Lamprosomatinae Lacordaire, 1848 77 2.7.4 Lamprosomatinae Morphology, Adults (Fig. 2.7.4.1 E – G). Length 1.8– 8 mm (largest species are in Lychnophaes and Lacordaire, 1848 the smallest in Cachiporra Chamorro & Konstan- tinov). Body between 1.5 times longer than wide Maria Lourdes Chamorro (subcircular) to as long as wide (rounded); pro- thorax basally as wide as combined elytral bases Distribution . Worldwide, approximately 250 but promptly narrowing anterad in dorsal view; species are classifi ed in 14 genera in four tribes: greatly convex with pronotal lateral margins at Cachiporrini (with a single monotypic genus approximately 90 ° angles to the anterolateral edge from Brazil), Neochlamysini (two genera from the of elytra in lateral view; anteriorly blunt, poste- Afrotropical Region), Sphaerocharini (one genus riorly inclined, in transverse plane semicircular from Brazil), and Lamprosomatini (ten genera). (ventrally fl at); smooth dorsal surface without pro- Within Lamprosomatini , only Oomorphus Curtis is nounced interstices. Color black to metallic, occa- widely distributed, with records from all biogeo- sionally multicolored, usually refl ective depending graphic regions. Oomorphoides Monr ó s is known on visual angle. from the Oriental and Australasian Regions. Lam- Head declined, anteriorly fl at, inserted into prosoma Kirby, Dorisina Monr ó s, Lamprosomatoides prothorax completely or up to frons with eyes Monró s, Lychnophaes Lacordaire, and Oyarzuna completely to barely visible from above. With- Bechyné are Neotropical endemics. Asisia Bezdek, out transverse occipital ridge or stridulatory fi le. Lö bl & Konstantinov and Scrophoomorphus Medve- Frontal region with or without median groove. dev are endemic to the Oriental region, and Xenoo- Eyes entire, not protuberant, fi nely facetted, with- morphus Monró s is restricted to the Afrotropical out interfacetal setae; canthus of each eye absent Region [Chamorro & Konstantinov 2011; Monr ó s to deeply developed. Antennal insertions not 1956, 1958, 1959; Seeno & Wilcox 1982]. exposed from above; subantennal grooves pres- ent. Frontoclypeal strengthening ridge absent or Biology and Ecology (Fig. 2.7.4.1 A – C). Lampro- weakly inversely V-shaped. Clypeal region trap- somatinae, like its sister-taxon Cryptocephalinae ezoidal. Labrum well-developed, subquadrate (collectively known as the Camptosomata), build or narrowly rectangular and appearing almost portable fecal enclosures as larvae. Both subfami- absent; anterior edge usually truncate, concave lies share similar life histories and oviposition or widely triangular. Antennae 11-segmented, behavior [Erber 1988; Chamorro, in press (this shorter than pronotum, weakly clavate to capitate volume)]. However, although larval cryptocepha- ( Cachiporra); scape subglobular to elongate and lines are largely saprophagous, larval lamproso- longer than pedicel. Mandibles usually deltoid, matines actively and sometimes destructively feed moderately elongate, gradually curved mesally, on the bark of live trees (Monró s 1956), including enlarged in some males; mola and prostheca their stems. Some species (i. e., Lamprosoma azureum absent. Maxillae each with digitate, setose galea, Germar) have been considered as potential biologi- quadrate lacinia, and digitate apical maxillary cal control agents of invasive plants (i. e., Psidium palpomere. Mentum transverse; ligula bilobed; cattleianum Sabine, the strawberry guava, Myrta- apical labial palpomeres digitate; mentum trans- ceae, an invasive species in the Hawaiian Islands verse. Gular sutures widely separated and short. and southern Florida, USA). Documented hosts Tentorium with anterior arms and bridge absent. include Bombacaceae, Combretaceae, Fabaceae, Cervical sclerites reduced. Melastomataceae, and Myrtaceae. Possible asso- Pronotum about 0.75 – 1.0 times as long as ciations with non-woody plants such as those in wide, broadly