depressus arrived in 1843 and the anobiid Lasioderma sericorne in 1848. ‘These detriti- 
vores are all associated with stored food products or wood. 

We observed an accelerating increase in the number of new records per year (fig- 
ure 8.5.2), from 0.1 p.a. between 1800-1849 to 3.5 p.a. during 2000-2007, with an 
intermediate level of 1.3 p.a. during the period 1900-1924. During this last period, 
33 new alien species were recorded, including 14 alone for the year 1900. This unex- 
pected increase coincides with the industrial revolution of the first developing Euro- 
pean countries (Cosseron and Faverjon 1991) (Great Britain, Belgium, France, and 
Germany) and with the increase in imports ensuing from it. 

8.5.4 Biogeographic patterns 

8.5.4.1 Origin of alien species 

Alien species come from all continents except Antarctica (figure 8.5.3) (arthropods 
most represented on this continent are Collembola and mites rather than beetles) 
(Schulte et al. 2008). The considerable periods of environmental stress in Antarctic 
(Benoit et al. 2009) limit the diversity of insects, even though a very few beetles do 
occur there (Vernon et al. 1999), such as the ground-beetles Amblysogenium pacificum 
and A. minimum. These factors explain easily the absence of invasives coming from 
Antarctic. 

About 82 aliens have origins currently considered cryptogenic. These are cosmo- 
politan species or distributed mainly in on one or more ecozones, with a tendency to 
become cosmopolitan. This is particularly the case with the cryptophagid Cryptophagus 
cellaris, a holarctic species which has become practically cosmopolitan following inter- 
national commercial exchanges (Delobel and Tran 1993). 

Asia is the most important source of aliens, with 58 species established in Europe 
(21%), comprising Dermestidae (13 spp.), Staphylinidae (8 spp.), Nitidulidae (6 spp.), 
Anthicidae (4 spp.) and Carabidae (3 spp.). These families are generally associated with 
stored products, crops, decomposing matter such as compost, and to a lesser extent 
with wood. The 16 other families number one or two species of aliens each. 


324 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Mean number of new alien species 
recorded per year during the period 
0 | 2 3 4 
| | 1 | J 

1800-1849 - 6 
1850-1899 i 22 

1900-1924 NS 23 
1925-1949 i 24 

1950-1974 i 34 
1975-1999 i 54 

2000-2007 - is 2° 

Time period 
Figure 8.5.2. Temporal changes in the mean number of new records per year of alien Coleoptera spe- 
cies of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coc- 
cinelidae, from 1800 to 2007. The number over each bar indicates the absolute number of species newly 

recorded per time period. 

About 35 aliens come from Africa and these comprise Nitidulidae (5 spp.), Cara- 
bidae (3 spp.), Histeridae (3 spp.), Hydrophilidae (3 spp.) and Tenebrionidae (3 spp.). 
Nitidulidae and Tenebrionidae have been transported through stored food products. 
The mode of introduction is unknown for Carabidae and Hydrophilidae. There are 
also 14 other families having one or two alien species, which are partly associated with 
stored food products and wood. 

The 55 aliens coming from the American continent (20% of the all alien species 
to Europe), include 24 species from North America and 31 species from Central and 
South America. From North America, the principal families are Dermestidae (7 spp.), 
Nitidulidae (6 spp.) and Tenebrionidae (4 spp.). Four species of Staphylinidae and four 
species of Ptiliidae derive from Central and South America. As for Asia and Africa, 
the neoarctic and neotropic aliens are mainly associated with foodstuffs and cultures. 
About 16 other families coming from America with one or two alien species have also 
been recorded in Europe. 

Relatively few aliens originate from Australia. The 25 species of Australian origin 
include Latridiidae (4 spp.), Ptiliidae (4 spp.) and Staphylinidae (3 spp.). These species 
have no economic impact. The 12 other families include one or two alien species each, 
among which are species of the stored food products (Ptinus ocellus, Anthrenus oceani- 
cus, Brachypeplus mauli) or living under the tree bark (Ptinella cavelli and P. errabunda). 

The aliens with a specifically tropical origin (Pantropical) are the least presented in 
Europe with 20 species, that is to say 7% of all exotic species to Europe. The families 


Coleoptera families other than Cerambycidae, Curculionidae... 325 

Africa 
12.7% 

Cryptogenic 
29.5% 

Asia 
21.1% 

Pantropical 
7.3% 

\ Australasia 
Central and South America \ 91% 

11.3% | 
North America 

9.1% 

Figure 8.5.3. Origin of the Coleoptera species alien to Europe of families other than Cerambycidae, 

Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae 

with the most species are Anobiidae (3 spp.), Bostrichidae (3 spp.) and Tenebrionidae 
(3 spp.). The eight other families have only one or two species each. These tropical 
aliens are associated with stored food products and fruits. 

During different time slices, the origin of alien species has increasingly diversified 
(figure 8.5.4). The number of ecozones represented has increased from three (Africa, 
Asia, Pantropical) during 1800-1849 to six since 1950-1974 (Africa, Asia, Australa- 
sia, Central and South America, North America, Pantropical). The geographic source 
has also varied temporally although Asia has always been both an important and early 
region of origin. This situation can be explained by the opening of the trade route 
between Europe and India by the Cape of Good Hope at the end of the 15th century 
(which was also the sole sea route before the opening of the Suez Canal in 1869) and 
the strong Western influence which followed, the opium wars and the East India Com- 
panies, which revolutionized methods and the extent of the trade with Asia. 

We highlight especially two ambiguous periods for biological invasions: 1850-1899 
and 1925-1949. During the first period, no new record of an alien from Africa was 
recorded in Europe. The same goes for the second period with a fall of the number of 
new arrivals detected from South America (nine in 1900-1924 and only two in 1925— 
1949). These phenomena may coincide with the Great Depression, the result of the 
economic crisis of 1929 (Cosseron and Faverjon 1991, Gravereau and Trauman 2001), 
which affected both the level of protectionism on trade routes and the overall volume 
of international economic exchange between Europe and its colonies. ‘The consequence 
for South America, Asia and Africa was “the crisis of dessert products”, coinciding with 
the fall of the purchasing power in Europe and North America. Thus in Brazil for exam- 
ple, in an attempt to control the market, coffee was burned in engines (Launay 1999). 


326 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Percentage of alien species 
per region of origin 
20 40 60 80 100 

o 

1800-1849 

1850-1899 [__| Cryptogenic 
(Africa 

1900-1924 PM Asia 
= Australasia 
(Peat] C & South America 

1925-1949 (I North America 
[___| Pantropical 

1950-1974 

1975-1999 

2000-2007 

Time period 

Figure 8.5.4. Temporal changes in the origin of the Coleoptera species alien zo Europe of families other 

than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae 

The late arrival of aliens to Europe from North America is remarkable (first record 
in 1935) and probably corresponds to weak exports of foodstuffs towards Europe (ex- 
cept cereals). For forest biotopes especially, the North American component of species 
is small and of limited economic impact in Europe (Dajoz 2007). 

8.5.4.2 Distribution of alien species within Europe and their range expansion 

The majority of European countries have been directly affected by alien species (fig- 
ure 8.5.5), but there is a very great mismatch in the number of species present in one 
country versus another. 

The archipelago of Svalbard, with an insect fauna of a meagre 230 species (Coulson 
2007), seems free from aliens. As in the case of Antarctica, the strong environmental 
contraints (harsh temperatures, shortened seasons and strong winds) have evidently 
limited the colonization of insects (Hulle et al. 2008) and geographical isolation has 
posed a barrier. For Macedonia there is a lack of readily accessible data (Tomov 2009), 
which has prevented us updating the situation there. 

The countries/islands most affected by aliens are France (126), Germany (107), It- 
aly (101), Austria (98), Great Britain (97), Switzerland (91), the archipelago of Azores 
(92), Denmark (89) and the Czech Republic (84). 


Coleoptera families other than Cerambycidae, Curculionidae... 327 

50°N- 

30°N 

Oo 

Number of alien species 

no data (ND 1-30) 0-62 I 101 - 126 

Co ss «5 ED 3-100 

Figure 8.5.5. Comparative colonization of continental European countries and islands by by the Cole- 
optera species alien zo Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomeli- 

dae sensu lato and Coccinelidae. Archipelago: | Azores 2 Madeira 3 Canary islands. 

The number of aliens per country is not significantly correlated with Global Do- 
mestic Product per capita (International Monetary Fund), latitude, nor longitude of 
the centroid of the country. In contrast, the number of aliens per country is signifi- 
cantly correlated with import (Spearman-Rho 0.650, P-value < 0.001) from 2003 to 
2008 (The World Factbook) and also more weakly with area (Spearman-Rho 0.432, 
P-value < 0.01). 

In spite of its geographical isolation (1500km from Europe, 1450km from 
Africa and 3900km from North America) and its small area, the archipelago of 
Azores has a large number of aliens. Since their historical discovery, the geographic 
position of the Azores has made the islands a strategic harbour for transatlantic 
ships, resulting in the introduction overall of several hundreds of taxa (Haggar 
1988, Heleno 2008). Twenty-four alien species have been recorded exclusively in 
the Azores archipelago. 


328 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Indeed, alien native species in Europe have colonized islands more than other con- 
tinental countries. The archipelago of Azores is the most affected with 126 alien species 
to Europe, followed by Great Britain (with 58 aliens), Faroe Islands (32 aliens) and 
Canary Islands (32 aliens). Perhaps surprisingly, Austria is the most important conti- 
nental country affect by alien native to Europe, with 13 species. 

8.5.5 Main pathways to Europe 

The most important pathways for accidental invasions of exotic species to Europe are 
those closely bound to international transport, whereas the most important processes 
relating to deliberate introductions are the biological control of agricultural pests and 
the pollination of crops (Ruiz and Carlton 2003). Rapidly developing international 
trade and the reduction of travel times by air to less than two days, have meant that a 
living insect can be transported almost any part of the world (Mouchet et al. 1995). 

Only three species have been introduced intentionally in Europe, two for biologi- 
cal control. The first is the cybocephalid beetle Cybocephalus nipponicus, originating in 
South Korea (Evans et al. 2005) and introduced into Italy for the control of cochineals 
bugs (Diaspididae) (Lupi 2002). The second species is Ripidius pectinicornis (Ripi- 
phoridae), a parasitoid of the german cockroach Blattella germanica (Falin 2001) which 
was released from culture and is now present in several European countries (Bétis 
1912). The third species is the tenebrionid Zophobas morio which has been used for 
bird and especially lizard food (Thomas 1995). 

About 98.9% of aliens have been introduced accidently in Europe. The exact path- 
way of introduction is difficult to establish. The introduction vector is unknown for 
123 aliens out of the total of 275. Theses aliens are essentially detritiphagous, saproxy- 
lophagous or predatory species. 

The first clearly identified means of importation is via stored products and crops 
(approximately 120 aliens, or 40%). This can be explained by the importance of the 
international stored products trade (cereals, fruits and vegetables) and the primary 
position of Coleoptera as pests of stored products (Delobel and Tran 1993). About 
20 Coleoptera have been implicated directly in the transport of woods. Some species 
have been found in wood derivatives such as Dinoderus minutus, a bostrichid intro- 
duced with furniture and bamboo-work (Lesne 1901). Few species have been identi- 
fied as transported with horticultural or ornamental products, despite the increase of 
economic importance of ornamental pot plants (Lawson 1996), in sharp contrast for 
example to the situation in Lepidoptera (see Chapter 11). However, the level may be 
underestimated for this route, as some Coleoptera tend to occur in compost and may 
pass unnoticed via the pot plant trade. 

The extruded starch products used as impact protection for fragile packing can 
even be a food source for stored grains pests (Fraga et al. 2009) as for Cryptolestes fer- 
rugineus, Lasioderma serricorne and Tribolium castaneum. Thus starch-packings could 
become a new vector of introductions in the future. 


Coleoptera families other than Cerambycidae, Curculionidae... 329 

8.5.6 Most invaded ecosystems and habitats 

The anthropogenic habitats most strongly colonized by coleopteran alien species (fig- 
ure 8.5.6), are buildings (50%), cultivated lands (20%) and forest habitats (10%). The 
large proportion of species associated with foodstuffs explains this relation. Conversely, 
the weak colonization of pseudo-natural habitats can be explained by the near-absence 
of phytophagous, and more particularly phyllophagous species among the coleop- 
teran families treated here. This result contrasts with the situation for other groups 
of predominantly phytophagous insects (Cerambycidae, Chrysomelidae, Lepidoptera: 
Chapter 8.1, 8.3, 11). 

In spite of the popularity of exotic species for the aquatic animal and plant trade 
(Leppakoski et al. 2002) and the fact that migrating waterfowl can transport aquatic 
invertebrates or their eggs (Figuerola et al. 2003), surprisingly no water beetle has been 
introduced into Europe, except for the dytiscid Megadytes costalis (again contrasting 
with the situation for Lepidoptera, the aquatic Pyraloidea: Chapter 11). This low im- 
portance of the aquatic route in Coleoptera is also observed in the United States, where 
only 2.2% of the invasive arthropods are aquatics (Pimentel et al. 2005). 

8.5.7 Ecological and economics impacts 

Most alien species do not become invasive in their new locations (Genovesi and Shine 
2003). It is often difficult to predict whether a new introduction will actually become 
established (Streito and Martinez 2008). However, the subset of alien species that are 
invasive may have significant environmental, economic and public health impacts and 
threaten the wholesale homogenisation of ecosystems (Sefrova 2005). 

Invasive alien species are now considered to be the second greatest cause of global 
biodiversity loss after direct habitat destruction (Simberloff 2001) and have adverse 
environmental, economic and social impacts from the local level upwards. 

The invasion of most Coleoptera treated here bears a direct relation to human pres- 
ence (synanthropic species). Their impact is essentially with stored foodstuffs which they 
can extensively damage (Sefrova 2005). Coleoptera damaging stored food products on a 
global economic scale are very few (Delobel and Tran 1993), but include several species 
of aliens in Europe, among which are Cryptolestes ferrugineus, C. pusillus, Lasioderma ser- 
ricorne, Oryzaephilus surinamensis, Rhyzopertha dominica, Tribolium castaneum, T: confu- 
sum and Trogoderma granarium. The impact of insect pests in a given situation can widely 
fluctuate depending on various parameters, in particular on production levels and the 
commercial value of those products infested both in time and in a geo-economic context. 
However, these synanthropic species are not known to have a direct effect on biodiversity. 

The situation for agronomic and forest species can be different. The buprestid 
Agrilus planipennis, recently recorded in European Russia, is a very good example. ‘This 
xylophagous East Asian species is presently causing significant damage to ash trees 

(Fraxinus spp.) in North America (Baranchikov et al. 2008). A. planipennis has killed 


330 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Percentage of alien species 
living in the habitat 
) 20 40 60 

B - Coastal habitats 

C - Inland surface water habitats 

D - Mire, bog and fen habitats 

E - Grassland and tall forb habitats 

F - Heathland, scrub and tundra habitats 
G - Woodlands and forests 

H - Inland without vegetation 

| - Agricultural and horticultural lands 

J - Buildings, houses 

X. Large parks and domestic gardens 

Habitats 

Figure 8.5.6. Main European habitats colonized by the Coleoptera species alien to Europe of fami- 
lies other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae. The 
number over each bar indicates the absolute number of alien coleopterans recorded per habitat. Note that 

a species may have colonized several habitats. 

over 15 million forest and ornamental trees in several US States in less than 10 years 
(Poland and McCullough 2006). It is alarming that European ash trees are not more 
resistant than those of North America (Baranchikov et al. 2008). Agrilus planipennis 
could become a serious pest in Europe with a dramatic economy impact as well as 
potentially for biodiversity associated with Fraxinus. 

Many species are associated with compost and even while their economical impact 
may be negligible (as mainly predators or detritivores), ecological disruption may still 
occur. This appears to be the case with the Staphylinid Lithocharis ochracea. This native 
beetle has declined, supplanted by the alien species L. nigriceps (Odegaard and Tom- 
meras 2000, Tronquet 2006). 

Even if the eradication of invasive species seems possible in Europe and in particu- 
lar for mammals (Genovesi 2005), the possibility of eradication of invasive Coleoptera 
appears much more remote. 

8.5.8 Conclusion 

On of the most striking consequences of globalization is the increase in the problem 
of invasive species (Perrings et al. 2005). The volume of international trade and travel 
is now so great, and the modes of entry so varied, that not all consignments or routes 
of entry can be screened (Levine and D’Antonio 2003). Three categories are particu- 
larly important to highlight for the coleopteran alien species treated here: synantropic 


Coleoptera families other than Cerambycidae, Curculionidae... 331 

| mm 

Figure 8.5.7. Habitus of some Coleoptera species alien to Europe. a Ernobius mollis b Tribolium casta- 
neum © Oryzaephilus surinamensis d Alphitobius diaperinus e Cryptolestes duplicatus § Dermestes lardarius 
g Gnathocerus cornutus h Rhizopertha dominica i Necrobia ruficollis j Trechicus nigriceps k Lyctus brunneus 
| Gibbium psylloides (Credit: Pierre Zagatti). 


332 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

habitats with essentially stored products, compost (probably that associated with orna- 
mental plants), and forest or wood-derived products. 

Acknowledgements 

We thank Alain Roques and David Roy for their useful comments on the manuscript 
and David Lees for linguistic improvements. 

References 

Aberlenc HP, Allemand R (1997) Acclimation en France de Ptilodactyla exotica, espéces a 
moeurs anthropophiles (Coleoptera, Ptilodactylidae). Bulletin de la Société entomologique 
de France 102: 93-102. 

Abood FE, Murphy RJ (2006) World distribution of Minthea rugicollis (Coleoptera: Lyctidae). 
Journal of Tropical Forest Science 18 (4): 250-254. 

Allemand R (2008) Anobiidae nouveaux ou méconnus de la faune de France (3e note) (Coleop- 
tera). Bulletin de la Société entomologique de France 113 (3): 397-402. 

Alonso-Zarazaga MA, Sanchez-Ruiz M, and Sanchez-Ruiz A (2003) Une nueva familia de 
Coleoptera para Espana: Acanthocnemidae. Bol. S.E.A. 32: 179-180. 

Anderson R (1997) Northern Ireland species inventories. Rove beetles (Coleoptera: Staphyli- 
nidae). Environment and Heritage Service. http://www.ehsni.gov.uk/pubs/publications/ 
Staph.pdf 

Anderson R, McFerran D, Cameron A (2000) The Ground Beetles of Northern Ireland (Co- 
leoptera, Carabidae). Atlas of the northern Ireland Flora and Fauna, Volume 1. Ulster 
Museum, U.K., Belfast. 256 pp. 

Angus RB, Wilson CJ, Maté JF. Hammond PM, Mann DJ (2003) Saprosites mendax (Black- 
burn) and S. natalensis (Peringuey) (Scarabaeoidea, Aphodiidae), two species introduced 
into Britain. Proceedings of the Second Pan-European Conference on Saproxylic Beetles, 2003, 
72-76. 

Arndt E (2006) Niche occupation by invasive ground-dwelling predator species in Canarian 
laurel forests. Biological Invasions 8: 893-902. 

Audisio P (1993) Coleoptera Nitidulidae - Kateretidae. Fauna d'Italia, vol. 32. Calderini, Italia, 
Bologna. 971 pp. 

Baranchikov Y, Mozolevskaya E, Yurchenko G, Kenis M (2008) Occurrence of the emerald ash 
borer, Agrilus planipennis in Russia and its potential impact on European forestry. OFPP/ 
EPPO Bulletin 38: 233-238. 

Baraud J (1985) Coléoptéres Scarabaeoidea: faune du Nord de l'Afrique, du Maroc au Sinai. 
Lechevalier, France, Paris. 651 pp. 


Coleoptera families other than Cerambycidae, Curculionidae... oi) 

Baraud J (1992) Faune de France: France et régions limitrophes . 78 . Coléoptéres Scarabaeoi- 
dea d’Europe. Fédération frangaise des sociétés de sciences naturelles and Société linnéenne 
de Lyon, France, Lyon. 856 pp. 

Barsevskis A, Bicevskis M, Valainis U, Savenkovs N, Cibulskis R, Kalnins M, Strode N (2004) 
Faunistic records of the beetles (Hexapoda: Coleoptera) in Latvia. 1. Acta Biol. Univ. Dau- 
gavp. 4 (2): 93-106. 

Batten R (1976) De Nederlandse soorten van de keverfamilie Mordellidae. Zoologische Bijdra- 
gen 19: 3-37. 

Beal RS, Kadej M (2008) Description of the Larva of Sefrania Bleusei Pic and assignment of 
Sefrania Sabulorum (Beal) to the New Genus Avaphonotos Beal and Kadej (Coleoptera: 
Dermestidae). Entomological News 119: 245-250. 

Bellés X (1985) Habitats and food habits of the Gibbiinae (Coleoptera: Ptinidae). Bull. Inst. 
Cat. Hist. Nat. 50: 263-267. 

Bellés X, Halstead DGH (1985) Identification and geographical distribution of Gibbium ae- 
quinoctiale Boieldieu and Gibbium psylloides (Czenpinski) (Coleoptera: Ptinidae). Journal 
of Stored Products Research 21: 151-155. 

Bengtson SA (1981) Terrestrial invertebrates of the Faroa Islands: IH. Beeltes (Coleoptera): 
check-list, distribution, and habitats. Fauna norv. Ser. B. 28: 52-82. 

Benoit JB, Elnitsky MA, Schulte GG, Lee RE, Denlinger DL (2009) Antarctic collembolans 
use chemical signals to promote aggregation and egg laying. Journal of Insect Behavior 22: 
121-133. 

Bercedo P (1997) El género Omorgus Erichson, 1847 en la Peninsula Ibérica (Coleoptera, 
Scarabaeoidea: Trogidae). Bol. S.E.A. 17: 29-31. 

Bercedo P, Arnaiz L, Coello P, Baena M (2005) Ozognathus cornutus (LeConte, 1859) new ano- 
biid for the Iberian fauna (Coleoptera: Anobiidae). Boletin Sociedad Entomologica Aragone- 
94372 DIBXDTA. 

Bercedo P, Becerra RG, Arnaiz L (2008) El género Calymmaderus Solier, 1849 nuevo para Ca- 
narias y descripcion de una nueva especie (Coleoptera: Ptinidae: Dorcatominae). Boletin 
Sociedad Entomologica Aragonesa 42: 33-35. 

Bétis L (1912) Faune entomologique Armoricaine, Coléoptéres Rhipiphoridés. Imprimerie 
Oberthur, France, Rennes. 40 pp. 

Borges PAV (1990) A checklist of the Coleoptera from the Azores with some systematic and 
biogeographic comments. Bol. Mus. Mun. Funchal 42 (220): 87-136. 

Borges PAV, Vieira V, Dinis F, Jarroca S (2005) List of Arthropods (Arthropoda). Jn Borges et 
al. (eds.): A list of the terrestrial fauna (Mollusc and Arthropoda) and flora (Bryophyta, Pteri- 
dophyta and Spermatophyta) from the Azores. Angra do Heroismo Horta, Ponta Delgada, 
163-221. 

Bouget C, Moncoutier B (2003) Contribution a la connaissance des Rhizophaginae de France 
(Coleoptera, Cucujoidea, Monotomidae). Bulletin de la Société entomologique de France 
108 (3): 287-306. 

Bouget C, Vincent R (2008) Les Latridiidae de la faune de France continentale et de Corse: 
mise a jour de la clé des genres et du catalogue des espéces (Coleoptera, Cucujoidea). Bul- 
letin de la Société entomologique de France 113: 101-120. 


334 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Boukal DS, Boukal M, Fikacek M, Hajek J, Klecka J, Skalicky S, Stastny J, Travnicek D (2007) 
Catalogue of water beetles of the Czech Republic (Coleoptera: Sphaeriusidae, Gyrinidae, 
Haliplidae, Noteridae, Hygrobiidae, Dytiscidae, Helophoridae, Georissidae, Hydrochidae, 
Spercheidae, Hydrophilidae, Hydraenidae, Scirtidae, Elmidae, Dryopidae, Limnichidae, 
Heteroceridae, Psephenidae). Klapalekiana 43 (Suppl.): 1-289. 

Bowestead S (1999) A revision of the Corylophidae (Coleoptera) of the west Palaearctic region. 
Muséum histoire naturelle Genéve, Suisse, Genéve. 203 pp. 

Brandt D (2006) More stamps and Story of the Faroe Islands. Postverk Foroya (The Faroese 
Postal Administration). Postverk Foroya (The Faroese Postal Administration), Denmark, 
Torshavn. 360 pp. 

Buckland PC, Panagiotakopulu E, and Sveinbjarnardottir G (2009) A failed invader in the 
North Atlantic, the case of Aglenus brunneus Gyll. (Col., Colydiidae), a blind flightless 
beetle from Iceland. Biological Invasions 11: 1239-1245. 

Bunalski M (1999) Die Blatthornkafer Mitteleuropas (Coleoptera: Scarabaeoidea): Bestim- 
mung, Verbreitung, Okologie. Frantisek Slamka, Slovakia, Bratislava. 80 pp. 

Callot H (2003) Cryptophilus obliteratus Reitter, 1878, espéece nouvelle pour la faune de France 
(Coleoptera, Languriidae). Bulletin de la Société entomologique de Mulhouse 59 (1): 5-6. 

Callot HJ (1993) Sur quelques staphylins capturés dans la Bas-Rhin et peut-étre nouveaux pour 
la Faune de France: Philonthus spinipes Sharp., Philonthus scribae Fauvel, Ontholestes haroldi 
Eppelsheim, Aleochara irmgardis Vogt. Bul. Soc. ent. Mulhouse Janvier-Mars: 13-15. 

Camerini G (2009) Factors affecting Lymantria dispar mortality in a willow wood in northern 
Italy. Bulletin of Insectology 62 (1): 21-25. 

Cho YB (2008) Korean species of the genus Bisnius Stephens (Coleoptera: Staphylinidae). En- 
tomological Research 38: 90-92. 

Cobos A (1986) Fauna ibérica de Coleépteros Buprestidae. Consejo Superior de Investiga- 
ciones Cientificas, Spain, Madrid. 364 pp. 

Coiffait H (1972) Coléoptéres Staphylinidae de la région paléarctique occidentale 1. Généra- 
lités, sous-famille Xantholinae - Leptotyphinae. Supplément a la Nouvelle Revue d'Entomo- 
logie 4: 1-593. 

Cosseron S, Faverjon P (1991) LEurope de 1815 a nos jours. Une histoire et une chronologie 
commentée. La Manufacture, France, Besancon. 641 pp. 

Costa Lima AM (1955) Insetos do Brasil. 9° Tomo, Capitulo XXIX: Coleopteros, 3a Parte. 
Escola Nacional de Agronomia, Rio de Janeiro 11 (9): 1-289. 

Coulson SJ (2007) Terrestrial and freshwater invertebrate fauna of the High Arctic archipelago 
of Svalbard. Zootaxa 1448: 41-68. 

Cuppen J (2003) Carpelimus zealandicus, een nieuwe kortschildkever voor Nederland (Coleop- 
tera: Staphylinidae). Nederlandse Faunistische Mededelingen 19: 35-40. 

Curtis J (1836) British entomology; being illustrations and descriptions of the Genera of In- 
sects found in Great Britain and Ireland. Vol XIII. pl. 578-625. Printed by Richard Taylor. 

DAISIE: European Invasive Alien Species Gateway. http://www.europe-aliens.org. 

Dajoz R (1977) Coléoptéres Colydiidae et Anommatidae paléarctiques. Faune de France et du 

Bassin méditerranéen. Masson, France, Paris. 280 pp. 


Coleoptera families other than Cerambycidae, Curculionidae... 335 

Dajoz R (2002) Les coléoptéres carabidés et ténébrionidés: écologie et biologie. Tec et Doc, 
France, Paris. 522 pp. 

Dajoz R (2007) Les insectes et la forét. Réle et diversité des insectes dans le milieu forestier. 2e 
édition. Tec & Doc, France, Paris. 648 pp. 

Darlington PJ (1964) Australian carabid bettles XIV. Perigona. Psyche 71: 125-129. 

De Laclos E, and Biiche B (2009) La Vrillette sans peine: quatriéme note (Coleoptera Anobii- 
dae). L’Entomologiste 65 (4): 207-213. 

Degiovanni A, Pezzi G (2007) Anthicus catalanus Bonadona, 1953 nuovo per l'Italia reperti di 
altre specie (Insecta Coleoptera Anthicidae). Quaderno di Studi e Notizie di Storia Natuale 
della Romagna 24: 69-77. 

Delobel A, Tran M (1993) Les Coléoptéres des denrées entreposées dans les régions chaudes. 
CTA/ORSTOM, France, Paris. 424 pp. 

Denux O, Augustin S, Berthelot A (2007) Biodiversité des Carabidae dans les peupleraies pi- 
cardes (Coleoptera). L’Entomologiste 63: 243-256. 

Du Chatenet G (2000) Coléoptéres Phytophages d’Europe. Tome 1. NAP, France, Vitry-sur- 
Seine. 360 pp. 

Duff AG (2008) Checklist of beetles of the British Isles. A.G. Duff, United Kingdom, Somer- 
set, Wells. 164 pp. 

Enckell PH, Bengtson SA, Wiman B (1987) Serf and Waif Colonization - Distribution and 
Dispersal of Invertebrate Species in Faroe-Island Settlement Areas. Journal of Biogeography 
14 (1): 89-104. 

Espanol F (1979) Géneros de Dorcatominae de la fauna europea (Col. Anobiidae). Nota 92. 
Misc. Zool. Barcelona 5: 33-42. 

Espanol F (1992) Coleoptera, Anobiidae. Fauna Iberica. Vol. 2. Museo Nacional de Ciencias 
Naturales, CSIC, Spain, Madrid. 195 pp. 

Evans GA, Dessart P, Glenn H (2005) Two new species of Aphanogmus (Hymenoptera: Cera- 
phronidae) of economic importance reared from Cybocephalus nipponicus (Coleoptera: Cy- 
bocephalidae). Zootaxa 1018: 47-54. 

Falcoz L (1929) Tableaux analytiques des Coléoptéres de la faune franco-rhénane (France, Hol- 
lande, Belgique, Région rhénane, Valais). Vol. II: Famille 33: Cryptophagidae. Imprimerie 
des Miscellanea entomologica, France, Narbonne.197 pp. 

Falin ZH (2001) Notes on the Occurrence of Ripidius pectinicornis Thunberg (Coleoptera: 
Rhipiphoridae) in the Continental United States and Hawaii. The Coleopterists Bulletin 55: 
194-197. 

Fauna Europaea Web Service: Fauna Europaea version 1.1. http://www.faunaeur.org. 

Ferrer J (2004) Espéces de Cynaeus Leconte 1851, introduites en Europe du Nord. (Coleoptera, 
Tenebrionidae, Ulomini). Nouvelle Revue d'Entomologie (N.S.) 21 (2): 103-104. 

Ferrer J, Andersson B (2002) Species of Cynaeus found in Sweden and Finland, with a note of 
the identity between Cynaeus depresus (Horn 1870) and Cynaeus opacus (Champion 1886) 
syn. conf. (Coleoptera, Tenebrionidae, Diaperini). Entomofauna, Zeitschrift fur Entomologie 
23 (12): 145-148. 


336 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Ferrer J, Martinez Fernandez JC (2008) Blaps mortisaga (L.) o la leyenda de la muerte, una 
especie introducida en Europa boreal y occidental (Coleoptera, Tenebrionidae). Boln. Asoc. 
esp. Ent. 32 (3-4): 245-261. 

Figuerola J, Green AJ, Santamaria L (2003) Passive internal transport of aquatic organisms by 
waterfowl in Donana, south-west Spain. Global Ecology and Biogeography 12: 427-436. 
Fikacek M, Boukal M (2004) Pachysternum capense, a new genus and species for Europe, and 
updated key to genera and subgenera of European Sphaeridiinae (Coleoptera: Hydrophili- 

dae). Klapalekiana 40: 1-12. 

Fraga FB, Alencar IDCC, Tavares MT (2009) Insect pests dissemination by extruded starch 
packages. Neotropical Entomology 38: 548-549. 

Franz JM (1958) Studies on Laricobius erichsonii Rosenh (Coleoptera: Derodontidae), a pred- 
ator on Chermesids. Part 1, Distribution, life-history and ecology. Entomophaga 3 (2): 
109-164. 

