Synopsis of warty leaf beetle genera of the World ( Coleoptera , Chrysomelidae , Cryptocephalinae , Chlamisini )

Th e 11 world genera of Chlamisini Gressitt are reviewed, diagnosed, and illustrated. A key for their identifi cation is provided. A replacement name is proposed, Kakita Chamorro-Lacayo & Konstantinov, nom. n., for Ceratochlamys Bokermann, 1961, a junior homonym of Ceratochlamys Habe, 1946 (Mollusca). Chlamisus rousei Medvedev, 1993 is designated as a junior synonym of Chlamisus straminea Suff rian,


Introduction
Chlamisini Gressitt constitute a relatively small tribe in the leaf beetle subfamily Cryptocephalinae Gyllenhal with approximately 500 species currently classifi ed in 11 genera (Blackwelder 1946;Monrós 1952;Seeno and Wilcox 1982).Th e distribution of species in the tribe is largely circumtropical.Th e majority of species and 10 of the 11 genera occur in the Neotropical Region, particularly in the Amazon Basin (Table 1).Th e only genus not represented in the Neotropical region, Hymetes Lacordaire, 1848, is known to occur only in the Oriental region.Of the 10 genera found in the Neotropics, four are endemic.Th e largest genus, Chlamisus Rafi nesque, has been reported from all biogeographic regions and it is the only chlamisine genus to be found in the Palearctic and Australasian regions.
Adult chlamisines are distinguished readily from other leaf beetles by their tuberculate or warty dorsal body surface (resembling caterpillar droppings), by their legs fi tting into depressions, and by the presence of antennal grooves adjacent to the prosternal process into which the short serrate antennae fi t.Hence, these beetles have the ability to tightly withdraw all appendages to form a compact cylinder and successfully mimic small fl ower buds, caterpillar droppings, or other forms of debris or excrement.Most are dark colored, but some are metallic or light with dark velvety spots.
Chlamisini, as well as other Cryptocephalines, have an interesting and unique life history.Each egg is individually covered by the mother, with a mixture of her own feces and rectal secretions to form a scatoshell, before releasing it into the environment (LeSage 1982(LeSage , 1984;;Erber 1988;Brown and Funk 2005).Th e individual plates that make up the scatoshell (egg-case) are thought to be unique to each species (Erber 1988).After eclosion, the larva retains this scatoshell.With each subsequent instar, the growing larva builds onto its inherited scatoshell with its own pliable feces (Brown and Funk 2005;Chaboo, Brown and Funk 2008;LeSage 1982LeSage , 1984)).Finally, pupation takes place within their cases.Th ese cases may not only aff ord larvae mechanical protection, but they may help to disguise them from predators by eff ectively resembling auxiliary plant buds.

Genus
Author 1801, nec Bolten, 1798) is the type genus of the tribe Chlamisini Gressitt, 1946, which is based on the valid name of the original nominal type genus (i.e., Chlamys).Nonetheless, the invalid name Fulcidacinae, based on the genus Fulcidax Voet, 1806, has been used previously (Jacobson 1924;Navajas 1944a,b;Chûjô 1940Chûjô , 1942;;Braga et al. 1999).To date, Chlamisus includes the majority of species in the tribe and has become a catch-all genus in need of a comprehensive taxonomic revision (Karren 1972;Monrós 1952;Reid 1991).Lacordaire (1848) proposed14 species groups in Chlamisus.
Th e second oldest name in the tribe is Fulcidax, a senior synonym of Poropleura Lacordaire, 1848.In addition to Poropleura, fi ve additional genera were proposed by Lacordaire (1848): Carcinobaena Lacordaire, Diaspis Lacordaire (= Diplacaspis Jacobson, 1924), Exema Lacordaire, Hymetes Lacordaire, and Pseudochlamys Lacordaire.Monrós (1948Monrós ( , 1952) ) established 2 genera, Melittochlamys Monrós for the fi rst 7 species included by Lacordaire (1848) in species group 1 of Chlamisus, and Aulacochlamys Monrós for several species in Exema.Bokermann (1961) proposed the genus Ceratochlamys Bokermann for an unusual species from Brazil; however, this name is a junior homonym of Ceratochlamys Habe, 1946 in Mollusca and a replacement name is here proposed.Th e most recent addition to the tribe, Neochlamisus Karren, was described almost 4 decades ago.Th e genus was established as a result of a comprehensive revision of North American Chlamisinae north of Mexico, and it consists of a number of species formerly included in Chlamisus and Diplacaspis (Karren 1972).Major chlamisine revisions have been regional, e.g., Monrós (1952) and Karren (1966Karren ( , 1972) ) for Argentina and North America north of Mexico, respectively.Th ese treatments comprise the most recent taxonomic activity in the tribe.
