The genus Cephaloleia Chevrolat, 1836 (Coleoptera, Chrysomelidae, Cassidinae) Author Staines, Charles L. Author Garcia-Robledo, Carlos text ZooKeys 2014 436 1 355 http://dx.doi.org/10.3897/zookeys.436.5766 journal article http://dx.doi.org/10.3897/zookeys.436.5766 1313-2970-436-1 4AE52FD68CF948DCAA79C15AD75FF7F1 Taxon classification Animalia Coleoptera Chrysomelidae Cephaloleia Chevrolat, 1836 Cephaloleia Chevrolat 1836 : 30. Type species: Hispa nigricornis Fabricius, designated by Staines 1991(1992) : 247. Chevrolat 1843 : 350 (noted); Blanchard 1845 : 182 (redescription); Orbigny 1845 : 60 (noted); Erichson 1847 : 151 (noted); Guerin-Meneville 1855 : 601 (faunal list); Baly 1858 : 39 (redescription), 1869 : 367 (noted), 1875 : 74 (noted), 1885 : 8 (distribution); Chenu and Desmarest 1870 : 341 (noted); Chapuis 1875 : 277 (redescription); Chenu 1884 : 341 (noted); Sharp and Muir 1912 : 567 (male genitalia); Maulik 1916 : 568 (museum list), 1932 : 294 (larva), 1933 : 935 (host plants); Uhmann 1930a : 232 (Costa Rica species), 1936a : 109 (noted), 1948a : 217 (noted), 1957a : 14 (catalog), 1964a : 402 (catalog); Lepesme 1947 : 529 (host plants); Guerin 1953 : 97 (faunal list); Buck 1958 : 146 (museum list); Beutelspacher and Batze 1975 : 159 (host plants); Wilcox 1983 : 136 (catalog); Gilbert and Smiley 1978 : 90 (noted); Seeno and Wilcox 1982 : 159 (genera); Machado-Allison et al. 1983 : 248 (noted); Jolivet 1988 : 14 (host plants), 1989 : 303 (host plants); Strauss 1988 : 95 (noted); Naeem 1990 : 31 (ecology); Staines 1991(1992) : 247 (type species), 1996 : 4 (Central America species), 1996(1997) : 13 (Nicaragua species), 1997 : 413 (Uhmann species list), 1999 : 240 (mimicry), 2002 : 731 (key to genera), 2004 : 311 (host plants), 2009a : 21 (redescription); Mariau 1994 : 254 (noted); Jolivet and Hawkeswood 1995 : 143 (host plants); Mexzon 1997 : 28 (host plant); Staines and Staines 1999 : 523 (Baly species list); Jolivet and Verma 2002 : 61 (noted); Erwin and Medina 2003 : 13 (predator); Arroyo et al. 2004 : 203 (host plants); Farrell and Sequeira 2004 : 175 (evolution); McKenna and Farrell 2005 : 118 (phylogeny), 2006 : 10949 (phylogeny); Strong and Sanderson 2006 : 10827 (phylogeny); Williams 2006 : 201 (noted); Chaboo 2007 : 44 (noted); Frank and Barreta 2010 : 8 (predator); Garcia-Robledo et al. 2010 : 50 (noted); Lawrence et al. 2011 : 13 (phylogeny); Sekerka et al. 2013 : 303 (noted); Schmitt and Frank 2013 : 57 (biology). Cephalolia Blanchard 1845 : 162 (misspelling). Gemminger and Harold 1876 : 3601 (catalog); Donckier 1899 : 547 (catalog); Weise 1910 : 82 (redescription), 1911a : 7 (catalog), 1911b : 9 (catalog); Bruch 1915 : 375 (faunal list); Handlirsch 1925 : 666 (classification); Uhmann 1936b : 481 (key), 1942 : 94 (morphology); Bryant 1942 : 205 (faunal list); Monros and Viana 1947 : 162 (Argentina species); Guerin 1953 : 97 (faunal list). Uhmannispa Monros and Viana 1947 : 172. Type species: Uhmannispa maculata Monros and Viana 1947 , by monotypy. Uhmann 1957a : 14 (synonymy); Staines 1995 : 863 ( Monros species list). Description. Body elongate, rather subparallel (rarely oval), flat or moderately convex. Head: small; eyes oval, convex, finely faceted, slightly prominent (Figs 19-20); labrum rather large (Fig. 19), anterior margin rounded; maxillary palps with palpomere 1 short, 2 oblong conic, 3 shorter than 1 or 2, 4 subequal in length to 2, truncate at apex (Fig. 19). Antenna: filiform, slightly thickened at apex. Pronotum: quadrangular, square or transverse; frequently widest just behind apical angle; usually margined laterally, sometimes canaliculate; basal margin bisinuate or occasionally biangulate. Scutellum: short; pentagonal or triangular (Fig. 21). Elytron: variable in form and color; with 10 puncture rows plus a short scutellar row; very narrowly margined (Fig. 22); one segment of abdomen exposed. Venter: prosternum strongly contracted between coxae, truncate at base; mesosternum short, transverse; metasternum larger; suture between abdominal sterna 1 and 2 often obsolete (at least in middle). Leg: short; femur dilated in middle; tibia short, dilated toward apex, obliquely truncate at apex; tarsi wide, short; claws divaricate (Figs 25-26). Larval morphology. In general, larvae of Cephaloleia Chevrolat are rounded oval, longer than wide, with even, regular margins formed by wide expansion of all segments from pronotum to caudal abdominal segment forming a scale-like shield (Figs 27-54); head and legs concealed by broadly flattened margins (Figs 27-54); margins can display setae (Figs 14-18); expansions extending far forward in front