Freude H, Harde KW, Lohse GA (1964) Die Kafer Mitteleuropas, Band 4: Staphylinidae I 
(Micropeplinae bis Tachyporinae). Goecke & Evers, Germany, Krefeld. 264 pp. 

Freude H, Harde KW, Lohse GA (1967) Die Kafer Mitteleuropas, Band 7: Clavicornia. Goecke 
& Evers, Germany, Krefeld. 310 pp. 

Freude H, Harde KW, Lohse GA (1969) Die Kafer Mitteleuropas, Band 8: Teredilia, Heter- 
omera, Lamellicornia. Goecke & Evers, Germany, Krefeld. 388 pp. 

Freude H, Harde KW, Lohse GA (1971) Die Kafer Mitteleuropas, Band 3: Adephaga 2, Palpi- 
cornia, Histeroidea, Staphylinoidea 1. Goecke & Evers, Germany, Krefeld. 365 pp. 

Freude H, Harde KW, Lohse GA (1974) Die Kafer Mitteleuropas, Band 5: Staphylinidae 2 (Hy- 
pocyphtinae und Aleocharinae), Pselaphidae. Goecke & Evers, Germany, Krefeld. 381 pp. 

Freude H, Harde KW, Lohse GA (1979) Die Kafer Mitteleuropas, Band 6: Diversicornia. 
Goecke & Evers, Germany, Krefeld. 367 pp. 

Freude H, Harde KW, Lohse GA (1989) Die Kafer Mitteleuropas: 1. Supplementband mit 
Katalogteil. Bd. 12, 346 S. Goecke & Evers, Germany, Krefeld. 245 pp. 

Gamarra P, Outerelo R (2005) Catalogo Iberobalear de los Aleocharinae (Coleoptera: Staphy- 
linidae). Boletin Sociedad Entomologica Aragonesa 37: 1-81. 

Gamarra P, Outerelo R (2009) Catalogo Iberobalear de los Micropeplinae y Proteininae (Co- 
leoptera: Staphylinidae). Boletin Sociedad Entomologica Aragonesa 45: 207-211. 

Genovesi P (2005) Eradications of invasive alien species in Europe: a review. Biological Invasions 
7: 127-133. 

Genovesi P, Shine C (2003) European Strategy on Invasive Alien Species. Council of Europe, 
Strasbourg, t-pvs (2003) 7 rev. 50 pp. 

Glavendekic M, Mihajlovic L, Petanovic R (2005) Introduction and spread of invasive mites 
and insects in Serbia and Montenegro. Plant protection and plant health in Europe: intro- 
duction and spread of invasive species, held at Humboldt University, Germany, Berlin, 9-11 
June 2005, 229-230. 

Gomy Y (2006) Contribution a l’établissement des catalogues régionaux: Histeroidea (Coleop- 
tera). V. L’Entomologiste 62 (3-4): 101-115. 

Gomy Y (2008) Contribution a létablissement des catalogues régionaux: Coleoptera Histeri- 

dae VII. L’Entomologiste 64 (6): 325-347. 


Coleoptera families other than Cerambycidae, Curculionidae... 337 

Gomy Y (2009) Contribution a l’établissement des catalogues régionaux: Coleoptera Histeri- 
dae VII. L’Entomologiste 65 (6): 313-328. 

Gravereau J, Trauman J (2001) Crises financiéres. Economica, France, Paris. 459 pp. 

Haggar JP (1988) The structure, composition and status of the cloud forests of Pico Island in 
the Azores. Biological Conservation 46: 7-22. 

Haines CP, Rees DP (1989) Guide pratique des types d’insectes et d’acariens qui s'attaquent au 
poisson traité. FAO Document technique sur les péches - 303. Organisation des Nations 
Unies pour l’alimentation et l’agriculture, Italia, Rome. 33 pp. 

Halperin J, Geis KU (1999) Lyctidae (Coleoptera) of Israel, their damage and its prevention. 
Phytoparasitica 27 (4) 257-262. 

Hava J (2003) World catalogue of the Dermestidae (Coleoptera). Studie a zpravy Okresnitho 
muzea Praha-vychod, Supplementum 1, Czech Republic, Praha. 196 pp. 

Hava J: A Catalogue of World Dermestidae. http://www.dermestidae.wz.cz. 

Heleno RH (2008) The impact of alien plants on native biota in the Azores: a food web ap- 
proach. PhD thesis. School of Biological Sciences, University of Bristol, England. 179 pp. 
Hemp C, Dettner K (2003) Description of larvae of the genus Formicomus laferte, and data 
on the life cycles of Omonadus floralis (Linne) and Notoxus monoceros (Linne) (Coleoptera: 

Anthicidae). Coleopterists Bulletin 57: 361-368. 

Hermann A, Baena M (2004) New records of Dermestidae (Coleoptera) for Spain and Europe. 
Boln. S.E.A. 34 (2004): 211-213. 

Holland JM (2002) The agroecology of carabid beetles. Intercept Limited, UK, Andover. 356 pp. 

Hulle M, Bonhomme J, Maurice D, and Simon JC (2008) Is the life cycle of high arctic aphids 
adapted to climate change? Polar Biology 31: 1037-1042. 

Hurka K (1996) Carabidae of the Czech and Slovak Republics. Kabourek, Czech Republic, 
Zlin. 565 pp. 

lablokoff-Khnzorian SM (1975) Etude sur les Erotylidae (Coleoptera) palearctiques. Acta Zoo- 
logica Cracoviensia 20 (8). 

Imperial Institute of Entomology (1930) The Review of applied entomology. Series A, Agricul- 
tural. The Imperial Institute of Entomology, England, London. 895 pp. 

International Monetary Fund: World Economic Outlook database. http://www.imf.org/exter- 
nal/pubs/ft/weo/2009/02/weodata/index.aspx [accessed October 2009]. 

Jeannel R (1942) Coléoptéres Carabiques, II. Collection Faune de France. Lechevalier, France, 
Paris. 600 pp. 

Kadej M (2005) Data on the occurence of some species of Dermestidae (Coleoptera) in Poland. 
Wiadomosci Entomologicene 24 (1): 21-31. 

Kirejtshuk AG, James DG, Heffer R (1997) Description and biology of a new species of Cybo- 
cephalus Erichson (Coleoptera: Nitidulidae), a predator of Australian citrus whitefly. Au- 
stralian Journal of Entomology 36: 81-86. 

Korge H (2005) Rote Liste und Gesamtartenliste der Kurzfliigelkafer (Coleoptera: Staphy- 
linidae) von Berlin. In Der Landerbeauftragte ftir Naturschutz und Landschaftspflege / 
Senatsverwaltung ftir Stadtentwicklung (Hrsg.): Rote Listen der gefahrdeten Pflanzen und 
Tiere von Berlin. CD-ROM. 


338 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Kreiss E-J, Schmitz A, and Schmitz H (2005) Morphology of the prothoracic discs and associ- 
ated sensilla of Acanthocnemus nigricans (Coleoptera, Acanthocnemidae). Arthropod Struc- 
ture & Development 34: 419-428. 

Kuschel G (1990) Beetles in a suburban environment: a New Zealand case study. The identity 
and status of Coleoptera in the natural and modified habitats of Lynfield, Auckland (1974— 
1989). DSIO Plant Protection Report no. 3. New Zealand Department of Scientific and 
Industrial Research, New Zealand, Auckland. 118 pp. 

Laibner S (2000) Elateridae of the Czech and Slovak Republics. Kabourek, Czech Republic, 
Zlin. 292 pp. 

Launay M (1999) Versailles, une paix baclée ? Le XXéme siécle est mal parti. Complexe, France, 
Paris. 188 pp. 

Lawson RH (1996) Economic importance and trends in ornamental horticulture. Ninth in- 
ternational symposium on virus diseases of ornamental plants, Israel, Herzliya, 17-22 March, 
1996. 226-237. 

Leppakoski E, Gollasch S, and Olenin S (2002) Invasive Aquatic Species of Europe Distribu- 
tion, Impacts and Management. Kluwer Academic, Nederland, Dordrecht. 583 pp. 

Leseigneur L (1972) Coléoptéres Elateridae de la faune de France continentale et de Corse. 
Bulletin mensuel de la Société linnéenne de Lyon, vol. 41, suppl. 1-380. 

Lesne P (1901) Synopsis des Bostrychides paléarctiques. L’Abeille 30: 73-136. 

Lesne P (1904) Supplément au Synopsis des Bostrychidae paléarctiques. L’'Abeille 30: 153-168. 

Levine JM, and D’Antonio CM (2003) Forecasting biological invasions with increasing inter- 
national trade. Conservation Biology 17: 322-326. 

Libungan L, Gislason GM, and Pérdarson T (2008) Varmasmidur — stersta bjalla 4 Islandi 
(English summary: The ground beetle Carabus nemoralis — biggest beetle in Iceland). Nat 
turufredingurinn 77: 15-18. 

Lima AMdC (1955) Insetos do Brasil. 9° Tomo, Capitulo XXIX. Coleopteros, 3a Parte. Escola 
Nacional de Agronomia, Série didatica, Brazil, Rio de Janeiro. 289 pp. 

Luff M (1998) Provisional atlas of the ground beetles (Coleoptera, Carabidae) of Britain. Bio- 
logical Records Centre, England, Huntingdon. 194 pp. 

Luff ML (2007) The Carabidae (ground beetles) of Britain and Ireland (Second Edition). 
Handbooks for the identification of British Insects. Vol 4 Part 2. Royal Entomological 
Society, England, Shrewsbury. 247 pp. 

Luka H, Nagel P, Feldmann B, Luka A, Gonseth Y (2009) Checkliste der Kurzfliigelkafer der 
Schweiz (Coleoptera: Staphylinidae ohne Pselaphinae). Mitteilungen der Schweizerischen 
Entomogischen Gesellschaft 82: 61-100. 

Lupi D (2002) Cybocephalus nipponicus Endrody-Younga (Coleoptera Cybocephalidae) on Di- 
aspis echinocacti (Bouche) in Liguria. Bollettino di Zoologia Agraria e di Bachicoltura 34: 
463-466. 

Machado A (1976) Introduction to a faunal study of the Canary Islands’ Laurisilva, with special 
reference to the ground-beetles (Coleoptera, Caraboidea). In: Kunkel G (Ed) Biogeography 
and Ecology in the Canary Islands. The Hague, Junk, Netherlands. 347-411. 


Coleoptera families other than Cerambycidae, Curculionidae... hey) 

Machado A, Oromi P (2000) Elenco de los Coleopteros de las islas Canarias. Catalogue of 
the Coleoptera of the Canary islands. Instituto de Estudios Canarios, Spain, Tenerife, La 
Laguna. 306 pp. 

Maddison DR, Schulz KS, Maddison WP (2007) The Tree of Life Web Project. Zootaxa 1668: 
19-40. 

Mann DJ (2006) Ptilodactyla exotica Chapin, 1927 (Coleoptera: Ptilodactylidae: Ptilodactyli- 
nae) established breeding under glass in Britain, with a brief discussion on the family Pti- 
lodactylidae. Entomologists Monthly Magazine 142: 67-79. 

Mannerkoski I, Ferrer J (1992) Cynaeus opacus Champion, a new tenebrionid beetle in Finland 
(Coleoptera, Tenebrionidae). Entomologica Fennica 31 (8): 95-97. 

Martinez M, Cocquempot C (1985) Sur la présence en France de Reesa vespulae, espéce indési- 
rable (Coleoptera Dermestidae). L’Entomologiste 41 (1): 21-25. 

Mazur S (1989) Note synonymique sur les genres Pactolinus MOTSCHULSKY, Pachylister 
LEWIS et Macrolister LEWIS (Col. Histeridae). Nouv. Revue Ent. (N.S.) 6 (1): 84. 

Mendonga E, and Borges PAV (2009) Distribution of the Exotic Arthropods from the Azores. 
Portugal, Univ. Azores, Angra do Heroismo. 87 pp. 

Mifsud D, Audisio P (2008) The Kateretidae and Nitidulidae of the Maltesse Archipelago (Co- 
leoptera). Bulletin of the entomological society of Malta 2008 (1): 15-37. 

Mitter H, Schuh R (2008) Catogenus rufus (F.) - neu ftir Europa! (Coleoptera: Passandridae). 
Koleopterologische Rundschau 78: 329-332. 

Moncoutier B (2001) Les “Clavicornes” de la faune de France (premiére partie). Le Coléopteriste 
43: 187-207. 

Moncoutier B (2002) Les «Clavicornes» de la faune de France (deuxiéme partie). Le Coléopte- 
riste 5: 7-33. 

Monzo C, Vanaclocha P, Outerelo R, Ruiz-Tapiador I, Tortosa D, Pina T, Castanera P, Urbaneja 
A (2005) Catalogacién de especies de las familias Carabidae, Cicindelidae y Staphylinidae 
en el suelo de los citricos de la provincia de Valencia, Espafia. Bol. San. Veg. Plagas 31: 
483-492. 

Mouchet J, Giacomini T, Julvez J (1995) Spreading of disease vectors and pests throughout the 
world by man. Cahiers d'Etudes et de Recherches Francophones/Sante 5: 293-298. 

Neculiseanu ZZ, and Matalin AV (2000) A catalogue of the ground-beetles of the Republic of 
Moldova (Insecta, Coleoptera, Carabidae). Pensoft, Bulgaria, Sofia. 164 pp. 

Newton A: Staphylinini Species Catalog Draft. http://www.staphylinini.org/staphylinini/Prod- 
ucts.html. 

Odegaard F, Tommeras B (2000) Compost heaps - refuges and stepping-stones for alien arthro- 
pod species in northern Europe. Diversity and Distributions 6: 45-59. 

Ortuno V, Toribio M (2005) Carabidae de la Peninsula Iberica y Baleares. Vol. 1. Trechinae, 
Bembidiini. Argania, Spain, Barcelona. 455 pp. 

Paulian R (1988) Biologie des Coléoptéres. Lechevalier, France, Paris. 719 pp. 

Paulian R, Baraud J (1982) Lucanoidea et Scarabaeoidea. Lechevalier, France, Paris. 477 pp. 

Peck SB (2009) The beetles of Barbados, West Indies (Insecta: Coleoptera): diversity, distribu- 
tion and faunal structure. Jnsecta Mundi 73: 1-51. 


340 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Perrault GG (1981) Le genre Lesitus (Foehlig) (Col. Carabidae) HI - le sous-genre Nebrileistus 
(Banninger). Bulletin de la Société linnéenne de Lyon 50 (7): 22-226. 

Perrault GG (1984) New record of Notobia (Anisotarsus) cupripennis (Germar) (Coleoptera: 
Carabidae: Harpalini). 7he Coleopterists Bulletin 38 (4): 334. 

Perrings C, Dehnen-Schmutz K, Touza J, Williamson M (2005) How to manage biological 
invasions under globalization. Trends in Ecology & Evolution 20: 212-215. 

Pimentel D, Zuniga R, Morrison D (2005) Update on the environmental and economic costs 
associated with alien-invasive species in the United States. Ecological Economics 52: 273- 
288. 

Poland TM, McCullough DG (2006) Emerald ash borer: Invasion of the urban forest and the 
threat to North America’s ash resource. Journal of Forestry 104: 118-124. 

Pollock DA, Ivie MA (1996) Anthicidae (Coleoptera) of the Virgin Islands. Florida Entomolo- 
gist 79: 230-240. 

Ratti E (2007) Silvanid beetles in Italy (Coleoptera Cucujoidea Silvanidae). Bollettino del Museo 
Civico di Storia Naturale di Venezia 58: 83-137. 

Ratti E: Coleotteri alieni in Italia. http://www.msn.ve.it [accessed 25.05.2007]. 

Reemer M (2003) Invasieve Arthropoda in Nederland: een eerste inventarisatie. European In- 
vertebrate Survey, Nederland, Leiden. 63 pp. 

Reibnitz J, Schawaller W (2006) Cynaeus angustus (Leconte, 1851) (Coleoptera: Tenebrioni- 
dae), eine neue Adventivart in Mitteleuropa ? Mitt. ent. V. Stuttgart 41: 153-154. 

Rogé J (2003) Prolifération de Lispinus impressicollis (Motschulsky, 1857) et de Symbiotes gib- 
berosus (Lucas, 1846), hétes dun biotope particulier, dans le département de l’Aude (11) 
(Coleoptera, Staphylinidae et Endomychidae). L’Entomologiste 59 (3): 65-66. 

Riicker WH (1995) Cartodere australica (Belon) und Cartodere norvegica (Strand) (Coleoptera: 
Latridiidae). Mitt. internat. entomol. Ver. 20: 59-61. 

Ruiz GM, and Carlton JT (2003) Invasive species: vectors and management strategies. /n In- 
vasive species: vectors and management strategies. Island Press, USA, Washington. 518 pp. 

Ruta R, Konwerski S, Fkrolik R, Lason A, Milkowski M (2006) New records of dermestid 
beetles (Coleoptera: Dermestidae) in Poland. Part 2. Megatominae. Wiadomosci Entomo- 
logiczne 25 (1): 21-28. 

Ruta R, Konwerski S, Kadej M, Herrmann A, Lason A (2004) Three species of dermestid 
beetles (Coleoptera: Dermestidae) new to the Polish fauna with remarks on dermestids 
introduced to Poland. Polskie Pismo Entomologiczne 73: 307-314. 

Ryndevich SK (2004) Review of species of the genus Cercyon Leach, 1817 of Russia and adja- 
cent regions. I. Subgenus Cercyon (s.str.) Leach, 1817. Cercyon lateralis-group (Coleoptera: 
Hydrophilidae). Annales Universitatis Mariae Curie-Sklodowska 59: 1-13. 

Santamaria JM, Gayoso A, Otero JC (1996) Los Laemophloeidae Ganglbauer, 1899 (Cole- 
optera) iberobaleares. Lista de especies y datos corologicos. Boln. Asoc. esp. Ent. 20 (3-4): 
107-114, 

Schaefer L (1949) Les Buprestides de France: tableaux analytiques des Coléoptéres de la faune 
franco-rhénane, France, Rhénanie, Belgique, Hollande, Valais, Corse. Famille LVI. E. Le 
Moult, France, Paris . 511 pp. 


Coleoptera families other than Cerambycidae, Curculionidae... 341 

Scheerpeltz O (1972) Studien an den palaarktischen Arten der Gattung Myrmecopora Saulcy 
(Col. Staphylinidae). Koleopterologische Rundschau 50: 93-109. 

Schmitz H, Schmitz A, Trenner S, Bleckmann H (2002) A new type of insect infrared organ of 
low thermal mass. Naturwissenschaften 89: 226-229. 

Schuh R, Mifsud D (2000) The cylindrical bark beetles of Malta (Insecta: Coleoptera: Zopheri- 
dae, Colydiinae). Ann. Naturhist. Mus. Wien 102B: 259-267. 

Schulte GG, Elnitsky MA, Benoit JB, Denlinger DL, Lee RE (2008) Extremely large aggrega- 
tions of collembolan eggs on Humble Island, Antarctica: a response to early seasonal warm- 
ing? Polar Biology 31: 889-892. 

Sefrova H (2005) Introduced and invasive insect species in the Czech Republic and their eco- 
nomic and ecological impact (Insecta). Acta Univ. Agric. Silvic. Mendel. Brun. 53 (17): 
151-158. 

Sefrova H, Lastuvka Z (2005) Catologue of alien animal species in the Czech Republic. Acta 
Univ. Agric. Silvic. Mendel. Brun. 53 (18): 151-170. 

Serrano J, Lencina JL, Andujar A (2003) Distribution patterns of Iberian Carabidae (Insecta, 
Coleoptera). Graellsia 59 (2-3): 129-153. 

Sharp D (1882-1887) Biologia centrali-americana. Insecta. Coleoptera. Vol. I. Part 2. Pub- 
lished for the editors by R.H. Porter, England, London. 824 pp. 

Shockley FW (2009) Endomychidae. Handsome fungus beetles. http://tolweb.org/Endomy- 
chidae/9169/2009.03.31 [accessed 31.03.2009] In: The Tree of Life Web Project, http:// 
tolweb.org/. 

Shockley FW, Tomaszewska KW, McHugh JV (2009a) An annotated checklist of the handsome 
fungus beetles of the world (Coleoptera: Cucujoidea: Endomychidae). Zootaxa 1-113. 
Shockley FW, Tomaszewska KW, and McHugh JV (2009b) Review of the natural history of 
the handsome fungus beetles (Coleoptera: Cucujoidea: Endomychidae). Insecta Mundi 72: 

1-74 

Simberloff D (2001) Biological invasions - how are they affecting us, and what can we do about 
them? Western North American Naturalist 61: 308-315. 

Smith RM, Baker RHA, Malumphy CP, Hockland S$, Hammon RP, Ostoja-Starzewski JC, Col- 
lins DW (2007) Recent non-native invertebrate plant pest establishments in Great Britain: 
origins, pathways, and trends. Agricultural and Forest Entomology 9: 307-326. 

Soldati F (2007) Fauna of France and Corsica, Coleoptera Tenebrionidae (Alleculinae ex- 
cluded). Systematic Catalogue and Atlas. Mémoires de la Société Linnéenne de Bordeaux 6: 
1-186. 

Sérensson M, Johnson C (2004) The first European records of the pantropical genus Bambara 
Vuillet, and a review of the immigrant featherwing beetles in Europe (Coleoptera: Ptilii- 
dae). Koleopterologische Rundschau 74: 287-302. 

Stoch F: Checklist of the species of the italian fauna. Version 2.0. http://www.checklist.faunai- 
talia.it [accessed 03.12.2003]. 

Streito JC, Martinez M (2008) Entomological actualities: new introduced pests insects (period 
January 2000 at June 2005). Zeme Conference Internationale sur les Ravageurs en Agriculture, 
France, Montpellier, 26-27 octobre, 2008, unpaginated. 


342 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

Tamisier J-P (2004) Clambus simsoni Blackburn, un petit coléoptére australien en pleine expan- 
sion en France (Coleoptera, Clambidae). Bull. Société linnéenne de Bordeaux 32 (1): 41-45. 

Telnov D (1996) Sixty three new and rare species of Coleoptera in the fauna of Latvia. Latvijas 
entomologs 35: 36-43. 

The World Factbook: Central Intelligence Agency, 2009. https://www.cia.gov/library/publica- 
tions/the-world-factbook/index.html. 

Théry A (1942) Coléoptéres Buprestides. Lechevalier, France, Paris. 221 pp. 

Thomas MC (1995) Invertebrate Pets and the Florida Department of Agriculture and Con- 
sumer Services. Florida Entomologist 78: 39-44. 

Tomov R (2009) Non-indigenous insects and their threat to biodiversity and economy in Alba- 
nia, Bulgaria and Republic of Macedonia. Pensoft, Bulgaria, Sofia. 112 pp. 

Trautner J, Geigenmuller K (1987) Tiger beetles, ground beetles. Illustrated key to the Cicin- 
delidae and Carabidae of Europe. TRIOPS Verlag, Margraf Distributor. Germany, Gai- 
mersheim. 488 pp. 

Tronquet M (2006) Catalogue iconographique des coléoptéres des Pyrénées-Orientales. Asso- 
ciation roussillonnaise d’entomologie, France, Perpignan. 78 pp. 

Turin H, Penev L, Casale A (2003) The Genus Cavabus in Europe. A synthesis. Pensoft, Sofia, 
Bulgaria. 511 pp. 

Valemberg J (1997) Catalogue descriptif, biologique et synonymique de la faune Paléarctique 
des coléoptéres Carabidae Latreille, 1806. Tome 1 Corpus. Mémoire de la Société Entomo- 
logique du Nord de la France, France, Villeneuve d’Ascq. 659 pp. 

Vernon P, Caron F, Davies L (1999) Annual activity of two endemic beetles (Carabidae) at the 
edge between fell-field and moorland on a sub-Antarctic island. European Journal of Soil 
Biology 35: 39-43. 

Vieira V, Borges P, Karsholt O, Wunderlich J (2003) The Arthropoda fauna of Corvo island 
(Azores): new records and updated list of species. Vieraea 31: 145-156. 

Vincent R (1999) Une nouvelle fois un Aridius (Coléoptére Latridien) va-t-il envahir l’Europe ? 
(Cartodere (Aridius) delamarei Dajoz, 1962). L’Entomologiste 55 (2): 53-55. 

Vorst O (2009) Cercyon castaneipennis sp. n., an overlooked species from Europe (Coleoptera: 
Hydrophilidae). Zootaxa 2054: 59-68. 

Vorst O, Van Ee G, Huijbregts H, Van Nieuwenhuijzen A (2007) On some smaller Latvian 
Coleoptera. Latvijas entomologs 44: 15-25. 

Weidner H, Rack G, Friedrichsen D (1984) Tables de détermination des principaux ravageurs 
des denrées entreposées dans les pays chauds. GTZ, Germany, schborn. 157 pp. 

Wittenberg R, Kenis M, Blick T, Hanggi A, Gassmann A, Weber E (2006) Invasive alien species 
in Switzerland: an inventory of alien species and their threat to biodiversity and economy 
in Switzerland. Federal Office for the Environment (FOEN), Switzerland, Bern. 155 pp. 

Yélamos T (1992) Data on Afrotropical Histendae (Coleoptera), with description of Hister 
sindarae n. sp. Misc. Zool. Barcelona 16: 37-43. 

Zahradnik P, Mifsud D (2005) Ozognathus cornutus (LeConte) - new record for the Palaearctic 
Region (Coleoptera: Anobiidae). Studies and reports of District Museum Prague-East, Taxo- 
nomical Series \(1—2): 141-143. 


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x (SOOZ) ‘JB 29 SstoryI) Sa ‘ha OISSLT AVS snoseyd adoyy) suvor1us1u 
S ‘(€QOZ) ‘Te 19 edezere7-osuoTy poom ‘Joaquin ‘LI “Ad YOO’ “WA ‘SAD eisepeqisny -odyd SNULAUIOGIUDIY 
S seplMaus0yULOy 
adomy ur; oS3urs satoads 
S9DUIIIFOY Wwyqey SOLUNOD poprauy proces 3s—) sane snyei¢ Ayrore,y 

“(II xtpueddy 20s) SINNA 
O} Jajor suoTIeIAdIgge JegeEp ‘(| xipuoeddy as) 9OTE OS] 0} Jofor suOTIeTADIQge sapoo ATUNO’) ‘oTUaSOIdATD FZ sdomng of udTTy W ssnIeIS ‘sepTaUIDN07) pur 077 

nsuas seprpawWosc1y’y ‘o7y] Nsuas sepruoTNoIN’ ‘seproAquiesay) URY}I J9YI0 saTjTuey Jo adorn o7 uate sotdads vsaidoajor ay} JO soMsTIOIOeIeY pure AsTT *4°S*S VIGUL 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

344 

(ZZ81 ‘12912 y]) 

(O€61) snoseyd SNIVAOULADUL 
AZojoworwy Jo sn msuy [ewodwy soo) WA ‘6661 -ovdyd SnAnNaUytg] 
sasnoyarem 
pue syjru paay 
jo asad sourur 
Jenuaiod ‘sanprsai 
Jo Jueiqeyur pue oyerjoduray, SNOIOA (ZZ81 JOHPY) 
ya3uaavos ATI] -PISY -hinep 1Lay]Ig SNISOLNAMNAST 
(S007) (6S81 
pnsjiy pue yrupesrye7Z ‘(CQ0Z) POLIOUy SNOIOA ‘YUODIT) SHINULOI 
‘Te 19 Opsoiog “(8007) PUeWOTTY Peo PEEP Sd ‘OU “LW SH ‘S007 YON “Hop SNYIDUSOZC) 
WN ‘HO 
‘NVO-Sd “TVd-Sa “Sa ‘IS 
(S007) ‘OU “AVW-Ld ‘OZV-Ld 
YAMS] PUL PAOLYIS ‘(60(0Z) syooq ‘Ld “Id ‘LW ‘“OIS<LI “UVS (O6LT 
sasiog pur eSuopuayy (6961) pyo ‘omarurny -LI ‘LI UD “AG ‘WOD-WA snoseyd ‘JOIAIQ) Wénauvjsva 
‘ye 19 apnary “(Z66T) Jouedsy poom os “Ud “ZO ‘AD “YH ‘Wd CLV] Ld ‘Z081 -o1fyd UNIQOIIN$ 
OPS ‘Neuyeise) 
ap a1iodeT] 
(S007) POLOWYy SnNOIOA UNUVDILAIULD 
‘Te 10 saslog (C861) SoTPPg sjonpoid poesois OZV-Ld ‘LW ‘UVS-LI {LI | esouryug, YON -TWnep UNIZIWY 

(6961) 

Te 19 epnoany ‘(S861) SPP 
(9002) ‘Te 9 St9quantp\ 
‘(SOOT) BYANIseyT pue PAOLJOS 
“(SOOZ) ‘Je 19 DFyepusarysy 
‘(6961) ‘Te 39 epnany ‘(Z661) 
Jouedsy ‘(¢Q0Z) ‘Je 19 sosi0g 

SIDUDIIFIY 

s1soU 
pig ‘sionpoid 

perois ‘sTyTu 

sionpoid 
perois ‘oooeqol 

venqeH 

dS ‘NVO-Sd ‘Sd “GVW 
(Ld “OZV-Ld “AC “MC “LV 

HO ‘Su ‘Ld “LI ‘AT 
‘DISSE ALY SaLP LISAH 
“ad “Md “ZO “Dd “LV “TV 

S31JJUNOD poprAauy 

umouyuy) 

Ld ‘8781 

adomy ut 
proses IST 

o1uas 

-o1dA17 

jeordonqns 
‘jeotdosy, 

SNOIOA 
-lnap 

snoseyd 
-ovdyd 

snjejg 

9S81 NeIppetog 
auiffy wnizayy 

COLT ‘snolqey 
IULOIIAIS DULLAPOISYT. 

satoads 
Ayrurey 


345 

Coleoptera families other than Cerambycidae, Curculionidae... 

