Aulacophora Chevrolat 1836

Autor: Reid, Chris, Halling, Luke, Beatson, Max
Rok vydání: 2021
Předmět:
DOI: 10.5281/zenodo.4678578
Popis: Aulacophora Chevrolat, 1836 Aulacophora Chevrolat, 1836: 378; Maulik 1936: 167; Wilcox 1972: 221; Anand & Cox 1986: 81; Barroga & Mohamedsaid 2002; Lee & Beenen 2015 = Raphidopalpa Chevrolat, 1836: 378; Gemminger & Harold 1876 (synonymy) = Rhaphidopalpa Rosenhauer, 1856: 327; Hincks 1949: 621 (synonymy) = Acutipalpa Rosenhauer, 1856: 327; Joannis 1866: 99 (synonymy) = Ceratia Chapuis, 1876: 100; Maulik 1936: 167 (synonymy) = Triaplatys Fairmaire, 1879: 113; Weise 1924: 9 (synonymy) = Orthaulaca Weise, 1892: 396; Maulik 1936: 167 (synonymy) = Cerania Weise, 1892: 396; Maulik 1936: 167 (synonymy) = Sphaerarthra Weise, 1896: 396; Maulik 1936: 167 (synonymy) = Pachypalpa Weise, 1896: 392; Maulik 1936: 167 (synonymy) = Paraulacophora Csiki, 1953: 131; Wilcox 1972: 222 (synonymy) Type species: Galeruca quadraria Olivier, 1808, by subsequent designation (Duponchel & Chevrolat 1842). This name is currently considered a junior synonym of the south-east Asian species A. analis (Weber, 1801) (Lee & Beenen 2015). Diagnostic description Generic diagnosis. The following diagnostic description is based on the Australopapuan species and the generic redefinition by Barroga and Mohamedsaid (2002). Body form elongate pear-shaped in dorsal view, length about 2x width, head, prothorax and elytra at shoulders varying slightly in proportions but generally successively wider by a factor of 1.5; dorsally sparsely and finely punctured (interspaces> puncture diameters), except pronotal anterolateral patch of secretory pits, and dorsally glabrous except trichobothria on frons and pronotal angles, semi-erect setae on clypeus, short erect setae on elytral lateral margins and some males with patch of setae posterior to elytral humeri; closely setose ventrally and on appendages. Head. Vertex smooth and impunctate, except a single trichobothrium above each eye, without longitudinal median groove; row of five pairs of long setae anterior to antennal sockets; frontoclypeus medially ridged and elevated from between antennal sockets towards anterior; anterior edge frontoclypeus truncate, shallowly concave or medially excavate; interantennal space 0.5–1.0x socket diameter, sockets level with anterior half of eyes and separated from them by 0.5–0.75x socket widths; eyes large and laterally prominent, width of head across eyes about 1.5x width of head at anterior eye margins; gena 0.5–0.7x eye length; postantennal calli distinct, slightly convex and approximately triangular, divided by short median groove, bases truncate and defined by deep postcallal groove extending laterally to posterior edge of eyes, sides defined by rim of antennal sockets, internal margins defined by grooves at base of frontoclypeal ridge; labrum with four pairs of discal setae, apical margin truncate; antennae 0.5–0.8x body length; all antennomeres elongate, 3–11 narrow and of similar width except some may be expanded in males; apical maxillary palpomere conical, shorter and narrower than preapical. Thorax. Pronotum transverse in dorsal view, widest anterior to middle, basal 2/3 straight edged and slightly contracted to base, anterior 1/3 strongly contracted to obtuse apical angles; pronotal lateral margins sinuate in lateral view; all pronotal sides with raised margins, margination broadest laterally and narrowest anteriorly (may be partly absent); anterior and posterior angles each with a trichobothrium, posterior angles about 90º; pronotum with transverse or sinuate groove at middle; prosternal process triangular, tapering to a thin elevated blade which terminates at about half length of procoxae; procoxae conical; procoxal cavities externally wide open, gap about length of hypomeral process; scutellum flat, approximately triangular with rounded apex; elytra usually at least slightly expanded from base to about 2/3 length, with rounded apex, often leaving pygidium partially exposed; elytra nonstriate, non-carinate, with or without transverse posthumeral depression; epipleura broad at base, abruptly contracted between meso and metacoxae and not defined beyond this point; fully winged; mesoventrite not covered by metaventrite, apex acutely pointed and base without procoxal rests; mesocoxae separated by much less than width of coxa; femora almost parallel-sided; tibiae with or without single apical spur, if present, length about half or slightly less than apical width of tibia metatarsus about 0.8x as long as metatibia; first metatarsomere as long as or slightly longer than metatarsomeres 2+3; tarsal claws bifid, inner teeth convergent and about 2/3 as long as outer teeth. Abdomen. Ventrites