Chrysomelidae: Gyllenhal 1813 81

New World (Coleoptera: Chryso- vas (Col., Chrysomelidae). – Revista Agronomia Noro- melidae: Cryptocephalinae). – Zootaxa 676: 1 – 46. este Argentino (Tucum á n) 2: 25 – 77. Chamorro-Lacayo, M. L., Konstantinov, A. S. – (1958): Notes on Lamprosomatinae (Chryso.). – & Moseyko, A. G. (2006): Comparative morphol- Coleopterists Bulletin 12: 29 – 33. ogy of the female genitalia and some abdominal – (1959): Los generos de Chrysomelidae. – Opera Lil- structures of neotropical Cryptocephalini (Coleop- loana 3: 1 – 337. tera: Chrysomelidae: Cryptocephalinae). – Coleop- Reid, C. A. M. (1990): Systematics of the Australian terists Bulletin 60: 113 – 134. Cryptocephalinae (Coleoptera: Chrysomelidae). Crowson, R. A. (1967): The Natural Classifi cation PhD. thesis. Australian National University, Acton. of the Families of Coleoptera. 187 pp. Classey, – (1995): A cladistic analysis of subfamilial relation- Hampton. ships in the Chrysomelidae sensu lato (Chrysom- Duckett, C. N., Gillespie, J. J. & Kjer, K. M. (2004): eloidea). Pp. 559 – 631 in Pakaluk, J. & S´ lipi n´ ski, S. Relationships among the subfamilies of Chrysome- A. (eds.) Biology, Phylogeny and Classifi cation of Cole- lidae inferred from small subunit ribosomal DNA optera: Papers Celebrating the 80th Birthday of Roy A and morphology, with special emphasis on the Crowson, Vol. II . Muzeum i Instytut Zoologii PAN, relationship among fl ea and the Galeruci- Warszawa. nae. Pp. 3 – 18 in Jolivet, J. S.-B. P. & Schmitt, M. – (2000): Spilopyrinae Chapuis: a new subfamily in (eds.) New Contributions in Biology of the Chrysomel- the Chrysomelidae and its systematic placement idae. Kugler Publications, The Hague. (Coleoptera). – Invertebrate 14: 837 – 862. Erber, D. (1988): Biology of Camptostomata Schmitt, M. (1996): The phylogenetic system of the Clytrinae – Cryptocephalinae – Chlamisinae – Chrysomelidae - history of ideas and present state Lamprostomatinae. Pp. 513 – 552 in P. Jolivet, E. of knowledge. Pp. 57– 96 in Jolivet, P. & Cox, M. L. Petitpierre & T. H. Hsiao (eds.) Biology of Chrysomeli- (eds.) Chrysomelidae Biology, Vol. 1: The Classifi cation, dae . Kluwer Academic Publishers, Dordrecht. Phylogeny and Genetics. SPB Academic Publishers, Farrell, B. D. (1998): “ Inordinate Fondness ” Amsterdam. explained: why are there so many beetles? – Science Sch ö ller, M. (2008): Comparative morphology of 281: 555 – 559. sclerites used by Camptosomatan leaf beetles for Fiebrig, K. (1910): Cassiden und Cryptocephaliden formation of the extrachorion (Chrysomelidae: Paraguays. Ihre Entwicklungsstadien und Schutz- Cryptocephalinae, Lamprosomatinae). Pp. 87 – 120 vorrichtungen. – Zoologische Jahrbuecher Systematik in Jolivet, P., Santiago-Blay, J. & Schmitt. M. (eds.) 12 Suppl.: 161 – 264. Research on Chrysomelidae . Brill, Leiden. Kasap, H. & Crowson, R. A. (1976): On systematic rela- Seeno, T. N. & Wilcox, J. A. (1982): Leaf genera tions of Oomorphus concolor (Sturm) (Col., Chryso- (Coleoptera: Chrysomelidae). – Entomography 1: melidae), with descriptions of its larva and of an 1 – 221. aberrant Cryptocephaline larva from Australia. – Suzuki, K. (1996): Higher classifi cation of the family Journal of Natural History 10: 99 – 112. Chrysomelidae (Coleoptera). Pp. 3 – 54 in Jolivet, Kimoto, S. (1964): The Chrysomelidae of Japan and P. H. A. & Cox, M. L. (eds.) Chrysomelidae Biology: Ryukyu Islands. III. – Journal of the Faculty of Agricul- Classifi cation, Phylogeny and Genetics . SPB Academic ture, Kyushu University 14: 141 – 164. Publishers, Amsterdam. Lacordaire, T. (1848): Monographie des col é opt è res subpentameres de la famille des Phytophages II. M é moires de la Socié t é Royale des Sciences de Liè ge 5: 890. Lee, C.-F. & Cheng, H.-T. (2007): The Chrysomelidae of Taiwan I. 199 pp. Sishou-Hills Observation Network Press, Taipei. 