deltoid, widest basally; ante- Compositae (i. e., Artemisia Linnaeus) have been rior sides concealed ventrad; posterior margin indicated; however lamprosomatines appear to slightly narrower or as wide as combined elytral feed on woody plants. Larvae of Lamprosoma chori- bases; lateral pronotal carinae not pronounced, siae Monró s carry cases that mimic the spines on entire to sinuate, not explanate or visible from the bark of their host, Chorisia spp.; the presence of above; anterior and posterior angles rounded or the case-bearing larvae may only be detected by fol- pointed; posterior angles bearing large seta; pos- lowing their feeding trails. Feeding by this species terior edge weakly to strongly produced medi- usually takes places in the evening or at night, and ally, not margined; disc entire, distinctly to the larvae may travel some distance from their well- weakly punctate. Prosternum absent directly in concealed resting spot, a location where the larvae front of coxae, deltoid laterally, on same plane remain for most of the day and where they may as visible section of mesoventrite; antennal eventually pupate (i. e., Lamprosoma chorisiae ). Lar- grooves present along prosternal process; pros- val lamprosomatines may be preyed upon by para- ternal process complete, usually parallel-sided sitic Hymenoptera, for example those in Cryptinae to deltoid, with apex (posterior margin) truncate Kirby (Ichneumonidae) [Caxambu & de Almeida ( Lychnophaes , Oomorphus ), concave ( Lamprosoma), 1999; Erber 1988; Kasap & Crowson 1976; Kimoto or pointed (Neochlamys Jacoby), or with apical 1964; Monró s 1949, 1956; Reid 1990]. bifurcation (Sphaerocharis Lacordaire). Notosternal 78 Maria Lourdes Chamorro

Fig. 2.7.4.1 Lamprosomatinae. A, Lamprosoma seraphinum (after Fiebrig 1910 & Erber 1988); B, Lamprosoma sp. (after Fiebrig 1910, Erber 1988); C, Lamprosomatinae egg (after Erber 1988); D, Oomorphus concolor larval head, anterior view (after Reid 1990); E–G, Lychnophaes globulosus (after Monró s 1956); E, dorsal view; F, ventral view; G, lateral view.

sutures distinct. Hypomera with variable surface absent, suture entire. Mesoventrite separated sculpture, almost always smooth and concave to by complete sutures from mesanepisterna; ante- accommodate basal sections of prolegs. Procoxae rior edge on same plane as metaventrite, large not projecting below prosternum, without con- section of the mesoventrite fl attened in Crypto- cealed lateral extensions; trochantins exposed cephalinae; in Lamprosomatinae mesoventrite within coxal cavity; procoxal cavities subcircular, more or less angled and only apex on same plane very narrowly (e. g., Neochlamys , Sphaerocharis ) to as prosternum; these sections of the mesoventrite widely separated (e. g., Lamprosoma ), open, with- restricted to small overall portion of the entire out lateral extensions. Scutellar shield deltoid, mesothorax); without procoxal or prothoracic medium sized (e. g., Neochlamys) to very small rests. Mesocoxae not conical or projecting, with (e. g., Oomorphus ), not elevated, anteriorly entire, exposed trochantins; cavities broadly separated posteriorly acutely pointed; rugosity present on (greater than widest diameter of each mesocoxa), anterior concealed part of mesoscutellum. Elytra subcircular, slightly oblique or not, partly closed about 1– 1.5 times as long as wide and 1– 3 times laterally by mesepimera and small portion of as long as pronotum; regularly (with eight to ten mesanepisterna. Metaventrite with discrimen as weak puncture rows) (e. g., Lychnophaes ) or irreg- long as entire sclerite (extending to metaventral ularly punctate (e. g., Asisia ); punctures small; process); katepisternal (transverse) suture absent; sutural striae fl at; disc smooth; humeri weakly metanepisternum moderately elongate, ante- developed, without abrupt basal edge; elytral riorly deltoid narrowing medially, broadening apices