Chalcolampra Blanchard, 1853

Leschen, Richard A. B., Reid, Chris A. M. & Nadein, Konstantin S., 2020, Generic Review of New Zealand Chrysomelinae (Coleoptera: Chrysomelidae), Zootaxa 4740 (1), pp. 1-66 : 18-20

Chalcolampra Blanchard, 1853

( Figs 2 , 3 , 18D, E )

Chalcolampra Blanchard, 1853: 328 . Type species: Chalcolampra convexa Blanchard, 1853 , by monotypy.

Allocharis Sharp, 1882: 98 . Type species: Allocharis marginata Sharp, 1882 , by monotypy. Synonymy by Daccordi, 1994:77 [as a subgenus].

Cyrtonogetus Broun, 1915: 343 . Type species: Cyrtonogetus crassus Broun, 1915 , by monotypy. Syn. nov.

Eualema Broun, 1903: 86 . Type species: Eualema walkeri Broun, 1903: 86 , by monotypy. Synonymy by Broun 1910: 77 [nec Daccordi, 1994:77; as a subgenus].

Diagnosis ( NZ species only). Body length greater than 3.0 mm and up to about 8.0 mm; elongate and moderately convex; colour uni- or bicoloured dark sometimes with a blue or green metallic luster. Antennae filiform. Procoxal cavities externally closed to very narrowly open. Hindwings present, absent, or reduced. Anterior edge of metaventrite curved to rounded or subrounded; metaventral lines parallel and not extending to middle of ventrite. First abdominal ventrite with subcoxal bead parallel to metacoxal cavity and not extending to middle of ventrite.

Description. Length 3.7–8.1 mm. Body broad and elongate, moderately convex; colour variable, generally uni- or bicoloured dark, sometimes with a blue or green metallic luster.

Head not forming a short muzzle, with genae extending a short distance beyond level of antennal and maxillary insertions, visible in dorsal view and retracted into prothorax with mouthparts directed anteroventrally; wide vertex or frons, nearly flat to feebly convex; postantennal calli present to weakly developed, but lacking pale areas. Eyes relatively well-developed, weakly or not protuberant, finely facetted and consisting of over 50 ommatidia; inner margin of eye without a small seta. Antennae long and filiform and reaching beyond hind margin of the pronotum, scape long and about 2 times longer than antennomere 2, club not well-developed; eye not contacting insertion, with distance between insertion and margin of eye less equal or less than diameter of insertion, distance between insertions about 2–5 times length of the scape. Clypeus somewhat trapezoidal to semicircular, anterior margin straight; frontoclypeal suture present; postclypeal lines present or absent. Labrum with curved sides and somewhat rectangular and typically transverse to semicircular, anterior margin emarginated or not, surface with 2–3 elongate setae per side, anterior margin at middle lacking distinct setal fringe, tormae long and thin, about 2–4 times longer than labral plate. Mandible with terebral edge simple. Maxillary palpus relatively long, palpomere 1 longer than wide, palpomere 2 not transverse and longer than wide, palpomere 3 about 2 times longer than wide or short to about as long as wide, greater to less than the length of palpomere 2, conical to cylindrical and often truncate to weakly acute (e.g., Chalcolampra speculifera ). Labium with relatively wide ligula, subequal in width to the lengths of palpomeres 1 and 2 combined, apex not divided at apex, palpal insertions separated by more than the width of the basal palpomere, palpi relatively short, palpomere 1 as wide as long, palpomere 2 longer as wide, palpomere 3 2–3 times longer than wide, conical to cylindrical. Mentum rectangular and transverse, width of mentum slightly longer or equal the length of labial palpomere 3. Intermaxillary process short and not extending anteriorly far beyond maxillary insertions, delimited behind by a ridge or a bead.