2.7.5 Chrysomelidae: Lee, J. E. (1993): Phylogenetic studies on the larvae of the Chrysomelidae (Coleoptera) from Japan. Cryptocephalinae – Japanese Journal of Entomology 61: 409 – 424. Gyllenhal 1813 Lee, J. E. & Morimoto, K. (1991): The egg and fi rst- instar larva of Oomorphoides cupreatus (Baly) from Maria Lourdes Chamorro Japan, with notes on the systematic position of Lamprosomatinae (Coleoptera: Chrysomelidae). – Journal of the Faculty of Agriculture, Kyushu University Distribution . Worldwide approximately 5300 35: 101 – 107. are classifi ed in 127 genera in three Monr ó s, F. (1948): Descripció n de diez nuevas especies tribes. Fulcidacini (most commonly and until de “Lamprosoma ” neotropicales (Col., Chrysomeli- recently treated under the name Chlamisini ), dae). Acta Zool ó gica Lilloana 5: 81 – 95. , and Cryptocephalini , with the high- – (1949): Descripci ó n de la metamorfosis de Lampro- est diversity occurring in the tropical and soma chorisiae Monr ó s y consideraci ó nes taxon ó mi- subtropical regions. Of the three tribes, Fulci- cas sobre Lamprosomatinae (Col., Chrysomelidae). dacini, the warty leaf beetles, is the least diverse – Acta Zool ó gica Lilloana 7: 449 – 466. with an estimated 500 species in 11 genera. – (1956): Revisi ó n gen é rica de Lamprosomatinae The only not present in the New World is con descripci ó n de algunos g é neros y especies nue- Hymetes Lacordaire (3 spp.) from Asia; four genera 82 Maria Lourdes Chamorro are endemic to the Neotropical region (Melittochla- tarsi hold and rotate each egg within a small fovea, mys Monr ó s, 13 spp.; Fulcidax Voet, 7 spp.; Kakita or depression, in abdominal ventrite 5. The pro- Chamorro & Konstantinov, 1 sp., and Carcinobaena cess of placing and arranging the shingles, or fecal Lacordaire, 1 sp.); three genera extend their range plates, around the egg to form the scatoshell may into the Nearctic region [Diplacaspis Jacobson, take approximately 20– 30 minutes. Eggs resemble 6 spp.; Karren, 17 spp.; Pseudochla- small seed pods. In the northern hemisph e105.] mys Lacordaire, 5 spp.] and three genera exhibit broader patterns of distribution into the Orien- Morphology, Adults (Fig. 2.7.5.1 A, I, L, M, N, tal, Australasian, Afrotropical and/or Palearctic Q, R, S). Length 0.8 mm– 10.0 mm, with the larg- regions [ Lacordaire, 26 spp.; Aulacochlamys est species occurring in Holarctic clytrines and Monró s, 32 spp.; and Rafi nesque, 400+ Neotropical Fulcidacini (e.g., Fulcidax Voet); the spp.]. The second most diverse tribe is Clytrini smallest species belong to Cryptocephalini ( Tria- with more than 1300 species in six subtribes and chus LeConte, LeConte, and some 62 genera. All 38 genera in the subtribe Clytrina , Gistel). Body 2 times longer than wide,1.5 times with the exception of Agassiz, occur in longer than wide (cylindrical), or rarely as long as the Old World; 11 genera are endemic to Africa, wide; in dorsal view parallel-sided with protho- and only Smaragdina Chevrolat is cosmopolitan. rax as wide as combined elytral bases; or rarely The subtribes Babiina (13 genera), Megalostomina rounded; anteriorly and posteriorly blunt for (seven genera), Ischiopachina (one genus), and most Cryptocephalini, some anteriorly and poste- Arateina (one genus; Brazil) are restricted to the riorly rounded, in transverse plane almost round; New World. The subtribe Eoclytrina (one genus) multicolored and patterned, particularly Crypto- and the incertis sedis genus Parantipa Medvedev cephalini, black with red humeri commonly in are endemic to Africa. Cryptocephalini consists Clytrini , brown, black, straw-yellow and some with of more than 3500 species in fi ve subtribes and 54 velvet spots in Fulcidacini , glabrous to pubescent, genera distributed worldwide. The most species particularly Clytrini , but some Cryptocephalini rich genus is Geoffroy with more (e. g., Mylassa St å l). than 1700 species. The subtribe Achaenopina (one Head not declined, anteriorly fl at; retracted genus) is endemic to southern Africa. Stylosomina into prothorax up to frons or almost completely, (three genera) is absent from the New World, with compound eyes completely to barely vis- whereas Pachybrachina (10 genera), Monachu- ible from above; without transverse occipital lina (16 genera), and