(6007) AOWOT, (S007) 

€D “WN “HO “AS ‘NVO 

“Sd ‘SH ‘IS “MS ‘SY “OU 
‘GVW-Ld “OZV-Ld ‘Ld “Td 
‘ON “IN ‘CW LW ‘NT 
ETD AL DISS LIeadys 
SLI ‘LI “AI “NH “AWORUD 
UD “AG UOO-Ud “UA 

eYyANIseT pure PAOTJIS “(69G[) ‘Te oidosyjueuds Td “Ad “Md ‘ZO ‘AD “WH otuas SNOIOA CLLI ‘snoqey 
19 apnasy “(G0OZ) ‘Te ® sa810g ‘poo po {| ‘Sa ‘vd “ad ‘AG ‘LV TV) ZO ‘OS8T) = -o1dA1Dp AIP} OD OlWY] SHUNT 

qd ‘vn 

‘HO “AS ‘NVO-Sa “TVd-SA 

‘Sa ‘IS “MS ‘Su ‘NU ‘OU 

‘GVW-Ld ‘OZV-Ld ‘Ld “Td 

‘ON “IN ‘CW ‘LW ‘AT 

‘ET ‘TI ‘AT ‘OISsLI UVS-LI 

(6007) ‘LI “Al ‘SI ‘NH “UD “Ad 

AOUIOT, “(600Z) SadIOg pue WOO-Ud Ua ‘Td ‘Od “Aa 
eSuopuayy “(8007Z) HX “(S007) speqasoa “Md “ZO ‘AO ‘UH “Dd orua8 SNOJOA (8¢Z] snoeuurT) 
‘ye 19 sod1og ‘([ BG) Uossuag poup ‘aisem| Of Tf} “Va “AG ‘AD ‘LV ‘AV “IV| Da ‘OP6T| = -ord Ag “ainep| OD Ang snungy 

(0002) 
TUIOIC) pue opeysey (6961) 
‘Te 39 apnoxy “(8007) HA 

(6961) ‘Te apnary 

SIDUDIIFIY 

inj 
‘sjonpoid posois 

sjonpoid posois 

veNqeH 

€D ‘NVO-SH LW ‘AT “Ad 
VN ‘HO “dS ‘Sd ‘IS “MS “SU 
‘NY ‘OU “Td “ON “IN ‘AT 
UVS-LI CLI ‘NH “Ad “YOO 
Ud “Ud ‘Td “Ad “Md °ZO 
UH “Od ‘Va “Ad ‘Ad “LV 

S31JJUNOD poprAauy 

uMOUxU al 

adoimy ut 
prosel IST 

oSuvri 
DATION 

snoJOA 
-1nap 

SNOJOA 
-1nap 

snjejsg 

T6LI Fezueg 
sadiawa snulgyg 

LER] WMS 
SNIIUIIIG SNUITT 

satoads 
Ayrurey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

346 

(6002) Aowey *(600Z) s°810g 
pue esuopuesypy ‘(000Z) WoIC 
pure opeypoeyy ‘(€Q07) JounIq 

pue durazy ‘(696[) ‘Je 19 epnary 

(Z00Z) 12Z9g pure ruuRAOIs9q 

(9661) STAT pure YooTog 

(S007 

pur eAoljog “elye 

) eyANise'T] 
WJ UT TuaTye 

1191109707) "19ey ‘(8007) HG 

(S007 

) eyANise'T 

pur PAaoyas (6961) [Pe 19 spnory 

(90072) ‘Te 39 
SiaquautA\ ‘(800Z) HN ‘(1861) 

uosisuag ‘(gQ0Z) PuewAaTTY 

SIDUIIIFJIY 

jorepoid 

ynpe ‘oseydoodur 
‘aseydninop 
‘Kedap [e1a3IA 

JaSUDAPOS BAIE] 
‘snoprydodoryiue 

sjonpoid 

perois “uiniteqioy 
‘s19asuT ‘slonposd 
jewrue porp 
sppy saneu 

ul sn3arieI9 
‘sionpoid 

perois “‘spaas 

sjonposd posois 

veNqeH 

HO “dS ‘NVO-SA 
“TVd-Sd ‘Sd “MS ‘NU ‘OU 
‘AVW-Ld ‘OZV-Ld ‘Ld 
“Td “ON “IN ‘LW ‘IT TI 
‘AT ‘OIS*LI “UVScLI CLI “AI 
‘NH “DAS-UD ‘NOTUD 
“AMOUD “UD “Ad “UOD 
Ud ‘Ud ‘Td “Ad “Md °ZO 
‘AO ‘UH “Od “Va CLV “TV 

CQVWsLd 
“LW SOAS-UD “UD ‘AO 

qd ZD 

LI “ad “Wd “Md °Z9 “UH 
dD 

‘VN “HO “dS ‘Sd ‘TS “MS 
‘Su ‘NY “OZV-Ld ‘Ld “Td 
‘ON “IN ‘CW ‘NT LT ‘AT 
‘LI “AI “OH “UD “AC “Ud 
‘Td ‘Qd “Ad “MC “ZO ‘AD 
UH ‘Od ‘Va “Ad ‘Ad “LV 

S31JJUNOD poprAuy 

LI ‘7861 

uMouUxyU al 

ZO “661 

ZO “S96T 

Ad ‘9161 

adomy ut 
proser 1ST 

jeordosy, 
-vIsSy 

POLIOWIY 

SRO 

POLIOWY 

SRO 

vIseyesIsny 

SNOJOA 
-lnap 

umouyun 

uMou yun 

SnOIOA 
-lnap 

snoseyd 
-odyd 

snoJOA 
-lnap 

snjejg 

(SCZT snavuury) 
sypLopf snpouougd 
TIGL 

Tq wheys1ogousaZ9 
sn1gqupy 

6rsl 
*919199U9S-91I9F ey 
SNILUIAD SNILYIU 

oeppryuy 

681 
‘IOIOS supngops 

SNIUABOUOSIAT 

(OS81 
‘aypsougy-ulgns)) 
190QUI SNUMLOIIAT 

IS8T natpjarog 

SNJIA SNULLT 

sotoads 
Ayrurey 


347 

Coleoptera families other than Cerambycidae, Curculionidae... 

sionpoid 
porois pur sdrys 
PABSSED PITIP ‘(N) 

JoJ0q ooquieq 

(LOGI) 2USzT ‘(696T) 
‘Te 19 apnary “(8007) Hd 

( susweUIO 
pur sjmMoq 
UsPOOM padre 
oruyia) Joquiny 

(‘elyeay UT TuoTye 1191102707) 
“IeY “(696 1) ‘Te 9 opnoasy 
eIOeOy 

(1061) 

auszT ‘(696[) ‘Je 39 epnely 

‘soo WNAF Tar0q 
wiois snoseyddjod 

(8007) H"d “(Z661) pnesreg 

poom 8umi103 

(8007) HG 

dD 
‘HO “AS ‘Sa “MS “(VW-Ld 
“IN “AC “Id “YH “Ad CLV 

as ‘Sd “LI “ad “3d 

Sd ‘OIS 
CLI UVSsLI “UOO-Us “WA 

umouyuy) 

umouxuy) 

uMouyuy) 

aD ‘961 

dD ‘7861 

jeordonqns 
‘eotdosy, 

vOLyy 

jeordonqns 
‘qestdosy, 

POLIOUy 

YON 

POLY 

snoseyd 
-ovdyd 

snoseyd 
-ovdyd 

snoseyd 
-o1dyd 

SNOIOA 
-lnap 
SNOIOA 
-lnap 

(SSSI “UoIseT[op\) 
sngpjoanofig 
snsapould, 

(0621 

JOIAT[Q) S#4nUsOI 
say doyauqsog 

CLLI ‘snoqey 
sngsvuou apdy 

aeprypinsog 
(ZG8T ‘au0oeT) 
snqvjava sntAvsay 

(LO6T ‘Aensursag) 
SISUAVIVU SAISOLUDS 

S9f199q 
uUoAspUuapoutl ‘S pue 

(T8G6T) pneseg pur uelyneg| szs10q jo ssus0q SNOJOA T68I “UINGyorg 
‘(800Z) HC ‘(Z661) pneseg| ut ‘poom Suni03 dD ‘1761 | elsepersny -Inep xDpuaul saqisoLdvy 
SnOJOA ZG8I “uRWayog 
(S861) pneseg sunp a OZViLd| wou, Igy) BEEP] OV sypomed smipoydy 
oepripoydy 

dD “HD “as 

‘NVO-Sd ‘SH “AS “GVW 

(9007) ‘Te 39 Bx9quozt <Ld ‘Ld “IN ‘CW CLI ‘IT 

(9661) A0URL “(0007) AT OISS EL VSL 11 
TUIOIC) pue opeyoeyy ‘(696 T) sonssT} ‘AH “Ad UMOoOrWs WA SnoJOA (6Z8[ [Nassvyyy aq) 
‘Te 9 opnoaly ‘(8007) HNC [e21992A U9I10I “Ad Md ‘ZO “Ad AG ‘LV! LI ‘vVyel “Tap S01Q04 SNULOIIJIIAIS 
adomy ut satoods 
sadUaIafaY yevIqey] SdIIJUNOD paperauy Prosar IST snqei¢ Ayrore,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

348 

(0002) FuICIO 
pur opewpey] “(986T) oqo) 

uMoOuUxU al 

POLIOUy 

YON 

snoseyd 
-ovdyd 

CLLI ‘sniotiqey 

DLOIID susasdng 

(8007) ‘Te 39 AoyTyouRIeg SNUIXDAT 

SPOOS JOISO “xITeS 

(1061) 2usaT ‘snoseydoycx 
poom 
uOTIONIISUOD 

povodut ‘(N) 
(LOGI) 2usa’T | J910q poom yIIDIy 

dd 

€00C 

Ad “S007 

uMouyxU ial 

vISy 

ROLY 

snoseyd 
-ovdyd 

snoseyd 
-o1dyd 

snoseyd 
-odyd 

QBgl reuse; 

stuuadiuryd snjisp 
sepnsordng 
(S68 I ‘useT) 
stuuadypuvg 
SNGHAISOG OAMALT 

T¢sl 
‘YyOsIey] asuayuaauas 

uojhxouly 

(900Z) ‘Je 29 S1oquai 
‘(600Z) AOWOT, ‘(S00Z) PXANISE'T 
pure eAaoszas ‘(600Z) sesI0g 
pue eSuopuayy ‘(0007Z) FOIE 
pure opeypeyy ‘(F0G6T) Use] 
‘(LOGT) 2US2T “(6961) ‘Te 39 
apnery (8007) BNC “(986T) 
80qo9 “(S00Z) ‘Te se810g 
(S007) 

eyANse'T] pur PAO (F061) 
2USIT‘(TOGTL) 2US2T ‘(696T) 

‘Te 19 opnary “(8007) Hd 

speosoo ATUTeUT 
‘sjonpoid posois 

(omut) sionpord 
Pposols “(eAvssec)) 
oojurl ‘ooquieq 

SIDUIIIFJIY 

reUqeH 

€D “HO “AS ‘NVO-Sd “TVd 
“Sd “SH “MS ‘OU “OZV-Ld 
‘Ld “Td “IN (LW ‘AT ‘OIS 
SLI UVS-LI “LI “AI “Das 
UD UD “Ad “UOO-Us 
Ud ‘Td “Ad “MC “ZO ‘AO 
YH “Od “Ad ‘Ad CLV “IV 
€D “AS “MS “Td “IN 
ISS AVSSL LT WS 
‘Ad “Ud “Md *ZO “Ad “TV 

SoTsJUNODS poperauy 

ZO ‘0061 

ZO “S96T 

adoimyq ut 
proses 3ST 

jeordory, 
-vISy 

jeordonqns 
‘estdosy, 

snoseyd 
-ovdyd 

snoseyd 
-ovdyd 

snjejg 

(Z6ZT ‘smiorsqey) 
VII Uu 7 uop 

vepsadozhgy 

(CLLI ‘sntotiqe.q) 

SNINUIUL SNAIPOUICT 

satoads 
Ajrure.y 


349 

Coleoptera families other than Cerambycidae, Curculionidae... 

ssurddosp 
ureiae ‘sjonpoid (TEST ‘uealoq) 
(Z661)| pezoas ur sapaoq ioyepoid snqpiospfiun 
SIaquizeA ) ‘(ZH6[) JouULal jo soyepaid uMouyuy) woUpy| /orusesed SNYILAJOULOY 
snojrydosdd 
(L661) ‘sJUSWUUOITAUS 
SIOqUIaTeA “(CQOT) PYANISe’] SNOTIeA UT oyeioduray, joyepoid (ZEST ‘SplgUsY1) 
pure eaoryas) *(966T) EN Joleperg Z) ‘O861 -eIsy| — /oyasered snidsv9 SnYI14S04AIq 
: saseJOls 
utster ‘synuevad 
(L661) SIoquizyeA “(Z861)| YIM poysodsuen vououry|  so0repord (CZLT ‘sniorqeq) 
saypnuruasiary pue Jouinesy ‘syiod ut TN ‘LI ‘AH “AC “WA “NMC | IN ‘0007 SRD orsered suayod SNUOLYIONT 
(p7eT Teun) 
(F861) yNeUEg snylursieuy POLIOUY snoseyd stuuadisdng 
‘(000Z) MHOI_ puke opeyoeyy Jo spaas uMouxuy) SRD -o1dyd DIQOLIONT 

(L861) 4[neeoc, “(O00 TOF) 
pure opeypepy ‘(9Z61) Opeypreypy 

uMouyu) 

POLY 

joyepoid 
jorisered 

FOBT “UOISETTON, 

snspaigqnu snqstaT 

(F861) yNesAg 

(1861) eA ‘(600Z) s°310g 
pue esuopusyy ‘(000Z) FWoIC 
pue opeypeyy ‘(9/61) pepe 
‘(Z00Z) BT (8661) HT 
‘(Zy6T) ouUrel (8007) HPA 
‘(S00Z) ‘Te 19 se8I0g “(9007) 
Ipuly ‘(QO0OZ) ‘Je 19 UOsIopuy 

(‘eIpeq] UT TuaTye TaO0sTO_D “MIeY 

SIDUDIIFIY 

SOABD STeT[99 

‘syiod ur [e101] 

veNqeH 

€D ‘NVO-SH 
‘OZV-Ld “AI UOO-Ud “UWA 

S311JUNOD poprAuy 

umouyuy) 

SH “9861 

adoimy ut 
proses IST 

POLY 

POLY 

jorepoid 
orisered 

snoseyd 
-ovdyd 

snjyejg 

(8781 ‘uealaq) 
snypurj duos 

SNUIISOULVT 
sepiquie’y 

(Z8ZI 

‘snioliqe..) vyvs.op 
StLG10G OKAY’) 

satoads 
Ayrurey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

350 

(FOOT) JorsTwRT, (8007) Hd 

snoseyd 

-oodur Sisoduroo 

‘PooMoly “ISIIOJ 

€D “AS “IN “Ad “Ud “LV 

AS ‘2861 

eIsepesIsny 

SNOIOA 
-lnap 

TOG] UANgypr[g 
TUOSUIS SHU) 

VN ‘Sa “MS OU “Td 
‘SIS WVS-LE LT AH 

oePIqUiey) 

(€CP8T Inojnq) 

WD WOO’ WA Na SNOJOA sap1ogwlgsoq 

(6007) AotuOT,| ~—_-FBuNY WO spo2q I] ‘ZO UH ‘Dd ‘VE ‘AP SLY] eMowyuQ, eisy| = -ainep| oy sng dnssoply 

aepHy 

EVs 

(euNey Ue oy} jeordonqns SnOJOA ‘aqny /uopuvjuou 

jo satoods 94] Jo AsIPpy :yIo1g uapie3 yeorurj0q LIX LI ‘YA | Umousuy, ‘yeotdosy, “lap Vv SNULAIYIO] 1G I 
(siod 

(TOOT) oHNOSUCY “(900Z) “oSeUUEP MJ) €D ‘HO “Td ‘LI SNOIOA (ZI81 3P9d) 

‘Te Jo SIOquaNI A ‘(8007) Hd sionposd posois If] ‘AH “Ad Wd Nd ‘LV “TV | UMouyuy vISy -Tnep V SIVA SNIPIULANT 

(900Z) ‘Te 39 SIOqUONIT\ 
‘(ZO61) S1quizfeA “(Z861) 
Jaynuruasiayy pur Jounesy 
‘(6007Z) AoW] “(€00Z) ‘Te 39 
OURIIIG “(QOOZ) UNTEIe pur 
nuvast[NdaN ‘(600Z) sedsJog pue 
eSuopusy ‘(000Z) FWOIC pure 
opeypeW “(Z007) BY *(9661) 
eHNH “(8007) BNC “(7961) 
uoisurpec “(S00Z) ‘Te 3 sadrog 

SIDUVIIJOY 

sinuvad ut osye 
‘suopies3 ‘orepoid 
“soduroo 

veNqeH 

€D “WN “HO 

“AS “NVO-Sd ‘Sd ‘IS “MS 
‘Su “AVW-Ld ‘OZV-Ld “Td 
‘ON “IN ‘CW ‘NT TI AT 
“LI ‘NH “AC “Ud ‘Td “AG 
‘ZO “WH “Od “VE “Ad CLV 

S91JJUNOD popeAuy 

dd ‘C06! 

adomy ut 
proses IsT 

jeordosy, 
-vIsy 

jorepoid 
orsered 

snjyejg 

aepruoyAsary 

(TEgT ‘uvalaq) 

sa I1dB1U SNIIYIALT 

satoads 
Ajrore.y 


2 

Coleoptera families other than Cerambycidae, Curculionidae... 

(‘elyeay UT TuoTye 1191109707) 
MEY “(000T) WoC pue 
opeysey “(800Z) HNC “(6661) 
peaisamog ‘(CQ0Z) ‘Je 19 sasiog 

(SNLIIOVM py 

‘DULAIPOISYT) 

(S007) dd1I ‘oD9eqo} 
eYyANIseT pur PAoras ‘(6/6]) ‘Te 
19 apnoly ‘(Q00Z) youareyD nq 
(9002) ‘Te 39 S1aquent “(S00Z) 
eyANIseT pure PAOTas “(6/6[) ‘Te 
9 apnosy ‘(000Z) youareyD) ng 
(8002) 

BPC (0007) eusveq ong 

UO S}DISUT 
uo Jorepaid 
eepriqouy: 
‘seprypoATIsog 
uo Joepaid 
aepInoAT] 

uo Jo1epaid 

€D “HO ‘NVO-Sd 
‘Sd ‘AVW-Ld “‘OZV-Ld 
“LI ‘WOO ‘Ud “UH 

Td ‘LI “Ad “ZO 

HoeZo: LV 

aD “Ud 

umouxuy) 

ZO “€961 

ZO ‘0661 

qD-EC6l 

vIseyesisny 

vIseyesisny 

SNOIOA 
-lnap 

ioyepoid 
orisered 

ioyepoid 
/orisered 
joyepoid 
orisered 

CsI dieys 

sywnbav snsadoy1scE 
aeprydoyAsory 

(SE8I 

‘1aqajoT) “anbnq 

SNAPJIOAIUDY |, 

(76LI 

‘ISSOY) SHqvqLatUuN 

SNUIISOSADT, 

(OF8T ‘UPUIMOQNT) 
SNADI SNIJUDAVT 

(6261) ‘Te 

jo apnery ‘(0Q0Z) JoUAIeYD Nq| sesnoy soquin pyo 

Sa “Ad “Ud “Ad “LV 

uMoOuUxU al 

POLY 

joyepoid 
orisered 

(L6L1 

JOZUP]) S2AV]IAINIS 

sndvdouadcE 

uorsres AIp 
Aaid ‘sauoq pyo 
Hype ep. 

‘(efos ‘eidod) 
UIIUOD [IO YITA 

(9002) ‘Te 3° 

BroquamA\ “(626 1) ‘Te 19 epnazy 
(9002) 

‘Je 19 Sroquai)\ “(600Z) AOWOT, 
‘(686T) Seoy Pue sourey] “(6Z61) 
‘ye 9 apna ‘(QOOZ) JousIVYD 
Nd ‘(S00Z) ‘Te 39 ses10g 

spas ‘aseydossou 
‘orepoid 

HO “as 
‘ON ‘IT ‘OH ‘Td XC ‘ULV 

HO 
“AS ‘OZV-Ld ‘Ld ‘ON ‘IT 
“ad ‘Td “Ad “Md “Dd CLV 

LI 9261 

IT “S€6l 

o1uas 

-o1dAr7 

jeordoriqns 
estdory, 

joyepoid 
orisered 

jorepoid 
orisered 

(8SZT snoeuulT) 
VIIVIOI VIGOAIANT 

(SZZT 1995 2) 
sadifna 11g 01aNT 

(9002) ‘Te 39 S1aquaat\ “(6007) 
sasiog pur eSuopuayy (6/61) 
‘ye 19 apna ‘(QOOZ) JousIVYD 

Nd ‘(S00Z) ‘Te 19 ses10g 

speurrue 
SutAesap ‘sou0q 
Pjo uo s01epaid 

SIDUDIIFIY 

vege 

HO “AS ‘OZV-Ld ‘ON 
UT ‘OH ‘Td “aad “Md ‘LV 

S31JJUNOD popeauy 

IT ‘9Z61 

adoimy ut 
prosel IST 

o1uas8 

-o1dAr7 

joyepoid 
orisered 

snjejsg 

(CZZ1 smioriqe.y) 
stpooifna VIGOLIANT 

sepHayy) 

satoads 
Ayrurey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

352 

(6007) soueL, 

‘(ZQ0Z) JoNNooUOYy “(000Z) 
TWOIC) pure opeypeyy ‘(Z961) “Te 
19 apnosy “(6761) ZO°fey “(800Z) 
BN “(S00Z) ‘Te 29 so810g 
(6007) AWOL, *(600Z) 

sosiog pur eSuopuayy ‘(000Z) 
TWIOIC) pure opeypeypy ‘(Z9G6T) “Te 
9 apnoaly (6761) Z07]eJ “(800Z) 
BN “(600Z) ‘Te 39 so810g 

syOosuT 
‘suuneqsioy 
‘sjonpoid posois 
‘snoseydoodur 

sumay Arp ‘Isuny 

aD ‘Wn 

"HO “AS “NVO-S8 ‘TS “MS 
‘OU ‘AVW-Ld ‘OZV-Ld 
‘Ld “Td “ON “IN ‘CW ‘LAW 
ALOIS LAr do 
“ad ‘Ud ‘Id “Nd °ZO “UH 
‘Od “Vd “Ad ‘Ad CLV “TV 

dD 

‘NVO-Sd ‘OU “AVW-Ld 

‘OZViLd “LW ‘AT ‘OISLI 
LL UD: Ws "ZO “Da “TV 

Ld “6€61 

SNOIOA 
-ninap 

SNOIOA 
-lMap 

(€9ZI 
‘jodoo¢) szuvyyaa 

snsvqdodlay 

CPS, wanig 
stuiffy snévqdoydhry 

(600) ACHOL, “(Z96T) 
‘ye 9 opnaly ‘(G761) Z09[e4 

Sypfuu ‘ore 

aq): Wl HOS. “IS 

MS ‘SU “OU “Td CLT ‘AT ULI 
“Ad ‘Ud ‘Td “Ad “Md °ZO 
‘Od “Vd “Ad ‘Ad CLV “TV 

SNOJOA 
-lnap 

8781 Tequey[Ay 
snnsUuvINnID 
snsvqdodlay 

(9002) ‘Te 39 Br9quont\ “(600Z) 
AOWOT “eIyeI] UT Malye 1191103707) 
"THaeY ‘(Z96T) ‘Je 29 epnosy 
(6761) 2097/8] “(8007) Hd 
(900Z) ‘Je 39 S19queI\\ “(S00Z) 
eyAnisey pure BAOIZIS ‘(O007) 
SPIOUIWIG] pur pseesopy ‘(/96T) 
‘Te 19 apnary “(8007) Hd 

SIDUIIIFJIY 

[evoreur ueyd pure 
poom Surkesap 

U] SsdIOF 
soseydoodur 
jeroieu 

queyd Surdesop 
uy ‘soduros 
soseydoodAur 

dd ‘CI 
‘TIX ‘D 

veNqeH 

€D “WN “HO “AS ‘NVO 

“Sd “SH “IS ‘NY “AVWLd 
“Td ‘ON “IN ‘CW ‘LW ‘NT 
LT ‘AT CLI “Ad “YOO 
Ud ‘Td “Ad “ZO “UH ‘AG 
€D “WN “HO “ds 

MS ‘OZViLd “Td ‘ON ‘GW 
(GT AT Lad ld “Aa Id 
‘ZO “WH “Ad ‘Ad CLV “TV 

S31JJUNOD popeAauy 

dD ‘ZE6l 

adomy ut 
prosel IST 

POLIOWY 

YON 

SNOJOA 
-lnap 

snoJOA 
-lnap 

snjyejg 

T88I ‘o]Aoureg 
ap noslig">) 
vipurdapqns 

SyaISOUaD’) 

LL8T PUIPY 

ISIN] DIADUOIY 

sepiseydoyddr+,y 

satoads 
Ayrurey 


353 

Coleoptera families other than Cerambycidae, Curculionidae... 

(0007) HHOIO 
pur oper “(8007) H"d 

(‘elyeay] UT TuoTye 1191102707) 
mey “(Z961) Te 39 epnary 
‘(6Z6T) Z07[R “(8007) BNA 

req stonod 

sjonposd posois 

€D ‘NVO-SA 

aD “AS 
“IN “AC ‘Ud “MC “ad ‘Ad 

dD ‘CI6I 

umouxuy) 

POLY 

POLIOWY 

YON 

jorepoid 
/orsered 

SNOIOA 
-ninap 

O¢6l 
IIe] saplowpigsvsv 
snundqsy 

sepreydas0qé,y 
(EPS 
UWTosJayuUry[) 
snn1Uusofiqva 
sng1yoUuapy 

(6007) 49°d 

(6007) 
AOUWOT “(Z96[) ‘Je 39 epnoiy 
‘(6Z61) ZO>78I (8007) HA 

Jopoay 
snsung ev Ajqeqoid 

sinu ‘sqmaj AIp 

NVO°Sd ‘Sd (LW ‘LI “Ad 
€D “WN “HO “dS ‘NVO-SA 
‘IS “MS “GVW:Ld ‘OZV-Ld 

“Td ‘ON “IN ‘CW ‘IT ‘AT 

UVS-LI “LI “Ad “UOO 

Ud “Ud ‘Td “Ad “Md °ZO 

‘Od “Vd “Ad ‘Ad CLV ‘dV 

LI “2661 

Dd ‘9S61 

otua38 

-o1dAr7 

snoJoA 
“nap 

snoJOA 
-ninap 

(ZZ81 ‘193312 y]) 

snatuodul snyain’) 

981 
“Zee IY snypuunf{qns 
snsvqdodhay 

(6007) Aowoy, *(6007) 
so8iog pue eSuopuey ‘(296 1) 
Tae PMP (GCG) AP Ted 

(2861) “TE 3° [oyu “(S007) 
‘Je 19 sasiog ‘([g6]) uosIsuag 

(6007) AOWOL, (S007) 
eyANIse'T pur PAIS ‘(/96T) 
‘Je 19 apnozy ‘(8007) ANG 

SIDUDIIFIY 

sjonposd posois 

veNqeH 

CVW-Ld ‘OZV 

cLd ‘AT ‘Ud “Od “Od ‘AG 
€D “WN “HO “aS 

‘IS “MS ‘SY “OU “Td “IN 
-LASEISAT OIssE Lal 
“Ad ‘Ud ‘Id “aa “Md “ZO 
‘Od “Vd “Ad ‘Ad CLV “IV 

S31JJUNOD poprAauy 

Dd ‘9S61 

ZO ‘0061 

adoimy ut 
proses IST 

SNOJOA 
-1nap 

snoJOA 
-lnap 

snjejg 

8781 PeyYaTAD 
snsopid sndvqdordta) 

€S6l 

‘JUMOIG -INojeg 

xoyvf snsvqdordiay 

satoads 
Ayrurey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

354 

(SOOT) PyANIseT pue 

BAOLJOS “ePHSIUTIIC] P]IOA\ JO SNOIOA COG6L ‘panel 
ansoyeie7) V ‘PArL “(€QOT) PAPH sjonpoid poesois CERO LW °ZO} ZO ‘VO0T| Blsepeasny -Tq19p V SNILUDIIO SNUILGIUP 
(sisou 
Areuuotssao01d 
(9007) ‘Je 39 S19quai aurd ‘siaaepeo 
‘(600Z) AOWOT “(SQOT) BYAnseT| ‘sadavy) snoseyd 
pure eaosag “elyely UT Ture -O190U PAIR] 
IaN0ITOD) mey “JePLSWIOC ‘aseydourjod 
PHIOM, JO onSoyere_y YW ‘eAvET] YNpe “or ‘sorqey 
“(COOT) PAPH ‘(6Z6T) ‘Te ‘somniruiny uo aida aig = A ad Fle TS o1ua3 SNOIOA HSCS ‘U0 DIT 
19 9pnox (8007) HMA] spay Snsauop} =H ‘If} <LI UVS<LI ‘NA ‘ZO‘Da) Td ‘Ss6r) — -orddrp -inep| Oo sadianyf snuasqqup 
summosnu 
UT SUOTID9T[OO UT 
(oepnsowsad SIIQe} USAOM 
POM Jo onsoyerey VW pure sole] SNOIOA 9/81 ‘AYS]OS 
"eARH ‘(6Z61) ‘Je 19 epnasy soded ‘poom| = J ‘T{ Td Ter) HRT GEST eIsy -unep| vy snplanyf snuaquupy 

(FOOT) ‘Te 29 PINY “oPpHsoauMI9q 
PHO Jo onsoyerey V 

"PARH “(6261) ‘Je 29 epnary 
(€00Z) JewWIaQyY “oepNnsautsaqd 

SIOMOP UO INpe 

SJaSUDABOS BATE] 

Td ‘AT LV 

AT‘I¥6I 

snoJOA 
-lnap 

I881 PuPY 
SNIISBINDI snuUaLgIUp 

PHO JO anBopereD Vy “ARH (€F81 
“(€COOT) PAPH ‘(6Z6I) Te snoJOA ‘adop{) syvsisny 

19 apnoasy “(800T) HAC say1ojo If aD “IN ‘UA} AD ‘ECG | eisepensny “3p V SNAIIOUIAGIUPY 
sepHsowisoq 

IZ6l 

‘esunox -Apospuq 

(‘eIyeI] UT TUSTTe 1301202707) “Ie yY ayeroduray, jorepaid snotuoddiu 
“(Z00Z) 14nTq ‘(S00Z) ‘Te 19 suvAy] sayeos Jo jorepoid LI ‘Z00Z -PISY /orsered snypgdaz0gk) 
adoimy ut satoads 

SIOUIIIFOY yevIqey S3IQUNOD papeAuUy psooas IST snqeig Ayrore,y 


355 

Coleoptera families other than Cerambycidae, Curculionidae... 

sTeaIao 
(9007) | pue snoseydorsou 
‘Je 19 Sroquaii\\ “(600TZ) AOWOT, BATE] 
“(¢00Z) fpey “(FO0T) PUSeEg ‘aseydourjod 
pue uueullopy “(€Q07) PAPH ynpe ‘soriqey 
‘(6Z61) ‘Je 19 aPpnesy “(800Z) uo Ayureur aq qy orua3 SsnOIOA IGLZT Wyeig 
HNQ ‘(S00Z) ‘Je 39 sesi0g spaoy Osamop| ‘Of ‘T{}| ‘HD “Td AT Nd ‘ZO ‘Dd| AD ‘8Z61 -o1dAr7 -inep ay LojonUN snuasaqyy 
SUOTIDIT[OI 
Jeorsojouro1ue 
(S00Z) | ‘ssurpying osye ang 
eYyANISeT Pure PAOTJIG “(FOOT) Te] (Too ‘snj “upys) 
9 vINY ‘(€00ZT) CARLY (8007) [eloveur uTs1I0 qos Tat orua3 SNOJOA €/6] ‘AsnueYyZ 
HN ‘(FOOZT) ‘Je 39 sPysaosreg -yeurrue jo asad LA) STONER Sih ASFA BT eZel -o1dAr7y -lnep oy 1QOUAIUUS SHUIDILYy 
(8981 
orerodurdy, SNOIOA ‘ANY 2Q JURsTNYY) 
(€QOTZ) PARYZ] ~~ stoNpord pesos [ Td | umouxua -PISY -mep| oy xu] snuasviyy 
oreroduray, SnoJOA Z68T ‘AYzZspyeasy 
(€OOT) PAPH sjonposd paroas { qs | uMouyu, -vISy -Inep Vv mjoI1gos snuasviyy 

(6007) 

AOW OT “elyeay UT TUSTTe 1193103707) 
Ney “(€OOT) PAPH ‘(6Z61) 

‘Te 19 apnary “(8007) Hd 

yerosoa 
ur ‘snoseydoiseu 

sjonpoid posois 

(€O0Z) PAPH 

SIDUDIIFIY 

veNqeH 

dp LAWL agg 

$31JJUNOD poprauy 

dd ‘2761 

umouxuy) 

adomy ut 
proses 3ST 

POLIOWIY 

SRO 

SNOIOA 
-lMap 

snOJOA 
-lnap 

snjejg 

(C6ZT “ssoqunyy) 
snqpiaspf snuasvuy 

9C6I 
‘Og suaasagndassaarp 
snuasvuyy 

satoads 
Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

356 

(‘oepnsotusiaqd 
PHIOM, JO onSoyere_D YW “eArET 
‘(€00Z) APH (8007) H"A 

souoqg peysns 

umouyuy) 

SNOIOA 
-ninap 

CSI TEM 

7 GIIA SIISIULLICT 

(6002) 

AOWOT “oephsawsad PjIo/\ Jo 
ansoyeie7) YW “PAR ‘(CQOZ) PAPE 
(6861) Say puE soureHy “(626T) 
‘Te 39 spnary “(6007) PuoueD) 

uonepaid s8sa 
‘(uJOD ‘sinuvad) 
sIONeUT [eI9SIA UT 
nq snoseydois.u 

IT ‘QH “Ud “Ad “Md “Dd 

Dd ‘0881 

snoJoA 
-ninap 

(SCZ ‘snovuurT) 
SNIADPAD] Saqsaulsacy 

(6007) 

AOWOT ‘(600Z) SedIog pure 
esUOpUayY “OePNssuJoq POA 
jo ondojerea VW “PAR ‘(€00Z) 
PARLE ‘(686T) Seay pure sourepy 
‘(6Z61) ‘Te 39 9pne4sy ‘(8007) 
BN (S007) ‘Te sassog 

onsowop 

AD ‘OZViLd ‘II 
‘AI “Al “Ud “Ad “Nd “Od 

dD ‘7981 

sNOJOA 
-lnap 

Z6L1 “uURpasny 
1G ISIAf SAISIULAAT 

spurysseis 

ysy peop 

‘sisou piiq ‘sasnoy 

(€007Z) APH 

(‘oepNsotusJaqd PIOA\ Jo 
ansoyeie7) VW “PAPE ‘(686T) SY 
pur sourezy ‘(6/6[) ‘Je 39 epnory 
(0007) 

TUIOIC) pue opeyseyy ‘(€00T) 
PARLE ‘(6/61) ‘Je 19 epnoly 
(9002) ‘Te 

qo SIaquantA\ ‘(GOOT) EYANIASe’T 
pure PAodjog ‘(O00Z) FHIOIO 

pure opeypeyy ‘(€QOT) PAPH 
(6861) SPY pue souTeH ‘(6/61) 
‘Te 19 apnary “(8007) Hd 

ur snoseydoiou 

sjonpoid posois 

snoseydoiou 

SIDUDIIFIY 

veNqeH 

‘Id 

dD “Td “AI “Ud “Ad 

€D “HO ‘NVO 
“Sd ‘OZV-Ld “Td “LW 
UT “ad ‘Ud “Ad “Od “LV 

$311JUNOD popeAuy 

umouyuy) 

Td ‘6161 

uMOuUxU al 

dD ‘8981 

adoimy ut 
prosel IST 

POLIOUYy 

S20 

ayerodura} 

-PISY 

SNOIOA 
-lnap 

snoJOA 
-1nap 

snoJOA 
-1nap 

SNOIOA 
-lnap 

snjyejg 

SORT uaaais 
SNIDUOLOI SASAUAIC 
CLLI ‘sniotaqey 
SNAOALUADI 
SAISIULAICT 

I8ZT ‘stonqey 
LOJONG SASIWAIT 

LEAP) 

IC] amy saqsauLsaq] 

satoads 
Ayrure,y 


357 

Coleoptera families other than Cerambycidae, Curculionidae... 