Pronotum transverse and convex to flattened, with a bead along all margins, or weakly indicated or absent posteriorly; posterior edge narrower than base of elytra; anterior margin emarginate and evenly concave, anterior angles weakly to strongly projecting, rounded to subacute; sides nearly parallel in basal half, curved, or sinuate, not strongly convergent anteriorly; posterior margin weakly convex or sinuate; posterior angles acute to subacute; disc moderately convex without sublateral groove, trichobothria present, coarse and dense punctures at middle of posterior margin absent; lateral carinae complete. Prosternum not or weakly vaulted at middle, without transverse notches in front of coxal cavities; prosternal lines absent (bead at inner margin of the procoxal cavities is present, but does not extend forward much beyond edge of coxa); prosternal process broad, short and extending a short distance behind procoxae, expanded laterally behind coxae and usually contacting an acute and elongate hypomeral process (sometimes a small gap is present); posterior margin straight, weakly convex or emarginated; procoxal cavities externally closed (or very narrowly opened). Notopleural suture distinct. Scutellary shield visible, triangular and somewhat transverse with rounded or acute apex. Elytra moderately convex, humeral calli distinct or absent, midbasal striae present or absent; surface smooth and punctation striate, moderately coarse and dense to fine and sparse; epipleura well developed, visible in lateral view, apex without ctenidium. Hind wings absent or brachypterous. Mesoventrite mostly hidden in ventral view, posterior portion between mesocoxae visible as a narrow strip; mesal part of mesoventrite with vertical surface confluent with prepectus. Meso- and metacoxae moderately to widely separated. Metaventrite typically shorter than abdominal ventrite 1 at midline (rarely longer, e.g., C. speculifera ), mesocoxal process broad to somewhat narrow with a straight, curved, or subrounded anterior margin, metaventral lines parallel and not extended to middle of ventrite, transverse metaventral (metakatepisternal) suture present but typically does not cross the midline. Metendosternite without or with a broad stalk (present in C. speculifera ) and laminae absent, widely spaced lateral arms with subapical anterior tendons. Legs with tibiae flattened to subrounded in cross section, meso- and metatibiae not clubbed with lateral edge curved and not irregular in outline, apically setose; tarsomeres 5-5-5 and variable, tarsomere 1 typically longer than T2, 1.5 times longer than wide parallel-sided or triangular, T2 triangular and may be wider than long and narrower and smaller than T3, T3 about as wide as long or longer and not deeply incised (bilobed), tarsomere 4 minute, T5 about 2–2.5 times as long as T3, appendiculate, weakly appendiculate, or simple; tarsomere 1 of male not greatly enlarged (but all tarsomeres may be, in general, enlarged).

Abdominal ventrite 1 variable in length but typically longer than ventrite 2 and 3 combined and always shorter than ventrites 2–5 combined, with a broad metacoxal process that is wider than long and with a weakly curved apical margin, ventrites 2–4 equal in length and each shorter than ventrite 1, ventrite 5 subequal to ventrite 4 or longer ventrites 3 and 4 combined with rounded or subrounded posterior margin; first abdominal ventrite with subcoxal lines parallel to coxal cavity that do not extend to middle of ventrite. Aedeagus variable, curved to weakly curved in lateral view and rounded, laterally or dorsoventrally compressed in cross section; apex in dorsal view acute to rounded or square-edged, may be expaned towards apex; flagellum present or absent. Ovipositor with paraproct 2 times longer than wide and lacking distinct baculus, coxite about 3 times longer than wide, stylus apically inserted and about 1.2 times the length of coxite. Spermatheca U-shaped, collum present spermathecal duct inserted onto base, spermathecal gland absent or very weakly sclerotised.

Comments. The species of Chalcolampra are widely distributed in Southeast Asia, Australia and New Zealand ( Reid 1993, 2006). There are 13 described species in New Zealand, and there are up to an additional 20 undescribed spp. from the South Island based on morphospecies sorted by Charles Watt contained in the NZAC. The first description of a New Zealand species was of Chalcolampra speculifera Sharp, 1882 ( Fig. 2C ), a species also described by Broun (1903) and placed in his new genus Eualema , but subsequently synonymised with Chalcolampra ( Broun 1910) . Meanwhile, Sharp (1882) named a new monotypic genus, Allocharis , based on A. marginata that he thought was closely related to the tribe Phyllocarites and was intermediate between Cyrtonus Latrielle and Gonioctena (without mentioning similarities to Chalcolampra ). Subsequently, Broun added 11 species to Allocharis and then erected a new monotypic genus, Cyrtonogetus Broun, 1915 ( Fig. 3B ). Allocharis was treated as a valid genus and the combination Chalcolampra speculifera presented in Withers et al (2015). Cyrtonogetus was retained as a separate genus by Daccordi (1994, see also 1996), and indeed there are differences between this species and other members of the group, as expressed by Broun (1915: 343): “The type of this genus is nearly related to Allocharis (Man. N.Z. Coleopt., p. 1306), but the body is not at all elongate-oval, being robust and oblong. The metasternum has the front margin distinctly elevated and truncate between the coxae, instead of being strongly, almost sharply rounded. The basal ventral segment is hardly as long as the following three, and its frontal suture is nearly quite straight. The posterior coxae are only a little farther apart than the intermediate. Tibiae gradually incrassate towards the extremity, each with a broad external groove there. Tarsi stout, basal two joints cordiform, 3rd entire and densely setose underneath, slightly emarginate, the claws of the terminal appendiculate at the base. In sternal structure it is similar to Caccomolpus , which, however, is composed of small subrotundate species with differently formed legs.” Maddison (2010) followed Daccordi’s classification, with Allocharis and Eualema listed as subgenera of Chalcolampra .

Reid & Smith (2004) treated the three New Zealand taxa Allocharis , Cyrtonogetus , and Eualema as valid and possibly the sister group to the New Caledonian endemic genus Zira Reid & Smith 2004 , but noted that the similarities were likely to be plesiomorphies within a larger group of taxa. The New Zealand and New Caledonian taxa share many similarities with Chalcolampra and are part of a larger taxonomic problem involving several genera. In New Zealand, these species form a morphological grade of completely winged forms to apterous forms that make up the bulk of species. A relatively robust and convex body with non-protuberant eyes characterises Cyrtonogetus and some Allocharis (e.g., A. subsulcata Broun ). The main difference between C. speculifera and most species of Allocharis is the posteriorly expanded and abruptly truncate elytra ( Fig. 2C ), but this feature is present, to a lesser degree, in some Allocharis species. The external closure of the procoxal cavities ( Fig. 3E ) varies from complete to narrowly open (see also Reid 1993, 2006). We believe it prudent to place the three New Zealand genera in Chalcolampra , treating Cyrtogonetus as a synonym. We also advocate dispensing with subgenera altogether until a broader study of the group is undertaken. Because of the difficulty in determining species we are unable to provide a key to the species.