Cryptocephalina (21 genera) ridge or stridulatory fi le. Frontal region rarely are found in all biogeographic regions. [Mon- with median groove. Compound eyes entire, not r ó s 1951, 1953; Crowson 1967; Seeno & Wilcox protuberant to strongly protuberant; fi nely fac- 1982; Masutti 1960; Reid 1990, 1991, 1995, 1998; etted, with interfacetal setae; weak to deep can- Moldenke 1970; Watts 2005; Sch ö ller 2002, 2007; thus present. Antennal insertions not exposed Chamorro-Lacayo & Konstantinov 2009; Bezd eˇ k from above; subantennal grooves absent. Fronto- 2010.] clypeal transverse strengthening ridge promi- nent to absent; clypeus variously shaped, usually Biology and Ecology (Fig. 2.7.5.1 C– F, J, K). A trapezoidal. Labrum well developed, square, apex detailed account of ( Cryptocephal- commonly truncate, but variably shaped. Anten- inae + Lamprosomatinae) biology was presented nae 11-segmented, longer than pronotum and by Erber (1988). Brown & Funk (2005) described fi lliform in Cryptocephalini (sometimes antenno- in detail case-making behavior and other aspects meres distally dilated and fl attened), shorter than of Neochlamisus life history ( Fulcidacini ). Crypto- pronotum and dentate in Clytrini , and clavate cephalini larvae are saprophagous, feeding on in Fulcidacini ; beginning of club variable; scape decomposing leaves in litter and on vegetation subglobular to elongate and longer than pedicel. growing on rocks, but also remove the outer layer Mandibles usually deltoid, moderately elongate, of twigs and fruits; some feed on fresh leaves and gradually curved mesally, rarely asymmetrical in seedlings. Adults are usually found on fl owers males (e. g., Chevrolat); from uni- to where they feed on pollen and petals, but also on quadridentate, lacking mola and prostheca. Max- fresh leaves. Camptosomata exhibit complex case- illae each with elongate, setose galea; lacinia vari- building and oviposition behaviors. Their unique ably shaped sometimes bilobed in Clytrini ; apical biology involves fecal enclosures provided by the maxillary palpomere digitate. Mentum trans- female to her offspring in the form of an egg-case verse; ligula bilobed; apical labial palpomeres or scatoshell, which the larvae retain and progres- digitate, apically truncate in some. Gular sutures sively build upon to create a portable dwelling. widely separated and short. Tentorium with ante- Each egg is individually coated by the female with rior arms and bridge absent. Cervical sclerites a mixture of her own feces and rectal secretions. reduced. During oviposition and during the coating process Pronotum about 0.75–1.0 times as long as wide, of each egg, the adult female rests her body for- widest basally; sides slightly rounded or sinuate; ward onto her fore- and mid-legs, while her hind base slightly narrower or as wide as combined Chrysomelidae: Cryptocephalinae Gyllenhal 1813 83

Fig. 2.7.5.1 Cryptocephalinae. A, Chlamisus perforatus Monr ó s adult, dorsal view, right side drawn without velvety spot (after Monr ós 1952); B, Pierce larva within fecal case, lateral view (after Erber 1988); C, Chilotoma scatoshell (after Medvedev 1962); D, Cryptocephalus bipunctatus (Linneaus) young larval case (after Erber 1988); E, Smaragdina scatoshell (after Medvedev 1962); F, Exema canadensis scatoshell (after Erber 1988); G, larva, lateral view (reproduced with permission of LeSage (LeSage 1985); H, Lexiphanes pupa, dorsal view (after LeSage, 1984); I, Cryptocephalus adult, dorsal view (reproduced with permission from Konstantinov); J, Labidostomis scatoshell (after Medvedev 1962); K, Metallactus scatoshell (after Erber, 1988); L, Megalostomis bubalus bubaloides adult (after Monró s 1953); M, Cryptocephalus sp., right maxilla; N, Cryptocephalus sp. prothorax, ventral view; O, Lacnabothra baccata larval head, anterior view (after Reid 1990 as ); P, Cryptocephalus sp. met-endosternite, dorsal view; Q-R, Fulcidax bacca (Kirby) adult, Q, ventral view, R, lateral view (after Monró s 1952); S, Hockingia curiosa Selman dorsal view (after Selman 1988).