Motivation for this study came from the realization that identifi cation tools for chlamisine genera, mainly for the Neotropical fauna, are outdated, incomplete, or diffi cult to use, particularly when dealing with very similar taxa (i.e., Chlamisus, Diplacaspis, Neochlamisus, Pseudochlamys, Fulcidax, and Exema).As with most Neotropical Chrysomelidae, identifi cation at all taxonomic levels below tribal, presents a major challenge, and generic diff erentiation of chlamisines is no exception.Yet, no single key off ers the tools to confi dently segregate species into genera.Th erefore, the objectives of this study were to provide comprehensive, accurate, and lucid identifi cation tools to world chlamisine genera and to identify potential areas of research.Th is was achieved by studying the type species of each genus (when available) and several congeners, and by comparing traditional and novel characters across all taxa.Th e product consisted of a dichotomous key to all world genera, as well as analogous taxonomic diagnoses and high resolution images/illustrations for each genus.
Even though we follow the classifi cation proposed by Lacordaire (1848) and our study is based on original generic concepts, we echo the sentiment expressed by Karren (1972) and Reid (1991) that there is a need for a complete reassessment of generic boundaries within Chlamisini, particularly Chlamisus, to refl ect natural groups.Th is study is the fi rst step towards a comprehensive revision of Chlamisini, which will center largely around taxa from the Neotropical Region.Characters identifi ed in this study will provide a basis from which to address generic concepts in the future.Complete disarticulation of key taxa from each genus and morphological comparison among the genera was beyond the scope of this study.

Materials and methods
Our treatment of all chlamisine genera is based on examination of the type species of each genus, when available.Additional representatives of each genus were examined (see material examined section under each generic treatment).All material examined is housed in the entomology collection of the National Museum of Natural History, Smithsonian Institution (NMNH).Th is institution holds the Monrós collection as well as many chlamisines identifi ed by Karren and Bokermann.Under each genus a list of the material examined is provided; it includes exact label data for each specimen [all labels are listed as they appear from top to bottom on the pin and each label starts with a lower case letter separated by a forward dash (/)].Genera are listed in alphabetical order.In addition to observations made of available specimens, the dichotomous key takes into account characters used by Monrós (1952), Karren (1972), and Riley et al. (2002).Terminology for morphological structures follows Karren (1972), Chamorro-Lacayo and Konstantinov (2004), andChamorro-Lacayo et al. (2006).
Remarks.Aulacochlamys resembles Chlamisus Rafi nesque in overall body shape, size and color; however, the six longitudinal carinae on the pronotum immediately dis- tinguish it from Chlamisus.Th e absence of spines on the fore-and midtibiae also diff erentiates most studied species of Aulacochlamys from Chlamisus; however, this character may be sexually dimorphic in Chlamisus (Karren 1972).