meeting at median suture, concealing all posterad sometimes becoming concave/angled tergites, or exposing or partly exposing strongly to accommodate hind femora (e. g., Lychnophaes , pigmented pygidium; epipleurae well-defi ned, Lamprosoma ). Metacoxae moderately separated, vertically oriented (visible in lateral view) or hori- horizontally oriented, extending laterally to meet zontally (hidden in lateral view), narrowing api- metanepisterna; plates absent. Metendosternite cally; elytral base (posterad of scutellar shield) with short stalk; long lateral arms usually wide never exposing metascutellum; elytral serration at base; laminae well-developed and broad or Lamprosomatinae Lacordaire, 1848 79 absent (e. g., Sphaerocharis ); anterior process short orro & Konstantinov 2011; Chamorro-Lacayo & (shorter than stalk), bearing variously separated Konstantinov 2004; Chamorro-Lacayo, Konstan- anterior tendons. Hind wings with long apical tinov & Moseyko 2006; Monró s 1956, 1958; Reid fi eld, anterior and posterior remnants of RP pres- 1990; Schö ller 2008]. ent; radial cell triangular, well-developed to api- cally open (radius-median crossvein missing); RP Morphology, Eggs and Scatoshell (Fig. 2.7.4.1 short or connected to posterior arm of radial cell; C). Egg oval, with chorionic stalk. Color milky- media (M) short; subcubital fl eck absent; three to white to yellowish-white. Surface of chorion four anal veins present and cross-veins forming micropustulate. Scatoshell cone-shaped, almost one to two anal cells; in some species, veins par- tassel-hat-shaped; resembling seeds, buds, horns; tially (anal region, e. g., Oomorphus fl oridanus Horn) may contain bark or plant remnants [Caxambu & or completely reduced (e. g., Oomorphus japanus de Almeida 1999; Lee & Cheng 2007; Lee & Morim- Jacoby). Legs usually short and stout, retractable oto 1991; Monró s 1949]. and hidden in dorsal view; trochanterofemoral joints strongly oblique with base of femora sepa- Morphology, Larvae (Fig. 2.7.4.1 D). Body obvi- rate from coxae; pro- and metafemora may be ously J-shaped in lateral view, with last fi ve abdom- keeled and bearing groove to accommodate tibiae; inal segments increasingly dilated and apical tibiae generally dorsoventrally fl attened and dis- segments bent anterad; head, pronotum, and legs tally widened, in some cases almost deltoid; inner strongly sclerotized; abdomen lightly pigmented. surface smooth and fl attened; tibial spurs absent; Larvae partly contained within bell-shaped case tarsi 5-5-5 in both sexes; basal three tarsomeres made of own feces and plant material, but head and usually as long as wide; densely clothed beneath legs exposed. Setation sparse, setae usually simple with adhesive microtrichia; penultimate tarso- and elongate. mere reduced and antepenultimate bilobed; pre- Head hypognathous, usually fl attened ante- tarsal claws simple, appendiculate or deeply bifi d; riorly, without distinct circular ridged margin; empodium not visible. entire head capsule in anterior view circular. Epi- Abdomen with fi ve free ventrites and tergites cranial suture Y-shaped, with long epicranial stem I– VII. Ventrite 1 almost more than twice as long ( = coronal suture); frontal arms enclosing a broad as 2, usually longer than ventrites 2 and 3 com- V-shaped frontal area extending toward dorsal bined, without postcoxal lines; intercoxal process stemmata. Median endocarina ending before junc- narrowly rounded to almost truncate. Functional tion of frontal arms and epicranial stem. Surface of spiracles present on tergites II – VII. Tergite VII head generally smooth but with numerous setae. forming strongly pigmented pygidium, some- Usually with three to fi ve stemmata clustered into times exposed (e. g., Sphaerocharis ). Anterior edge two groups on each; total number of stemmata of sternite VIII in males without median strut. may vary, but on each side always segregated into