(9007) ‘Je 19 sIOquANITA\ 
‘(SOOT) PYANISe'T pure PAOIJIS 
“eryel Ul TuaTye 1191109]07) 
T17eY ‘(¢861) 30dwoenbs07- pue 
zane ‘(€00Z) PACH “(6Z61) 
Te? sprett 48000) BPC. 

SUOTID9T[OO 
uunasnur UT pure 
ssovjd onsawop 

€D “HO “dS ‘ON ‘AT 
“LI “Ad “Ud “Ad “Md °ZO 

aD ‘ZZ61 

POLIOUIY 

YON 

SNOIOA 
-lnap 

(6€61 “VOIP 
avyndsan vsaay 

(‘oepnsotusaqd PIoA\ Jo 
ansoyeie7) VW “PART ‘(€QOZ) PAPE 

(£007) PAPH “(6Z61) 

Tae eprett (8000) BPO 
(‘oepNsowsJad PIOA\ Jo 
ansoyeie7) YW ‘PAR ‘(€Q0Z) PAPE 

sjonpoid posois 

IN ‘1G 

uMOUxU al 

uMouUxyU al 

umouyu) 

jeordoay, 
-vISy 

jeordoriqns 
eotdory, 
eOLOWy 

SRO 

SNOJOA 
-lnap 

SNOIOA 
-1nap 
SNOIOA 
-lnap 

(IEP:4SI161 

‘MOITV) 40709144 
DULOUOPBLG 
8esI 
d][Aquayy-uranry) 
saduynf snurgdi¢C) 
(Z88T ‘oudef) 
IULOY SISJQQONT 

(£007) 
eaeH “(6Z61) Te 39 spner 

(S00Z) ByAnase’y] pur eaowyas 
‘(Q00Z) FUIOIE) puke opeLpeyy 
‘(€007Z) PAPH “(6861) SPY Pur 
sourey] (6261) ‘Te 19 apnaqy 

(900Z) ‘Je 39 Ss19quai)\ 

‘O8188 “(6861) Seo pur sure] 
‘(6Z61) ‘Te 39 epnany ‘(8007) 
BN (S007) ‘Te 9 sassog 

SIDUDIIFIY 

SNOIOAIIIOP 
(sJouIOy UIOD) 
JaneUr [eI939A UT 

nq snoseydoisou 
“SSOlIqey 
‘sionpoid yewrue 

uo ‘SOMsourop 
(sJouUIOy 
UJOS)I9}7eUT 
[e2999A UT ING 
snoseydoioou 
‘soriqey ‘sionpoid 
jewrue uo 
‘Onsoulop 

veNqeH 

€D “HO ‘NVO-Sd 
Td {Rly al. teas Ze) aL 

dD 
‘HO ‘OZV-Ld ‘Ld “Td “LI 
UT “dal “Ud “Od “LV “TV 

S911JUNOD popeAuy 

umouxuy) 

Td ‘6161 

Td ‘TZ81 

adomy ut 
proser IsT 

ayeroduray, 

-vISy 

POLIOUIY 

SRO 

snOJOA 
-lnap 

snoJOA 
-Inap 

snoJOA 
-lnap 

snjeysg 

098T Ays[NYsoy 

XDAO SIISIUAICT 

OPST ‘Neupaisez op 
suode] snuviansad 

SIISIULALICT 

GLLI 9D 2d 

SNJDINIDUL SIJSIUAICT 

satoads 
Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

358 

(9002) ‘Te 39 S19quant\ *(6007) 
AOUWIOT “(CQOZ) PYANAseyT pur 
PAOIJES “(€0OT) PAPH “(6L61) 

speorao Aqpersodsa 

€D “HO “Td “OIS 
<LI ‘UVS-LI “LI “AI “NH 

SNOJOA 

8681 

‘SUIOAW WNLAVUDAS 

‘Je 19 apnoazj ‘(800Z) HNq| ‘s~onposd pasoas “Ad “Md “ZO “Dd ‘LV “TV! AD ‘S681 “3p DULAIPOSOLT. 
sdsem 
(900Z) ‘Je 19 Bs9queIT A “(600Z)|  AreITOS Jo sisdu (€8/T 
AOUIOT “(€QOZ) PAPH ‘(6Z61T)| Ul pue suoneniis q ‘HD snOIOA Qsqiop]) Wnuquis 
‘Te 39 apnory “(8007) HA oFsoWOp ‘LT ‘AT “Wd SC ‘Dd SLV| Dd ‘PO6T “Tap DULLIPOSOLT 
(900Z) ‘Te 29 sIOquaNIIA\ ‘(SO0Z) SUONIDIT[OO 
eYANISe'T PUP PACTS “(900Z) ‘TP uinasnut UT (6P81 

19 vay “(€00Z) PACH “(6261) Te 
9 apnoly ‘(FOOT) ‘Te 39 SPysAasreg 
(£007) PAPH *(6Z61) 

‘ye 9 apnoasy “(GQOT) PYANIse'T 
PueRAogeg “(800c) 9d 

(€002) 

pue suonenis 
onsswop 

sjelioreur TeuTUe 

HO “dS “Td 
dl CAC NG ZO 

aD ‘LI ‘Ud 

Td ‘176 

LI “S€6r 

POLIOWY 

$2850 

oyeroduray, 

-vIsSy 

SNOJOA 
-lnap 

SNOJOA 
-lnap 

‘JOIJOS) wngsndup 
DULAAPOSOALT. 

GEST ‘AYs|NYOsIOY 
SHITDAJUOI sviapopdg 

PACH ‘(6Z61) ‘Je 39 epnosy sinuead SNOJOA C/81 ToI3yY 
“elyely UT TUdTTe 1J9IOJTO7) “11k Y ‘spaas porois LI ‘8S6I “ep tuaplay saporg14og]. 
£981 
‘TJayoequoipoy 
(£007) spnueul Joyepoid vipyidvI0fns 
eAeHY ‘(6Z6[) ‘Je 29 epnory Jo saseo 332 uMoUyUy) | eoUpy ‘eIsy| — /oisered DSSOJBDULNDYT. 
(€007) SNOIOA CET JOY 
PACH ‘(6Z61) ‘Je 39 opnosy uMOUUL) POLY -Inep tuaphay sadojay 
(F007) SUOTIDI][OO 
‘ye Jo PINY “oPpNsouJoq PIO, Jeorsojouro1ue 
jo ondojererd YW “PARP ‘(€00Z) ‘sy]IS MOpUIM snoJOA 6681 
PAPH] ‘(800Z) fapeyy pue [eog ‘souoq ysy Td ‘8661 POLIT -TWnop OI 2asnag vIuvLaS 
adomy ut satoads 
sadUaIafaY yeqviqey] satl1JUNOD paperauy psodaI IST snje1Sg Ayrore,y 


359 

Coleoptera families other than Cerambycidae, Curculionidae... 

6007 
sasiog pur eSuopuay es vlafy1UouL POLIOUTY snoseyd (zajoyosyps‘q) 
"Te 19 sassog ‘(966) ses10g | saprowagqupslug’) uMouyuy) S29 -ovdyd snausod snsapouo’y 
snoseyd OZG6I ‘Sogo 
uMouyun fal AC | TC“ Z61 voy -ovyd| oy | ntojdvg snsogdotpuvy 
sepliowwyy 
vououry|  so0repord CLL] ‘snioriqey 
(8007) Hq jorepoid uMoUxuL) SD] /nisesed syvjsoa sakpooayy 
seprsn iq 
(66Z1 
(6261) SnOIOA ‘JOZINIID) 407 0I1s4aa 
‘Je 19 opnosy “(600Z) Tueue7D uonepaid ssso uMoUxuL) -l0p DULLIPOSOALT. 
4sod 
(‘eIypeq] UT TUaTye T91.0IT07) | JofeuT <asnoyarem 
IVY “(GOOT) BYANase’] ur sionpoid 
pure eaoyag “aeprnsousod porois jeuTue 8/8] 
PHIOM, JO onSoyere_D YW “eArEy pure yeqoS3aa SNOJOA ‘UOTE azquisva 
‘(€00Z) PAPH “(8007) HMC] — Aap Aue aeoym dD “AS ‘AT ‘SLI ‘Td “ZO} €D ‘8261 “Tap puLaposo4y 

(S007) 
eYAN se] pure PAOIJag “eITeI UT 
TudT]e 1193103]07) Mey ‘(€O0T) 

eaeH] (6261) ‘Te 39 apnary 
(S007) ByAnase’y] pur 

PAOIJAS “IPISIUIIIC] PjIOA\ JO 
onSoyeie) V “eAeH ‘(€00T) PAPH 

(€O07Z) PARH 

(eyfea] Ul TuaTye HonoajoD “ey 
‘(FOOT) PUseg pue UUPLIIOT] 
‘(€OOT) PAPE “(8007) HG 

SIDUDIIFIY 

WOTDaT[OO 
UT S]OOSUT 

sjonpoid posois 

sionposd posois 

sionpoid posois 

‘aseydoypdsd 

veNqeH 

aS “IN {LI ‘Ud “ZO ULV “TV 

€D “Td ‘LI “TV 

S31JJUNOD popeAauy 

ZO ‘0061 

ZO “S007 

umouxuy) 

AD ‘9S61 

adomy ut 
pr0sel IST 

POLOWY 

SO 

POLIOWIY 

528-0 

POLIOWY 

YON 

SNOJOA 
-lnap 

SNOIOA 
-lnap 
SNOIOA 
-lnap 

SNOJOA 
-1nap 

snjejg 

I881 FeuPYy 
SAPLOULOIVBIUL 

DULAIPOSOALT. 

996I ‘99T 29 
ovyD Wnsoqas15uo] 
DULAIPOSOLT. 

COST Ieporasyy 

AUDINSUL DULAPOSOLT 

ySsl 
QMUODIT wnsnjaut 
DULAIPOSOLT. 

satoads 
Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

360 

(Z661) Sow PA 
‘(€661) URIL pur [Pqopeq 

AutOr) *(8Q0Z) AUIOr) 

AUIOry “(1 /6]) ‘Je 19 9pnay 

(6007) 
‘(9007) 

ooruew ‘sinueod 

‘spod pur yreq 

jopun Joyepoid 

SIDABP¥D ‘SID9J 
rorepaid Ay 
Anynod ‘Sorueur 

umMouyuy) 

umMouyu) 

POLY 

POLY 

ioyepoid 
orsered 

joyepoid 
/orsered 

(OZ8T ‘[hasrepy) 
yakvus xisopdiq 
TEGT IPIEYOY 

SNIDIAYS: UaIap 

SNPJIUOIIIOY") 

ur suzy do1s 

“UN OGIAT. POLOWy rorepoid (LIST ‘qmy4eg) 
(SOOT) ‘Je 39 sosi0g uo Jorepaid { OZV-Ld| umouxu) SD pnisered|) y | sashpopsos sdoutssvy 

(9002) 

‘Je 19 Sroquaii/\\ “(600TZ) AOWOT, 

‘(600Z) sesiog pure esuopusyy 
“(LZ6T) ‘Te 39 epnany “(G007) SNINIUNIVACT HO ‘OZV syuas} = sorepard (PEST “UosyoIIy) 
‘Je 19 saBs0g ‘((Q66]) Sed10g| ‘sadavyz ‘sIaAeped S| STTTSAT. A et |) ATEEGGI -oidAry| = ommsered) =) oytund sdou12v’) 
deplIoIsip] 
(SOOT) BYANIseyT pure vaoljag suoosysnur Sd “Id SNOJOA C/8] 
‘(SL6T) ueLIOZUYY-YOIO|qL] ts ia LT WOO dd NAZI “Ee? ‘yo101_y 191d aun 
seprpAorq 
poom paiesap CER] 
(®6007Z)| ‘sued eutorpow o1uas sNOJOA ‘snd sussaudap 
‘ye 19 Aappoys ‘(9€8T) sBaND ‘sun Jj Arp ‘Inoy Of ‘[ Wi Md) UA ‘EPst -o1dA17y “Tap Dd snaaupvivdojofy 
(6007) AOWOT, one ‘yeroleur CEST 
“(®600Z) ‘Je 29 AAPpouS (Z00Z) queyd Surdesop HD| YOON o1ua3 SNOIOA ‘oqny wnvjinDvs 
JarNooUOYy ‘(CQOZ) ‘Te 39 ses10g uo ‘sn3unj uo| of ‘{ ‘T| “Ad UOORA WA “Dd ‘LV “WA ‘ZE6T -o1dAr7 -amep} snaaupivdojopy 
sepryppAwopuy 
LL81 ‘dreys 
(6Z61) ‘Te 39 epnesy ‘(8007) HG uMouyun AD ‘1861 | eiseperusny snqvyns snavdsuny 
adomy ut satoads 
SdIUIIIFOY WweIIQGey SoIIJUNOD popeauy pr0sel IST snjqe1¢ Ayrure,y 


361 

Coleoptera families other than Cerambycidae, Curculionidae... 

(F007) Yoraepuay 
‘(600Z) sesiog pure eSuopusyy 

‘(TZ61) ‘Je 39 epnery ‘(Z007Z) ‘Te jorepoid (JOST “ureysreyy) 
19 TeyNog ‘(CQOZ) ‘Te 9 sasiog n OZV-Ld ‘ZD | Umouxua eeisy| = /omisered| oy sdanadtu uokosay 

SPUPSSeIS JOM 

(900Z) ‘Je 29 Sroquaiu\\ “eIyeI] ‘SJUSWIUOITAUS 

UT TUaTTe 1199109707) “19eY ‘(0007) prumny snores qa ‘HO “AS 

SRIOUIWIG] pur pseesopy ‘(1 Z6T) uy Jovepord| =| “6.4 | ‘SH “IN ‘IT ‘LI “AG ‘Wa ‘Td|) = LI “Z| avesodwey| s01eposd €/8 | ‘dreys 
‘Te 29 epneary (8007) HPA ‘asoduros ‘€H) “HH “NC “ZO “A “LV “IV ‘OS6I -eIsy|  jolysered| = snqpuyutyy Wola’) 

(F007) Yoracpudy ‘(Q00Z)| URNS avo ‘sIIMIy 
TWOIC) pure opeypeyy ‘(Z00Z) ‘Je| Sur0s ‘poomeds NVO-Sa PCR “WorseTjon, 
12 TeyNog ‘(CQOT) ‘Je 19 sasiog Sutsodurosep | ‘OZV-Ld ‘LI ‘ZD UH SLY] usouyun eouyy| uMouyun| y snyvuinbut uokosay 

(‘elyeay] UT TUoTye 193109707 “Ie Yy 

(0007) PHOTO 
pure opeypeyy ‘(8007) AWOL 

(600Z) sesiog pur esuopusyy 

SIDUDIIFIY 

Sa09RJ ‘SIDABPED 
BoIe POpeAUI UT 
onueUI pure METIS 
‘asueI JATIVU UT 
vyundC Surdedop 

[los Apues 
‘SaddPJ ‘SIDARPPD 

vege 

Sd ‘Ld ‘OIS 
<LI ‘UVS+LI CLI “Us “WH 

NVO-Sd ‘Sd “Ud 

OZV-Ld ‘LI 

Sd “Wd ‘AD 

S311JUNOD poprAuy 

LI ‘786 

uMOUxU 7 

umouxuy) 

Ud ‘PZ6I 

adomy ut 
proses IST 

POLIOWY 

SRO 

“eOLOury 

YON 

POLIOWY 

S20 

POLY 

SNOIOA 
-1nap 

SNOIOA 
-lnap 

jorepoid 
orsered 

joyepaid 
/orsered 

snjejg 

seprprydospAyy 

PEST “UOsyoy 
suasny snutadvs 

COBT “Hessel 
LLIN] SNIVULOLDT 

(LISI 

qmydeg) sesuarpiswg 
snsmvI0dd ET 

LIST ‘myAeg 
snypjaundig AIST] 

satoads 
Ayrurey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

362 

(9007) 

‘ye 19 SIoquaty\ “(600Z) AOWOT, 
‘(S00Z) exANIseT pure PAOTJIS 
(9661) ‘Je 39 eleweIUeS *(600Z) 
sod10g pue eSuopusy “(800Z) 
BN ‘(S00Z) ‘Te 22 sasiog 

(9661) ‘Je 19 elrewelUeS 

(Z881-Z881) dreys 
(FOOT) [eynog pure yooryly 

yreq Jopun 

‘sjonpoid posois 

sjonpoid posois 

yreq yeo Jopun 

erewpaio1dyd 10 
‘s3a7eM ppoy ued 

HT 

€D “WN “HO “dS 

‘Sd ‘SU “AVWsLd “OZ V-Ld 
Lal Td AA LT A OISELI 
‘NH ‘UD “AC ‘Ud ‘Id “AG 
‘ZO “WH “Od” “Ad ‘Ad “LV 

Td ‘NH 
‘ad “Ud “MC “ZO ‘Ad “LV 

ZO ‘SL81 

Ud “0661 

LI “6761 

o1uas 

-o1dA17y 

POLIOUy 

SRO 

joyepoid 
orisered 
‘snOIOA 
-lnap 

snoJOA 
-lnap 

uMouyuUn 

(LEST ‘suaydars) 

snauianssal 
saqsajoqday) 

(FEI 

PEM) susvoy dnp 
saqsagoqday 
seprso;ydounry] 

PHS Iues[nyy 
laqsafv] BULOSO] aT 

(‘elyeay UT TuoTye 

HoNoZ]OD “HIeY “(000Z) WOO 
pure opeyse (4007) [exnog 
pure yooryly “(Z00Z) ‘Te 39 Jeynog 
(6007) ¥P°d ‘(600Z) s°810g pur 
eSuopuayy ‘(SQ0Z) ‘Je 9 sasiog 

JoyeM SUIPUrIS UT 

JoIeM SUIpUrIS UT 

NVO°-Sd ‘LI UD 

uMouUxU ial 

uMoOuUxU ial 

POLIOWY 

YON 

uMouxyuUn 

uMouxuUn 

(F681 
‘ues[nyy) asuadva 

UNULIISHYIV 

666[ ‘UssueT] 
1duvys wnusagsocE 

(600Z) sed10g pue 
vouOpuEeY ‘(000Z) HWOIO pue 
opeypeyy ‘(SQ0Z) ‘Te 19 sasiog 

(9007) ‘Te 39 S3aquaut “(S00Z) 
eyANAse’'] Pue PAOIJIG ‘(000Z) 
SeIOUIWIG] pur pseesopy ‘(1 Z6T) 
‘Te 19 apnary ‘(8007) Hd 

SIDUDIIFIY 

eAuayy 

UT [IAd0M vURURG 
uo Jorepaid 33a 
‘syueyd yi 

oem Surpuras 

‘orrydowsy) 
s}USWIUOIIAUS 
prumy snorea 
uy ‘Soreposd 
“soduioo 

veNqeH 

NVO'Sd 

‘Sd ‘AVW-Ld “‘OZV-Ld 
LI “UD “AC “Wd “AD “WH 
€D “HO “AS 

‘ON “IN ‘LI ‘NH “AC “Ud 
‘Td “MC °ZO “Ad “LV “TV 

S91JJUNOD poprAauy 

umouyuy) 

LI ‘OS6T 

adoimy ut 
pr0sel IST 

POLY 

ayeroduray. 

-PISY 

jorepoid 
orisered 

jorepoid 
orsered 

snjyejsg 

(Z6LT ‘sniolsqe.y) 
ayvUuIuLopqn 
uinusaqsopMqvCq 

HRB] ‘dreys azzgns 
uninadordts) 

satoads 
Ayrure,y 


363 

Coleoptera families other than Cerambycidae, Curculionidae... 

(900) ‘Te 39 Broquaqt 
“(SOOZ) ByANIseyT pue PAOTJOS 
‘(Z961) ‘Je 9 epnany “(300Z) 

FNQ ‘(800Z) Wa0urA pur 1o3no0g 

(SOOT) ByANAseT pue PAOLJIS 

(€00Z) 10]12D 

(9002) ‘Te 39 

Braquait\ “(6007Z) se810g pur 
eSuopuayy ‘(SQOZ) ‘Te 19 sasiog 

(900Z) ‘Te 29 S19quantT Ay, 

‘(SOOZ) PYANIseT pur PAOIJOS 
(9661) ‘Je 19 elIeEUIEIURS ‘(800Z) 
HN ‘(S00Z) ‘Je 9 sassiog 

(900Z) ‘Je 39 S19quai)\ 

‘(966I1) ‘Je 39 BlyeUeIUeS 

(6007) Aowo], 

‘(S00Z) ByAMasey pur eaosjas 
(9661) ‘Je 19 elIeWIEIULS ‘(Z700Z) 

winiiyeqioy UT 
punoj ‘snsunj uo 

spoo, ‘sumay Arp 
‘Spoas SHPULLVULD 

mmoop ‘ures3 

‘adeydoypdsd 

eUsS TA 
‘snoseydooAur 

‘sjonpoid posois 

oidoryjueuds 
JeoyM ‘uress 
‘symay AIp 
(snuwey 
-O1eS) POOM 
Arp ‘mop usoo 

aseulep ‘ured 

POH) AG ad Zo) 

dd Wi Md ‘Lv 

HO ‘OZV-Ld “LW ‘IV 

€D “WN “HO “aS 

‘Sd ‘SU “AVWsLd “OZV-Ld 
Ad" Td: OISSLL AV Sel 

LI ‘NH “UD “Ad “Ud ‘Ta 
“Id “ZO “MH “Ad “LV “TV 

HO ‘NVO°S4 ‘Sd “OZ V7Ld 
‘Ad “MOOS “Ud “Ad “TV 

OZV-Ld 
“LW “UVS-LI CLI “Ad “Wa 

ZO “6S6I 

ZO ‘0061 

Ad ‘7861 

umouyur) 

Ud ‘GO6I 

UA “T661 

jeordonqns 

SNOIOA 
-nInap 

SNOIOA 

-ninap 

SNOIOA 
-ninap 

SNOIOA 
-ninap 

SNOIOA 
-ninap 

SNOIOA 
-lnap 

SNOJOA 

(TZ8T “Woasey[o ) 
1U0SIDN vIMaIsIpy 
sepHplney 

CZ81 

TOINDY 12GISALA 
DYLOUOXDAVY J 

PL8 [FUNDY 
SNIVLIIN]GO 

sngigdogdhay 

(1V8T 499H) 

sasaqus snqigdoqdda) 

oepriinsury 

(OZ8T ‘eyJPAnNos5y) 
snaiaang saqsajoqgdday) 

(VET ‘sAnD) 
mands sagsajoqqhay 

(ZT81 Tequgyss) 

yarNoouoyy ‘(GQOZ) ‘Te 19 ses10g ‘ordosryaueuds M025 Da Ke EY “Tv ZD“SZeL| ‘jeardoryp: lop snpisnd saqsajoqday 
aD ‘VN “AS ‘Sd ‘SU 

(9661) SiMe ‘GVW-Ld ‘Ld “Td “LW “OIS (ZS61 “MOH 

‘ye 19 BIZeUTeIURS “ele UT TUATye joseydoyodsd “TE AM Saal a ora SNOJOA 2Q ]2016) saptonjisnd 

HWoNOITOD “Mey ‘(800Z) HNd| ‘swnposd posois Td MC ‘ZO MH “Ad ‘IV! ~LI‘8Z61 -o1dA17y -lnop saqsajoqday 

adomy ut satoods 

S9IUIIIFIY WwIIQGey So1IJUNOD popeauy prosel IST snje1¢9 Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

364 

(8007) # Ad 
‘(800Z) JUS.UT A pur JosNog 

(6007) 

AOUIOT “(966T) 40UTSL “(Z00Z) 
armnoouoyy ‘(600Z) SesI0g pur 
eSuopusy ‘(Z9G61) ‘Je 39 epnosy 
‘(8007) BNC ‘(800Z) 2U20UTA 
pur 1o8nog ‘(CQ0Z) ‘Je 19 sasiog 

yreq ourd 
‘sasnoy ‘sja10y 
‘sasnjal yeqrosoa 

poomory 
‘Avy ‘suINIF UNOI 
‘snuuny Isotoj 

HO “AC ‘Ud “Od ‘Ad “LV 
€D “WN “HO “dS ‘Sa 

MS ‘SU ‘OU “OZV-Ld ‘Ld 
“Td ‘ON “IN “AW ‘CW ‘NT 
LEAL DIssLl avs 

LI ‘NH “UD “Ad “YOO 
Ui Ud ‘Td “Ad Md “ZO 
UH “Od ‘Va “Ad ‘Ad “LV 

Ud “8061 

dD ‘6881 

otua3 

-o1dA17y 

SNOIOA 
-ninap 

SNOIOA 
-lnap 

(S¢ZT ‘sneouury 
syplgsaual VIAVIIJAO’) 

(L781 
jequayAy )wyvsuoja 
VIAVIIJLO’) 

(9661) A°UraL “(800Z) BA 
‘(800Z) JUS.UTA pur IosNog 

(6007) AowoL, 

‘(600Z) sesiog pure esuopusyy 
‘(O00Z) FHIOIC) pure opeypryy 
(8007) HNC ‘(800Z) UI0UTA, 
pur 18nog ‘(¢Q0Z) ‘Je 19 sasiog 

snp 

‘sulewial ‘sn Jj 
Arp ‘soduroo 
‘snoseydooAur 

Aey 
‘onze asoduros 
‘snoseydoosur 

aD “AS “ON ‘AT ‘Ud “AP 
aD ‘Wn 

"HO “AS ‘NVO-SH “TVd-SA 
‘SA ‘IS “MS ‘Su ‘NU ‘OU 
‘GVW-Ld “OZV-Ld ‘Ld “Td 
‘ON “IN ‘CW LW ‘NT 
EPL AE DISS LT Ay Sort 
Lal SI SAS 
UD “Ad UOO- Ud “UA 

‘Td “Ad “MC “ZO ‘AD “UH 
‘Od “Va “Ad ‘Ad CLV “TV 

dD ‘6881 

ad ‘0S81 

vIseyesIsny 

SnOIOA 
-lnap 

SNOJOA 
-lnap 

(L781 ‘peyuays5) 
DIIDAYS: uUoz AIP OIAD’) 

(6E8T ‘poomisay\) 
LafIpou aapoqsy’y 

(666) 2U20UTA, 
‘(800Z) 2Us9UIA pure iasnNog 

(€OOT) OWE (8007) HAG 
‘(800Z) 2Us9UTA pure iasNo0g 

SIDUIIIFJIY 

Aeoap pero 

‘snoseydoodur 

yreq Japun 

‘snoseydooAur 

rege 

Refel 
dD “HO “AS “GVW-Ld “IN 
‘Ad “Ud Md “ZO “Ad (LV 

SorsJUNOS peperauy] 

Ud “961 

Ad ‘0007 

adoimy ut 
prosel IST 

POLIOWY 

528-29 

vIseyerisny 

SNOJOA 
-lnap 
SNOJOA 
-lnap 

snqeI1¢ 

(0961 ‘zoleq) 
LT 

(ZZ81 ‘FouIey) 
VIVIISU[IQ AdaPOIAV’) 

satoads 
Ajrure.y 


365 

Coleoptera families other than Cerambycidae, Curculionidae... 

sasiog pur eSuopuaypy (2961) 
‘ye 19 opnoaly ‘(8007) HNC 
(GO0Z) ACWOT “(SQ0T) ByANase’T 
Pur PAorjog ‘(ZTOOT) JoHNoouOpy 

(2961) ‘Te 39 apnoxy 

HNQ ‘(800Z) Ws0urA pur 13no0g 

(SQOZ) BYANase’] pue vaosyag 
‘(800Z) JUS.UTA pur JosNog 

© 

(6007) 

umouyun 
yeroqeur 

yueyd Surkesap uo 
(g00Z)| ‘snsuny uo asead 
‘elreqiay ‘speatao 
sjuouredde 
‘sIe]]a9 ‘sOOI 

‘synaspooy 

€D “HO “AS “LW ‘AT “AI 
‘ad ‘Ud “ZO ‘O”d “Ad “LV 

Wd “Md °ZO 

uMOUxU al 

Ud ‘OSs 

ZO “0061 

eIseyeLIsny 

snoJOA 
-ninap 

SNOIOA 

-ninap 

SNOIOA 
-ninap 

L881 

“UO]IG SNIL]VAISND 
SNIPIAGYIVT 

(OS8I ?qny) 
uni Dy asaUatgy 

(ZZ81 933124) 
V1 DNAS (a) DUJasIUuatg] 

(9661) AOUpPL, 
*(ZOOZ) JaHNOsUOPY ‘(800Z) HAA 
‘(800Z) JUS.UTA pur JosNog 

$9o0J1 plo “SsoSSOU 
‘snoseydoodur 

dD ‘AT Ud 

dD ‘Z061 

SNOIOA 
-ninap 

(P88 1 FuI2y) 
SNBAV DYBLIUIICT 

(900Z) ‘Je 39 S1oquai 
‘(6007Z) ACWOT, (6007) sesI0g 
pure eSuopusyy ‘(Z96[) ‘[e 39 
apnory (8007) TE0UTA pure 
yasnog ‘(CQ0Z) ‘Je 19 sasiog 

Aayreq 
‘UIOO ‘TelIayeU 
queyd Surdesop 
uo ‘sn3unj uo 

HO ‘OZV 
Ld ‘U1 “AC “Od ‘Ad “LV 

IT “2661 

snoJOA 
-ninap 

(S6Z1 [I™yAeq) 
DIDAAIS DIADIIIA 0") 

(9007) ‘Te 9 

SIaquant\ ‘(Z96[) ‘Te 19 epnory 
‘(800Z) JUS.UTA pur JOsnNog 
(900Z) ‘Je 29 sIOqUONIA\ ‘(600Z) 
AOWOT ‘(600Z) sadiog pure 
eSuopus| ‘(Z9G61) ‘Te 39 epnasy 
‘(8007) BNC ‘(800Z) U20UTA 
pure 1o8nog ‘(CQ0Z) ‘Je 19 sasiog 

SIDUIIIFJIY 

yerorew 
queyd Surdesop 
uo ‘sn3unj uo 

‘syueyd jeuroipow 
‘o00eqo} 

[eloqeuw 
queyd Surdesap 
‘sjeasao ‘saotds 
‘ovoed uo I3unJy 

veNqeH 

HO 
UT ‘OH “AC ‘Ud ‘Ad ‘LV 

HO ‘OZV 
“Ld “AC “Wd “Od ‘Ad CLV 

S31JJUNOD poprAauy 

dD ‘Z681 

Ud ‘PZ81 

adoimy ut 
prosel IST 

SNOJOA 
-lnap 

SNOIOA 
-lnap 

snjyejg 

(L781 ‘TeyueTAy) 

suagsaqgnd VIAVIIILO) 

(ZEST “WJOWOD) 
van VILVINLO) 

satoads 
Ajrore.y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

366 

PyEM oy? ur Ajares 
(8007) Hd ‘spred Joquin 
sosnoy 

ur sod poom ‘(N) 

SNUIXDAY ‘snILANC) 

(‘eIyea] UT TuATTe 1393309]0>) 
‘ey (6961) ‘Te 39 apna 
(900Z) ‘Te 29 S1oquant Ay, 

“elyel Ul Tale HoqO0s[0D “WeYy} sasnoy ul poom 

Ud ‘Td CLV 

HO “ad ‘Ud ‘LV 

umouyuy) 

Id “SE6I 

Ad “9661 

POLIOWY 
YON 
POLIOUIY 

YON 

snoseyd 
-ovdyd 
snoseyd 
-ovdyd 

TI6[ ‘useT 
SISUIUIS SNIMT 

SC8I IUD] “T 
"( sqqootund sngokT 
COST ‘IU0DIT 

"T *[sypossava sng 

(9007) ‘Te 29 Sioquont A “(C00Z) 
eyANise'T] pur BAO “(600Z) 
so810g pue eSuopucyyy “(S00Z) 
"Te 19 DPpopuaar]sy “(6961) "Te 

19 apnary ‘(CQOTZ) ‘Je 19 sasiog| poomdes ‘sorueur 

HO ‘Su ‘Ld “LI ‘AT 
UVS-LI CLI “UD “AC “Ws 
MNGZO- De Ad LV “Ty 

Ud ‘OSs 

snoseyd 
-odyd 

(O€8T ‘suaydars) 

snauunsg sngaT 

(900Z) ‘Te 29 S1oquant py, 

“elpel] UT TUaTTe Ta0sjo | ~pyay ur poomdes snoseyd LOGI ‘OUSIT 
"Mey “(6961) ‘Je 19 spn ‘s10OIJ JosuIs HO Wa ‘Ly | umouyuy POLY -o1dyd snupoidfy sng] 
sepnokT 

[erorew LL81 

(900Z) ‘Je 29 S19quaiu\ | IUeTd SurAedap uO o1ua3 SNOJOA TONY SyVIUILLO 
‘(800Z) 1USIUIA PukIesnog| ‘sn3uny uo ‘O11 HO “Id “AG ‘UA “MC ‘LV| ad ‘€661} = -ordAr_ “LENOp DIXNvaUsY 
SOAR] VIGIL 

peop ‘sasnoyarem SNOJOA unolg szuuadissas 

(8007) H9q ‘mers UO Suny qD “AC! AC ‘8761 | ersepersny lop snuvyrgdorayy 

(900Z) ‘Te 39 B19quo1t MW ‘(600Z) 

AOWIOT, ‘(ZO0T) JoTINOIUO|, 
‘(Z96T) ‘Te 19 ePpnery “(Z861) ‘Te 
19 TPPYOU “(8007) HN ‘(800Z) 

JUd.UTA, pure J98NOg ‘(CQ0Z) 
‘Je 19 sasiog ‘([g6]) uosIsuag 

Jeroiew 
queyd Surdesop 

uo ‘snsunj uo 

‘orq7e “sreypao 
‘s][Puu /speatao 

SIDUDIIFJOY 

veNqeH 

aD “HO 
‘OZViLd ‘II ‘AT “ACG “YOO 
“Ud “Ud “Od “Od ‘Ad “LV 

$31JJUNOD poprAuy 

Ud “TS81 

adoimy ut 
prosel IST 

SNOIOA 
-lnap 

snjyejsg 

(Z9Z| ‘shoeuurT) 

SNINUIUL SNIPIAIVT 

satoads 
Ayrure,y 


367 

Coleoptera families other than Cerambycidae, Curculionidae... 