not laterally bordered by raised bead; ventrite I with rounded to broadly triangular intercoxal process; apex of male ventrite V with rectangular lobe defined on either side by slots; apex of female ventrite V truncate or concave; penis symmetrical or slightly asymmetric, apex entire, base without recurved lobes; ovipositor with well-developed apodeme (tignum) at base of sternite VIII; palpi one-segmented; spermathecal receptaculum falcate or C-shaped, transversely grooved at base and with swollen collum at insertion of duct and gland. Diagnosis of the Aulacophora indica species-complex Head capsule, pronotum and elytra entirely pale (brownish-yellow when dead, orange or red when alive) (Figs 2–9), one Japanese species, A. bipunctata (Olivier, 1808), with small black spots; male first antennomere approximately pyramidal in shape, expanded from base to middle, with a flat area on most of its anterior face (Figs 10–17); male antennomeres 3–11 not modified (Figs 18–25); male with deeper transverse prothoracic groove than female (Figs 28–29); elytra without posthumeral transverse depression, males of some species with lateral patch of erect setae posterior to humeri (Figs 18–25); all tibiae with single apical spur (Fig. 24); female pygidium usually elongated beyond apex of apical ventrite (Figs 44–57); apex of male last ventrite with elongate-oval cavity on elongate-rectangular middle lobe, which is defined on either side by narrow slots (Figs 32–43); male tergite VIII thickened and modified into either a transverse bar or a pair of posteriorly directed prongs (Figs 83–93); penis asymmetricin dorsal view, except A. wallacii (Figs 94–108). Hostplants where known are predominantly cucurbits (Cucurbitaceae) but may also include legumes (Fabaceae). The group of species designated here as the Pale Pumpkin Beetles (PPB) or A. indica species-complex (Appendix 1) is a group of externally similar species that are difficult to separate. The following described species belong in this complex, based on examination of material, or descriptions: A. abdominalis, A. bipunctata, A. cornuta, A. foveicollis (Lucas, 1849), A. indica, A. kotoensis Chujo, 1962, A. relicta, A. sulaksonoi Mohamedsaid, 2009, A. wallacii and A. wilsoni. Although the Javanese endemic A. bipunctata has small elytral spots, it otherwise conforms to the definition given above and its male genitalia are similar to those of A. indica (Barroga & Mohamedsaid 2002). The A. indica species-complex is approximately equivalent to the genus Rhaphidopalpa Rosenhauer, 1856, as defined by Weise (1892) using male and female secondary sexual characters. By including only males with setose humeri, Rhaphidopalpa was constrained to be a smaller group than the definition of PPB included here. Weise erroneously abandoned the name Aulacophora because it had been used for a plant genus, described A. indica under five different names in Rhaphidopalpa (Barroga & Mohamedsaid 2002; Lee & Beenen 2015), and at the same time made this species (under a sixth name, A. similis (Olivier, 1808)) the type of his new genus, Orthaulaca Weise, 1892. Using the current nomenclature for all the named species he listed, Weise’s Rhaphidopalpa only definitely included A. foveicollis and A. indica. In a morphological analysis of the nearby Sunda fauna, Barroga & Mohamedsaid (2006) identified a clade which included A. indica and A. cornuta, together with A. bipunctata, A. flavomarginata, A. lewisi Baly, 1886 and A. mouhoti Baly, 1886, defined by the thickening of male tergite VIII. Aulacophora mouhoti, described from the Malay Peninsula and Vietnam, is a poorly understood pale species defined by a variable female secondary sexual character and apparently only reliably known from females (Baly 1886; Barroga & Mohamedsaid 2002, 2006). We consider it unlikely to be a distinct species and as it is outside the region of interest it is not discussed further. Aulacophora lewisi has black elytra, simple male antennae and a short female pygidium (Lee & Beenen 2015) and is therefore excluded from our definition of the A. indica species-complex, although there is molecular evidence for its placement in this group (Fig. 1). Aulacophora flavomarginata has mostly black elytra, modified male antennomeres 3–5 and the female pygidium is not produced (Barroga & Mohamedsaid 2002); this species is also excluded from our definition of the A. indica species-complex. The thickened and modified male tergite VIII does not define the indica species-complex, but may have been a significant precursor for development of the other male and female abdominal characters shared by the group. Aulacophora nigroscutata Baly, 1886, described from North Maluku (eastern Indonesia), is