A study on the Neotropical Anthaxiini (Coleoptera, Buprestidae, Buprestinae)

Abstract Revision of the Neotropical genera of the subtribe Anthaxiina Gory & Laporte, 1839 (Coleoptera, Buprestidae, Buprestinae, Anthaxiini). Five new genera are described: Anthaxita gen. n., Charlesina gen. n., Cobosina gen. n., Marikia gen. n. and Sanchezia gen. n. Genus Agrilaxia Kerremans, 1903 is divided into two subgenera: Agrilaxia and Costiptera subgen. n. and the genus Bilyaxia Hołyński, 1989 is divided into three subgenera: Bilyaxia, Paraguayetta subgen. n. and Tomasia subgen. n. One new species is described: Anthaxita peruviana sp. n., and two informal species-groups are suggested within Agrilaxia (Costiptera subgen. n.): Agrilaxia (Costiptera) modesta (Kerremans, 1897) species-group and Agrilaxia (Costiptera) occidentalis (Kerremans, 1900) species-group. Lectotype is designated for Agrilaxia mrazi Obenberger, 1932. A key of all genera/subgenera is provided and all treated taxa are illustrated.

Head large, as wide as anterior pronotal margin; frons flat, vertex almost 3 times as wide as width of eye; antennae very short reaching anterior third of pronotal margins when laid alongside; clypeus short, transverse, anterior margin weakly emarginate.
Elytra about twice as long as wide, subparallel at anterior two thirds, regularly tapering at posterior third; elytral apices narrowly rounded, very finely serrate; elytral epipleura very narrow, narrowing posteriorly, reaching posterior fourth of elytral margin; subhumeral carina well defined; humeral swellings small; basal, elytral, transverse depression of elytra deep, narrow, reaching scutellum but interrupted by small tubercle near humeri; elytral sculpture very fine, homogeneous.
Etymology. The genus name Anthaxita gen. n. (feminine) indicates the strong similarity to the genus Anthaxia.
Differential diagnosis. Except for the characters mentioned in the key, the genus Anthaxita gen. n. differs from other Neotropical genera of Anthaxiina by the combination of the following characters: pubescence of ventral surface, very wide vertex, very fine sculpture of pronotum and elytra, serrate posterior margin of pronotum, shortened elytral epipleura, missing suture between abdominal ventrites 1-2, short lateral pronotal carina, posteriorly acuminate prosternal process, concave anal tergite (pygidium), narrowly attenuate elytral apex and by the absence of the "agriloid" carina, lateroposterior pronotal depressions, and prescutellar pit. The male is unknown.
Distribution. Peru. Note. Among all Neotropical Anthaxiini, the genus Anthaxita gen. n. is most similar to the Holoarctic species of Anthaxia, particularly to the subgenus Melanthaxia Rikhter, 1949.
Description of female holotype. Head wide, large, as wide as width of anterior pronotal margin; clypeus twice as wide as long, separated from frons by shallow, transverse impression, anterior margin widely, shallowly emarginate; frons flat with two shallow, rounded depressions above antennal insertions; vertex flat, 2.8 times as wide as width of eye; eyes relatively small, elliptical, not projecting beyond outline of head; antennae very short, finely serrate from fourth antennomere; scape pyriform, slightly flattened, 3.5 times as long as wide; pedicel suboval, 1.5 times as long as wide; third antennomere weakly triangular, 1.5 times as long as wide; antennomere 4 sharply triangular, 1.3 times as long as wide; antennomeres 5-10 trapezoidal, 1.3 times as wide as long, terminal antennomere rhomboid, somewhat longer than wide; sculpture of head consisting of small, fine, very dense polygonal cells with microsculptured bottoms.
Elytra 2.2 times as long as wide, regularly convex, subparallel, regularly tapering at posterior third; humeral swellings slightly projecting beyond outline of elytra; each elytron narrowly rounded apically, with quite indistinct serrations; subhumeral carina well defined, flat, reaching apical third of lateral margins; elytral epipleura narrow, not very well defined, becoming narrower posteriorly, reaching posterior fifth of elytral margins; elytral sculpture very fine, consisting of tiny, transverse, zig-zag rugae which are much denser on humeral portion of elytra than that on disc.