elytral bases; lateral pronotal carinae present or margined or not; disc entire in Cryptocephalini absent, rarely weakly explanate; anterior and and Clytrini to greatly sculptured and sometimes posterior angles rounded or pointed, bearing large with plateaued projections in Fulcidacini . Proster- seta; posterior edge straight to medially produced, num in front of coxae usually narrow and shorter 84 Maria Lourdes Chamorro than shortest diameter of a single coxal cavity, dons. Hind wings with long apical fi eld; each wing fl at to moderately convex, sometimes produced containing small distinct sclerite and anterior and to conceal mouthparts. Prosternal process com- posterior remnants of RP; radial cell triangular, plete, usually parallel-sided; males with median well-developed to apically open (Radius-Median projection in some Cryptocephalini (e. g., Bassa- cross-vein missing); RP short to indistinct; media reus Haldeman), apex (posterior margin) truncate, (M) short; subcubital fl eck absent; three to four broadly produced (e. g., Pachybrachis Chevrolat), to anal veins present and cross-veins forming two bilobed (Cryptocephalus ), fl at on same plane with anal cells (distal 2AC and basal 1AC). Legs gen- mesothorax or not. Antennal grooves present erally well-developed; short and stout with along prosternal process in Fulcidacini . Notoster- tibiae bearing grooves to accommodate tarsi in nal sutures distinct. Hypomera of different sur- some Fulcidacini ( Carcinobaena ); trochanterofem- face sculpture, almost always smooth and concave oral joint strongly oblique with base of femur to accommodate forelegs. Procoxae not project- separate from coxa; pro- and metafemora may be ing below prosternum, without concealed lateral enlarged, particularly in males; inner and exter- extensions; trochantins exposed within coxal cav- nal surfaces of tibiae may be keeled and bear- ity. Procoxal cavities subcircular, very narrowly ing groove; tibial spurs present in some taxa; (e. g., Laicharting) to widely separated (more tarsi 5-5-5 in both sexes; penultimate tarsomere than widest diameter of coxa, e. g., Stegnocephala reduced and antepenultimate bilobed, all usu- Baly), open to closed, without lateral extensions. ally wider in males; tarsomere 3 densely clothed Scutellar shield variously shaped to seemingly beneath with adhesive microtrichia; pretarsal absent (i. e., Adiscus Gistel) abruptly elevated or claws simple to deeply bifi d; empodium not not, anteriorly convex to emarginate, posteriorly visible. concave in Fulcidacini to obtusely angulate in Abdomen with fi ve free ventrites and six ter- some Cryptocephalini ; stridulatory device pres- gites. Ventrite 1 more than twice as long as 2, ent on concealed part of mesoscutellum. Elytra usually longer than ventrites 2– 4 combined, about 1– 1.5 times as long as wide and 1– 3 times without postcoxal lines; intercoxal process nar- as long as pronotum; regularly (with eight to ten rowly rounded to almost truncate. Functional distinct puncture rows and elevated or fl at sutural spiracles present on tergites I – VI. Tergite VI striae) or irregularly punctate, punctures of vary- forming strongly pigmented pygidium, always ing size; disc greatly sculptured in Fulcidacini ; exposed; anterior edge of sternite VIII in male humeri well-developed or not, abrupt basal edge without median strut. Ventrite 5 in females with usually present; elytral apices often meeting at variably-sized apical fovea. Males with segment median suture, concealing all tergites, but inde- IX membranous and spiculum gastrale Y-shaped. pendently rounded, exposing greatly pigmented Aedeagus of cucujiform type; tegmen Y-shaped; pygidium; epipleura well-defi ned, disappearing struts (remnants of tergite IX) either present or apically; stridulatory device ( Clytrini ) present on absent; penis fl attened to rounded, slightly to underside of elytra as basolateral rounded patch strongly curved apically; apically and/or laterally and apicolateral narrow strip; elytral base (poste- usually with tufts of setae. Sternite VIII in female rad of scutellar shield) rarely exposing metascutel- lacking spiculum ventrale. Ovipositor short, rigid lum; elytral serration entire to acutely serrate and and oval with distinct proctigeral, paraproctal, interlocking in Fulcidacini . Mesoventrite sepa- and coxital baculi; paraprocts deltoid, slightly rated by complete sutures from mesanepisterna; shorter than undivided coxites, sclerotized or less anterior edge on same plane as metaventrite, pigmented proximally, fl attened, digitate lobes of without procoxal or prothoracic rests. Mesocoxae variable form, apically setose; styli absent. Sper- not conical and