Ventrite 5 with well-developed stridulatory fi le two discrete groups, either above or below anten- on distal border in Lamprosomatini and females nae; total number on both sides in some cases with very weak apical fovea. Males with segment unequal (i. e., Oomorphus concolor with fi ve on one IX membranous and spiculum gastrale Y-shaped. side, three on another; Lamprosma fi ve and four)]. Q: I believe this is a Aedeagus of cucujiform type; tegmen Y-shaped; Frontoclypeal suture absent. Labrum fused to typo and should be struts (remnants of tergite IX) either present or frontoclypeus, lacking median anterior projection “Lamprosoma”. absent; penis fl attened to rounded, slightly to (present in some Cryptocephalinae ); clypeolabral strongly curved apically; tufts of setae usually fusion line weak; anterior clypeolabral margin absent. Sternite VIII in females lacking spiculum generally concave to sinuate. Antennae elongate, ventrale. Ovipositor short, rigid and oval; proc- approximately two-thirds the length of head (i. e., tigers deltoid; paraprocts narrow to widely deltoid, Oomorphus ) to short (i. e., Lamprosoma ), composed of approximately of equal size to coxites, sclerotized two or three segments; inserted apically on anten- or less pigmented proximally, fl attened; digitate nomere 2 and slightly larger in size than anten- lobes of variable form, apically setose; styli digi- nomere 3; sensorium basally strongly sclerotized; tate and setose. Spermatheca strongly to moder- apical part weakly sclerotized and narrowing. Man- ately sclerotized, variably shaped, usually J-, C-, dibles symmetrical, adentate to tridentate, without or S-shaped. Rectal sclerites (“ Kotpresse ” ) pres- mola; each with pronounced globate basolateral ent in female; ventrally with large, variously- condyle. Ventral mouthparts retracted; maxillary shaped chitinpolster (e. g., Lamprosoma , Cachiporra, articulating area slender to apparently obsolete. Sphaerocharis, Pseudolychnophaes , Oomorphoides) or Cardines moderately oblique, divided; stipites dis- with median chitinpolster and two lateral, usually tinctly longer than wide; outer lobe (mala) digitate, narrow sclerites (e. g., Xenoomorphus ); dorsally with setose; inconspicuous inner lobe fused to stipites, large variously-shaped chitinpolster (e. g., Lam- bearing two or three stout setae; maxillary palps prosoma , Oomorphoides , Cachiporra ) or with median three-segmented, inserted on well-developed pal- chitinpolster and two lateral sclerites (e. g., Pseu- pifer (described as palpomere 1 of four by Monró s dolychnophaes, Sphaerocharis , Xenoomorphus) [Cham- [1949]); palpomere 3 with digitiform sensillum 80 Maria Lourdes Chamorro in groove on outer surface. Labium consisting of Lee 1993; Reid 1995, 2000; Schmitt 1996; Far- narrow prementum and prominent mentum and rell 1998). Earlier studies hypothesized Fulcida- submentum; paramental sclerites present; ligula cini ( = Chlamisini ) and Lamprosomatinae to be broad, simple, membranous, apically bearing three sister taxa (Monr ó s 1960; Suzuki 1996; Kasap & pairs of spiniform setae; shorter than two-seg- Crowson 1976), or Lamprosomatinae as the mented palps; palpomeres of equal length or seg- sister-taxon to a monophyletic Clytrini + Cryp- ment 1 longer and usually broader than 2; palpiger tocephalini (Lee 1993). However, these relation- present, transverse to elongate. Hypopharyngeal ships remain uncorroborated. Lamprosomatinae sclerome and bracon absent. are hypothesized to be monophyletic (Kasap & Pronotum sclerotized; prosternum without Crowson 1976; Duckett et al . 