dorsally pale, with an excavate male abdomen, but is excluded from the PPB because the female abdomen is structurally quite different (Baly 1886, 1889). Two other species of Aulacophora in the region, or nearby, A. coffeae (Weber 1801) and A. bicolor (Weber 1801), may be entirely pale but they have male first antennomere simple, male last abdominal ventrite simple and female without elongated or swollen pygidium. However, these two superficially similar species are included in the key so that they can be separated from the A. indica species-complex (PPB). Further away from the region of interest, there is an entirely pale species in Sulawesi, A. unicolor Jacoby, 1883, but this lacks the male characters of the PPB complex and has distinct creamy-white antennae and is therefore also excluded from consideration (Perkins et al. 2019). The circum-Mediterranean curcurbit pest Aulacophora foveicollis (Lucas, 1849) clearly belongs in the PPB complex and is particularly similar to A. abdominalis and A. indica. Aulacophora foveicollis differs from these species by the following combination of characters: entirely pale, male pronotum with deeply arcuate central depression divided by a median ridge, penis similar to A. abdominalis but with strongly undulated profile, male tergite 8 elongate with short paired apical projections separated by semicircular concavity, female pygidium produced and sometimes bilobed at apex, apex of female sternite 7 semicircularly excavate (Maulik 1936; Berti 1990; Borowiec 2013). Excluded species names (Appendix 1) There is clearly a problem of identity with the four names coined by Boisduval (1835) for pale species of Galerucinae collected in the region and none have been identified with any certainty by subsequent authors (for example Allard 1888; Weise 1924; Lea 1924, Wilcox 1972). We have been unable to find type material. The best described, Galleruca relicta Boisduval, 1835, is discussed below, under A. relicta. The other three Boisduval names represent poorly described and unrecognisable species from either New Guinea (Galleruca flaveola Boisduval, 1835), Australia (G. punctata Boisduval, 1835) or the “Pacific Ocean” (G. scutellata Boisduval, 1835). Galleruca flaveola has been treated as a possible senior synonym of A. aruensis Weise, 1892 (Weise 1924; Wilcox 1972), itself a junior synonym of something else (see below under A. abdominalis). The description of G. flaveola is so poor (only “lutea; abdomine, pectore pedibusque nigris”: Boisduval 1835: 558) that is impossible to identify to genus. None of the Aulacophora species in the region have entirely black legs, therefore, if accurately described, G. flaveola is unlikely to be an Aulacophora. Galleruca punctata is also not identifiable to genus. If it is an Aulacophora it is most likely a junior synonym of either A. abdominalis (Wilcox 1972; Kimoto 1990) or A. relicta, and therefore of no taxonomic significance. Both Galleruca flaveola and G. punctata should be discarded, as nomina dubia, like other unidentifiable Boisduval names (Reid 2006). Galleruca scutellata has been treated as a senior synonym of the single Australian Aulacophora species with a black scutellum, A. wilsoni Baly 1888, because of Blackburn’s tentative synonymy of these two names, which ignored the notable difference in type localities (Blackburn 1890; Lea 1924; Wilcox 1972; Kimoto 1990). Blackburn noted that Boisduval’s description was “perhaps unworthy of attention” (Blackburn 1890: 361). Aulacophora wilsoni is a scarce species of high rainfall forests in southeastern Queensland and northeastern New South Wales, a species unlikely to have been encountered by collectors in the early 19 th century, and there are no pale species of Aulacophora with a black scutellum in the Pacific region. With this combination of colour and type locality, G. scutellata is more likely a senior synonym of Candezea palustris (Perroud & Montrouzier, 1864), as originally suggested by Allard (1888) and discussed by Baly (1889) and Lea (1924), the latter accepting A. wilsoni as a valid Australian species. Candezea palustris is an abundant species in the West Pacific (Beenen 2008), surprisingly not otherwise noted by Boisduval. The name Candezea palustris is stable and designates an important pest of sweet potatoes in the region (Kimoto et al. 1984). In the interests of nomenclatural stability, we think it best to discard Galleruca scutellata and also treat this name as a nomen dubium until it is formally supressed. The names Galleruca cristovallensis Montrouzier, 1856 and G. flavescens Montrouzier, 1856 are also attached to miserly descriptions, although both are regarded as valid species (Wilcox 1972). Type material of these species