(900Z) ‘Je 19 S19quai 
‘(CQOZ) BYANAse'T] pure eAaosjas 
“elyel] UT TUOTTe 1191109]07) 

‘Tey ‘(SQOT) ‘Te 19 sasiog 

(9661) AOUlPSL, 
‘(SGG6I) PUTT ‘(9Z6T) UaNeEg 

(6661) SID pure unodyepy 
(9007) Aydinyy pue pooqy 

SIDUIIIFJOY 

sjonposd parois 
‘sadei3 porp 
‘OZIeYY ‘Telayew 
queyd Surdesop 
uo ‘sn3unj uo 

"ISOIOJ 
ourd ur szugsaaqzs 

VIN ABUY JO SIOMOY 
UO ‘sasnoyusass 

WIeM ‘VIPUL 
DhapqIV) 

(vasogs 

‘vissvquooyy 

DIIyar] ‘xvquog 

“piuuaslapy 
‘sndAvI0q APF 
DyIZfV) YE 
uy} Joieais 
JO S[aAaq] Yoseds 

YIM Joaquin pue 

$991} SNOJIJIUOD 
JO Jea[peorq 
‘poompiey 
posod-aprm 
yore ‘apjioaq 
Surpeaj-Taquiny 

reuqeH 

Ho 

‘OZV-Ld ‘LI “Ud “ZO CLV 

IN ‘AI “AG 

SoTsJUNOS poperauy 

ZO “€861 

IN ‘IZ6I 

uMOUxU ial 

adomy ut 
prosel IST 

POLIOWY 

YWON 

POLIOWY 

SRO 

jeordosnqns 
‘yeotdosy, 

SNOIOA 
-lMnap 

snoseyd 
-odyd 

snoseyd 
-ovdyd 

snqeI1¢ 

OC8T ‘aU0DaT 
SNIVIYVG SNBLVNT 

sepiseydojo04/ 

C16] ‘uordureyy 
puviayqqvo 

DUAIS! Tap OW 
oePHIPPIOW 

(SS8T Toye) 
SYJOIIONA vagqUulyy 

satoads 
Aprorey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

368 

(€007) SNOIOA 
Je 19 VIIIA “RIPE UT TuaTye soyreues3 lp 
INOITO7) "IeY ‘(600Z) sessog| ‘samnay Arp ‘spoas OZV ‘snoseyd ISS] ‘ueWYyog 
pue eSuopuayy “(€661) OIsIpny SIPULADULDT, iat “Td'kd “DIS-LECEI “TY | LT ZZ61 ROLY -ouyd| y  |suswunf{ smprqdodivy 
SNOJOA 
Sa a Saal 9S61 
ppy ur ‘OZV-Ld ‘DIS-LI UVS-LI jecrdonqns| ‘snoSeyd ‘uosqoq zuvuaadf 
(0661) S°810g “(EG6T) OISIPNY| ozreur ‘symay Arp fT) ‘LID YOO NAC “TV LIT ‘9Z6t| — “Teordory, -ouyd) oy smaydodiny 

(6007) ACW], 

‘(SOOZ) PYANISLT pue PACIZIG 
‘(LQ0Z) JeRNOOUO/Y ‘(800Z) 
OIsIpny pue pnsyIT “(600Z) 

HO ‘Sa 
OZ “Ted (LIN OISSLI 

SNOJOA 
-lnap 

(Z6LT ‘sntorsqe.j) 

sodiog pue eSuopueyy “(S00Z) SP2u a UtOS UVS.LI LI UOO-Ud “UA vopoury|  ‘snoseyd bpcsits A 
Te 29 so8sog “(€661) CIsIpNY| _‘sionposd pesos ‘Aa AC ‘ZO “DE ‘LV“TV| ZO‘006T] SRO} -odyd snrqdodar’) 
NVO 

“Sd “IVd-Sda “Sd ‘TIS “Su UE 2 
‘GVW-Ld “AW “LW ‘OISsLI “HERep POST ‘Avrnyy 
UVSLI ‘LI UD UOO-U ‘snoseyd snavaasauafiq 
(S007) OlsIpny pure pnsyfy siMIJ Ud}0I Sa AS SH“ “Wa “aT POLY -odyd smiqgdodivy 
(600) s9srog pue 7961 
eSuopuayy ‘(CQ0Z) ‘Je 19 sasiog yreq Jopun SNOIOA ‘Adssey) 29 Joupser) 
(0661) S9310g ‘(€66T) OIsIpny| ‘sionpord posois CVW-Ld ‘OZV-Ld eIseperIsny “lp ynvu sndaddqgovag 
(1007) JoHNoouO|y yO8I 
‘(800Z) OISIpny pur pnsjpy SNOJOA ‘AelIny Jayjosap 
“elpei] UT TUDTTe 1aOITO7) “WIRY symsy Surkesap POLY -lnop snydaddqgavag 
seprnpHin 

(900) ‘Te 39 Sroquaqt 
‘(600Z) ACWOT, ‘(600Z) S2sI0g 
pure eSuopuspy ‘(Z96[) ‘Je 2° 
apnory ‘(GQOZ) ‘Te 19 sedi0g 

SIDUIIIFJIY 

one ‘sypru ‘Aeoop 
“o1seM ‘Tela eu 
queyd Surdesap 

uo ‘snsunj uo 

veNqeH 

HO ‘OZV 
<Ld (LW ‘11 ‘OIS<LI “UWS 
TLL AG Ma “De LV 

S31JJUNOD poprauy 

Dd “SS6I 

adoimy ut 
pr0sel IST 

SNOJOA 
-lnap 

snjyejsg 

(SCZT ‘snovuury) 
DIAOIAIAS vavy dy, 

satoads 
Ayrure,y 


369 

Coleoptera families other than Cerambycidae, Curculionidae... 

(6007) AOWOT, “(8007Z) OISIPHY 

pure pnsyiyy ‘(600Z) sesiog pue} sasnoy ‘uvaueisay 
eSuOpUs| ‘(000Z) FWOIC pur | -Ipeu ul s1oop3no 

opeypeyy “(S00Z) ‘Te 19 ses10g 
(6002) Aowe] *(S007) 
eYAMISL'] Pure PAOIJIG ‘(800Z) 
OIsIpny pue PNsyIV “(600Z) 
sasiog pur eSuopuayy ‘(SQ0Z) 
Te 39 so8s0g (E661) OFSIPAW 
(9007) ‘Je 2° dIOQUIIIT 
(6002) ower, *(S00Z) 
eyANise'T] pur BAOIas “(€00Z) 
doueee (O00C) Sepa pur 
preesapQH ‘(600Z) sesr0g pur 
vsuopueyy “(000Z) MHOC pure 
opeyoryy (8007) HX ‘(S00Z) 
"[e 19 sads0g “(€66]) OIsIpny 

adosne yer1uao UT 

‘symmay AIp 

symay Ap 

snoseydoides 
sodurod ‘speaz39 
Sonsowop AyUTeU 

VN ‘NVO°Sd “TV4-SH ‘Sa 

‘IS ‘NU ‘OU “OZV-Ld ‘Ld 
ELA DISSE aVSsLI- Sas 
UD “DANAUD ‘NOUS 
“AMOUD UD UOO-UA 
Ud ‘AO ‘UH “Od “Vd “TV 

OZV-Ld ‘LW ‘IT 
‘DIS:LI “UVSLI WLI UOD 
Ud “Ud “AC *ZO ‘Od “LV 

€D “HO 

“AS ‘NVO-Sd ‘Sd “AVW-Ld 
‘OZV-Ld “Td ‘ON “IN 
“LW ‘OIS<LI “UVS-LI “LI 
UW) “Ad UOO-Ud “Us ‘TA 
Md “ZO ‘Od “Ad ‘Ad “LV 

umouyuy) 

ZO ‘0061 

dD ‘8eol 

jeordonqns 
‘qeotdosy, 

POLIOWY 

SRO 

jeordoay, 
-vIsy 

SNOJOA 
-1nap 
‘snoseyd 
-odyd 

SNOJOA 
-1nap 
‘snoseyd 
-ovdyd 

SNOJOA 
-lnap 
‘snoseyd 
-o1dyd 

POST ‘AeLIN 
sodau snjtqdoduv’y 

eVsl 

‘UOSYDIIT SMqv]LINU 

smppydoday) 

8581 ‘AYspNYosioy] 

SMPJOUIBADUL 

smppydodayy) 

(0007) FOI 

pue opeyseyy “(€661) OlsIpny 
(900Z) ‘Je 19 sIOquaNIT\\, 
‘(600Z) ACW “(S00Z) 
eYAMISL'] Pure PADIS ‘(800Z) 
OIsIpny pue pnsytV “(600Z) 
sod10g pue esuopusyy “(S00Z) 
‘Te 19 saB10g “(€661) OISIPHY 

SIDUDIIFIY 

soleus 
‘symmay Ap ‘ozreul 

suroosYysnur 

‘punois uo siimdy 

‘soreuess UT 
speosso ‘sqmmay AIp 
‘sadei3 Surkeoop 

veNqeH 

NVO-Sd “UD “AC “Ud “WH 

HO" Sa “OZV 

“Ld “Td “LI LT “OISsLI 
YVS:LI ‘LI “AG “YOO 
UWA “ZO “Od ‘Ad CLV “TV 

S31JJUNOD popeAauy 

LI ‘0081 

adoimy ut 
prosel IST 

POLIOUIY 

SO 

Jeordoay, 
-eISy 

SNOJOA 
-lMap 
‘snoseyd 
-odyd 

SNOJOA 
-1nap 
‘snoseyd 
-ovdyd 

snjejsg 

POST ‘AeLIN\ 

snausy smpiqgdodiv) 

(8SZI ‘snoeuurT) 
snaaqqiuay 

smpiydodayy) 

satoads 
Ayrurey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

370 

(£661) OlsIpny 

sums ‘saTqriasoa 
‘orepoid 

9t99q YIeG 

HO “ad 

ad ‘ZZ61 

POLIOWY 

YON 

Joyepaid 
orisered 
‘snoseyd 

-ovdyd 

(0621 
‘JOIATO) suqvisuf 

S nyt Yo OLGIS 7) 

(elpea] UT uate Looayod “wey 
‘(QOO0Z) OlsIpny pur pnsfIpy 
‘(OO0Z) FHIOIG) puke opeyprjy 
(6007) AOWO]T, “eryeay UT TuoTye 
IaNOJTO7) “13eyY ‘(800T) OCISIPNV 
pure pnsfIVv “(000Z) HOTS 

pure opeyseyy “(€661) Olsipny 

simay Arp ‘Oorueul 
‘snoseydoosur 

suroosysnur 
‘(Snuna) SYMIF 

HO ‘NVO°SA ‘Sd ‘CW 
‘OIS:LI “LI “AC “Ud CLV “TV 

NvO'Sd 
‘AVW-Ld “CW “LW ‘OIS 
SELEY SSE CLP as “Ty 

LI ‘0061 

jeordoay, 
-vIsy 

POLIOUY 

S20 

SNOJOA 
-1nap 

snoJOA 
-lnap 

6Y8I ‘reUrTe,y 
stapjnz0 vavandy 

EPRI “UosySIIA 
yjoam vavandy 

(‘elyeay UT TuoTye 

TaNOJjO7) “Wey “(€G66[) OlsIpny 

(SOOT) ‘Je 39 sesi0g 

ozreul 

Sd ‘Ld “DIS<LI “LI “Ud “TV 

OZV-1d 

POLIOWYy 

SRO 

SNOJOA 
-lunap 
‘snoseyd 
-o1dyd 
SNOJOA 
-lnap 
‘snoseyd 
-ovdyd 

6961 

‘uosqoq smzqdvaz 

smppydodavy) 

EPS] “UosyoY 

snstaans srqiqdodiv’y 

(£661) OISIpny 
(S007) eyANIse’T 

pure PAodjag ‘(Q00Z) OISIPNY 
pue pnsfl “(€661) OSIPry 

SIDUIIIFJIY 

sunp Anjnod 
‘punois uo 

sumsy ‘symnay AIp 
(uIO09 

‘OzTeUl) SoTIeURIS 
‘s1oopino 

siMsj Ud}01 

vege 

IS ‘SU 

‘OZV-Ld ‘OIS-LI “UVS 
SLIGL Md Zo) dey 
‘SA ‘Ld “LW “OIS-LI 

UVSsLI LI “AUYO-UD “UD 
WOO Ud Ud “Md “ZO ‘AO 

S$931JJUNOD poprAuy 

ZO “€861 

adomy ut 
prosel IST 

Jeordoay, 
-vISy 

Jeordoay, 
-eIsy 

SNOJOA 
-lnap 
‘snoseyd 
-ovdyd 
SNOJOA 
-1nap 
‘snoseyd 
-ovdyd 

snjyejg 

8581 ‘Ayspnyosioy] 

snpjasoid 
smppydodavy) 

eVsl 

“UOSYSIIY $212]0SgG0 

smppydodiy) 

satoads 
Ayrurey 


371 

Coleoptera families other than Cerambycidae, Curculionidae... 

(6007) AotuOL, (Q00Z) FOI 

pure opeypeyl “(€661) OSIpny 

(eqpeay uy 
Tule Wonosjod “mMey (6007) 
sod10g pue eSuopusyyl “(S00Z) 

Te 39 so8s0g “(€661) OFSIPAW 

(6007) sasiog pure 

eSuopueyy “(S00Z) ‘Te 39 sesI0g 

(6007) 
so81og pue vsuopusyy “(C00Z) 

"Te 19 sad10g “(€66T) OlsIpny 

(6007) 
so81og pue vsuopusy “(C00Z) 

‘Te 19 sassog “(€G6]) OIsIPHY 

(8007) 
OIsIpny pue pnsyIV “(600Z) 
so810g pue eSuopucyyy “(S00Z) 

Te 39 sed10g “(€66T) OlSIPNY 

SaTquiodoA 

pur sama Arp 
siMyy JoyIO 

pure sorsoqmens 
‘yyeq yeo Jopun 
soqsayye's 1OOsuT UT 
symsj SUIMIOI 
pure Surdesop 

UOLIIED ‘sIOARPEd 

deFINS [fos oy 
uo Ye souoqg PI° 

Wn ‘NVO-Sa “TVE-Sa 
‘Sa ‘IS ‘Su ‘AN ‘AVN 
“Ld ‘Ld “AW ‘LW ‘OIS-LI 
VSL ‘LI ‘DASAD 
‘ANAAD ‘NOD “AYO 
AD UD WOO UI 
‘kO YH ‘Dd ‘WE ‘LV “IV 

HO ‘NVO 
-Sd ‘IS ‘OZV-Ld ‘LI “Ws 

OZV-Ld ‘LW 

LI ‘9Z61 

LI ‘0061 

OZV-Ld 
“SOOT 

Jeordosy, 
-vISy 

POLIOWY 

SO 

POLY 

SNOJOA 
-lnap 
jorepoid 
jorisered 
‘snoseyd 
-odyd 
SNOJOA 
-ninap 

SNOIOA 
-ninap 

SNOIJOA 

-ninap 

snoJOA 
-lnap 

(86Z1 ‘sntorge.) 
sypsauuny snsogdos) 

(Sz8I ‘fes) 
VIVUIULAD VIOPIIAIS 

(1S8I ‘uUeUayog) 
SYVIgu vYyoUuad 

(CZLI ‘sntotiqe.q) 
VaPlOIstp VILSOULC) 

(9¢Z| ‘snoeuurT) 
U0]0I VIISOUC) 

(€8ZT FeTTeYDS) 
DIADULDI DINPUINT 

(8007) SSIpRYy PUe Pas 

‘(800Z) HNC ‘(€661) OISIPNy 
(900Z) ‘Je 39 BI9quaNIA\ ‘(600Z) 

AOUWIOT ‘(CQOZ) PYANAseT pur 

PAOIJIS ‘(CQOT) JoWIoy “eIyeI UT 

TuaTe Wono0sjod “Mey (6007) 

sasiog pur eSuopuayy ‘(SQ0Z) ‘Te 
19 MYOpusaryry “(CG6T) OSIpny 

SIDUIIIFJIY 

uayjod 
DASIU DIOLIVT 

sums ‘saTqriasoa 
‘orepoid 

apiseq yseq 

veNqeH 

aD (LW 

aD ‘Wn 

‘HO “dS ‘TS “MS ‘Su ‘NU 
‘OU “Td “AW ‘CW {LT ‘TI 
“LI ‘NH “UD “Ad “Wd “ZO 
‘UH “Od “Va ‘A ULV “TV 

SotsJUNOS poperauy 

umouyul) 

dd ‘0S6r 

adoimy ut 
prosel IST 

POLIOWY 

YON 

snoseyd 
-o1dyd 

jorepoid 
orisered 
‘snoseyd 

-o1dyd 

smq¥I1¢ 

(COST “weYysie) 
SIULOIYNA SagIasyaAy 

(S€sI 
‘Aes) snqvusistapynb 

SNJIGIOAG ISTE) 

satoads 
Ajrare,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

372 

(9002) ‘re 

1a Biaquant Ay ‘(POOT) Uosuyo[ 
pue uossuaios “ele UT TuSTye 
Noo) “WEY “(000Z) WoO 
pur oper “(8007) H"d 

(4007) wosuyof 

pure uossuszos ‘(8007) Hd 
(9007) ‘Te 2° 

BIoquat\ ‘(FOOZ) UOsuYoOf pure 
UWOSSUIIOS “(QOOZ) SPIOUIWIGT pur 
preesapQH ‘(600Z) sesr0g pur 
eSUOpUs| ‘(68G61) ‘Te 39 epnasy 

ssourisqns sTuesI10 

Sumo “soduroo 
‘sunp ‘sie1uqey 
otuasodoryiue 

[eloreur sTuesI0 

SUNOI Id] 
‘SUIAOU ssei3 

MOAISUNY 

dD 
‘HO ‘NVO°S8 ‘Ld ‘LI “Ud 

€D “HO “dS ‘GVW 

dD ‘Z861 

POLY 

POLIOUIY 

YON 

SNOJOA 
-lnap 

snoJOA 
-lnap 

TL6[ ‘Uosuyof 
IVUAaGYaVIIJUDS 

SIYITAIOAIYT 

(ZZ8T ‘smoyuey) 
14dasol st I1AJOLIY 

“(LZ61) ‘Je 29 epnely ‘(800Z) ‘snoseydordes -Ld ‘OZV-Ld ‘ON “IN “AI POLIOUTY SNOIOA (ZS8I ‘UIPE) 

BN ‘(S00Z) ‘Te #9 sas10g ‘asoduros “Ad Ud ‘Id IC ‘ZO ‘LV qVIon “HNP SLADINSUL SIYILATOLIY 
(FOOT) UosuYyo{ puke UOssuaIOS 

‘(€QOT) JOWPI2y ‘(TZ6T) POLOWYy SNOIOA (TZ8T ‘smoyeyyl) 

‘Te 39 opnozyz “(8007) Hd asoduros €D “AS ‘ON “IN “AC “AC| €5D ‘9961 GuON ueele PIPL DEP LOL Fe 

seprypd 

(600Z) s2s10g pue POLIOUY snoseyd PPS] JOWIOYspyl 

eSuopuayy ‘(GQOZ) ‘Te 19 sasiog UIONIIIMS I OZV-Ld| UMouyuA YON -ouyd| oy sngyod SnLIVIVY J 

oeploryeyg 

(800Z) YNYIS pur JPY 

SIDUDIIFIY 

SIO} UIIOATI UT 

elaidoajo7) Suls0q 
-poom Jo Joyepaid 

veNqeH 

S31JJUNOD poprAuy 

LV *Z007 

adoimy ut 
proses 1sT 

POLIOUY 

YON 

jorepaid 
/orisered 

snjejg 

(86ZT ‘sniorsqe.q) 
snf{ns snuasoqv’) 

oeplipursse 

satoads 
Ayrurey 


DTS 

Coleoptera families other than Cerambycidae, Curculionidae... 

(900Z)|  sauownsedde ut 
Te 39 Stoquanyy, “(900Z) UUEY | stueld ‘esnoqusars oorf SNOIOA L761 ‘uIdey 
‘(Z661) puewaly pur suapoqy Ul VUavIWIG gill HO ‘IS ‘LI “Ud) LI ‘TZ61 eOLyy -unep| oy VIIIOXA VIMIIVPO)L 
auprAoepornd 
SNOIOA Cgq] “UoueITY 
SAU EN ee see dS ‘ON ‘Td vIsy “lop suosugol Maury 
POLIOWY SNnOIOA (9961 ‘seqkq) 
COTE) MOSEL BYE BORER cand Rasa) ad YON “nop vasnf nivquivg 

(F007) yorrepury 
(FOOT) Uosuyo[ 
pure uossuszos ‘(8007) Hd 

OI] ISdIOF 
$9011 peop Jo 
yreq 1y3n Jopun 

dD ‘L961 

jeordonqns 
‘estdosy, 

eIseperisny 

SNOJOA 
-lnap 
SNOJOA 
-lnap 

(9901 ‘seqdq) 
VIAOJUOI DADQUDG 

ZL6| ‘Uosuyof 
IVLONDY VIJGUMT 

(FOOT) UOsuYoO[ puke UOssuaIOS 

(joodzaarq 
‘uopuoT ‘3°9) 
SOTIID [eISeOd 
adie] punose 
seare popoom 
ur ssunano 
sseio ul deay 

dD ‘6761 

vIseyersny 

SNOJOA 
-lnap 

(S281 ‘smoyzaeyy) 
1uU0suts DIU 

(F007) 

uosuyof pur uossuaIgs ‘(€00Z) 

JoWINyY ‘(686[) ‘Je 39 epnoiy 

S90I] Prop 

Jo saroads jsour Jo 

yreq 1y3n Jopun 

dD “IN “AI “Ad 

AD “S76I 

vIseperisny 

SNOIOA 
-lnap 

CL6[ ‘uosuyo[ 
UPUNgvLLa DIAUL 

(FOOT) UosuYyof pue UossusIOS 
(F007) UOsuYo{ pue UOssudIOS 
“elypel] UT TUaTye 1191103]07) 
TNeY ‘(0OOT) SeIOUTUTGT, pue 
preedop@ (6861) ‘Je 39 apnary 

SIDUDIIFIY 

SIOJTUOS 

pue SOAPO[-PEOIG 

Peep Josie 
1ysn Jopun 

SIOAIBUNY 
‘snoseydosdes 

“soduroo 

veNqeH 

dS “MS “ON ‘AT ‘NH 
“Ad ‘Id “MC “ZO ‘Ad “LV 

S311JUNOD popeAuy 

AD “9€6I 

Id ‘YZ61 

adoimy ut 
prosel IST 

eIseyersny 

SNOIOA 
-lnap 

SNOIOA 
-lnap 

snjyejsg 

(€68T “unoIg) 
YJaays VIJaUly] 

(PERT ‘smoyeyyl) 
voiuodvl vavasoavg 

satoads 
Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

374 

(Z00Z) HPL ‘(6007) se810g pue 
eSuopuayy “(000Z) OID pur 
opeyseyy ‘(C00Z) ‘Te 19 sed10g 

(9002) ‘Te 39 B9quoat\ “(6007) 

AOWOT, ‘(CO0T) eyAnisey pure 
BAQoS (O00) SEPULO Pur 
preesopQ ‘(600Z) sesi0g pur 
eSuopuayy ‘(GQOZ) ‘Te 19 sasiog 

(SOOZ) ‘Je 39 sesi0g 

(C861) pnesreg 
pur ueipneg ‘(600Z) sed10g pure 

jorepoid 
PAIR] ‘SIOABPLD 
‘yreq ‘syuejd peop 

sseueue ‘eurueq 

Peg 
ur aseydormayo 

“soduroo 
‘sjonpoid pos0is 

-snoseydoides 

SISOIOJ 
UI s]saU JUIPOI 
‘suTOOTYysnul 
poreanns 
aseurep ‘Anjnod 
‘sayqeis ‘one 

:orrydodosyiue 

snonploop 

NVOS4d 
‘OZV-Ld “IN “AC ‘NC “Ad 

HO “AS ‘OZV 
“Ld “Td (LW “LT “AC ‘Td 
“Ad “MC “ZO ‘Od ‘Ad “LV 

Ad ‘TI61 

ZO “SL81 

OZV 
Ld “S007 

jeordoriqns 
‘estdosy, 

POLIOWY 

SRO 

jorepoid 
/orisered 
‘SnOJOA 
-1n0p 

SNOIOA 
-lnap 

SNOIOA 
-lnap 

snoseyd 

(PHS I ‘aT[AoUsV] 

-ulsgn)) zsuzpavlsap 

vy diowmdhay 

(TEST ‘PIEA\) 
DUudalpv Si NAIASDY 

SepIUeATIS 

(7equeT}AD) 

SNIUUNAG snuajsy 

ovpisurdyes 

IPS] URWIMAN 

eSuopuayy ‘(CGQOZ) ‘Je 19 sasiog snoseyddjod -ovdyd vaiuodul vytdog 
td ce Seg A 

908 ‘sloqunyy, 

(6961) ‘Je 2° opnoiy oidoryueuds joyepaid stusootuioed 

‘(100Z) UNE “(ZIGI) SH9q| ‘Plousesred wez07q f IN ‘LI ‘Td “MC ) Usouuy eisy|  jomusered| oy snipidigy 
seprioydidny 

snoIOA P61 Wd 

asnoyusals ACTECEL -lnop sadiaqny YIIVPOTIU 

adoimy ut satoads 

S9UIIIFOY WwIqey SOIUNOD poprauy proses IST snyei¢ Ayrare,y 


BRD 

Coleoptera families other than Cerambycidae, Curculionidae... 