projecting, with exposed tro- matheca strongly to moderately sclerotized, vari- chantins. Mesocoxal cavities narrowly to broadly ably shaped, usually J-, C-, or S-shaped., Rectal separated (greater than widest diameter of meso- sclerites ( “ Kotpresse ” ) present in female; ventrally coxa); subcircular, slightly oblique or not; partly sclerite complete, without median section (two closed laterally by mesepimera and slight portion lateral sclerites), or with median section thick- of mesanepisterna. Metaventrite with discrimen ened (three sclerites apparent), variously shaped, as long as entire sclerite (extending to metaven- laterally extended into plates; dorsal sclerites tral process; katepisternal (transverse) suture complete as longitudinal strip (one sclerite), with- absent; metanepisternum moderately elongate out median section (two lateral sclerites), or with and moderately narrow; sometimes bearing white two lateral and one median section (three scler- setae. Metacoxae narrowly to widely separated, ites); additional sclerites and chitinpolsters pres- horizontally oriented, extending laterally to meet ent [Monr ó s 1951, 1953; White 1968; Reid 1990; metanepisterna; plates absent. Metendosternite Chamorro-Lacayo et al . 2004, 2006; Sch ö ller 2008] . with stalk short to elongate, lateral arms long, laminae well-developed and broad; anterior pro- Morphology, Eggs and Scatoshell (Fig. 2.7.5.1 cess short to moderately elongate (almost as long C – F, J, K). Oval, with or without chorionic stalk. as stalk), bearing variously separated anterior ten- Color milky-white to yellowish-white. Surface Chrysomelidae: Cryptocephalinae Gyllenhal 1813 85 of chorion micropustulate. Scatoshell of vari- bearing three pairs of spiniform setae; shorter ous shapes and with various surface sculpture; than two-segmented palps; palpomeres of same resembling seeds, buds, etc. A detailed descrip- length, segment 1 broader than 2; palpiger trans- tion of fecal case architecture is given by Chaboo verse to elongate. Hypopharyngeal sclerome and et al. (2008) [LeSage & Stiefel 1996; Lee & Cheng bracon absent. Gular sutures separated. Tentorial 2007] . bridge narrow. Prothorax as long as meso- and metathorax Morphology, Larvae (Fig. 2.7.5.1 B, G, O). Body combined or slightly shorter; with distinct sclero- strongly J-shaped in lateral view, with last fi ve tized plates; prosternum without armature. Meso- abdominal segments bent anterad; head, pro- and metathorax with asperities; egg-bursters on notum, and legs strongly sclerotized; abdomen tubercle with long seta and spine present in fi rst lightly pigmented. Larvae partly contained within instar. Quadrate sclerites associated with legs pres- bell-shaped case made of own feces and debris, but ent. Legs well-developed, fi ve-segmented, of equal head and legs exposed; setation sparse. size; pretarsus (tarsungulus) long, acute, claw-like; Head hypognathous; anteriorly usually fl at- strongly curved in Fulcidacini or arched, with sin- tened to concave, sometimes with distinct circular gle seta; mesocoxae relatively widely separated. ridged margin; entire head capsule resembling Spiracles annular uniforous, not placed at ends of fl attened cork (usually in Cryptocephalini and spiracular tubes. Clytrini ) or in anterior view oblong in some Fulci- Abdomen more than twice as long as thorax, dacini, to circular in Cryptocephalini and Clytrini . with segments of similar length. Terga without Epicranial suture Y-shaped, with long epicranial sclerotized plates, not extending laterally beyond stem (= coronal suture); frontal arms enclosing edges of sterna. Asperities present throughout. a broad U-shaped frontal area, or straight and Segments IX simple, not enclosed by sternum VIII. frontal area V-shaped; extending to antennal Last fi ve to six segments curved anterad. Anal open- insertions anteriorly. Median endocarina end- ing transverse. Spiracles similar to those of thorax; ing before frontal arm-epicranial stem junction spiracular-peritreme diversely-shaped among taxa. or absent1 . Rugosity and microsculpture present Chaetotaxy of head and body diagnostically and on surface of head in some Clytrini and Crypto- taxonomically important; setae from simple and cephalini . Six stemmata clustered in two groups elongate to clavate distally with minute spicules on each side: two pairs of posterior stemmata and appearing serrate; in Fulcidacini, distal setae above antennae, one pair of anterior stemmata on tibia elongate, apically fl attened, enlarged, and below antennae. Frontoclypeal suture partially curved. Reid (1990 and references therein) offer visible or absent, lateral fourth or weak depression detailed discussion