is probably no longer extant. The first was described from San Cristoval Island (now Makira, Solomon Islands) and has a transverse pronotal groove, parallel-sided elytra, is 4.5 “lignes” long (10 mm) and is “testacée, lisse” (smooth and brick-red coloured) (Montrouzier 1856: 70). This species is too large and wrongly coloured for a PPB, so is not discussed further here. Gallerucella flavescens was described from both Woodlark Island (now eastern Papua New Guinea) and San Cristoval Island (Solomons) and also has a transverse pronotal groove, but has expanded elytra, is 3 “lignes” long (6.8 mm) and is “entièrement jaune, sauf le dessous du corps qui est rembruni” (entirely yellow except the underside which is darkened) (Montrouzier, 1856: 71). This vague description probably applies to a species of PPB but it is impossible to say which one, and in any case all the evidence from the material to hand is that the currently valid species names for any PPB in the New Guinea and Solomons region are senior to anything described by Montrouzier. In the interests of nomenclatural stability, we think it best to discard Galleruca flavescens and treat this name as a nomen dubium. Another Montrouzier name, Galleruca artensis Montrouzier, 1861, described from New Caledonia and overlooked in revisions of that fauna (Beenen 2008, 2013), is equally impossible to identify from its four lines of description. It was originally compared with G. argyrogaster Montrouzier, 1861 (synonymised below with A. abdominalis), from which it only differed by pale antennae and legs and an unusual golden elytral spot in living specimens (Montrouzier 1861: 394). Galleruca artensis is probably a junior synonym of Aulacophora abdominalis, but it could possibly be a pale form of A. deplanchei (Perroud & Montrouzier, 1864), discussed below. Type material of G. artensis is probably lost. Since there is no reasonable way of determining the species, we think this name should also be set aside, as a nomen dubium. Entirely pale species in the region that do not belong to A. indica species-group (Appendix 2) (i) Pale colour morphs of Aulacophora deplanchei (Perroud & Montrouzier, 1864) and A. bicolor (Weber, 1801). We have only had access to typical maculate forms of A. deplanchei, including a photograph of the lectotype, so the following discussion is partly based on the confusing literature. Aulacophora deplanchei (Perroud & Montrouzier, 1864) was described from New Caledonia with black maculae on the elytra. It was then redescribed by Beenen (2008) but he later entirely rejected his own redescription without explanation (Beenen 2013). His redescription appears to have been an accidental ‘copy and paste’ of his description of A. montrouzieri Beenen, 2008 in the same paper (a preoccupied name, later renamed A. xavieri Beenen, 2013). Beenen (2008) noted that some specimens of A. deplanchei were entirely pale dorsally. This potentially makes them members of the PPB species-complex. The original description of A. deplanchei did not mention structures of the first antennomere, male ventrite 5 and female pygidium, but Beenen associated this species with the maculate species A. xavieri, which was described in detail and lacks all of the PPB diagnostic attributes. Our examination of specimens of A. deplanchei shows that the male antennae are simple, the male apical ventrite is short and flat and the female pygidium is not extended. The male genitalia of A. deplanchei are certainly similar to A. xavieri and dissimilar to any member of the PPB complex treated here (Beenen 2008). We therefore exclude A. deplanchei from the PPB complex, but as it has entirely dorsally pale morphs we include it in our diagnostic key. The presence of rows of long setae on the male antennomeres suggests that A. deplanchei might be related to both A. bicolor (Weber, 1801) and A. coffeae (Hornstedt, 1788). Aulacophora perroudi Baly, 1888, described from New Caledonia and overlooked by subsequent authors, is probably a synonym of A. deplanchei as it has dark antennae, dark elytral humeri and female pygidium rounded (Baly 1888: 177). Furthermore, the female pronotal groove is described as absent medially, which is almost true of the small amount of material to hand. However, in the absence of type material and the possibility that there is a valid spe
Published as part of Reid, Chris, Halling, Luke & Beatson, Max, 2021, Revision of the Australopapuan and West Pacific species of plain pumpkinbeetles, the Aulacophora indica species-complex (Coleoptera: Chrysomelidae: Galerucinae), pp. 1-73 in Zootaxa 4932 (1) on pages 6-13, DOI: 10.11646/zootaxa.4932.1.1, http://zenodo.org/record/4545448
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