of the head for a distance much greater than the width or length of the head (Figs 27-54), beyond the thorax at the sides to a width greater than 1/2 the width of the body proper and beyond the abdomen at the sides to a width wider on each side than the width of abdomen proper, width at caudal end nearly as great as at anterior end; expansions narrowly laminate; segments more or less distinct with spiracles (Fig. 13), sides plicate; elevated along central longitudinal medial line which is wider after the middle to the prothorax and narrows on tergites 7-9. Divisions between the head and the prosternum and abdominal tergites 7-9 are not clearly defined. Dorsal surface convex. Head retracted (Figs 11, 15); antenna with three antennomeres (Fig. 17). Legs consist of two distinct segments plus base; ending with a single strong recurved claw (Figs 12, 16). Taxonomic position. Historically Cephaloleia has been placed in the tribe Cephaloleiini Chapuis, 1875 ( Staines 2002 ). The tribe Cephaloleiini has been synonymized with the tribe Imatidiini Chapuis, 1875 ( Monros and Viana 1947 , Borowiec 1995 , Staines 2002 ). However, Lawrence et al. (2011) demonstrated that the true author of Imatidiini is Hope (1840) . This makes Imatidiini a senior subjective synonym of Cephaloleiini ( ICZN 1999 , Art. 23.1). The tribe Imatidiini contains 17 genera ( Staines 2002 ). Cephaloleia can be distinguished from the other genera by the following combination of characters: antennae with 11 antennomeres; mouth not projecting forward; elytra subparallel; body not cylindrical; apical margin of pronotum truncate or weakly rounded in middle; base of elytra without carina; last three abdominal sterna not hirsute; and pygidium generally exposed. Species excluded from Cephaloleia . Three species currently in Cephaloleia need to be assigned to other genera. Cephaloleia bipartita Pic, 1926c belongs to Hybosispa Weise, 1910 due to the pronotum lacking a seta in any angle, the antennae being inserted into pits and the deep excavation of the frons. Cephaloleia minasensis Pic, 1931d and Cephaloleia viridis Pic, 1931d belong to Stenispa Baly, 1858 due to the antennae being inserted into shallow pits which are divided by a longitudinal keel, the shape of the basal two antennomeres, and the cylindrical body shape. The species Cephaloleia lalli (cited in McKenna and Farrell 2006 ) is not a valid name (ICZN Art. 15). Requests for this specimen or additional information were not responded to. Remarks . Most Cephaloleia species are generally similar in appearance. Some species are easily recognized by the body shape or color pattern. Other species can only be distinguished by the sculpture of the head. Important sculpturing is the degree and strength of punctation on the vertex (Fig. 20) and the presence, absence or shape of sulci or carinae (Fig. 20). The sulci or carinae sometimes continue between the antennal bases and onto the frons. Characters on the antennae are also important. The relative lengths of the first three antennomeres and the presence or absence of triangular projections on antennomeres two to four distinguish a number of species. Antennal projections are not used in the key for some species since the presence or absence of projections is a sexual character in these species. If the pronotal margin is canaliculate (channeled or grooved) or not is extremely useful with some species. Another useful character is whether the elytra have a declivity from behind the humerus at puncture row 7. Also on the elytra the arrangement of the apical punctures is useful in species determinations. Male and female genitalia were examined and found not useful for species determinations but the shape of the last sternite is useful for gender identification (Figs 23-24). There are three groups of species which differ from the general pattern of the genus but do not clearly belong to other genera or justify erecting a new genus so are retained in Cephaloleia . The barroi-group ( Cephaloleia barroi Uhmann, 1959c and Cephaloleia sandersoni Staines, 1996) have a convex, rounded body similar to the genera Demotispa Baly, 1858 and Stilnapsis Weise, 1905b. The gracilis-group ( Cephaloleia gracilis Baly, 1878, Cephaloleia formosus Staines, 1996, and Cephaloleia vagelineata Pic, 1926c) are much more flattened than other Cephaloleia and have the elytral apex truncate. The humeralis-group ( Cephaloleia humeralis Weise, 1910, Cephaloleia obsoleta Weise, 1910, and Cephaloleia uhmanni Staines, 1996) resembles members of Stenispa but differ in several characters. Species hypotheses included in this revision are based on