The problem with Chalcolampra is broader than the New Zealand issue, and the genus may not be monophyletic. Reid (2006) suggested that Phyllocharis Dalman and Chalcolampra might be placed in synonymy, though larval characters may support their separation ( Reid 1991): this may be true in New Zealand when comparing larval forms (C. Wardhaugh, pers. obs); one form with a well sclerotised circular plate at the apex of the abdomen and others lacking this structure. Meanwhile, no adult synapomorphies have been recognised for Chalcolampra , and the genus appears twice in a key to Australian chrysomelines by Reid (2006) based on differences of the tarsal claw ( Reid 1993). Furthermore, unpublished genetic data support non-monophyly of the genus (J. Jurado, pers. com., 2009). The likelihood that the New Zealand species of Chalcolampra are paraphyletic, or of multiple origins, is indicated by two distinct larval morphologies. Clearly a complete overhaul of the genera of this group is required.

All New Zealand forms have a more or less parallel-sided body form, are relatively large (over 4 mm), have eyes that are well-developed (though they may or may not be protruding), and the antennae on the larger forms are filiform to subfiliform, lacking a club or have broadened apical antennomeres, which distinguishes the genus from other native New Zealand taxa. Host records are few for New Zealand Chalcolampra . Species in Australia feed on Senecio (Asteraceae) , Prostanthera (Lamiaceae) and Veronica (Scrophulariaceae) ( Reid 1991, 2006; Holtkamp & Hosking 1993). Members of Australian Chalcolampra s. str. feed mainly on Asteraceae and Pittosporaceae (Jurado- Rivera et al. 2009) and the New Zealand species C. speculifera has been collected from Pseudopanax (Araliaceae) , which is also the host for species of Promechus Boisduval , the only other chrysomeline known to feed on Araliaceae ( Jolivet & Hawkeswood 1995) , a rare host for any chrysomelid ( Reid & Beatson 2018). The larva of one species of Chalcolampra , similar to C. speculifera , has been observed feeding on the composite Olearia colensoi ( Wardle et al. 1971) and was illustrated by Reid (1995). It blocks its daytime retreats with an anal plate formed by terga VII–IX. Other New Zealand Chalcolampra feed on Veronica (Scrophulariaceae) . A species like C. tarsalis Broun has been collected from the subalpine species V. albicans in Northwest Nelson, where the larva feeds on leaves day and night, though adults are nocturnal (C. Wardhaugh, pers. com., 2018). Larvae of this group lack the anal plate and have been recorded previously from Veronica ( Purdie 1884; as Allocharis marginata Sharp ) and were illustrated and briefly described by Hudson (1934; as Allocharis robusta Broun ). Lastly, two species similar to C. crassus have been collected from Celmisia brevifolia , C. haasti and under Celmisia leaves ( Asteraceae ).

Distribution. South Island.

Included species. Chalcolampra crassa ( Broun, 1915) , comb. nov. (from Cyrtonogetus ), C. fuscipes ( Broun, 1917) , C. limbata (Broun, 1893) , C. marginata ( Sharp, 1882) , C. media ( Broun, 1917) , C. morosa (Broun, 1893) , C. nigricollis ( Broun, 1917) , C. picticornis ( Broun, 1917) , C. praestans ( Broun, 1917) , C. robusta ( Broun, 1917) , C. speculifera Sharp, 1882 (= Eualema walkeri Broun, 1903 , syn. nov.), C. subsulcata ( Broun, 1917) , C. tarsalis ( Broun, 1917) .

Figures

FIGURES 2A–M. Dorsal habitus of Chalcolampra spp.; A, Chalcolampra sp.; B, Chalcolampra sp., length 4.8 mm; C, Chalcolampra speculifera Sharp, length 8.7 mm; D, Chalcolampra sp.

FIGURES 3A–F. Habits and structures of Chalcolampra spp.; A, Chalcolampra sp., dorsal habitus; B, Chalcolampra crassa (Broun), dorsal habitus of holotype; C, Chalcolampra sp., lateral view, length; D, Chalcolampra speculifera Sharp, detail of head; E Chalcolampra sp., detail of head; F, Chalcolampra sp., ventral view of anterior portion of body.

FIGURES 18A–E. Subantennal grooves (ventral view of heads, A–C) and prothoracic glandular ducts. A, Maurodus cinctiger; B, Maurodus lepidus; C, Maurodus maculatus; D, Chalcolampra sp., dorsal view through cleared prothorax; E, same, detail.