Remarks.Th is genus resembles Melittochlamys in overall body shape; however, the modifi ed legs of Carcinobaena immediately distinguish it not only from other chlamisines, but from most other genera in Chrysomelidae.Diagnosis.Length 3.09-7.72mm, width 1.90-5.36mm.General body shape cylindrical.Body usually not metallic in color.Frons in canthus of eye usually without yellow spots (face may be entirely or partly yellow, with yellow area extending into canthus, but elytra do not have velvety spots).Frons glabrous or (rarely) covered with dense hairs.Pronotum and elytra usually glabrous or (rarely) covered with dense short hairs.Antenna serrate beyond 3 rd or 4 th antennomeres, 2 nd antennomere slightly widened, globose, 5 th antennomere nearly as large as 6 th .Pronotum medially elevated, with various bumps and short carinae.Pronotal base opposite mesoscutellum (posterior pronotal lobe) with well diff erentiated notch.Prosternum posteriorly narrowed, posteriorly much narrower than anterior margin.Anterior margin of metasternum concave.Mesoscutellum short, transverse.Metascutellum not exposed.Sutural serration of elytra usually incomplete (suture entire immediately following mesoscutellum).Elytral tubercules well developed.Males without spines or spinulae on fi rst ventrite.Tibiae slightly curved, slightly fl attened, with sharp dorsal edge.Fore-and midtibial apices with spine.Tarsal claw appendiculate.Male ejaculatory guide (part of internal sac of aedeagus) symmetrical, without sheath.Apex of spermathecal duct as wide as rest of pump.
Remarks.Chlamisus is close to Exema and Neochlamisus.From Neochlamisus it can be separated by the following characters: body usually not metallic in color; frons in canthus of eye usually without yellow spots (face may be all or partly yellow, with yellow area extending into canthus, but elytra do not have velvety spots); and male ejaculatory guide symmetrical, without sheath.From Exema it can be separated by the following characters: males without spines or spinulae on fi rst ventrite; 5 th antennomere nearly as large as 6 th ; sutural serration of elytra usually incomplete (suture entire immediately following mesoscutellum); and prosternum posteriorly pointed (narrowed), posteriorly much narrower than anterior margin.
Remarks.Th is genus may be distinguished from other Chlamisini by the shape of the prosternum, and an exposed metascutellum in combination with the appendiculate tarsal claw.According to Karren (1972), South American species of Diplacaspis may not possess bifi d claws or a posteriorly expanded prosternal process between the mesocoxae.However, the only two species available for our study [D.batesi (Baly) and D. prosternalis (Schaeff er)] share character states consistent with the diagnosis.
Remarks.Th is genus may be distinguished from Chlamisus by the following characters: males with spines or spinulae on fi rst ventral abdominal segment; 5 th antennomere much smaller than 6 th ; sutural serration of elytra complete (in Chlamisus males without spines or spinulae on fi rst ventrite and 5 th antennomere nearly as large as 6 th ; sutural serration of elytra usually incomplete); ejaculatory guide short, single, tubular, and sclerotized (elongate and paired in Chlamisus).Lacordaire (1848) established Exema for species of Chlamisus that have a very abrupt antennal "club", the fi rst serrated antennomere (6 th ) is much wider than 5 th .
Exema was designated a junior synonym of Chlamisus by Gressitt and Kimoto (1961) based on their study of Oriental species; however, Exema continues to be treated as valid (Karren 1966(Karren , 1972;;Riley et al. 2003;Seeno and Wilcox 1982).
Remarks.Th is genus can be separated from all other chlamisine genera by the anteriorly elongate metasternum projection (concave in other chlamisines), by the subcylindrical 3 rd , 4 th and 5 th antennomeres, and by the absence of spines on the fore-and midtibiae.
Th ree species are known from India and Java (Monrós 1952).
Remarks.Only a single female specimen represents this genus.Whether the laminar projection on the head, which distinguishes this genus from all other chlamisines, is sexually dimorphic is unknown.In leaf beetles, sexually dimorphic characters on the head, such as projections and enlarged mandibles, are usually present only on the male (e.g., Labidostomis Germar; Pseudochlamys Lacordaire; Normaltica Konstantinov) (Konstantinov 2004;Konstantinov and Korotyaev 2004).On the other hand, all female cryptocephalines have a modifi ed 5 th ventrite for the purpose of scatoshell coating.It is possible, that this laminar projection on the head is only present on the female also for purposes dealing with coating of individual eggs.Females, during the coating process, strike a pose in which the entire weight of their bodies rests on their forelegs and possibly their head.Th is laminar projection may be helpful in balancing their body.One thing is certain, observation in the fi eld and continued collecting eff orts to discover the male are necessary to understand the function of this modifi cation.