(900Z) ‘Te snosoarsuny (€€G] “UOIOWeD)) 

a) sriaquait AX ‘(000Z) SeIOUTWOT, ‘orepoid HO ioyepoid vsauagopnasd 
pue preesopQ “(600Z) ‘Te 39 BYNT ‘asoduros I) “AS ‘ON “AC ‘Td “MC ‘LV]| Id ‘8861 eisy|  jomusered| oy puojoly 
oeprurpAydesg 

(‘eIpeI] UT TuaTye TaNOsTOD “NIeY 

(‘elyeay UT Tuorye 

Tanogpo*) "Wey “(Z007) Wey 
(‘elyeay UT TuaTye 1191109707) 
Mey “(600Z) sasrog pue 
eSuopusyy “(S00Z) ‘[e 19 ses10g 

(900) ‘Te 39 Broquaqp\ 

“(GO0OZ) ACWOT, “(GQOT) ByANAse’T 
Pur PAOas (6007) S9810g 
pue esuopueyy ‘(000Z) MWoIO 

P[Py Ur soar peap 
jo yreq Jopun 
‘sionpoid posois 
‘SOTUPLT ‘OTT 

OIS<LI ‘UVS-LI 

HO ‘NVO 
“Sd ‘SU “OZViLd ‘Ld “ON 

uMouyuy) 

uMouUxU al 

uMOUxU ial 

POLY 

vIseyesIsny 

SNOIOA 
-lnap 

SNOIOA 

-1nap 

snoJOA 
-1nap 

9Z8T Tuy 

SIOIIIAL SNUBAIIS 

9Z8T Tuy 

is1may snuBATLIS 

(O88I ‘unolg) 
SNYIAAV] SAUBATLS 

(8SZI ‘snovuurT) 

pure opeypeyy ‘(SQ0Z) ‘Je19| — sxonpord posois TA TAT A ae “Ya snoJOA SISUIUDULANS 
oppopuraryy “(S00Z) ‘fe 39 sessog| ——_‘a8eydoypdsd “Ad “NC ‘ZO ‘Da ‘AM SLV| Ld ‘¥681 “HEAP snpny dav2tC) 
(900Z) ‘Je 39 S1oquai 
‘(600Z) AOWOL, “(S00Z) PyANase’y] 
pur vaosyag “(600Z) SeB1og pue HO ‘NVO-Sd ‘OZV (6881 
eSuopusy ‘(000Z) FWOIC pure sjonpoid poesois -Ld ‘Ld ‘ON ‘LW ‘AT ‘SNH jeordonqns snOIOA ‘PPANV]) 4070 I4IUL 
opeypeyy “(G00Z) ‘Te 39 sedI0g ‘oseydoypdsd “AA Nd ‘ZO ‘Dd AP ‘LV] ZO ‘7961) — “feordory, “Hnep snjrq daveic) 
OSGI ‘peosyey 
SPoos VIGIVIIPVZV sNOJOA SNIDUIULNID 
(8007) Hd ‘ynwoes09 AD “0861 “Ainop sryay Pavziuc) 
(Z00Z) JHNODUO| (FELT “UuEpPsny) 
“(Z00Z) 23eY ‘(600Z) sed10g pue SnoJOA SMUAOIIADII 
eSuopueyy “(SQ0Z) ‘Te 19 ses10g | —_ sionpord porois OZV-Ld ‘Usd “AC} Us “9061 72 P. SHULQISEENT 
adoimy ut satoads 
SdUIIIFOY WwIqey SoLNUNOD poprauy psoas IST snyei¢ Ayrare,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

376 

(9007) SOULIP, yesl 
yonbuoiy ‘(600Z) ses10g pure pues ‘sayored OZV-Ld o1uas snoseyd ‘susydars suqvauyig 
eSuopuayy ‘(CQOZ) ‘Te 19 sasiog Jeaseoo Asseis| ¢y ‘Tq OZV-Ld ‘SOOT -o1dA17 -ouyd| snuyadiv’y) 
(IZG6] ‘uorsureD) 
(9007) 2onbuogy, PD) AS‘ON| Ad‘ o1ua3) = joyepoid stuuaduny, 
(0007) SEIU], pur preESopQ soduro| CF ‘TG| “AG ‘UA ‘TANG ‘AG ‘TV) = ‘TH6T) «= ordi} ppptsered) pugyaquagog 
(9002) 3onbuogy, 
“ered, UT TuaTTe TaNo0ajorD “IeY 
‘(000Z) seFouTUNG], pure preesapy dD 
‘(6002) ‘Te 9 PNT “(S007) ‘HO “AS ‘NVO-Sd ‘ON ‘LI AC| eaeseduray,| — roxepord (pZ81 ‘dreys) 
23105 (8007) HMC “(800Z) CYD| orepasd asoduros| = of 1] “AC “Ud ‘Td “MC SLV “Tv) ‘Id ‘0S61 -eIsy|  jorysered| oy snag sniustgy 
ayerq Soyer) vououry|  sorepord (CS6I ‘euRIOUIS) 
satoodg rururpAydeag “UOIMINT Of ‘J JIS-LI {LI “ZD “TV | Usouyuy YON orsered V mupod SnIUSI 
(SOOZ) ‘Te 19 OZUO/y SIAOIS SNIUO Jeordoniqns joyepoid (6S8I ‘zaeery) 
‘(SOOZ) Cfe79INE pur vireurery ‘s[e1999A Surkesap ‘al Sa ‘LI) Sa ‘7007 ‘eotdosy, orisered V VIVUOLINUL VIIGIP 

(6007) 

sodiog pue eSuopueyy (0007) (8681 
TUIOIC) pue opeyoeyy ‘(C00T7) jorepoid ‘AYS[NYISIOY[) 
‘Te 39 soBI0g ‘(06G61) S9810g NVO-Sd ‘OZViLd “LI “TV| LI ‘S661 | sy WrFy) — /oprsered Stuuadiinpp viaqay 
(6002) se810g pue 
eSuOpus| ‘(000Z) FHWOIC pure (sJo17e5) orua3 joyepoid (TZ81 “WorseTfoA\ ) 
opeypeyy ‘(SQ0Z) ‘Je 19 sasiog| vag uo s01epeid NVO-Sd ‘OZV-Ld | umouyuy -oidAry| = onsered suodfipyyiu snygouy 
sa[qras UT 
(600Z)| (sayy) ex2xdiq 
sosiog pur eSuopuapy ‘(hZ61)| snoydeyss0p>A9 o1uas jorepoid CCQ] ‘snpy 
‘Je 19 apna ‘(CQOZ) ‘Te 29 ses10g jo soepaid OZV-Ld ‘LV| umouyug | -oiddrg| —ontsesed vynsagnd pavgs0ay 

sory 
joyepoid “poomeas 
Sutsodurosep 

(8007) Ha 

SIDUDIIFIY 

veNqeH 

S311JUNOD poprAuy 

umouxuy) 

adomy ut 
prosel IST 

POLIOWIY 

YON 

jorepoid 
orisered 

snjejsg 

EVR] “Wisysouuryy 
DULINLADUL DIOP 

satoads 
Ayrurey 


DIT. 

Coleoptera families other than Cerambycidae, Curculionidae... 

(9007) ‘Te 19 Br9quait 

(‘eI[eI] UT Ture 193109[07) "3eyY 
(600Z) ses10g pue 

eSuopusy ‘(000Z) FWOIC pure 
opeypeW “(S00Z) ‘Te 19 se810g 
(9007) 

yonbuoly ‘(600Z) ses10g pue 
eSUOpUs|| ‘(000Z) HWOIG_ pue 
opeysryy ‘(GOOZ) ‘Te 19 sed10g 

(600Z) ‘[e 39 RANT 
‘(C00Z) 28I0yf (E007) UeddnD 

(Z00Z) ‘Te 39 3SIOA, ‘(8007) 
BN ‘(S00Z) ‘Te #9 sas10g 

(6007) ses10g pue 
eSuopuayy ‘(GQOZ) ‘Te 19 sasiog 

(9002) 
yonbuoly *(GQ0Z) ‘Je 39 sesi0g 

(9002) 39nbuoyy, *(S007) 
‘Te 19 sa810g ‘((966[) ses10g 

SIDUDIIFIY 

Sunp ope9 

syueq Apues 
sayoeoq 

pues ‘syueq JOAl 
‘sute|dpooy 
sayoeoq 

pues ‘syueq JOAI 
‘sure|dpooy 
sayoeoq 

pues ‘syueq JOAl 

‘sure|dpooy 

sayoraq 
pues ‘syueq JOAI 

‘sure;dpooy 

vege 

HO 

NVO 
“SH “AVW-Ld ‘OZV-Ld 

NVO‘SH ‘OZV-Ld 

€D “HO “aS “AC “ad “LV 

$31JJUNOD poprAuy 

umouxuy) 

LI ‘7861 

uMmouyuy) 

adomy ut 
proser IsT 

vIsSy 

vOLIOWY 

SRO 

POLIOUIY 

YON 

vIseyessny 

jorepoid 
/orsered 

umMouyun 

umMouyun 

jorepoid 
orisered 

uMouxyuUn 

umMouyun 

jorepoid 
orisered 

joyepoid 
orisered 

snoseyd 
-ovdyd 

snjejsg 

(L9G6T ‘ouueef) 
uuuas snqsaqdoqdaT 

L861 

‘a0vq sesuadnippyns 
VIOISIIC 

(GE8I ‘Uosypity) 
snyaqgaynd 
snsodoadoy 

(Z9Z1 ‘shoeuurT) 
sapiogdyis vat’) 
(0061 

‘dreys) snaipuvqpaz 
snuyadivy) 

(GE8T “Uosysi4y) 
sylaqns snuyadavy) 
(ZO8T 
‘JSIOYUIARIL)) 
snyisnd snupadivy 
(O€81 

“UTS YIOUUPY[) 
SIIIVAS SNULY AULD’) 
(9081 
“‘ISIOYUIARIL)) 
SnUutIt440I 
snuyaduvy) 

satoads 
Ayrurey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

378 

(900Z) ‘Te 29 S1oquant Ay, 
“(SOOZ) ByANIseT pue PAOTJOS 

“elyely UT TUdTTe 119OJTOT) “111eyY snoseydoides HD “AS ‘ON ‘NT ‘IT SLI SNOIOA FOG] ‘Paney 
‘(600Z) ‘Te 39 EAN “(C007) e810yy ‘asoduroa Of ‘I) “Ad Ud ‘Nd *ZO “Ad ‘LV | MC “SZ61 BIsy TSP; OV sogvadtur snjaytxC 
(9007) ‘Je 39 S1oquaiu 
‘(€00Z) Fwy “(000Z) add 
SpIOUNLU], pur preesapg ‘(600Z) ‘HO “AS ‘NVO-Sa ‘AVN 
sodrog pue vsuopusyy “(000Z) oidoryiueudg “Ld ‘OZViLd “Td ‘ON 
UOIC pue opeyseyy “(600Z) ‘Te ‘snoroarsuny “IN ‘DIS EE LT Pao 
19 EAN “(GOOZ) AS10y “(FZ6T) ‘Torepord UW) “AC ‘UOO-Ud “Us ‘TA BOLOUNYy SHOTOR COSI “ZPD 
‘Te 39 apna ‘(CQOZ) ‘Te 39 sad10g asoduios) of I] “Ad SIC ‘UH ‘Vd “AD ‘LV| Ua ‘8S81 SRO “BEep| Vy vated vI0340) 
(Zc81 
(9007) 2onbuosy, ‘AyspNYyOsIOY[) 
“(€O0Z) 98x “(600Z) ses10g pue OZV-Ld (ctIsy syportssanduun 
eSuopuayy (S007) ‘Te 19 sass0g Cl OLE ZS ‘CoOT| 40) eONyy| UMouyuN| Vy snawaun] 
6061 
(ZZ61) syueyd jorepoid ‘oping sadiaasq 
zyjad1s2y49¢$ ‘(Z661) UOsJopuy | Jopun purs 13M UT qD “Al “Uy | uMouyuyp orisered viodorauAtpyy 
(9007) JonbuoLy:, (OF8 | ‘UosyoITy) 
‘(Q00Z) HUOIG pue opeyoepy gD “AS ‘NVO-SA ‘AY SNOJOA puu1guos 
(G00Z) 28101 “(800Z) HNC poompeap ‘AVW:iLd ‘OZV+Ld “Ad | Ad ‘0261 “BEnep sappy dasosauLaly 

(900Z) Jenbuogy, 

‘(CQOZ) BYANAse’] pure eAaosjas 
‘(000Z) SEIOWUIG], pure prevSapgy 
(6002) ‘Te 19 BINT “(S00T) 2810 
‘(H9G61) ‘Te 19 epnasy (8007) 
HNq ‘(S00Z) ‘Je 39 sesi0g 

‘lorepaid 
“soduroo 

SIDUIIIFJIY 

veNqeH 

€D “WN “HO “dS ‘Sa 

MS ‘OZV-Ld “Td “ON “IN 
‘AT (LI ‘NH “AC “Ud ‘TA 
“Ad “MC “ZO “Ad “LV “TV 

$91JJUNOD popeAauy 

ZO ‘TI6I1 

adomy ut 
prosel IST 

Jeordoay, 
-vIsy 

jorepoid 
/orsered 

snjyejsg 

(6681 ‘aeery) 
SIIABIU SIAVYIOYIT 

satoads 
Ayrure,y 


6 78) 

Coleoptera families other than Cerambycidae, Curculionidae... 

(0661) TP49sMy “(8007) HG 

snoyrydoyey 

umouyuy) 

vIseyerisny 

SNOJOALNOp 
‘orepoid 
orisered 

(0061 ‘dreys) 
Lojonun sndjvdosay, 

(9002) ‘Te 

qo Blaquait\\ ‘(900Z) Ionbuosy, 
“ered UT TuaTTe aNo0ajoD “"IeY 

snosoarsuny 

HO 

‘(000Z) SPIOWWIG] pur preesopy ‘orepoid “AS ‘NVO-Sd ‘ON ‘LI “Ad POLIO F861 ‘asyo'_| 
“(600Z) ‘Te 39 BINT ‘(S00Z) 28104] ‘asoduros Wd “Md “ZO “Ad ‘LV “TV| AC ‘SZ61 YON | uMouyun VIVABIULUAT DSNIGIIAT 
888i ‘dreys 
Ty| usouyuy visy| uMouyUN SNIIAIGIS SNUIYIVT, 
(9002) JenbuolL, “(6007Z) AowOL, 
‘(SOOT) BYANIse'T pur PAOIJIS SNOIOATIOp 
“elyely UT TUaTye 1191103]07) Mey SIOARPeO lie) fleece kL ‘orepoid PL8I ‘dreys 
‘(600Z) Te29 PANT (C661) OPED} = “oT eyqeas ut Wd Md °ZO ‘Dd “LV “TV] LI ‘0861 jorsered sadiuads snyquojigg 
(9007) ‘Te 39 Sioquont pA (9007) €D “VN “HO “aS “NVOD 
ronbuosy, “(600Z) AOWOL, (S007) -SH ‘Sd ‘IS “MS ‘Su ‘OU 
YATE] puUE PATA “(000Z) ‘GVW-Ld ‘OZV-Ld ‘Ld 
SpIOUMUIG], pure preesapy “((00Z) ‘ON “IN “AW GW ‘NT LT 
JUIOIC) pue opeypeyy “(6002) ‘Te ‘AT UVS“LI LI ‘NH “UD yL8I 
19 PANT “(C00TZ) 2810 “(ZZ61) “Ad ‘Ud ‘Id “Aa “Md “ZO orerodurai| — Jorepard ‘dreyg snjnduviza 
WEHIOD “(SO0Z) ‘Te 39 sed10g | 1orepard ‘asoduros UH ‘Dd ‘VE “AG ‘LV IV] LI ‘0761 -eIsy| — /otasered SnyqUOHIG 
(9007) ‘Je 39 S19quai)\ 
‘(900Z) Jonbuosy “eryeay UT rare speqadaa 
HWonosjor "mIeY ‘(00OT) HHOIE, Surkeoap ‘soon (LZ81 ‘Akeay 
pure opeyoryy ‘(6007) ‘Je 39 PYNT]  smoryea Fo simsz HD ‘NVO-SH SnoJOA ORA)) sesuagupulys 
‘(C00Z) 28103, (8007) NAG wares Sunios ‘Sd ‘Ld ‘OISsLI ‘LI “AG “UdA] LI ‘8861 ) esepensny “Tep vgusoaojy dna 
adoimy ut satoads 
SIDUDIIFIY WeIQGey S31JJUNOD poprAuy pr0ser 1ST snqeI1¢ Ayrore,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

380 

(2661) PHY 
pure rysoyiouuepy ‘(FO0Z) Joe 

sdeay D1SeM 

AS ‘8861 

POLIOUy 

SRO 

SNOIOA 
-lnap 

OZ8T ‘UIOH 
snssasdap snavul’) 

(2007) 

Heplos “(900Z) JoyfeMmeyos pue 
ZUUQIOY “(TOOT) UOssiopuy 
pure JoqoJ (F007) Jou 

(9007) ‘Te 39 S1quart\ “(600Z) 
AOUWIOT “(Q00T) SeIAWIWUIGT pur 
preesopQ ‘(6961) ‘Je 19 epnary 

(O00Z) MUOIG_ pure opeyrpy 
“(6961) ‘Te 3° 9pnaxg “(8002) 
Hnq ‘(S00Z) ‘Je 39 sasiog 

(9007) 

‘Je 19 S1aquaii\\ “(600Z) AOWOT, 

‘(6961) ‘Te 39 epnesy “(8007) 
BN ‘(S007) ‘Te 9 sas10g 

SIDUIIIJOY 

sdeay o1sem 
‘snoseydoides 
sduin3s 

pyo yzeq Jopun 
‘SUMIJ UIIOI UT 
onsswop AyuTey 
sodurod ‘sanpIsai 
jo asad sourur 
syuns} UT 

Isunj UO sIoopyno 
‘sionpoid posois 
‘SONPISaI JO 

sad Jourw 

sisou 1eq 

/pliq pure syunsy 
ua]0J UT sIoopyno 
‘solsem pure 

sadoej UO spaoj 
‘sasnoy UdyxTY 
jo ueIGeyUT 
UOWIUWIOD ‘sonpIsas 

jo qsod sourur 

vege 

as “Ad ‘Ud ‘Td 

HO “AS ‘OU 
‘ON ‘IT ‘NH ‘UD “Ad “Wd 
‘Id “NC “UH “Dd “LV “TV 

ae) 
‘NVO-SA “LI “Ud “Ad “Nd 

aD “HO 

‘NVO-SH ‘ON “AW OLIN 
ITAL AH AG aed 
Ud Ud “Ad “Nd “Od ‘LV 

SoTsJUNOS poprauy 

AS ‘8861 

Dd ‘OF6! 

uMOUxU ial 

AW ‘T76I 

adoimy ut 
proses IST 

POLIOWY 

SO 

jeordonqns 
‘yestdosy, 

jeordonqns 
‘eotdory, 

SNOIOA 
-lnap 

SNOIOA 
-1nap 

snoJOA 
-1nap 

SNOJOALNOP 
‘orepoid 
orisered 

snjyejg 

(LC8I ‘eIU099T) 
sngsnduv snavul’y 

(€Z81 
‘kes) snquiaspfig 

sn3pydongdp 

(I8ZI ‘snisisqe.y) 
snqve1avy 

smqongdy 

(L6Z1 

‘Jozue) snutsadvip 
smqongdy 

seplmuoligouey, 

satoads 
Ayre. 


Coleoptera families other than Cerambycidae, Curculionidae... 

(9007) 

Je 19 Biaquout\ ‘(600Z) ACWOT, 

‘(CQOZ) BYANAse’] pue eaosjas 
‘(6007) sasiog pue eSuopuayy 
‘(O00Z) FHIOIC) pue opeypreypy 
‘(6961) ‘Je 29 epnany “(300Z) 
BN ‘(S00Z) ‘Te 39 sass0g 
(6007) ACWOT, 

< (S00Z) exANIseT pure PAOIJIS 
‘(O00Z) HIOIC) pue opeyprypy 
‘(6961) ‘Je 29 epnany “(300Z) 
Hnq (S007) ‘Je 39 sadsog 
(000Z) TOAD pue opeysey 
‘(6961) ‘Je 29 epnagy “(300Z) 
HN ‘(S00Z) ‘Te 39 sass0og 

(S00Z) ByAnase’y] pur eaowyas 

sjonposd paroas 

snoseydoodur 
‘spear ATUTRUT 
‘sjonpoid posois 
snoseydoodkur 
‘spear ATUTeUT 
‘sjonpoid posois 

cf ‘tf 

tH) 

‘NVO-SH ‘OU “OZV-Ld ‘Ld 
‘ON “AW CLW ‘IT ‘AT ‘OH 
UD “AG UOO- Ud “UA 

“dd “Md “ZO “Dd “LV “IV 

dD 
‘NVO-S4 ‘OZV-Ld ‘LW 
UD WA Md *ZO ‘UH ‘Dg 

aD ‘NVO 
“Sd “IT UD “Ud “Id “UH 

AW UD Ud “ZO UH 

ZO ‘0061 

IT ‘9Z61 

ZO ‘VEOI 

o1uas 

-o1dA17y 

vOLyy 

POLY 

SNOIOA 
-lnap 

SNOJOA 
-lunap 

snOIOA 
-1nap 
SNOJOA 
-lnap 

(L6L1 

‘\sqia}) Wnauvisys 

wengyoqid], 

(POST “UOIsETTON\) 
snssandapqns snso]0q 

(SPg] ‘UURUISST/\) 
1BANGAZIVL SNLOJYT 

(9C8T ‘ourewuTe.J) 
oypkydvaiag vigde] 

(900Z) ‘Je 39 S1oquai 
‘(6007) ACWOL, “(S007) 
eyAnase'] pur PAOIas “(()00Z) 
TOC) pue opeypeyy ‘(S00Z) 
‘Te 19 OPpopuaarysy ‘(696 T) 

‘Je 19 apnoazy ‘(8007) ANG 

(6007) Aowo], 

‘(O00Z) THIOIC) pue opeypryy 
(9002) ‘Te 39 SIoquai 
“(SOOT) PYANISe'T pur PAOIJIS 
‘(600Z) sediog pue eSuopuayyy 
‘(O00Z) HIOIC) pue opeypreypy 
‘(6961) ‘Te 39 epnazy ‘(8007) 
HN ‘(S00Z) ‘Te 39 sass0g 

SIDUIIIJIY 

sasnoyorem 
UT sjeasoo 
‘sjonpoid posois 

sasnoyorem 
UI SUTeIS [edII9 

SOSNOYOIEM 

UT SUTeIS [edII9 

reuqeH 

€5 ‘HO ‘NVO 
“Sa ‘SU ‘OISLI ‘LI “Wd 
“Aa MC ‘ZO “Dd ‘LV “IV 

NVO 
“Sa (LI ‘WOO-Ud “Ud “TV 

dD ‘HO ‘NVO-Sd ‘OZV 
“Ld “LW ‘AT SOIS-LI “UVS 
SLI ‘LI “AC “Wa “Ad “ZO ‘LV 

SoTsJUNOS popeauy 

LI *ZZ61 

ZO ‘0061 

adoimy ut 
prosel IST 

POLIOWY 

$280 

SNOIOA 
-lnap 

SNOJOA 
-lnap 

SNOIOA 
-lnap 

smq¥I1¢ 

OSI ‘ssnoysIIeEA 
aVZMLO SNIMAYIVT 
(LOST ‘snioriqe.q) 

SnSOIXDUL 
SN4aIOYIVUL) 

(S6ZI 

‘snIoliqe.]) sajnu.soo 
SNLIIOGIVUL) 

satoads 
Ajrarey 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

382 

(SOOZ) ‘Je 39 sesi0g 

(S00Z) ByAnase’y] pur eaoyas 

Lvdsyp vidquvuuy 
339 Joyepoid 

sjonpoid 
Pporois ‘Io1T 
‘snoseydosou 

‘adeydoypdsd 

Le aC "ZV 

ZO “T96T 

POLY 

ioyepoid 
orisered 

SNOIOA 
-lnap 

y8sl 

TOIVTONY SMUVIIOADUL 

Se apt OG? Udy 

(Zest “Bn[y) 
snyisnd sasaqvr04doT 

sepHissoSosy, 

SNOIOA (SZLI ‘snioriqey) 

(2661) Ope°49q Sa ‘2661 | Pisepeaisny “lop SNSOLIGNS SNZLOWC) 

(POST ‘AvaTe) 

SNOJOA SNIDULADIQUS 

(L661) Ops29g of ‘If Sa} Sa ‘Z661| Pisepensny “HMep] SNBLOULC) 

depIsosy 

siod ajndar POLIOUIY SnOJOA (OLLI ‘sntoriqey) 

(S6G6I) SewOYT,| OF poo; se pasn uMOUyU) CR) -hinep o1sou svgoydo7 
(900Z) ‘Je 39 S1oquai pA 
‘(600Z) AOWOL, “(CO00T) BAnase’] 
pure eaodyag “elyei] Ur Ture 

TTONOZTOD “WEY ‘(000Z) dD ‘HO ‘NVO-Sa CC6I 

TUIOIC) pue opeyoeyy ‘(696T) ‘ON ‘U1 ‘AT SLI ‘NH “AG SnOJIOA “rerSooquaik¢) 

‘Te 19 apnary “(8007) Hd 

‘Je 19 Sraquait\\ “(600Z) ACWIOT, 

(9007) 

‘(COO7Z) BYANAseT pure eaoyas 

‘(O00Z) HHIOIC) pue opeyprypy 

‘(6961) ‘Te 39 epnezy “(8007) 
BN ‘(S00Z) ‘Te #9 sas10g 

SIDUVIIJOY 

sjonposd posois 

sjonpoid posois 

veNqeH 

da NG- ZO OEe LY Iv 

€D “HO ‘NVO 
“Sd ‘OZV-Ld ‘ON ‘11 ‘AT 
-LI‘NH “UD “AC ‘Ud “Ad 
Nd 7S WH Oa y “ly 

S91JJUNOD poprAuy 

dad ‘Z76l 

ZO ‘0061 

adomy ut 
proses 3ST 

jeordon 

POLY 

-lnap 

snoJOA 
-lnap 

snjejg 

ALOJINAISIP UNTOGIAT 

8981 
TeA np urjanboef 

unsn{uoz UNTOGIAT 

satoads 
Ayrurey 


383 

(‘eIped] UT TuaTye TaN0s[OD “NIeyY 

sasnoyusai3 

pryps0 

aD ‘Sd 
“LI “Ud “ZO “Ad “LV “TV 

LI ‘T061 

otuas 

-o1dAr7y 

umouyun 

(6S81 ‘TEA 1d 
uronbef) sag2adxauz 
SNAIULOUN T 

Coleoptera families other than Cerambycidae, Curculionidae... 

(GO0Z) AOWOT “(SOQ0T) ByANase’T 
pure evAosjas ‘(600Z) ses10g 

pure eSuopusyy ‘(CQ0Z) ‘Je 29 
oPpPPuUrar|yy “(COOZ) ‘Te 32 sa810g 

SIDUDIIFIY 

(8007) Ha 

(OOOZ) PUSHY pue ynyos 

yoquin 

Prey Ur ye 
‘sjonpoid posois 
‘SNOJOATUI®S 

‘adseydoypdsd 

veNqeH 

HO ‘Su 
‘OZV-Ld ‘Ld ‘IT ‘AT ‘LI 
‘Ad “dd “MC “ZO ‘Dd “LV 

S31JJUNOD poprauy 

dD ‘7961 

uMouUxyU ial 

Ld “€081 

adomy ut 
prosel IST 

eIsepesIsny 

POLY 

POLY 

uMouyun 

umouyun 

SNOIOA 
-1nap 

snjyejg 

THR] “UOSYsIIy 
snsoursyjnf 
SNAIULOUNM T 

(C981 
‘AyS[NYISIOY[) 
snyn{na sniadossuy 
sepiiaydo7 

(QSZI ‘snoevuurT) 
SNIUVILANDUL 

SIL OLQIUIT 

satoads 
Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

384 

(TH6T) ATPL, 

“(6PG61) FJORDS “(6ZG6T) ‘Je 19 epnosy snoseyd (COST IStIT) 
‘(9861) S©qO’) “(GQ0Z) ‘Te 19 sadI0g snosanc) -ovkyd| = smjngsnduv snjsy 
sepnsoidng 

POLIOUTY 
(6007) se810g yijou SsnOIOA (96Z] JezanaID) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sassiog sunp OZV-Ld| umouyuy ‘eIseIny -inep| susapva snsogdosnyg 
(LOLI 
(6007) se810g SNOJOA | ‘SNOBUUTT) SHLAvUDIS 
pure eSuopusyy ‘(GQ0Z) ‘Te 19 sassiog uMmouyuy) | eoLIFY YON lop SNUAIISOUVIV) 
TEP POM: 
SOULE (ZZZI 
(6007) se810g ueq[odoursos snosoA| 97905) saiivotsof 
pure esuopusyy ‘(GQ0Z) ‘Te 19 sassiog Aeoap [e1o3aA umouyus) ‘odomy “lop snpvuoug 
(6007) ses10g (P78 “TeurIs5) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sassiog punoss Aakepo umouyus) adoimy] uMouyun)  syzung snurpojay 
(CPST Ipruyps) 
(SOOZ) ‘Je 39 seds0g spunois Apues uMouyuy)| = OAvaRyeg| UMOUYUN | s777guyISUL SNULOIIPLOT) 
Ad isthe bea 
$1001 uvourdo} SNOJOA (FSR ‘NoIpyatog) 
(8007) HG Arp ‘a3seydoyodsd -Ipsyl nap | saprogqis snatsavyds 
SNOIOA ZEQT wm 
(000Z) twOI_G pur opeysryy sionpoid pasois NVO-Sq| Umouyua odomy lop snignp snug 
(900Z) ‘Je 39 Sioquani HD uOIsaI (FCI ‘UOIsETTOAN\) 
‘(OOO0Z) THIOIG, pue opeypeyy ‘(766 1) sommUINs pue s3aI1 ‘NVO-SH “MS “GVW ueouesyo} snoseyd saprouynd 
Jouedsy ‘(G00Z) 24yng pur sopry 9q Poavo[peolg PpooM -Ld “Td ‘NH “AC ‘LV| Usouyuy -IPeV -o1dyd snsauLosyc) 

(600Z) sediog pue esuopusyy 

‘(000Z) PWOIE puke opeyseyy sSIM} snoseyd PLLI 1224) 29q 
‘(Z661) Jouedsy ‘(CQ0Z) ‘Te 19 sedI0g ‘SoINUUIN, UIPOOM NVO-Sd ‘OZV-Ld| UMowyu -o1kyd | wngpjaund uniqouy 
ZepHqouy: 
satgadg 
SIIUIIIFOY [2 eee S3IIJUNOD popeauy | psr0da7 IST eee WIZ Ayrore,y 

‘({] xtpuoddy 208) SIN 0} Jojor suorrerasigge yeugqepz *(] xipuaddy 22s) 99 TE OS] 0} JOJOI SuOTIeTADIQQeE sapood AT1UNOD ‘sepTaUINN07) pur O77 

nsuas sepyourosds1y’y ‘o7v] Nsuas FepruoTNoIN ‘sepioAquiesa_y ULYI Joyo sorrurey Jo adommg wz uate saroads viaidoajo>) dy} JO somstiaioeIeYS pur AST] °7°G*S VIGUL 


385 

Coleoptera families other than Cerambycidae, Curculionidae... 