Distribution. Peru. Note. The holotype was originally pinned, so the right elytron and corresponding portion of the abdomen are somewhat damaged (see Fig. 1). Description. Medium-sized (5.5 mm), black, elongate, matt, entirely asetose; lateral margins of elytra with green tinge.
Head as wide as anterior pronotal margin; clypeus narrow, trapezoidal, anterior margin deeply emarginate; frons widely grooved; vertex flat, twice as wide as width of eye; eyes broad elliptical, not projecting beyond outline of head; antennae short and robust, scarcely reaching midlength of lateral pronotal margins when laid alongside; scape broad pyriform, twice as long as wide; pedicel subcylindrical, 1.4 times as long as wide; third antennomere short, 1.2 times as long as wide; antennomeres 4-10 short, trapezoidal, nearly twice as wide as long; terminal antennomere rhomboid, 1.5 times as wide as long; sculpture of head consisting of fine, simple, sparse punctures on frons and weakly defined, dense cells on vertex.
Pronotum twice as wide as long, with wide, deep lateroposterior depressions, without distinct sculpture; lateral margins weakly S-shaped, posterior angles right angled; "agriloid" carina well defined, rather sharp, as long as lateral pronotal carina; basal tubercles weakly defined, prescutellar pit small, deep, well defined. Scutellum small, cordiform (Fig. 31), flat, twice as long as wide, resembling the scutellum of the click-beetles of the subfamily Cardiophorinae.
Elytra elongate, 2.3 times as long as wide, widely sphenoidal with shallow, longitudinal depression at posterior fourth; humeral swellings small but well defined, basal, transverse depression deep, reaching scutellum; elytral epipleura strongly reduced, very narrow, not reaching apex of elytra; subhumeral carina well defined, reaching apical portion of elytra; elytral sculpture consisting of fine microsculpture and very small, lustrous grains. Anal ventrite with fine but sharp, lateral serrations; all tarsi distinctly widened; tarsal claws simple, strongly hook-shaped.
Etymology. The new genus Charlesina gen. n. (feminine) is dedicated to my friend and colleague, Charles Bellamy (Sacramento, California), one of the best world specialists in the taxonomy of Buprestidae, with my warm thanks for his life long cooperation.
Differential diagnosis. The monotypic genus Charlesina gen. n. strongly differs from other Neotropical Anthaxiina except for the genus Sanchezia gen. n. Both genera share some general characters like black, flattened body without distinct sculpture of dorsal surface, widened, robust antennae and tarsi, well defined lateroposterior pronotal depressions and "agriloid" carina. The genus Charlesina gen. n. strongly differs from the genus Sanchezia gen. n. by the deeply grooved frons, well defined humeral swellings and basal, transverse, elytral depression, by the narrow, reduced elytral epipleura, long, well defined subhumeral carina and by the entirely asetose body.
Length: 5.8 mm; width: 1.7 mm. Distribution. Brasil (Rio de Janeiro, Rio Grande do Sul). Note. The locality data given in the description (Obenberger, 1932) slightly differ from those on the syntype (see above); although the number of syntypes is unknown, there is no doubt that the species was described from this specimen which is hereby designated as the lectotype (see above). Cobos (1972), although treated this species in the genus Anthaxia subgen. Agrilaxia, noticed that this species should be probably treated in the genus Cylindrophora. This opinion was accepted by Bílý (1997) and Bellamy (2008). After the re-definition of the monotypic genus Cylindrophora (C. maulica) Molina, 1782)) and its transfer to the subtribe Curidina (Bílý, 2004), Agrilaxia mrazi (together with several other species of the former Cylindrophora) was formally attributed to the genus Bilyaxia. This act was overlooked by Bellamy (2008) who still treated Charlesina mrazi in the genus Cylindrophora sensu Cobos, 1956and 1972. Cobos (1972 described Anthaxia (Agrilaxia) mrazi ssp. cyaneobscura Cobos, 1972 on a single male from south-eastern Brasil (Rio Grande do Sul). This subspecies differs from C. mrazi mrazi only slightly by its colouration and by the more robust aedeagus (Figs 130, 131 in Cobos 1972 Description. Small (4.25 mm), subcylindrical, rather convex, subparallel, entirely asetose.