Th e name Ceratochlamys Bokermann, 1961 is preoccupied by an available name in Mollusca, Ceratochlamys Habe, 1946.A replacement name, Kakita Chamorro-Lacayo & Konstantinov is provided and Ceratochlamys Bokermann is invalid as a junior homonym.
Etymology.Kakita, modifi ed from its correct spelling "caquita", is a Spanish word for "small feces or excrement".Th e name alludes to the resemblance these beetles have to small pieces of excrement.Th e name is feminine.Distribution.Central and South America (Monrós 1952).

Melittochlamys
Remarks.Some species of Chlamisus (e.g., Chlamisus achalay Monrós, 1952 andChlamisus perforatus Monrós, 1952) also have velvety spots on the elytra while others have a broad prosternal process.Monrós (1949) broadened his own defi nition of the genus to include species that lack velvety spots on the elytra, have a broad, parallelsided prosternal process, and a globous, oval body shape with the pronotum dorsally smooth and continuous with rest of body.Melittochlamys can be separated from all other chlamisine genera by the nearly rectangular prosternal process; the process is more or less triangular in other chlamisines.
Remarks.Neochlamisus was proposed to include a few species, formerly placed in Chlamisus and Diplacaspis, based on their similarity in the male and female genitalia (Karren 1972).Degree of exposure of the metascutellum varies greatly, from broadly exposed to completely concealed by elytra.Neochlamisus can be separated from other chlamisine genera by characters of the male genitalia.Th e male ejaculatory guide is asymmetrical, with a sheath.Among external characters, color of the frons in combination with the presence of velvety spots usually allows for recognition of Neochlamisus (frons on canthus of eye usually with yellow spots, or if without yellow spots, then elytron with two velvety spots).Distribution.North, Central, and South America (Karren 1972).
Remarks.Pseudochlamys can be distinguished from other chlamisines by the following characters: head not completely retracted into prothorax; mandibles enlarged in males, normal in females; fore-and midtibial apices without spine; prosternum strongly and abruptly constricted beyond anterior margin; and prosternal process more than ¾ as long as prosternum.
Five species are included in this genus.
.00 mm.General body shape subglobular.Antenna serrate beyond 3 rd antennomere, 3 rd antennomere only slightly dilated distally.Pronotum without median elevation, relatively smooth and continuous with rest of body, without well developed median longitudinal sulci.Pronotal base opposite mesoscutellum (posterior pronotal lobe) with or without notch.Prosternum not acutely narrowing posteriorly, prosternal process broad and parallel-sided.Anterior margin of metasternum broadly concave.Mesoscutellum quadrate.Metascutellum not exposed.Sutural serration of elytra completely absent or weakly developed.If sutural elytral serration present, well developed beyond middle of suture towards the apex.Elytral tubercules not well developed, frequently with velvety, discrete spots and microsculpture different from rest of body surface.Tibiae slightly curved, convex dorsally, with sharp edge dorsomedially.Fore-and midtibial apices without spine.Tarsal claw appendiculate.
mm.General body shape cylindrical.Body usually yellowish.Frons glabrous, canthus of eye as yellow as rest of frons.Pronotum and elytra glabrous.Head not completely retracted into prothorax; mandibles enlarged in males, normal in females (Figs.4 K, L).Antenna serrate beyond 3 rd antennomere, 2 nd antennomere slightly widened, globose, 5 th antennomere as large as 6 th .Pronotum medially elevated, with small protuberances.Pronotal base opposite mesoscutellum (posterior pronotal lobe) with well diff erentiated notch.Prosternum strongly and abruptly constricted beyond anterior margin; prosternal process more than ¾ as long as prosternum.Anterior margin of metasternum concave.Mesoscutellum short, transverse.Metascutellum concealed by elytra.Sutural serration of elytra complete.Elytral tubercules poorly developed, their microsculpture not diff erent from rest of body surface.Tibiae slightly curved, more or less cylindrical, with sharp dorsal edge and one more less developed ventral ridge.Fore-and midtibial apices without spine.Tarsal claw bifi d or appendiculate. Diagnosis.