on chaetotaxy nomenclature visible (usually Fulcidacini ). Labrum fused; cly- for Camptosomata ( Cryptocephalinae + Lamproso- peolabral fusion line weak; anterior clypeolabral matinae) [Monr ó s 1951, 1953; LeSage 1982, 1984 margin concave to acutely sinuate. Antennae a, b, 1985, 1986; LeSage & Stiefel 1996; Reid 1990; short, appearing two-segmented in both Clytrini Medvedev 1998; Agrain & Marvaldi 2009; Jerez & and Fulcidacini; three-segmented in Crypto- Briones 2010]. cephalini and sensorium similar in size and width to three antennal segments combined; sensorium Morphology, Pupae (Fig. 2.7.5.1 H). Exarate, weakly sclerotized, located apically on antenno- white to yellowish. Head not visible from above. mere 2 in Fulcidacini and Cryptocephalini , absent Pronotum bell-shaped. Setae on tubercles pres- in Clytrini . Mandibles symmetrical, tri - quad-, or ent or absent on head (epicranium), pronotum, quinquedentate, without mola; each with pro- mesanotum, metanotum, femora, and abdominal nounced globate basolateral condyle. Ventral segments I– V, VI. Shape of tergites VI– VII variously mouthparts retracted; maxillary articulating area shaped. Posterior podothecae bare. Laterally poste- slender to apparently obsolete. Cardines moder- riorly directed projections on tergite VII absent or ately oblique, undivided; stipites distinctly lon- fl eshy usually with subapical seta and or spinose. ger than wide; outer lobe (mala) digitate, setose; Elytrothecal lobe and urogomphi present or absent inconspicuous inner lobe fused to stipes bearing [LeSage 1984; Reid 1990]. two or three stout setae; maxillary palps three- segmented, each with well-developed palpifer; Phylogeny and Taxonomy. Cryptocephal- palpomere 3 with digitiform sensillum in groove inae tribes and genera have remained essen- on outer surface. Labium consisting of narrow, tially unchanged since their establishment by poorly defi ned prementum and prominent men- Chapuis (1874), with the exception of incorpo- tum and submentum; paramental sclerites pres- rating the former subfamilies Fulcidacini and ent; ligula digitate, simple, membranous, apically Clytrini under Cryptocephalinae following the results of a higher-level phylogenetic analysis of Chrysomelidae by Reid (1995). The monophyly of Cryptocephalinae is well supported based on 1 Agrain & Marvaldi (2009) indicated it being absent. morphological (Reid 1995, 2000; Schmitt 1996) 86 Maria Lourdes Chamorro and molecular analyses (Farrell 1998; Duckett Chaboo C. S., Brown C. G. & Funk D. (2008): Fecal case et al. 2004; Gomez-Zurita et al. 2007, 2008) of architecture in the gibbosus species group of Neoch- higher-level relationships in the Chrysomel- lamisus Karren (Coleoptera: Chrysomelidae: Cryp- idae. Tribes within Cryptocephalinae are mor- tocephalinae: Chlamisini). – Zoological Journal of the phologically uniform and each is undisputedly Linnean Society 152: 315 – 351 . monophyletic. However, relationships of the taxa Chamorro-Lacayo, M. L. & Konstantinov, A. S. within the subfamily have not been confi rmed (2004): Morphology of the prothorax and pro- by any published phylogenetic analysis. The dis- coxae in the New World Cryptocephalini (Coleo- sertation by Reid (1990) inferring the phylogeny ptera: Chrysomelidae: Cryptocephalinae). – Zoo- of mostly Australian Cryptocephalinae remains taxa 676: 1 – 46. Chamorro-Lacayo, M. L., Konstantinov, A. S. & unpublished. Approximately half of all genera Moseyko, A. (2006): Comparative morphology of of Cryptocephalini (21 genera in four subtribes) the female genitalia and some abdominal struc- are restricted to Australia and surrounding areas. tures of Neotropical Cryptocephalini (Coleoptera: However, generic synonymies may reduce the Chrysomelidae: Cryptocephalinae). – The Coleopter- total number of Australasian genera to approxi- ists Bulletin 60 (2): 113 – 134. mately 15. Evidence suggests Lamprosomatinae Chamorro-Lacayo, M. L. & Konstantinov, A. S. (2009): to be the sister-taxon to a monophyletic Crypto- Synopsis of warty genera of the World cephalinae (Reid 1995, 2000; Gomez-Zurita et al . (Coleoptera, Chrysomelidae, Cryptocephalinae, 2008), a clade known as the Camptosomata . Ear- Chlamisini). – ZooKeys 8: 63 – 88. lier studies hypothesize Fulcidacini and Lampro- Chapuis, F. (1874): Histoire Naturelle des Insectes: Genera somatinae as sister taxa (Monró s 1960; Suzuki des Col éopt è res, Vol. 10. Famille des Phytophages . iv + 1988, 1994, 1996; Kasap & Crowson 1976), or 455 pp. Libraire Encyclop é dique de Roret, Paris. Lamprosomatinae as sister-taxon to a monophy- Crowson, R. A. (1967): The Natural Classifi cation lum Clytrini + Cryptocephalini (Lee 1993); how- of the Families of Coleoptera . 