Head small, mouth parts small, only slightly visible, partly retracted into head; frontoclypeal portion of head rathed reduced, clypeus very short, widely transverse, with straight anterior margin; frons strongly convex; vertex about twice as wide as width of eye; eyes rather large, elliptical, not projecting beyond outline of head; antennae very short, reaching anterior third of pronotal margins when laid alongside; antennomeres 4-11 obtusely triangular to trapezoidal, wider than long.
Anal ventrite wider than long, slightly truncate apically, with very fine, lateral serrations; legs simple, male metatibiae not modified; tarsal claws thin, simple, as long as half of terminal tarsomere; aedeagus (Fig. 30) very short, widely spindle-shaped, with the maximum width at the posterior third of the parameres, which are sharply pointed; medial lobe wide, pointed apically, without lateral serrations.
Etymology. The genus Cobosina gen. n. (feminine) is named in honour of my late colleague and well-known specialist in the taxonomy of the Neotropical Buprestidae, Prof Antonio Cobos Sánchez (Almería, Spain).
Differential diagnosis. Although the genus Cobosina gen. n. resembles very much some European species of the genus Anthaxia (see below) it shares some principal characters with the Neotropical Anthaxiini: completely asetose body (only a few species of Anthaxia from south-eastern Asia are entirely asetose), prolonged scutellum which is longer than wide, narrow and shortened elytral epipleura, "agriloid" carina, posthumeral carina (the latter two rather reduced), obtusely rounded elytral apex and simple, spindle-shaped aedeagus.
The genus Cobosina gen. n. differs from other genera of the Neotropical Anthaxiina by the small, non-prognathous mouth parts which are partly retracted into the head, reduced, very short clypeus which is not separate from the frons (most of the Neotropical Anthaxiina are more or less prognathous), by the asetose protibiae (very often with the brush-like, stiff, pale setae on inner margin of protibiae in other Anthaxiina, namely Agrilaxia - Fig. 33) and last but not least by the clearly Anthaxia-like appearance.
Cobosina willineri (Cobos, 1972 Note. The species was formally attributed to the genus Bilyaxia by Bílý (2004). Already in the original description (Cobos, 1972) noted that this species is an extraordinary and isolated element of the Neotropical Anthaxiini being very similar to the European species of the Anthaxia (Anthaxia) funerula Illiger, 1803 species-group. After having studied the holotype and further, male specimen of this species from NMPC I found that A. (B.) willineri ought to be removed from Bilyaxia, and a new genus should be proposed for this species.
I failed to find any difference between the sexes and also no significant variability since only two specimens were available for study.
Length Description. Medium-sized (4.0-5.5 mm), flat, black, lustrous, entirely asetose. Head as wide as anterior pronotal margin; clypeus broadly trapezoidal, anterior margin finely emarginate; frons flat to weakly depressed, vertex flat, 2.5 times as wide as width of eye; eyes reniform to elliptical, not projecting beyond outline of head; antennae filiform, slightly overlapping midlength of lateral pronotal margins when laid alongside; pedicel as wide as scape, 1.5 times as long as wide; antennomeres 4-10 sharply triangular to trapezoidal, terminal antennomere prolonged, 2.5 times as long as wide.
Aedeagus ( Differential diagnosis. The genus Marikia gen. n. somewhat resembles by its dark colouration, flattened body and narrowly acuminate elytra some small species of the tribe Melanophilini Bedel, 1921. Among the Neotropical Anthaxiina it is most similar to the genus Anthaxita gen. n. from which it differs (except for the characters given in the key) by the lustrous and flat, entirely asetose body, narrow clypeus, longer and slender antennae, indistinct pronotal sculpture, long, sharp "agriloid" carina, triangular scutellum, posteriorly widened prosternal process, uneven and somewhat caudiform elytra, strongly elevated elytral suture and by the rudimental subhumeral carina.