2004; Chamorro & armature. Meso- and metathorax with asperities Konstantinov 2011). Relationships among 11 of throughout; two pairs of egg-bursters on tuber- 14 lamprosomatine genera were recently analyzed cles with long seta and spine associated with each based on morphological characters of the adults present on meso- and metathorax (possibly also (Chamorro & Konstantinov 2011). Recognition of on abdominal segment I) in fi rst instar. Quadrate four tribes was confi rmed with Cachiporrini sister sclerites associated with legs present. Legs well- to ( Neochlamysini + ( Sphaerocharini + Lampro- developed, fi ve-segmented, slightly unequal in somatini )). With descriptions of numerous new size, hind legs longer than middle legs and these species and genera, Monró s (1948, 1956, 1958) longer than forelegs; tarsungulus long, acute, considerably enhanced our knowledge of the claw-like, with single seta. Spiracles bicameral, not diversity of this group of insects. However, given placed at ends of spiracular tubes. the recent discovery of a new tribe from Brazil and Abdomen more than twice as long as thorax, countless new species remaining to be described, with segments of similar length. Terga without our understanding of the group will continue to sclerotized plates, not extending laterally beyond improve. edges of sterna. Segment IX simple, not enclosed by sternum VIII. Last fi ve to six segments curved anterad. Anal opening transverse, laterally less pigmented sclerites bearing dense patch of setae. Acknowledgments Spiracles similar to those of thorax [Casari 2008; Caxambu & de Almeida 1999; Kasap & Crowson I thank R. G. Beutel, M. Buffington, A. Konstan- 1976; Monró s 1949; Reid 1990]. tinov, R. A. B. Leschen, and A. Norrbom for valu- able comments and suggestions on an earlier Morphology, Pupa. Exarate, yellowish-white to version of this manuscript. Special thanks are bright orange. Body tapering posteriorly. Head extended to C. A. M. Reid for granting permis- not visible from above. Antennae free, not hidden sion and making available his larval illustration under prosternal grooves. Pronotum bell-shaped. of Oomorphus . The invitation by C. Chaboo and Setae inserted on tubercles present or absent the editors to contribute this section is greatly on head (epicranium), pronotum, mesonotum, appreciated. USDA is an equal opportunity pro- metanotum, femora, and abdominal segments vider and employer. I– V and VI. Shape of tergites VI– VII variable. Pos- terolaterally directed projections on tergite VII absent. Elytrothecal lobe and urogomphi absent [Caxambu & de Almeida 1999; Monró s 1949; Reid Literature 1990]. Casari, S. & Teixeira, E. (2008): Immatures of Lampro- Phylogeny and Taxonomy. The subfamily was soma amethystinum Perty, 1832 (Chrysomelidae, originally proposed by Lacordaire (1848) as the Lamprosomatinae). – Zootaxa 1713: 39 – 46. “ Lamprosomid é es ” section (tribal-level) of the Caxambu, M. G. & de Almeida, L. M. (1999): Descricã o “ Clythrides ” (subfamily-level). Classifi cation of dos estagios imaturos e redescri ç ã o de Lamprosoma Lamprosomatinae benefi ted from generic revision azureum Germar (Chrysomelidae, Lamprosoma- by Monró s (1956), which resulted in numerous tinae). – Revista Brasileira de Zoologia 16: 2432 – 2256. Chamorro, M. L. (2013): Chrysomelidae: Crypto- taxonomic changes, most notably the recogni- Q: I cephalinae Gyllenhaal, 1913. In R. A. B. Leschen tion of Sphaerocharini as a tribe of Lamprosomat- believe & R. Beutel (eds.) Handbook of Zoology . Walter inae after more than 150 years of uncertainty the page de Gruyter, Berlin. and for the subsequent recognition of the tribe range will Chamorro, M. L. & Konstantinov, A. S. (2011): Neochlamysini . need to Cachiporrini, a remarkable new tribe of Lamproso- be added Evidence suggests Lamprosomatinae to be the matinae (Coleoptera, Chrysomelidae) from South in the sister-taxon to a monophyletic Cryptocephal- America. – ZooKeys 78: 43 – 59. proofs. inae , a clade collectively known as the Campto- Chamorro-Lacayo, M. L. & Konstantinov, A. S. (2004): somata (Kasap & Crowson 1976; Suzuki 1996; Morphology of the prothorax and procoxa in the Lamprosomatinae Lacordaire, 1848 81

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