214 pp. Classey, ever, these relationships remain uncorroborated. Middlesex. Circumscription of generic limits with precise and Duckett, C. N., Gillespie, J. J. & Kjer, K. M. (2004): reliable diagnostic features is lacking. Modern Relationships among the subfamilies of Chryso- revisions and cladistic analyses are sorely needed melidae inferred from small subunit ribosomal to address over- and under-splitting of natural DNA and morphology, with special emphasis on groups [Crowson 1967; Reid 1995; Suffrian 1866, the relationship among the fl ea beetles and the 1863]. Galerucinae. Pp. 3 – 18 in Jolivet, P. A., Santiago- Blay, J. A. & Schmitt, M. (eds.) New Developments in the Biology of Chrysomelidae . SPB Academic Publish- ing, The Hague. Acknowledgments Erber, D. (1988): Biology of Camptosomata Clytrinae – Cryptocephalinae – Chlamisinae – Lamprosomat- I am thankful to Drs. Alexander Konstantinov, inae. Pp. 513 – 552 in Jolivet, P. A., Petitpierre, E. & Laurent LeSage, and Chris Reid for permission to Hsiao, T. H. (eds.) Biology of Chrysomelidae , Kluwer reproduce their illustrations. I thank Drs Alexan- Academic Publishers, Dordcrecht. der Konstantinov, Richard A. B. Leschen, R. G. Beu- Farrell, B. (1998): “ Inordinate fondness ” explained: tel, Sonia Scheffer, and Allen Norrbom for valuable why are there so many beetles? – Science 281: 555– comments and suggestions on an earlier version 559. of this manuscript. The invitation by Dr. Caroline Gomez-Zurita, J., Hunt, T., Kopliku, F. & Vogler, A. Chaboo and the editors to contribute this section is P. (2007): Recalibrated tree of leaf beetles (Chryso- greatly appreciated. melidae) indicates independent diversifi cation of angiosperms and their insect herbivores. – PLoS ONE 2: e360 (1 – 8). Literature Gomez-Zurita, J., Hunt, T. & Vogler, A. P. (2008): Mul- tilocus ribosomal RNA phylogeny of the leaf bee- Agrain, F. A. & Marvaldi, A. E. (2009): Morphology of tles (Chrysomelidae). – Cladistics 23: 1 – 17. the fi rst instar larva in the tribe Clytrini, with two Jerez, V. & Briones, R. (2010): Mylassa crassicollis new descriptions in the subtribe Megalostomina (Blanchard, 1851) (Coleoptera: Chrysomelidae: (Coleotpera: Chrysomelidae: Cryptocephalinae). – Cryptocephalinae): biology and description of Zootaxa 2147: 59 – 68. immature stages. – The Coleopterists Bulletin 64 (1): Bezd eˇ k, J. (2010): Chrysomelidae: Cryptocephalinae: 31 – 38. Fulcidacini. Pp. 76 in L ö bl, I. & Smetana, A. (eds.) Jolivet, P. (1952): Quelques donné es sur la myr- Catalogue of Palearctic Coleoptera, Vol. 6. Apollo Books, mecophilie des Clytrides (Col. Chrysomeloidea). Stenstrup, Denmark. – Bulletin de l’ Institut Royal des Sciences Naturelles de Brown, C. G. & Funk, D. J. (2005): Aspects of the Belgique 28: 1 – 12. natural history of Neochlamisus (Coleoptera: Kasap, H. & Crowson, R. A. (1976): On systematic rela- Chrysomel idae): fecal case-associated life history tions of Oomorphus concolor (Sturm) (Col., Chrys- and behavior, with a method for studying insect omelidae), with descriptions of its larva and of an constructions. – Annals of the Entomological Society of aberrant cryptocephaline larva from Australia. – America 98 (5): 711 – 725. Journal of Natural History 10: 99 – 112. Chrysomelidae: Cryptocephalinae Gyllenhal 1813 87

Lee, J. E. (1993): Phylogenetic studies on the larvae – (2000): Spilopyrinae Chapuis: A new subfamily of the Chrysomelidae (Coleoptera) from Japan. – in the Chrysomelidae and its systematic place- Japanese Journal of Entomology 61 (3): 409 – 424. ment (Coleoptera). – Invertebrate Taxonomy 14 (6): Lee, C. F. & Cheng, H. T. (2007): The Chrysomelidae of 837 – 862. Taiwan I. 199 pp. Sishou-Hills Insect Observation Schmitt, M. (1996): The phylogenetic system of the Network Press, Taipei, Taiwan. Chrysomelidae – history of ideas and present state LeSage, L. (1982): The immature stages of Exema of knowledge. Pp. 57– 96 in Jolivet, P. & Cox, M. L. canadensis Pierce (Coleoptera: Chrysomelidae). – (eds.) Chrysomelidae Biology, Volume 1: The Classifi ca- The Coleopterists Bulletin 36: 318 – 327. tion, Phylogeny and Genetics. SPB Academic Publish- – (1984 a): Egg, larva, and pupa of Lexiphanes sapona- ing, Amsterdam. tus (Coleoptera: Chrysomelidae: Cryptocephali- Sch ö ller, M. (1999): Field Studies of Cryptocephalinae nae). – The Canadian Entomologist 116: 537 – 548. Biology. Pp. 421 –436 in Cox, M. L. (ed.) Advances in – (1984 b): Immature states of Canadian Neochlami- Chrysomelidae Biology . Backhuys Publishers, Leiden. sus Karren (Coleoptera: Chrysomelidae). – The – (2002): Taxonomy of Cryptocephalus Geoffroy – Canadian Entomologist 116: 383 – 409. what do we