(L661) SI9quIZTeA, 

odoin 

venus) 

‘(600Z) sed10g pure esuopusyy ‘UOoIsal (Z8Z1 
‘(Z007Z) HT ‘(S00Z) ‘Te 19 sadsiog aq ‘GVW-Ld ueoUPIIO} yoyepaid | ‘sniorsqe,y) saqvjouzg 
(8007) H"C ‘(000Z) ‘Te 39 UOsIopuy spoas aeaovidy ‘OZV-Ld ‘TT “AI ‘S| Usouyuq, -Ipeyy | /onsesed snyqopposiupy 
surequnoU 
‘adoing snoseyd S78 Swag 
(000Z) ‘Je 39 UOsIapuy spoas svaovog| ‘1g ‘bq Jemue7 -ouyd| = vsvanuou vip 
Spur] d1seM 
(Z861) ‘spaas svaoeNpsed snoseyd (Z6LI sezueg) 
‘Je 19 TJo3xpuy ‘(1 861) uosisuag 29 vousoduros uMouyuy)| — soseaeTEg -o1dyd SnIYND vivULy 
adoiny 
jenuay) 
‘uoIsaI ecsl 
spios ueouesioy snoseyd| ‘eIIIA “q°5 29 BIA 
(Z00Z) H"T (8661) BMT (8007) HMA | + Apus ‘spoas avaov0q uMOUUL) “Ipe -oikyd) -y vigoqiup vivupy 
(2661) B19quIaTeA “(6007) 
sasiog pur eSuopuayy ‘(Q00Z) WOO snoseyd (PLLI 1904) 
pure opeypeyy ‘(GQ0Z) ‘Te 19 sesi0g spoas asvaovog NVO-SH ‘OZV-Ld) umouyup}| ondsearyeg -odyd IC) vauav pavupy 
(Z66T) S22quIITeA ‘(Z00Z) adoiyq|  sorepord] 71 QT ‘pruazyos1jnq 
BVT “(ZHGT) Pruurel “(8007) HAA aD| @5D ‘0081 fenusy}| = /onisered smayvavd xvqy 
sepiqeie’) 
(wenqupy) adomy (VOLT ‘sniorsqe.f) 
(6261) ‘Je 39 epnesy ‘(Z86T) SSOUI UO spr2j JseayINos snoseyd VIVIAISIULIS 
‘Je 19 TJo3xouy ‘(186]) uosisuag ‘oddaas stdosyreuds Oy] umouyuy}| 2 JenueD -o1dyd VIADIO[TULS 
. sepryssdg 
(Z¥6T) (PLZT 1995) 
AIDUT, “(6FG61) JFIeYIS “(6Z61) ‘eI snoseyd 3(]) Ywuimnsv 
(C¥6I) 
ABUL, “(6PGT) PFPLYPS (6007) S°310g BGT ‘snoeuurT 
pure eSuopusyy ‘(6/61) ‘Je 9 epnesy snoseyd VIVINIVULUALOU 
‘(9861) S©qO7) “(GQ0Z) ‘Te 19 sasI0g SIOJTUOD ral -odyd susasdng 
I sated 
SdIMIIIFOY ae ae SaIIUNOD popeauy | pr0de7 IST Ceecel sWIsaY Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

386 

(2661) S29quIZTeA “(6007) 

sasiog pur eSuopuayy ‘(Q00Z) OI | suswUOJTAUS Apues POV joyepoid (ZG6LI ‘sniorsqes) 
pure opeyperyy ‘(SO0Z) ‘Te 39 sasiog ‘plye ‘souojs Jopun NV0O-Sd ‘OZV-Ld| UMouyuy | YON ‘ureds orisered snyngojaund SNUINT 
Joyeposd (C6LI ‘Jozueg) 
SOTIOISO S[| UMouyuE)| UeTIoqIsomy)  /onisered SNIDUIULAAY SNISIIT 
odoin 
UI2ISoM (ZI8T ‘prayosynqd) 
Spur] o1seM ‘yeI1U99 ioyepoid SNIDUIDADULO{NA 
(Z00Z) BVT (8661) BMT “(8007) HNC | —_‘saseroy ‘surerunow dD ‘TH61 ‘usaiseq| — /onsered SnISIIT 
(C181 Praryosynq) 
(600Z) ses10g pur esuopusyy sounp ‘spyoy uvouedIo} snoseyd snpuansuysstp 
‘(000Z) FWOI_ pur opeyseyy| ‘syred ‘sjros Arp ‘spoas umouyus) -Ips| -ovdyd srpdavey 
Poul) snoseyd (8C8T ‘Jetanbo7)) 
(OO0Z) FHIOI_G pure opeyprpy souojs Jopun ‘spoos umouyuy)| ‘yea ‘ured -oukyd| = szpouaf sadtupsy 
(snojrydoso) 
sureqUuNoU soyepoid (O6ZI ‘qmyAeq) 
‘SIOALI SUOTE 64 uMmouyuy)| ueLoqisomy| /oriseszed sypoas snigdvdy 
IGJUuvUaAy_) ‘XILV) (SCZ ‘snoevuurT) 
‘sSoq pur siaats Suoye joyepoid snyqtdvI1Aqy 
(OO0Z) FHIOIG pue opeyprpy skedop [e1939A UT dC ‘6A uMmouyuy)| ueoqisomy| /otseszed Sp1dqaMaqE 
(€00Z7) suapses onqoIeaKye jorepaid] = 49/7 T2INW VO 
‘ye 9 ULINT (8007) ‘Te I uesunqr] ‘spjey ‘spue[poom ee ol umouyus) ISO /orisered SUJVAOULIU SNGVAV’) 
(€007) rorepaid CZLI ‘snioniqey 
‘ye 19 ULM] “(Z007) BMT ‘(8007) Hd SISIIOF cy q5)| GD ‘9Egt| uewoeqisomy|  jontisesred SNXIQUOI SNGVAV’) 
odoin 
(€00Z) spo [eua_) pue joyepoid QC/| ‘snoeuurT 
ye 19 ULINT, (Z00Z) BMT (8007) HNd| 3s240} ‘pyey ‘[ros Ap uMouyuy), usJasayy | — /omuisered) = smgvqjaaupa sngvivy 
(€00Z) sosnyj[ou JoIwpaid odoimny Jorpaid QC/[ ‘snoeuurT 
‘ye 19 ULINT “(Z00Z) BMT ‘(8007) Hd ‘spury a1sem ‘sureyd uMmouyuy) usoIsay\ | /orasered SNIVAND SNQUAV’) 
saseq 90} 
“yreq ‘syunI) poyjay JouTUl jorepoid (SZLT ‘snioriqe.y) 
(Z00Z) H"T ‘(8661) BMT ‘(800Z) BNA Japun ‘ros Apues umouxuy)| visy ‘edomg| jonisered| = smgvuny snasyyey 
adomy Ut sa1ad¢ 
SddUdIIFIY soy] yeviqey] SaIUNOS popeauy | proser sT | odues oaneN | = swTsay Ayre] 


387 

SNAPI") pue SIIQY 

Coleoptera families other than Cerambycidae, Curculionidae... 

UI saTJaT[es apiaoq ysreq sorepoid} (QT ST ‘yeyuaTAy) 
(0007) HWOIG pure opeypepy | Ul seq prumy Jopun uMOoUyuU() onorejpop{| = /omisered vuvu vIhGIVT, 
(661) 
Broquia|eA “(Z00Z) H"'T ‘(8661) (8SZI ‘snoeuurT) 
HT ‘(0007Z) MH0I-E pur opeyoeyy o1OTeIeye jorepoid snuypgsgdoonay 
‘(800Z) H"C ‘(000Z) ‘Je 19 UOsIopuy saTqeas ‘sTeT[Io aD ‘NVO-SH “AI) Usouyuyp Sa} /oTaTsered snapogds 
(6781 “uvalaq) 
SppPy pareanyno joyepoid snjnasn1sUuo]go 
(Z00Z) BVT ‘“(866T) BVT (8007) HAG | ~—- -Wou UT satuoTOo UT dD ‘6Z81 adomg} — /ojsered snsyvqlas 
(600Z) Sesiog pue esuopusyy| sdoq sJeou ‘spurysser3 jorepoid (96ZI Sezueg) 
‘(800Z) HNC ‘(S00Z) ‘Te 19 sasi0g JOM UT 91SeM umouyuy) adomy} = onisesred| styusaa sngousosaggy 
spoom prumny Joyepoid (OZ8I Inojnq) 
(Z007Z) BH" ‘(8661) HT ‘(8007) HNC | ‘YsesF UT sou0js Japun orsered | sngpysis9 snqougsoLay gy 
ado (TIS ‘praryosignq) 
sous Jemusy pur} soxepoaid SNIVISNSUD 
(Z007) BMT ‘(8661) HYT ‘(8007) HN | JUAN YUM porerosse UIDyVWON | = /onsesed SNYINSOAIDT 
JOUTUI Jojepaic (Z6LT ‘sniorsqe.y) 
(000Z) ‘Je 19 UOsIopuy S1SdIOJ UT SdOq Jesu Al ‘0061 | eisy ‘edomy| jonisered| vynggn2 sngsqoopigg 
(6007) s2810g POLIFY YON royepoid (96ZI ‘snioriqe.s) 
pure eSuopuayy ‘(GQ0Z) ‘Te 19 sadiog IS¥OD ‘SIDAII SUOTE umouyus) ‘odomy| —/orsered sadiq]y sngsupLDg 
SIOAII SUOTR ‘SqIoy Joyepoid (ETSI ‘Aes) 
‘sJUsTUOIAUS prumy uMOUyUL) onorearyeg| = /omisered | smpoovugay snmosploc 
(SOOT) CIGUOL pue SOAP] ‘SIOATI eISY IOUT Jolpaid (S6ZT JIAO) 
OUNNO ‘(000Z) TWOIO pur opey~pepy|  Suoye ‘soyssreur Ares uMOUyU) ‘osdomy| —/orsered sniava sng dvjong 
(LZZ1 
snonploop soyepaid| — ‘9z905)) suyngnumu 
(Z007Z) BH" ‘(8661) H"T ‘(8007) HG Jopun ‘qos Arp qd ‘9261 onorearyeg | — /oisesed SISA OLIN 
9L81 “HeTNIg 
ssoq adomy}  soxeposd RF] Op psreyporg 
(L661) SI9quIZTeA, ‘TeIOT] Jesu ‘IseM WN ‘NH ‘LV| UsMouxuQ usoynosg} = /omsesred| — snavjyve suspuudT 
adomy Ut sa1ad¢ 
sadUaTaFaY soy] yeviqey] SaIUNOD popeauy | proser sT | oduesl oaneN | swTsayy Ayrore,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

388 

Poly you SnoIOA C/81 FoNIDyY 
(Z961) ‘Te 9 epnory 

“(6ZG61) ZO77&] “(Z86T) ‘Te 29 puy feel ag SNOIOA OFP8T “UOsyoITy 
‘(COQOZ) ‘Je 19 sadIOg ‘(T861) UosIsuag aseydoodur of OZV-Ld ‘OA| UMouyuQ -nInop syndy viapULory 
; aeprseydoydAry 

ssulqqno 

sseis pure isodwios 

uapies ATperodso 
(600Z) sesiog pure esuopucyy ‘(6661) ‘sooeyd wireM UT snoloaA) (/Z8T ‘fequafAy) 
peaisamog ‘(CQ0Z) ‘Je 19 sadiog| sureuror yueyd Apjouw uMoUyu) onorearyed op | s7jvdaqu] snsapoI1sag 
oeprydoyAro) 
(6261) SsUIpying ‘spriqoue POLFY YON Joyepoid (9¢Z| ‘snoeuurT) 
‘Je 19 spnaiy “(CQOT) ‘Je 19 sess0g oearey Aoid Jaquin uMoUxUL) ‘odomy| —/orsered sou. of1dCQ 
(6Z61) spliqoue POLFY YON joyepoid (ZE8T “uINIS) 
‘Je 19 opnaiy “(CQOT) ‘Je 19 sads0g 4aid ‘s3urpying uMoUyu) ‘odomy| —/orsered snaigsauop of1dCE) 
UOISIY (O6Z1 
(SOOT) BYANIASe'T ueouesyo} yoyepaid | IatAt[Q) ausooupLsas 
pur eaoljas ‘(6/6T) ‘Je 19 epnesy — ZO ‘0661 -Ipsyy | /onsesed unydouq 
: aepla[D 

S80] UdOI suOUTe 

ssour ut ‘durnis eRISy IOUT 

U9}10I UT slIqop qd ‘HD “AS “IN “AI ‘adoing snOIOA LIGI Fey 
(8007) HIG ‘SnIVIAT AsOTTOY UT ey ‘NH “Ad ‘Nd “TV| Usouxuy usJoyInos -ainep| sajynpyyod snquyy 
avpiquyyy) 

IIOATURIS 
‘SJUDWIUOIJIAUS Poul) (SZ8I 
Apues ‘pire ‘souois COL snoseyd ‘uevalaq]) snqypso2 
(0007) WOI_ pure opeyoeyy Jopun surequnow uMmouyuy) | YON ‘ureds -ovdyd SNJJIUIAIYISIIGIST 
(Z00Z) B"T ‘(8661) HYT puryoqy Ul IsoduroS Joyepord IEQI ‘Ueslaq 
‘(OO0Z) ‘Je 39 UOsIOpuy (8007) Hd UT ‘yeIOI] Teou jomsered | — sgvjougns snqgoaay 
I sa19adS 
sadUaTaFaY soy] yeviqey] SaIUNOD popeauy | proser sT{ | oduesr oaneN | swTsayy Ayrore,y 


389 

Coleoptera families other than Cerambycidae, Curculionidae... 

(961) CyB 
‘ye 9 apnoasy ‘(GZ61) Z07R ‘“(Z86T) POV SnosOA | WINIS sapuansuystp 
‘Je 19 TJo3xouy (1861) uosisusg | sionposd pasois ‘syrur ai Od | UMouxuy | eisy ‘odomy “0p snsvqdodhay 
(6007) sesiog pur eSuopuayy (C6LI 
‘(Z9G1) ‘Je 39 ePpnery ‘(676I) Zoe snoIOA “‘ISqIo}{) suqvjUuap 
(8002) HMC “(S00Z) ‘Te 19 se810g sumay Arp “Moy CVW-Ld ‘OZV7Ld spomeefed | -AEep sndvqdosdlay 
odoin 
(Z961) Te penusy SNOIOA OFRT “UOsyoITy 
19 apnayy (6761) Z07]eJ (8007) HIG aseydoodur ‘UIOYION lp DPIDANY DILVULOIY 
SNOJOA OER ‘suaydaig 
(6761) Z0>]e4 “(800Z) HIG aseydoodur uMOUxUL) adomy -19p VIIVISAL DIADULOIPY 
adoiny 
usayInos SNOIOA L9G] ‘uosuyo[ 
(8007) HG aseydoodur uMouxul) ‘yerU9_) “lp IPUDAIS DIADULOIYY 
(L961) “Te EOlAry Yaou SOE (S61 “T™]4ed) 
19 apnaty (6761) ZO>]eJ “(800Z) HIG aseydoodur umouyus) ‘adoing lop vyisnd pisvuLoly 
odomy L881 
UJOYION snosoa | Saniay xvogzy9und 
(8007) HG aseydoodur umouyus) ‘yerU9Z) lop VIADULOIYY 
BOLI you SNOIOA ZOST Weysiepy 
(6761) 20772 ‘(8007) HG aseydoodur uMouyuy) ‘odoin “ep DINPIIU VIAVULOIPy 
(Z961) ‘Je 39 epnosy SNOIOA OPQ] ‘UOSypIIY 
(6761) Z07]B] “(SQOZ) ‘Te 19 sasi0g dD ‘OZV-Ld| Usmouyuq, eIseIny -inep DpUnNUL DILDWUOLy 
JSIIOJ JOJTUOD ureds 896] 
Ayurew ‘peoiqe ‘adoingy SnoIOA| “pues 29 UOsUYyO[ 
(8007) #2G SIIqap pooM ud}I01 qD ‘9Z61 yeqqua7) -Inep 19840] VIADULOILy 
sdoiny SNOIOA LSS “WOISeTTON, 
(6761) ZO>]e4 “(800Z) HIG aseydoodur umouyus) usOyON lop 1dojsig VisvULoty 
(Z961) Je] 390q Surseurep synpe snoioa| (gOgT ‘fequayjé5) 
19 apnoaty (6761) ZO7]eJ (8007) HAG osye ‘oseydoodur uMoUuxul) sdomy “Ip sadiasnf{ piavuolpy 
jooq SUISeWep si{Npe snoloaA| (gO FeyuoYIsS) 
(6Z6T) Z077eJ “(8007) HIG osye soseydoodur uMoUyul) sdomy “Ip VIVISN DIADULOIET 
adoimy ut satad 
SIIMIIIFOY SOP] WIIQGey SaIUNOD popeauy | proser sy] | oduesr oaneN | swTsay Ayre] 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

390 

Cassn 

OLY 

‘adoinq 

édATIeU) 
ueqyod SnOIOA| FOG] Teneqsuer 
(900Z) ‘Je 29 S1oquaii yy) onsourop umouxyuy) -OUISO’) -lnep 1SSOA SNUABDIAT 
eISY IOUT SCSI “ZaeeIy] 
(9007) ‘Te ‘adoing SNOJOA snqvjnovultaponb 
qo SiaquantA\ (6261) ‘Je 19 epneszy onsowrop umouxyuy) usayinos -lWnep snuasvuyy 
SsnOJIOA QC/]| ‘snoeuury 
(6261) ‘Je 19 epnesy sjelioqeu jeurue umouyus) adomnyq -INep onjad snuasayy 

UOIBOI 
ueouedIoy snoIoA| C¢gy ‘uURUIOpyey 
(6261) ‘Te 39 ePpnesy ‘(8007) Hd oNsauOp If qd ‘HO| umouyuy “IPI -unep| snauunsg snuasaypy 

PISY IOUT] 
‘adoiny SNOIOA (O6ZI S14) 
sionpoid pasois Te Tel. Yq) umouyuy usayinos -ainep | sawiaspfig snuasvupy 
sepHsowsoq 

(6002) 

sas1og pur eSuopuayy (676) Z07e] SNOIOA S6LT TmyAed 
‘(800Z) HNC ‘(S00Z) ‘Te 19 sasi0g siseq xIyes ‘punoss qD ‘OZV-Ld “A]| UmouyuQ adomy -ineap| sajnqops snuasigdy 
Svs 
(Z96T) ‘Je 39 apnosy $1sou SNOIJOA “UNINIG 2p1ULGIS 
(6761) 20772] “(SQOZ) ‘Je I9 sesiog| vdsa, pure speurueuT UMOUYU) eiseing -ninap snsvqdoday 
(961) (8SZT 
‘ye 9 apnoasy ‘(GZ61) Z07TB “(Z86T) SNOIOA| ‘“snaeuUrT) szI77UvIS 
Te 19 TJo3xouy ‘(186]) uosisuag symsy Arp ‘ures3 uUMOUyUL) adoing -ninap snsvqdodiay 
(6007) se810g pure eSuopusyy CPS 
‘(Z9G1) ‘Je 39 epnery “(676I) ZOOTRY POLIFY YON SNOIJOA “WIINIS SNIVULSDS 
‘(CQOZ) ‘Je 39 sasIOg ‘(066 1) sesI0g uUMOUYU/) ‘adoing -ninap snsvqdodlay 
satgad¢ 
SdUdIIJOY yeuqey SaIUNOD popeauy | proser sT | oduer oaneN | swIsayy Ayrore,y 


391 

Coleoptera families other than Cerambycidae, Curculionidae... 

(600Z) sesiog pure esuopucyy *(7Z6T) adoiny snoseyd (TPS ‘uosysiry) 
IMdusIasIT ‘(CQOZ) ‘Te 19 sass0g sqniys C4 OZV-Ld| UsouyuQ usJoyInos -ovkyd |  snosgorp snjouvjapyy 
adomy 
uJOYWON, (LOST ‘sniorsqe.y) 
(ZZ61) IMousIasr] ‘ye11U99 snoseyd SYD PLOYLsoULave 
“(000Z) JOUGIeT ‘(C00Z) ‘Te 19 sasI0g oye10d ‘syeat39 $1003 Ca OZV-Ld| UsouyuE UIOISI\\ -o1dyd snogipy 
aepliaqeyy 
(vsn 
01 poyiodur)} soxepaid| 9Pg] Toneyuosoyy 
(6261) ‘Je 19 epnery ‘(8CG[) zuesy ioyepaid pryde adoma} = orisesed | 12u0sqo14a snigonsvT 
sepnuopoisg 
(6007) sesiog pur esuopuay/y 
‘(OOO0Z) THIOICG, pue opeypreyy ‘(6/61) sonpoid NVO SnOJOA O6ZI ‘wyeig 
‘Je 19 opnoaiy ‘(CQOT) [e319 sasiog| ewrue uo sNsauop -Sq ‘OZV-Ld ‘AT| UMouyuQ OOILTOHL Tap | saqvjnpun sajsausaqe 
(OO0Z) HHIOIG_ pure opeypep ‘(6Z61) sjonpold SnOIOA QC/[ ‘snoeuurT 
‘Je 19 opnaiy ‘(CQOT) [e319 sesiog| ewrue uo sNsawOp NV0O-Sd ‘OZV-Ld| UMouyuQ adomnyq op} snuzinu sajsausaqe 
adoiny SNOJOA O61 EM 
(6261) ‘Te 29 epnezy ‘(800Z) HVA sionpord pasos dD Iv] wsowuy eaue) — -Paep| ——_avajo snuaapuy 
(6261) SnOIOA (19Z[ ‘snoeuurT) 
‘Je 19 opnaiy “(CQOT) ‘Je 19 sass0g UOTIDa][OO UT s1asuT SL OOIeTOHL “Tap | wns0asnu snuasgqup 
UOIsaI 
(9007) ‘Te SIOMOP UO siNpe ueouedIo} SnOIOA OF8] ‘UOSyIY 
qo BIaquantA\ (6261) ‘Je 32 epnesry| suonNda]Too ur s1osuT HO ‘IV| UMouyuQ Ips -Inep snaysaf snuacqqupy 
UOIBOI 
ueoueIIO} SnoIOA I88[ THIPY 
(6261) Je 29 ePpnesy ‘(8O00T) HNC | syeuurue poynas ‘sunys GD ‘IV! dD ‘E861 -Ipoy[ 3seq -INep| snjy.0j09 snuaagiup 
UOIBOI 
(FOOT) Puseg uvoUuesIoy SnOIOA (Z8Z1 ‘snioriqe.y) 
pure uuewsay ‘(6/6[) ‘Je 19 apne sionpoid pasois qD “Aq| uMouyu~ -Ipoy Map | snqwisspfisg snuasvupy 
[3 SNOIOA I88I FoHPDY 
sionpoid pasois { eA ‘POLIFY YON -hinep xaduats snuasviyy 
I sa19adS 
sadUaTaFaY soy] yeviqey] SaIUNOS popeauy | proser sT{ | oduesr oaneN | swTsayy Ayrore,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

92 

(SLL ‘snioiiqe,y) 

(6007) se810g sliqap poopy “Toews soyepoid SYD PIO GLOuLaVvYy 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog orues10 SutAesop of OZV-Ld| umouyuy adomyg| onisered uol1a’) 
adomy 

(6007) se810g soroyseas Jenuey}| sorepoid 6Z8T ‘susydaig 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sasiog UO poameas SUTNOI q OZV-Ld| Umouyua UIYUON | — /oratsesred snssadap uols1a’y 

seprrydorpAyy 

(6007) sasiog | wnupy “sadaj ‘SIaAeped snOIOA 6061 IpreIppig 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog ‘surdeoop ysy gq OZV-Ld| Umouxzuy adoinq -lIMap | suaasagiugns snutsdvy 

(6007) sestog | wnap ‘sadaq ‘sIaaeped Jojepaid (OGZT “eql9sS) 

pure eSuopuayy ‘(GQ0Z) ‘Te 19 sadiog ‘surdesop ysy gq umouyuy)|  omoseaeyed| = jomisered | sugutsgszuas snutsdvs 

6F8 AYsNyos 0] 

(6007) SesIog | wnup ‘sadaq ‘sIOAEped jorepoid snynasniunyd 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog ‘surdesop ysy g umouyuy)|  omoseaeyed| = jomisered snutsdvs 

UNAPT ‘SIDIJ ‘SIDAEPEI jorepoaid| (CQg] ‘uueWOP) 

(SOOZ) ‘Je 39 sads0g ‘surdeoop Ysy n uUMOUyU) adomy| = /omisered | suaasaynsapa snutsdvs 

(6007) sasiog | wap ‘sadaq ‘sIOAeped OT eIsesOIUa_ royepoid (86ZI ‘snioliqe.s) 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sasiog ‘surdeoop ysy fal uMOUyU) -ornd| = /otaisesed | sugvuzwunov snutsdvs 
UOISOY 

uvoueiio1|  so0repord (Z9Z| ‘snoeuury) 

(6861) Inzeyy | —- [esoN] Ju “SuNp Moo Ig uMmouxyuy) -Ipsyy| 9 onusesred| = solv sagsyouovyy 

UOISIY (ZO8I 

(6007) se810g [los Apues ‘skeoap ueoue II} soyepoid| —SaSiy]]) saavipiusp 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sadsog | feIoSaa ‘sadaq ‘sIOARped Le umouyus) -Ipsyy | /onsesed snsmvI0dh ET 

(600Z) s2910g rs Peel Jolepoid (CHET ‘oqny) 

pure eSuopuayy ‘(GQ0Z) ‘Te 19 sasiog fal umouyuy)| yanos adornsa| = jomisered| wngound sngtsovqvyEy 

sasiog pur eSuopuayy ‘(Q00Z) WOO jorepoid| (€Qg] ‘uURUTTOF]) 

pure opeypeyy ‘(GQ0Z) ‘Te 19 sasi0g SUNP asIOY ‘Mod al NVO-SH‘OZV-Ld| UMouyuyQ| onoresryeg) jomisered| = srusootud1u snqwuoy 

———— ee ee Sep 

I sated 

SIIMIIIFOY SOP] = SaIUNOD popeauy | proser sT{ | oduesI oaneN | swTsay Ayrure,y 


393 

Coleoptera families other than Cerambycidae, Curculionidae... 

JOOI polIp ‘spaas]Io 

os 

aD ‘WN 

UOIsay 

(SOOT) ByANAseyT pure vaojfas ‘sinu ‘sjonpoid “AS “Td ‘NH “AC ‘TA ueouedIoy SNOIOA (PEST “PIEA) 
(8007) HMC ‘(S00Z) ‘Te 2° so810g uress pure uress SCT ZO “aAG EY Ie | ZZ ZOE -Ipsyp -ainep | sasuadva sagsajoqdry 
seproo;ydounry] 
(0S81 
UOISIAY ‘JOIIMUASITY]) 
(SOOT) BYAMase’] pur q) ‘HD uvauesiay snoseyd sngqugsaa 
PAOYPS “(8OOT) HNC “(€661) O'SIPNY | MuwUrT ‘wnugLUy ZI ‘a| ‘TT “AC “ZO “AG SEV] AD ‘6Z6T] PP SAK -o1dyd smposardlyaveg 
UOIBIY (F98T 
(8007) Hd uvouedIo} snoseyd| ‘Aermypj) 2u7q.4127Uv 
‘(C00Z) ‘Te 19 seBI0g “(€6G61) OISIPNY | YuwuTT ‘wnugLUy ZI ‘d| dD “IN ‘NT ‘IT “Ad] AD ‘9Z6I aA -o1dyd smposandyavig 
seplyaiaeyy 
(9¢Z| ‘shoeuurT) 
joyepoid SAP1OIVGVAVIS 
(SOOZ) ‘Je 39 seds0g a uMOUxUL) eiseingq|  /onisered uniypisavgds 
des yuryd 

uo pure ‘Iduny JoeUr adomy ISZI ‘sniorqey 
(6007) se810g oruesi0 SutAesop jenua) ioyepoid unqvjnqsndiq 
pure eSuopuayy ‘(GQ0Z) ‘Te 19 sadiog ‘sunp yeurureur q uMOUyU/) ‘uloIsoy\ | — /onaisesed unipiuavyds 

odoin snoseyd 

usoynos -odyd 
_ ‘uloisom| — Joyepard (LZZI ‘I01s10.) 
(SQOZ) ‘Je 39 sedr0g sJayeM SUIpuRs UT d‘id umouyus) genus} orusered| = supiay sasvqoojapy 
(6007) s9s10g odoin (Z6LI ‘snistzqe) 
pure eSuopuayy ‘(GQ0Z) ‘Te 19 sadiog prydoyey g uMOoUyUs) JOAO [[V} uMouyun £0]091g snagooug 
(6007) s9810g (T9ZT* snoeuury) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sasiog fal uMouxyuy) adomy} umouyun| saznbsinb uodosay 

omnueu pue UOTE 
WIOIJ popsooal Osye sdomy 

inq ‘satToArqsoy oBIe] yesusZ joyepoid (SO8I ‘yeyuay]Ar)) 
(600Z) 1SIOA, jo Sunp ur Ayurew n uMouyu) UIYUON | — /ontsered snjajosqo UolI41a) 
satad 
SdIUAIIF2Y ae eeeeey SaIIUNODS popeauy | pr0de7 IST Greece JUIISIY Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

394 

CEsI 

J1SseM OZV-Ld SNOIOA] “ITA “Gf “D2 eA 

(SOQ0Z) ‘Je 39 seds0g Je1999A Isou sjour a OZV-Ld “CO0Z adomy -lWINep |W 407091g vuLojouopy 
depIuUoJOUO[ 

(OPS ‘neuyaise> op 

snosoa| o10de'7) sypurdupu 

(800Z) HNq| purjpoom jorrxosdes 5) q5| uMouyuaQ adomy “Ip SNSADIJOUIXY 
sepliyoryep| 

depIpLoy ouseied 
‘snoyooroy ynpe 

uMouyus) PISeINy 

‘orepoid 
/orisered 

(T8ZT ‘Sey[ed) 
SIIQUIADA SLAQUIy 

a 

IEPIOP 

odoin 

urayINos 
‘yerqua7) SNOJOA ZERIT vosysy 
(8007) HNC ‘(1861) uosisueg Isunj A Oy] UMouyuQ UIDISIN\ -ainep| — susouss1jnf{ sdoqw’) 
J dEpIpolsT 

so1ep 
‘IMopP ‘oye ‘ero yeu 

queyd SutAesep SNOIOA (OS8T 10231247) 
(80072) Hd uo ‘sn3unj uo (a uMoUyUL) | UrIseayiJOU -hinep 1GJ0lBLIQ SAY. 

UOIsaI 
(8007) H2C ‘(800Z) JU29UTA, - gq ‘OZV uevoUurdIIoy SnOIOA (JOST “ureysreyy) 
pue yosnog ‘(SOOZ) ‘Je 29 sasiog Inoy ‘syuryd Arp If -Ld ‘OIS-LI “AG! dD ‘6881 -Ipey -hinap sypootfna DIJasaUuat 

adoimy 
(sorqe ‘“Itf-se[snop) usJoyliou SnOIOA) /9g] ‘Ays_NYydsIO/] 
(L9G) ‘Je 19 epnosy astperoads sJayruo0s +3) dq “Td | usouyuaQ jesus, Wap | W.10491gV VIAVINILO’) 
SNOJOA (OFGI ‘puess) 
(S66T) J3PMY “(SO00Z) ‘Je 19 sesiog| -yseq snjndod sopun dil AVWW-Ld ‘OZV-Ld| umouyug adomy Map| vI1daas0U asapoqy’) 
2EPHPESE | 
sa1gads 
SIDUIIIFOY se need SdIIJUNOD popeauy | p10daI IST Pate JUIISIY Ayrore,y 


(6007) s9810g 

298 

pue eSuopusyy ‘(Z96[) ‘[e 39 epnery OZV-Ld sdomy snoioa| (%g/] “siaqunyy) 
‘(S00Z) ‘Te 39 sasr0g “(€G61) OISIPNY suroorysnta raat JE OZV-Ld ‘C0OT| = UIOyION “nep| — vavandig vawindy 
OZV-Ld SNOJOA (QGZ[ ‘snoeuury) 

(S00Z) ‘Te 39 sads0g “(€G6T) OISIPNY OZV-Ld ‘C007 BIsy ‘adoing “Lap varsav vavandy 
(6002) s9s10g uorder | snoroarnap CHE “uOsysIT 

pure eSuopucspy ‘(Z96T) ‘Je 39 epnoiy uvoueiio| ‘snoseyd snqvuststaponb 
“(G00Z) “Te 9 ses10g “(€G61) OISIPRY sumay Arp If} | OZViLd “AC ‘LV| AC ‘0002 “TpPeW -oityd surg dodun) 
sep yaPpHin 

*SIOI1 PoAey| 

-peolg jo SOYIUPIG 

peop uo Surmoss 

UNINSAT Y UNIAY 

(8002) 1a sn8ung oy ya 

00 Te 
|e! 

POLY YON 

‘soured 

dD “C661 

uJoy NOs 

SNOIOA 
lop 

(Z881 J9312y) 
srusooyyf snisviny 

Coleoptera families other than Cerambycidae, Curculionidae... 