similarities with morphological characters of type specimens. However, molecular analyses suggest that some Cephaloleia species are not monophyletic but a complex of cryptic species ( McKenna and Farrell 2005 ). Therefore, future studies will need to combine traditional taxonomy with ecological and molecular data to elucidate species boundaries. Figures 1-10. Cephaloleia beetles and their host plants 1 Cephaloleia alternans in a Calathea ( Marantaceae ) rolled leaf. Madre Selva Station, Dto Loreto, Peruvian Amazon Scale bar equals 1 cm 2 Cephaloleia placida eggs attached to a Renealmia alpinia ( Zingiberaceae ) leaf. La Selva Biological Station, Costa Rica. Scale bar equals 1 mm 3 Cephaloleia dilaticollis first instar larva feeding on Renealmia alpinia ( Zingiberaceae ). La Selva Biological Station, Costa Rica. Scale bar equals 3 mm 4-10 Examples of Cephaloleia host plant families, Scale bars equal 10 cm: 4 Heliconiaceae ( Heliconia imbricata ). La Selva Biological Station, Costa Rica 5 Zingiberaceae ( Renealmia costaricensis ). La Selva Biological Station, Costa Rica 6 Costaceae ( Costus malortieanus ) La Selva Biological Station, Costa Rica 7 Marantaceae ( Calathea leucostachys ). Braulio Carrillo National Park, Costa Rica 8 Cannaceae ( Canna bangii ). Machu Picchu pueblo, Dto Cuzco, Peru 9 Bromeliaceae ( Pitcairnia arcuata ) Braulio Carrillo National Park, Costa Rica 10 Orchidaceae ( Oerstedella exasperata ) Quijada del Diablo, Prov. Chiriqui , Panama. (Figure 10 from Sekerka et al. 2013 ). Figures 11-14. Larva of Cephaloleia erichsonii . (La Selva Biological Station, Costa Rica) 11 Head 12 Leg 13 Spiracle 14 dorsal view of setae in the lateral margin. Figures 15-18. Larva of Cephaloleia puncticollis . (Corcovado National Park, Costa Rica) 15 Head 16 Leg 17 Antenna 18 dorsal view of setae in the lateral margin. Figures 19-26. Adult structures and sexual dimorphism in Cephaloleia beetles 19 Head and mouth parts ( Cephaloleia belti ) 20 Head, dorsal view showing carina ( Cephaloleia dilaticollis ) 21 Scutellum ( Cephaloleia belti ) 22 impressions and setae on elytron ( Cephaloleia belti ) 23-24 typical sexual dymorphism in last abdominal segment ( Cephaloleia dilaticollis ): 23 Female 24 Male 25-26 Sexual dymorphism in setose attachment pads ( Cephaloleia dilaticollis ) 25 Female tarsa with bifurcate setal tips 26 Male tarsa with bifurcate and discoidal setal tips. All specimens were collected at La Selva Biological Station, Costa Rica. Scale bar equals in all panels equal 200 μm . Figures 27-34. Cephaloleia larval stages 27-28 Cephaloleia belti , first instar, dorsal and ventral views 29-30 Cephaloleia belti , second instar, dorsal and ventral views 31-32 Cephaloleia dilaticollis , first instar, dorsal and ventral views 33-34 Cephaloleia dilaticollis , second instar, dorsal and ventral views. All specimens collected at La Selva Biological Station, Costa Rica. Scale bars in all panels equal 1 mm. Figures 35-42. Cephaloleia larval stages 35-36 Cephaloleia dorsalis at La Selva Biological Station, Costa Rica, first instar, dorsal and ventral views 37-38 Cephaloleia dorsalis , second instar, dorsal and ventral views 39-40 Cephaloleia histrionica at Braulio Carrillo National Park, 1500 m. elevation, Costa Rica, first instar, dorsal and ventral views 41-42 Cephaloleia histrionica , second instar, dorsal and ventral views. Scale bars in all panels equal 1 mm. Figures 43-50. Cephaloleia larval stages 43-44 Cephaloleia placida at La Selva Biological Station, Costa Rica, first instar, dorsal and ventral views 45-46 Cephaloleia placida , second instar, dorsal and ventral views 47-48 Cephaloleia puncticollis at Corcovado National Park, Costa Rica, first instar, dorsal and ventral views, specimen preserved in alcohol 49-50 Cephaloleia puncticollis , second instar, dorsal and ventral views, specimen preserved in alcohol. Scale bars in all panels equal 1 mm. Figures 51-54. Cephaloleia larvae 51-52 Cephaloleia erichsonii at La Selva Biological Station, Costa Rica, second instar, dorsal and ventral views 53-54 Cephaloleia orchideivora at Quijada del Diablo, Prov. Chiriqui , Panama 53 first instar, dorsal view 54 second instar, dorsal view (Figures 53-54 from Sekerka et al. 2013 ). Figures 55-59. Cephaloleia pupae 55 Cephaloleia belti (La Selva Biological Station, Costa Rica) 56 Cephaloleia dilaticollis (La Selva Biological Station, Costa Rica) 57 Cephaloleia dorsalis (La Selva Biological Station, Costa Rica) 58 Cephaloleia placida (La Selva Biological Station, Costa Rica) 59 Cephaloleia puncticollis (Corcovado National Park, Costa Rica). Scale bars in all panels equal 2 mm.