know? (Coleoptera: Chrysomelidae: – (1985): The eggs and larvae of Pachybrachis pecans Cryptocephalinae). – Mitteilungen des Internationalen and P. bivittatus , with a key to the known immature Entomologischen Vereins 27 (1 – 2): 59 – 76. stages of the Nearctic genera of Cryptocephalinae – (2007): A new genus of Cryptocephalinae from (Coleoptera: Chrysomelidae). – The Canadian Ento- Madagascar (Coleoptera: Chrysomelidae). – Linzer mologist 117: 203 – 220. Biologische Beitr ä ge 39: 693 – 702. – (1986): The eggs and larvae of Cryptocephalus qua- – (2008): Comparative morphology of sclerites used druplex Newman and C. venustus Fabricius, with a by Camptosomatan leaf beetles for formation of key to the known immature stages of the Nearctic the extrachorion (Chyrsomelidae: Cryptocephal- Genera of Cryptocephaline leaf beetles (Coleop- inae, Lamprosomatinae). Pp. 87 – 120 in Jolivet, P., tera: Chrysomelidae). – The Canadian Entomologist Santiago-Blay, J. A., Schmitt, M. (eds.) Research on 118: 97 – 111. Chrysomelidae, Vol. 1. Brill, Leiden. LeSage, L. & Stiefel, V. L. (1996): Biology and immature Seeno, T. N., & Wilcox, J. A. (1982): Leaf Beetle Gen- stages of the North American clytrines Anomoea era (Coleoptera: Chrysomelidae). – Entomography 1: laticlavia (Forster) and A. fl avokansiensis Moldenke. 1 – 222. Pp. 217 – 238 in Jolivet, P. A. & Cox, M. L. (eds.) Selman, B. J. (1962): Remarkable new chrysomeloids Chrysomelidae Studies, Volume 3: General Studies . SPB found in the nests of arboreal ants in Tanganyika Publishing Academic Publishing, Amsterdam. (Coleoptera: Clytridae and Cryptocephalidae). – Masutti, L. (1960): Ecologia ed etologica del Crypto- Annals and Magazine of Natural History 5: 295 – 299. cephalus pini L. – Bolletino di Zoological Agraria e Bachi- – (1988): Chrysomelids and ants. Pp. 463 – 473 in coltura 3: 143 – 178. Jolivet, P. A., Petitpierre, E. & Hsiao, T. H. (eds.) Medvedev, L. (1998): To the Knowledge of the North Biology of Chrysomelidae. Kluwer Academic Publish- American larvae of Clytrinae (Coleoptera, Chryso- ers, Dordcrecht. melidae). – Latvijas Entomologs 36: 36 – 43. Suffrian, E. (1863): Zur Kenntniss der sü dameri- Moldenke, A. R. (1970): A Revision of the Clytrinae of kanischen Cryptocephalen. – Linnaea Entomologica North America North of the Isthmus of Panama . 310 pp. 15: 77 – 342. Stanford University Press, California. – (1866): Zur Kenntniss der s ü damerikanischen Monr ó s, F. (1951): Revisi ó n de las especies argentinas Cryptocephalen. – Linnaea Entomologica 16: 1 – 483. de Chlamisinae (Col., Chrysomelidae). – Acta Zoo- Suzuki, K. (1988): Comparative morphology of the logical Lilloana 10: 489 – 671. internal reproductive system of the Chrysomel- – (1953): Revisi ó n sistem á tica de las especies de Cly- idae (Coleoptera). Pp. 317 – 355 in Jolivet, P. A., trinae de la Argentina, Paraguay, Uruguay y Chile Petitpierre, E. & Hsiao, T. H. (eds.). Biology of (Col., Chrysomelidae). – Acta Zool ó gica Lilloana 14: Chrysomelidae. Kluwer Academic Publishers, 5 – 274. Dordcrecht. – (1960): Notas sobre Chrysomelidae (Coleoptera). – – (1994): Comparative morphology of the hindwing Acta Zool ó gica Lilloana 17: (1959): 2 – 24. venation of the Chrysomelidae (Coleoptera). Pp. Reid, C. A. M. (1990): Systematics of the Australian Cryp- 337 – 354 in Jolivet, P. H., Cox, M. L., Petitpierre, tocephalinae (Coleoptera: Chrysomelidae). i-xviii + 887 E. (eds.) Novel Aspects of the Biology of Chrysomelidae . pp. Australian National University, Canberra. Kluwer Academic Publishing, Dordrecht. – (1991): A new genus Cryptocephalinae from Aus- – (1996): Higher classifi cation of the family Chryso- tralia (Coleoptera: Chrysomelidae). – Entomologica melidae (Coleoptera). Pp. 3 – 54 in Jolivet, P. & Cox, scandinavica 22: 139 – 157. M. L. (eds.) Chrysomelidae Biology, Vol. 1: The Classi- – (1995): A Cladistic analysis of subfamilial relation- fi cation, Phylogeny and Genetics. SPB Academic Pub- ships in the Chrysomelidae sensu lato (Chrysomel- lishers, Amsterdam. idae). Pp. 559 – 631 in Pakaluk, J. & Slipinski, Watts, J. R. (2005): A new genus and species of S. A. (eds.) Biology, Phylogeny, and Classifi cation of cryptocephaline leaf beetle (Coleoptera: Chryso- Coleoptera: Papers Celebrating the 80th Birthday of melidae) from Costa Rica. – Insecta Mundi 19 (3): Roy A. Crowson . Muzeum i Instytut Zoologii PAN, 139 – 142. Warsaw. White, R. E. (1968): A Review of the genus Cryptocepha- – (1998): Two new genera of Cryptocephalinae (Cole- lus in America north of Mexico (Chrysomelidae: optera: Chrysomelidae) from Australasia. – Journal Coleoptera). – United States National Museum Bulletin of Natural History 32: 1169 – 1179. 290: 1 – 124.