(L961) snOJOA PCR “UOIsETTON 
‘Je 19 spnaiy “(CQOT) ‘Je 19 sass0g ould uo ‘xupvUvy €r) HO ‘LV| umouyuy -ainep| —sastavucpy snurbiag 
sepiseydojoo4/ 

spuris 
(8007) vaItg -nUuoqI0LpuacT sorepaid L781 ‘eyuatAy 
HNq ‘(€00Z) aBNosuopy pure ioSnog ioyepoid +3) qD)| FD ‘E861 adomy} = onisered| szpuvus snovgdoziqy 
(6007) sesiog 33PIO}IS SNOJOA ZEST eqny 
pure esuopusyy ‘(GQ0Z) ‘Te 29 seds0g | Apped ‘sonprsor Apped oy -rinap | stooturds vusojouopy 
LZE8t oqny 
snOoIOA vypjoaaofisponb 
(SOQOZ) ‘Je 39 sass0g sures SutAesop of ‘{ -ninap DULOJOUOT 

31SEM 
(6007) se810g Je1o80a soseydooAur SNOIOA E61 “1SqI9}q 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sassi0g joseydosdes o[ *[ -ainep| — sadia1d vusojouopy 
(6007) ses10g snoroa| (/78T ‘fequayjé5) 
pure eSuopuayy ‘(GQ0Z) ‘Te 19 sadiog DISPAA [PIIS9A of ‘[ -TMap | syjostsug] vuLoJoUoPT 
I satad 
SOIUIIIFOY SOP] Lat SaIIyUNOD popeauy | pr0de7 IST eee JUIIS2Y Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

396 

(6007) ses10g (COLI 

pure eSuopusyy ‘(6661) Pysyeung SNOJOA ‘Jodoos) snazsdyy1 

“(SOQOZ) ‘Je 39 sesI0g ‘(Z7661) pneseg Ip sn3pydoy1uc 

IEPIOVqesedg 

(6007) ses10g snoroa] (OST ‘TeyueT[Ay) 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sassiog uMouxyuy) -ainep| wnpusnd unipiuag 

(6002) s9810g SNOIOA (PPST VOqITY) 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sasiog uMouyun uMOUxUL) -ainep | vjostnf xtadounapy 

SISdIO} ISU 

isoduroo ‘sasseored PD “AS ‘ON “IN snoIoA| (Z/8T ‘sMoyuey[) 

(161) ‘Te 29 epnazy (Z007) Hd] ‘uny Sumos “Bunp O{ Sa] “AI “AC ‘Td ‘IC ‘Lv] aS ‘7E61 odomg}| = aap] wavudoo srqotagosoy 

seprypd 

SISUIAAD 

SNgIUOS dTASTYMWosS snoseyd (Z6LI sezueg) 

(SOOZ) ‘Je 39 sadr0g MOTIPA JO spas ji OZV-Ld| UmouxzuA adomy -ovkyd | smasnss09 sndIVIVY 

seproryey 

(600Z) s95I0g poom istour SNOIOA (Q¢Z[ ‘snoevuurT) 

pure esuopuayy ‘(GQ0Z) ‘Te 29 sadsog | ‘soyseaq UO poomILIp — -ainep| wnuvjaus sapdaavpy 

depLIsWepeC 

istyeToads SnOIOA S88 Tay 

(800Z) HNC ‘(€66T) Olsipny |} seiAurJ91se8 snoasida | dD)! dD ‘F007 eIseIny -inep SNISNPV SNIPVIOT 

06ZT 

(SOOT) BYAMse'T] pur eaosjag sdomy SnoIoA | ‘Issoy yyjnsvuoany, 

‘(Z9G1) ‘Te 39 epnosy “(€661) orsipny sayeIgaiJ9a sau0q iE | Z| ZO ‘0061 usJoyInos -inep VINPUING 

OZV-Ld adomy snoseyd CPST “WINS 

(SOOZ) ‘Je 39 sasIog ‘(€66]) OIsIpny viavivs viadany Ca ‘Vi qD ‘OZV-Ld ‘/98[ | UsloiseayInos -ovkyd| — snupout sagsadsyay 
(600Z) sesr0g pur eSuopuayy 
‘(OO0Z) FHIOIEG pur opeyoepy 

“(Z961) ‘Je 19 ePpnelyz ‘(8007Z) Hd SUIpoo5 snoseyd (CZZI ‘sniorqe.q) 

‘(COOZ) ‘Je 39 sasdIog “(€G61) OlsIpny | -uarjod ‘ovaovsos ‘odes Il] NVOD-SdH ‘OZV-Ld odomy -o1fyd snauaw saqsasnayy 
(6002) s9810g 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sasiog SNOJOA CHQ] ‘UOsYysIy 

‘(0661) 89810g ‘(€66]1) OISIPNY (i eIseiny “nop yynsuol vavindy 

I sated 

SIIMAIIFOY SOP] Leeatl SaIUNOD popeauy | pioser sy] | oduesl oaneN | ows Ayrure,y 


397 

Coleoptera families other than Cerambycidae, Curculionidae... 

(vyjaq) e1aidiq (T9ZI ‘snoevuury) 

(600Z) sesIog pure esuopucsyy ‘(FZ6T) snoydeysso0poA9 Jo joyepoid vypynisndig 
Je 19 opnany “(GQOZ) ‘Te 39 saB10g | sprortseredoyoe Areuyos I OZV-Ld| umouyuQ| onoseorfeg| — /orusesed pavyooyy 
seprurAydess 

(600Z) sesiog pur euopusyy (2961) SNOIOA] (SORT eyuay|ay) 

‘Je 19 opnaiy “(CQOT) ‘Je 19 sass0g 

aseydoodur 

addons 

-ninap 

sniqnp snpuigds 
Se PIParacs 

(6007) see10g shoves. (O6ZT J9IATO) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog ‘ai OZV-Ld| UMouyzuQ odoing -IMNOp | saqwyuapiun snuvayis 
seprurAyiS 
adomy 
usoisey 
‘UIOYION snoseyd (QSZI ‘snovuury) 
(SOQOZ) ‘Je 39 sads0g sagovIpodousys ‘yerus7 -ovdyd voedo vadApy 
odoin 
Jemusoyinos| soyeposd (CZZI ‘snioriqe.q) 
sppey soxepoid yreus umouyur)| 29 UIaIsAA\| /olisered| wyvsiaav] viuvUIgY 
PEPIUCHS 
(C781 
snoIOA| ‘azuNY 29 JoT[NY) 
(L861) Uosisueg SOABIT ‘SOSSOUT uMouxyuy) -UN9p| — szuyqoo snugo1uasy 
seprusewpAdg 
IsOduro9 AS ‘ON odoimy SNOJOA (S¢ZI ‘snoeuurT) 
(6661) PIseung ‘(Z661) prezeg ‘snoseydordes ‘LT ‘AH ‘Id “IG) Md ‘08st usoYINos “ainep| — syusootspu sagauc) 
(6002) s9810g 
pure eSuopusyy ‘(6661) Fysjeung OZVe la SNOJOA (Z9Z1 ‘snoeuurT) 
“(SOOZ) ‘Je 39 sasi0g ‘(Z7661) pneseg sunp sdomy -ainep| ova snévgdoquc 
(6002) s9810g 
pure eSuopueyy ‘(6661) Fysjeung SsnOIOA (6SZT Seqeryos) 
“(COOZ) ‘Te 39 sesiog ‘(Z7661) pneseg sunp aq adoinq -ainep | sninvy snévgdoquc 
sated 
SdIUIIIFOY WwIIQGeyY SaIUNOD popeauy | pioser3sT] | oduesr oaneN | swTsay Ayrore,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

398 

(600Z) sesiog pur esuopusyy 

eloidiq 

yews jo SPATE] 

‘(000Z) TWIOIG pur opeypeyy “(FZ61)| SUTT[E“p-|los JonUuOS OZV-Ld joyepoid (SCSI ‘ZieeIy) 
‘Je 19 apnary “(CQOZ) “Je 19 sasiog| —eo1oTorq ‘“Joxeposd A! NVWOD-Sa ‘OZV-Ld ‘SCOOT odomy} = /omisesed VIADILOD DIAG 
(O€8I 
(600Z) sesiog pure esuopucyy ‘(FZ6T) joyepoid “UTS YIOUUPY[) 
‘Je 19 spnaiy “(COOT) ‘Je 19 sads0g prusered| nuadouvssys nyagpy 
(6007) ses10g 
pure eSuopueyy ‘(FZ6[) ‘Je 39 epnory NVO-Sa sorepoid (OLS TeyuaTfAs) 
“(CQOZ) ‘Je 39 se8IOg ‘(066 ]) se8I0g ‘GVW-Ld ‘OZV-Ld| UMouyuQ adomy| jonisered) vuvjuauvuy nagay 
(0007) FuIOIO 
pue opeyseyy ‘(7Z61) ‘Je 19 epnoiy NVO-Sa! OZV-Ld jorepoid (TER ‘suaydars) 
‘(S00Z) ‘Te 19 sedIOg “(0661) se810g ‘AVW:Ld ‘OZV-Ld ‘C007 adomg) = /oysered YINIIUY maysy 
OZV-Ld rorepoid LOGI ‘eaney 
OZV-Ld ‘COOT|  osearyed| == jortsesed SIOILINIV VAY 
POLITY YIION 
(6007) se810g ‘IOUTJ PIS ioyepoid (ZG8I Zeer) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sadsiog ‘sdomy} orusesed| suosfynoads snjlgouy 
(6007) sesiog UPU[OCGOUISOD Jolepoid | (ZO IssIOYUIAPIF)) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sadsiog ‘sdomy} omisered| = saynpyiu snjlgouy 
(G00Z) ses10g 
pure eSuopusyy ‘“(FZ6[) [Te 39 epnory rorepaid | (7Q8T sroyUsAeI4)) 
“(CQOZ) ‘Je 39 sasIOg ‘(066 1) se8I0g uMmouxyuy) orisered SYDUD DYISIMYy 
sayquas 
ul (vasmpyy) erordiq 
(VZ61) ‘Je 39 epnosy ‘(Z861) snoydeys0joAo jorepoid 6E8I I99H 
‘Je 19 TJo3xouy ‘(1 86]) uosisuag jo soyepaid uMOUyU/) odomy| /onisered vsivds DABYIONY 
erednd Aj uo osye 6h81 
(600Z) sesi0g pur esuopusyy ‘(7Z61)| pure sosseur Ap aeow OZV-Ld jorepoid “Ty Tayoequaipay 
‘Je 19 opnaiy “(CQOT) ‘Je 19 sass0g Sutdedap UO prj onorearyeg | = /oisesed | stwsostavja vavgooay 
I stad 
SODUIIIFOY ae eet SaIUNOS popeauy | proses sT{ | oduesr oaneN | swsayy Ayrure,y 


399 

Coleoptera families other than Cerambycidae, Curculionidae... 

(F261) OZV-Ld sorepaid (Sg I Zeery) 
‘Je 19 opnaiy “(CQOZ) ‘Je 19 sass0g n OZV-Ld “SOOT odomy} = jonisered| wanynsuvia vagy 
eISY IOUT 
(600Z) ses10g pur esuopusyy NVO-Sa ‘adoinq|  Jorepoid ZOQ | Weysrepy 
“(HLOL) ‘Te 9 epnary ‘(0661) SesI0g A} ‘AWW-Ld ‘OZViLd usypnos | —_/ojsered VpIpios wayyy 
(600Z) sest0g pur eSuopuay ‘(F/6T) pea] sours Jolepaid | (PH | Tovoamuasolyy) 
‘ye 9 apna] ‘(GQOT) ‘Je 39 sasiog rT Jj. & CVW-Ld ‘OZV-Ld ‘ODDOIOJ /orisered stagsnqod vIagIy 
(6007) sesiog pure esuopusyy ‘(F/61) OZV-Ld sodomy Jorepaid (GE8 | UOsysiry) 
‘Je 19 opnaiy “(CQOT) ‘Te 19 sads0g rar ‘SCOOT UIYUON | —/orisesred vI111GO DIBYIY 
(6261) Je 9 epnezy ‘(Z86T) SNIJUD DIO adomy royepoid (Z¢g | ‘UoswoYy ) 
‘Te 19 TJo3xouy ‘(186]) uosisusg snpidiayy wuny fal uMoUxuL) UIDyVWON | = /onsesed SIULOIASIU VIBYIPT 
(600Z) Sesr0g pur eSuopuayy 
‘(OOOZ) THIOICG) pue opeypryy ‘(F/61) ) * | hl joyepoid (9G8 [ ‘Z1eeIy) 
"Te 39 opnary “(GOOT) ‘Te 19 sesi0g N|  NvVO-Sd ‘OZV-Ld jousered vusiu moqiy 
(600Z) sest0g pur eSuopuay ‘(F/6T) odoiny (O€8I 
‘ye 9 opnoary “(Z86T) ‘Je 39 [Payouy uJOYIION, ioyepoid “Ua YyIJOUUPY[) 
‘(CQOZ) ‘Je 19 sadIOg ‘(T861) UosIsuag ae ae Sd ‘OZV-Ld ‘Od| Sa ‘€007 qenuay| jomisered| szuuadipiany vagy 
(PL61) 
"Te 39 opnorzy “(Z86T) “Te 39 [pug rorepaid O€GI ‘SUTTEA 
‘(800Z) HN ‘(1861) uosi8usg ysoduros “3sau priq of uUMOUyU) adoms| = onisered 1poomavy vIaYIY 
(FZ61) ‘Je 29 Opneary “(Z86T) joyepoid (OIGL ‘Aased) 
‘Je 19 TJo3xouy ‘(186]) uosisuag n umouyus) odoms| —onisered VIAVBIAS VIBYIT 
(600Z) sesrog pur eSuopusyy 
‘(000Z) FOIE pur opeypey] “(FZ6T) 
‘ye 19 opnoaly “(Z86T) ‘Te 39 [Payouy NVO-Sad ‘GVW rorepaid | (998 [ Is1oyUaAeIF)) 
‘(CQOZ) ‘Je 19 sasIOg ‘(T86T) UosIsuag | spyey ioIseO Toreposd Il <Ld ‘OZV-Ld ‘Qd| UMouyuy adomy| onisered ounf viag1py 
(6002) s9810g 
pure eSuopusyy ‘(FZ6[) [Te 39 epnory OZV-Ld jorepoid (PEST ‘Joysory[) 
‘(S00Z) ‘Te 19 sesI0g “(Q6G6I) se810g] 3s9U [eUNTUL pue pIiq a OZV-Ld ‘C007 adomg} — /ojsered vstarp visyty 
adomy ut satad 
S2IUIIIFOY SOP] Ss, SaIIUNOD popeauy | pr09e7 IST Ceecel JUIISAY Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

400 

(QZZ1 F9TRW 

(6007) se810g uvourdyoy ‘C) stusoonavaf 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog Aedop ‘O1SeM of -Ipeyyomns| uMouxun snurg dost) 
odoin Z98 | “‘UOsmIOYT, 

(600Z) sesiog pure esuopucsyy ‘(FZ6T) uJOYIION vIvULvYy 1g 
‘Je 19 9pnaiyj ‘(CQOT) ‘Je 19 sads0g n qenusy| UuMouyun vuapy dod) 
(OS8I 

(6007) se810g jorepaid| ‘oqny) wnupusagy 

pure eSuopuayy ‘(GQ0Z) ‘Te 19 sadsiog royepaid soduroo of ‘J odomq} = /omsesed SNYIUOLGYY 
(600Z) SesI0g pur esuopucsyy ‘(FZ6T) (JOST stoyUsAvIL)) 
‘Je 19 opnaiy “(CQOT) ‘Je 19 sess0g ral viseing| UMOUyUN|  smjnq1sd1U sntigvy 
(600Z) sest0g pur eSuopuay ‘(F/6T) osdomny OIGT Arrp ey 
‘Je 19 spnaiy “(CQOT) ‘Te 19 sads0g fal ujoyinog} uMouyUN|  snucufus sngoajdnq 
(9007) jelioqeur sodomy SNOIOA FIG FUPY 

‘Je 19 S1oquaiu pK “(600Z) ‘Je 39 BxN’T | ueyd Sumos aAsoduros of HO “Aq ‘LV) umouxyua uJOYyINOsS lop isapazsag snydvpy 
(600Z) sesiog pur esuopucsyy ‘(FZ6T) OZV-Ld (6€8 | ‘Wosyoisy) 
‘Je 19 opnaiy “(CQOT) ‘Je 19 sass0g OZV-Ld “SOOT adoinq| uMouxun vivaynd vgd’y 

(6002) se810g 
pure esuopusyy ‘(GQ0Z) ‘Te 19 sassiog 

OZV-1d 

OZV-1d 
“SOOT 

(WYN 

omur) sodomy 

uMouxuUn 

(8SZI ‘snovuurT) 
snsopjixpur snjty doar) 

(600Z) Sesiog pue eSuopusyy 
“(0002) 1010 pure opeyseW “(pZ61) 
"ye 19 opnony “(GOOT) “Te 19 sass0g 

NVOSSa. 
‘AVWiLd ‘OZV-Ld 

umouyuy) 

adomy 
UJOYION 

uMouxuUn 

(ZO8 [ IsJOYUaAPI)) 
VANISQO VIIVPLO’) 

ck kl 

(6007) ses10g OZV-Ld POLY (908T “IsroyuaarsTy) 

pure eSuopusyy ‘(CQ0Z) ‘[e 39 sasiog ‘SOOT | ‘eIsy ‘odomy uMOoUyUN|) YUL00GIUvx snip) 
euesing (S81 ‘uordag) 

umouyuy)| ‘eisysouryy| uMouyun| ysopnynd vinjstqovag 

(6007) sesrog OZV-Ld| FV GION Jojepaid | (TOSI “IsoyUaAPI‘)) 

pue vsuopusyy “(SQ0Z) ‘Te 39 ses1og| —__ Jorepaad ‘asoduros Of I OZV-Ld ‘COOT] eIsy ‘edomg]} — /aysered SnpIpsos sniustg 
(OOOZ) FHIOIEG pur opeyepy 

‘(PLOL) ‘Te 19 apnany “(Z861) NVO sorepaid (96g I z1eery) 

Je 19 ]Poyouy_ ‘(1861) uosisusg | -Sa‘AVW-Ld ‘O4| UMouyuQ adoms| = onisered VIVIOULL DIAG 

. satgad§ 

SdDUIIIFOY [Fe etry SoIUNOD poprauy | psJooos IsT Peel auIsay Ayrore,y 


401 

Coleoptera families other than Cerambycidae, Curculionidae... 

(600Z) sesiog pure esuopucyy ‘(F/6T) 
‘Je 19 opnaiyj “(CQOT) ‘Je 19 sass0g 

(600Z) sesiog pur eSuopuayy 
‘(000Z) RUCIO pue opeyeW “(FZ61) 
‘Te 19 apneny “(GOOT) ‘Te 39 sesI0g 

(LOT) ‘Te 9 apnoary “(Z86T) 
Je 19 TPoyouy ‘(1861) uosiusg 

speio3oa uanol 

(OO0Z) HHOI_G pur opeypepy 

(9002) 
‘Je 19 S19quai pA “(600Z) ‘Je 29 BINT 
(6002) se10g 

OZV-1d 

NVO-Sd ‘OZV-Ld 

Od 

OZV-~Ld 
‘C007 

uMOuUxU al 

uMOUxU al 

uMouyuy) 

odoing 

adoing 

adoing 

adoing 
snseone7) 

‘adoimy 
uJOyINOSg 

umouyun 

umouyun 

uMouyun 

uMouyun 

uMouxuUn 

(OFS ‘Uosyoiry) 
ppuin viodosauuyy 

(0G8 | JaveMuasaTy) 
vypsns BLodosauayy 

(SE8 [‘smoy1eY) 
stusonqasg puavyAy 

(E81 

‘suoydars) s77092431Uu 
snsodoja yy 

LG81 

TeA np urjanboef 
wgatapur snidadosoyyy 
(ZOST “sroyusarss)) 

pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog uUMOUyU) eiseingq | UMOUYUN| vyaIvLGI0 stAvgIOYIT 
(SAV 

(600Z) ses10g pure esuopusyy ‘(FZ6T) OZV-Ld ‘qyout) (€Egt ‘suaydars) 

‘Je 19 spnaiy “(CQOZ) ‘Je 19 sass0g ‘SOOT adoimy] uMouyun) — sappisnd snursydaT 

_y 

- 

speqoSaa UaNOI a 
fi) 

al 

eLIOqIS 
‘odomy 

(908 “sroyusarss)) 

(FL6T) ‘Te apnoary “(Z86T) [euaD pue auuadianf 
‘Je 19 TJo3xouy ‘(1 861) uosisuag | spyoy yom ‘sorrur ‘s3oq Od | UMouyuA uIDyVON | uMouyUN UNIGOLYIVT 
(600Z) Sediog pure esuopusyy 

‘(000Z) HHIOIO pur opeypeY “(/Z6T) NVO-Sd| OZV-Ld (SC8T SsnoysaIeA\ ) 

‘ye 19 apna “(GQOZ) ‘Je 29 sasiog ‘dVW-Ld ‘OZV-Ld “C007 sdomy umouyun vaqund VIOAM IE] 
adomnyq 

(600Z) ses10g pure esuopusyy ‘(7Z6T) OZV-Ld yenusd 198 | ‘UOswOYT 

‘Je 19 spnaiy “(CQOT) ‘Je 19 sass0g “SOOT ‘ulYyUON | uMouyun|  sadzaypyf vysacqojvE] 

(1S81 Ary 29 

adomy quesinyy) se4jvdisuoy 

(8007) HG dD ‘6861 usaisay\ | UMOUxUN SNGYIDUZOLPOLT 

sated 

SdIUIIIFOY SOP] SaIUNOD popeauy | pioser3sT{ | osuesl oaneN | swsayy Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

402 

(S007) ‘Te 39 sa810g 

‘COOT | OUT) eIseINy 

CN 

Joyepoid 
orisered 

(ZOST “sroyusarss)) 
snuuzquos sngqUojIgg 

(£861) 

ioyeposd 

(JOST “ssoyusarss)) 

ia] 
‘Te 19 TJo3xouy ‘(1 861) uosisuag a uUMOUyU/) onorejopxy| = /oatsesed | sagoyvqdaa snqquojig J 
(6007) s9810g (Zzgt ‘Tequsy}Asy) 
pue eSuopusyy ‘(g007) HC ‘(S00Z) snqvjoundunasvd 
ye 19 so810g ‘(/661) uosIopuy Nn gq ‘OZV-Ld “AI) aI ‘vsst umouxun snypy dom 
(6007) sesr0g 90ST IsIOYUsAeID) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sasiog fal umouyun sngdjnas snjaqhxcC 
(O€81 
adoing ‘Way IOUURY[) 
(PZ61) ‘Te 39 epnosy ‘(Z86T) yenusZ VOGALOULIVY 
‘Te 19 TJo3xouy ‘(186]) uosisusg fal uUMOUyU/) ‘uIDyWON | uUMouyUN ppoddxe 
(Z861) (68ZT ‘Tyed) 
‘Je 19 TJo3xouy ‘(1 86]) uosisuag Aedap [e1o3IA of umouxyuy) odoimy] UuMouyuUN) ana wunyvuc 
(Z861) snseoneo PERI ‘susydaig 
‘Je 19 TJE3xouy ‘(1 86]) uosisusg | sjewruweurosrw sisou (hal uMouxyuy) ‘adomy| umouyun | wnqvapoxa untvUC 
odoin 
(Z861) [enusy) 29 (9081 “AstoyUsArsr)) 
Je 19 TPo3xouy ‘(1 86]) uosisusg ssoq qd uMmouxyuy) uIYyUON| uMouyun| waosn{ unsgdoC_ 
(6007) sesiog pue esuopusyy ‘(F/61) OZV-Ld odoimny Jolepaid | (908 | ‘IsIOyUIAPIF)) 
‘Je 19 spnaiy <(CQOT) ‘Je 19 sass0g ioyepaid aur “SOOT UIIYyVON | — /osered puyjisnd Y{03YQC) 
adomy 
(PZ6L) ‘Je 39 Opnoary ‘(Z86T) Jemuey (TERT ‘suaydais) 
‘Te 19 TJo3xouy ‘(1 86) uosisusg uUMOUyU/) ‘uIDYyON | uMouyUN vivid VaVIC) 
(908 “sroyusarss)) 
OZV-Ld snynsaz04d 
(SOQOZ) ‘Je 39 sass0g “SOOT adomy} uMouyun snqUsigoayy 
(8781 
(6007) se810g ‘Ipneq) saplo1qo1g1yy 
pure eSuopuayy ‘(GQ0Z) ‘Te 19 sadiog umouxyun SNIUSIQOAaNT 
sated 
SOIUIIIFOY SOP] WwIIQGeyY SaII}UNOD poprauy JUIIS2Y Ayrure,y 


403 

Coleoptera families other than Cerambycidae, Curculionidae... 

(6002) s9810g 
pure eSuopueyy ‘(F/6[) ‘]e 39 epnosy 

odomy 

(O€8I 

“UT YyIOUUPY[) 

“(CQOZ) ‘Je 19 sasIOg ‘(066 1) se8I0g uJDyVION | uMouyuUN}| vasv4saq vLodoaojgg 
(600Z) sestog pur esuopuay ‘(F/6T) (JOST stoyUsAarss)) 
‘Je 19 spnaiy “(CQOZ) ‘Je 19 sass0g umouxyun sasaq pLodoaojgg 
OZ8I 
(600Z) sesiog pure esuopucsyy ‘(7Z6T) ‘Tpneg szuusofisnsuv 
‘Je 19 spnaij “(CQOZ) ‘Je 19 sass0g umouxyun vsodoaojgy 
(6007) (ZOST “stoyusarss)) 
sasiog pur eSuopuay ‘(Q00Z) WOO (WYN jorepoid SYLIDAQULN 
pure opeypeyy ‘(GQ0Z) ‘Te 19 sasi0g NVO-Sd ‘OZV-Ld omur) sodomy} — /omisered sngqUojtg 
(OI8T TeyutAD) 
(6007) se810g BOY YON |  sJowpoid sntavyinbsinb 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog ‘eiseing | —/orasered sngqUontg 
(6007) sesiog Jolepoid (9¢Z| ‘snovuurT) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog adomy| jomisered) = sngyjod snqquojtg gy 
(VOLT FTW 
(4861) elaqig|  Jorepoid Q) smqpulsivUL 
‘Je 19 TJoyxouy ‘(1 86]) uosisusg umouyus) ‘sdomq| —/orsered SNYIUOIIG 
cE81 
(6007) se810g oyepad | ‘susydarg s1u.10918U0) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sasiog eiseingq| /onisesed sngqUojtg 
(Z861) Jojepaid | (TORT “IsoyUaArI4)) 
‘Je 19 TJo3xouy ‘(1 86) uosisusg umouyuy)| onoreoryeg| = /oraisesed | suzuvyauur sngquojig gy 
cL8I 
OZV-Ld snseoneo| joyepoid| = qaane,y smqvugsaual 
(SOQOZ) ‘Je 39 sads0g ‘adomy| —/orsered SNGIUOIIG 

(600Z) sediog pue esuopusyy 
‘(OOO0Z) THIOICG, pue opeypryy ‘(7/61) POLIFY YON rorepaid | (7QBT sroyUsAes4)) 
‘Te 39 epnary “(GOOZ) ‘Te 19 ses10g NVO-Sd ‘OZV-Ld ‘eiseng | /oyusered | snaproasyp snqquopiqg 
I satad 
SdIMIIIFOY << SaIUNOS popeauy | pioser3sT{ | oduesr oaneN | swisay Ayrure,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

404 

6¢81 
jorepoid TOIH S14qUaa1sUoy 
(SOQ0Z) ‘Je 39 seds0g fal OZV-Ld| Umouyu te orisered snuyoygquvy 

(600Z) sest0g pur eSuopuayy ‘(F/6T) 
‘ye 19 opnary “(Z86T) ‘Je 39 [Peyxouy adomyq ioyepoid (S6ZT JIAO) 
‘(SOOZ) ‘Te 29 sasiog ‘(1861) uosisuag “‘saSsOU ‘SdUOIS asad OZV-Ld ‘O} UMouyuyp | son /orusered SIAVIUL] SNUIJOGIUDX 
(9007) : ZOOLN i eer (776 [O19 pod) 
‘Te 19 siaquait A “(6007Z) ‘Te 39 BYNT isoduros “pury o1seM — qD ‘HO “AA Nd {LV} Usouyu uJoYyINOS -IMNIP) = 2G IAVUL YIOANIIAGT, 
OZV-Ld (SAV (ISZI ‘smisiiqey) 
(600Z) sesiog pue esuopusyy n ‘COOT| Uy) edoinq| uMouyun| saynpyzu snsoddqgovy 
(9¢Z| ‘snoeuurT) 
(6007) se810g snuyauLoshaga 
pure eSuopuayy ‘(GQ0Z) ‘Te 19 sadiog n eiseiny | uMouyUN snsodhgavy, 
(Z861) (VN ZO8T “IsToyUsArsD) 
‘Te 19 TJo3xouy ‘(186]) uosisuag n umouyun SNIDUBIS SNULYIVT 
(Z861) SNOIOA) ZO8T IsIoyUoARID) 
‘Je 19 TJo3xouy ‘(186]) uosisuag nN -inep SUJOINID] SNULGIVI, 
(ZO8T ‘o[fAoureg 
(FZL61) op nosiig) 
‘Je 19 opnaiy “(CQOT) ‘Je 19 seds0g n umouyun|  sanbuidosd sniuns 
(908T “sroyusarss)) 
(PZ61) ‘Je 19 epnoy uMouxyuy) odomy) UuMouyuN) — s2V77a0419 YqUss0ar) 
(6007) s9310g OZV-Ld odomay | roxepord| (QT “Wosypiry) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog ‘CO0Z usoyinos| —/o1asesed snsouinad snuLay 
(Z861) Joyepaid) = (ZO8T “ueYsieyy) 
Je 9 TJo3xouy ‘(1 861) uosisusg umouyuy | pemusy ‘sdpy| — /ontsesed | snuzjaosau snipanc) 
“(COLI ‘sntorqey) 
(GOOZ) CfeI93INE pur esreurery “(/86]) SnOIOA sn.saqqyIvlg 
‘Te 19 TJo3xouy ‘(1 861) uosisusg speqoSaa UaNOI uMoUuyuy) | = TOIBaeTeY Ip SNULIJOAT 
sated 
SIDUIIIFOY se ened S2IIJUNOD papeauy | psr0da7 IST eae JWIISAY Ayrore,y 


405 

Coleoptera families other than Cerambycidae, Curculionidae... 

WOISoI 6081 
(O00Z) WOO uesue IO} snosoa | ‘aypiane’y] saprozpogdv 
pure opeypeyy ‘(GQ0Z) ‘Te 19 sasi0g sionpoid pasois uMOUxUL) -Ipoyy lop SIJBISMGIVAT. 
(000Z) FWoIG_ pure opeyoepy dq) ‘NVD SNOJOA TG6LI ‘snioliqey 
‘(800Z) HNC ‘(S00Z) ‘Te 19 sasi0g sionpoid pasois -Sq ‘OZV-Ld “AI) Usouyuy adoing “lop SNANISQO O1AQIUIT 
UOIdaI 
uevouedIo} SNOJOA B6LI ‘snioqey 
(000Z) HWoIC pur opeyseypy uMmouxyus) -IPey -ainep| — sngvzaund snanpay 
snoIOA (66ZI J9zueJ) 
uUMOUyU/) -inep zu1d snaj144.007) 
S1SIIOF SNOIOA (OGZT ‘snoriqe.y) 
(80072) Hd poavaypeolsgq pro uMOoUxyU/) -nep SIAVIUY] SNIIJLOY 
uvouedIo} 
-IPe SNOIOA POST ‘afoneyT 
(8007) HG uUMOUyUL) ‘adomy -ninap vpuosmnus sd J 
ado 
(S007) ZoepueuIay penusy SNOJIOA (SCZ ‘snoevuurT) 
ZIUTLYY pure JoqIBJ “(8007) HG SNOJOAIIIOp q5| umouyuy| pur ussiseq -nep vovsisoUuL SdDIG 
SNOIOA TOS] “Wreysieyy 
(8002) HC ‘(S00Z) ‘Te 19 ses10g dD ‘OZViLd | wHouwyun odomy SLAs P miafigiay sdvyg 
UOIBOI 
(SOOT) PYANISe'T uvourdyoy SsnOIOA (SCZ ‘snoevuurT) 
Pur vAodjag “(CQOZ) “Te 19 sass0g ZO ‘8881 “IP “HINep sv518 sda 
: depIUOLIqouey, 
(Z861) Jojepaid | (TORT “IsoyUaArI‘)) 
‘Je 19 TJo3xouy ‘(1 861) uosisuag OA] UMouyuA, adomy} = /omisered | sussaudap snuospojy 
(ZOST “weysiey) 
(4861) jorepoid snuu1guog 
‘Je 19 TPo3xouy ‘(1 861) wosisuag | sreyjao ‘suapses ‘sisoroy} Tf ‘Z] YF ‘D uMOUxUL) odomy| onisered snumospoly 
. I sa1gads 
SIDUIIIFOY soY] yeuqey saIUNOS pepeauy | proser3sT | odues oaneN | swIsayY Ayrore,y 


Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 

406 

ueouesoy (OGZT J9IATO) 

(8007) #"G fal q5| uMmouyua -Ipsyy| umMouyun}| ausosyna uniuojny 

seproydo7 

(6007) s°s10g pur eosuopuayy SNOJOA (Z9Z] ‘snoeuury) 
“(CQOZ) ‘Je 39 sesIOg ‘(066 1) sedI0g | -anep AaQUIS XOLT 
IEPISOIT, 

(6261) ‘Te 29 apneryz ‘(G00Z) sesI0g OZV-Ld SNOIOA (99Z| ‘snovuurT) 
pure esuopuayy ‘(GQ0Z) ‘Te 19 sadiog ASaIO} UT “YIeq 13) OZV-Ld ‘SOOT adomy -TNMap | sap1oqsauLsap SNISOLGT 

SIDUIIIFIY 

vor | sense | snunoy popes 

adomy ur 
prosal sq | oSues oaneyy 

WISI 

seppsolyy 
sa19adS 
Ayyure,y