Distribution. Ecuador. (Cobos, 1956) No difference between both sexes were observed and also no variability in the colouration apparent.
Head wider than anterior pronotal margin, eyes large, elliptical, projecting beyond outline of head; clypeus short, wide, anterior margin straight; frons flat with small, rounded cells with tiny central grains (central portion of frons with rather sparse cells), vertex flat, 1.6 times as wide as width of eye, with fine, transverse rugae; antennae reaching posterior fourth of lateral, pronotal margins when laid alongside; pedicel triangular, antennomeres 3-10 widely triangular to trapezoidal, much wider than long.
Pronotum tapering anteriorly, 2.2 times as wide as long, with fine, medial, longitudinal depression and large, deep lateroposterior depressions reaching anterior fourth of pronotum; "agriloid" carina well defined, long, reaching beyond midlength of lateral margins; prescutellar pit missing; pronotal sculpture consisting of very fine, rather indistinct and transversely widened, polygonal cells without central grains. Scutellum very small, triangular, 1.5 times as long as wide.
Etymology. The genus Sanchezia gen. n. (feminine) is named in the honour of my late colleague Prof Antonio Cobos Sánchez (Almería, Spain), the well-known specialist in the taxonomy of the Neotropical Buprestidae.
Differential diagnosis. The genus Sanchezia gen. n. is quite unmistakable, characteristic genus which is somewhat similar to the genus Charlesina gen. n. (see above). It differs from other taxa of the Neotropical Anthaxiini by the rather non-Buprestid appearence, very fine, nearly velvet texture of dorsal surface, robust antennae with antennomere 3 triangularly enlarged, very wide elytral epipleura and other characters mentioned in the key.
The genus Agrilaxia was described by Kerremans (1903) and 19 mostly Neotropical species were included without designation of a type species. Agrilaxia flavimana Gory, 1841 was subsequently designated by Chamberlein (1926) as the type species of the genus. Later on, the genus Agrilaxia was synonymized by Théry (1930) with Anthaxia Eschscholtz, 1829, ressurected again as a valid genus by Obenberger (1930), then finally treated as a subgenus of Anthaxia by Cobos (1972); this concept was followed also by Bílý (1984Bílý ( , 1993. Bílý & Bellamy (1999) upgraded Agrilaxia again to the generic level which was followed by Bílý (2000Bílý ( , 2004 and Bellamy (2008).
Even after separation of some species as independent genera (see above), the genus remains still rather heterogeneous. I propose that first, the small and rather homogeneous group of species with short, green or violet, wedge shaped body should be split as the separate subgenus Costiptera subgen. nov (see below).
Diagnosis. Body short, wedge shaped, ventral surface with short but distinct, white pubescence, metepisterna and metacoxae often with a patch of white tomentum; dorsal surface green, golden green or violet-green, sometimes pronotum black or violet with two longitudinal, green stripes. Head large, often wider than anterior pronotal margin, weakly prognathous; clypeus short, transverse, anterior margin straight; frons widely, deeply grooved; vertex shallowly depressed, about twice as wide as width of eye; antennae short, rather robust, reaching midlength of lateral pronotal margins when laid alongside; sculpture of frons consisting of small, dense, polygonal cells with small central grains.
Etymology. The name of the new subgenus Costiptera subgen. n. (feminine) is composed of the Latin substantive "costa" (carina, rib) and Greek substantive "pteron" (wing) to stress one of the main characters of the subgenus.
Differential diagnosis. The subgenus Costiptera subgen. n. differs from the nominate subgenus particularly by shorter, wedge shaped elytra (usually 2.5 times as long as wide), pubescence of the ventral surface and by the set of characters given above and in the key.
Note. Two species groups can be separated within the subgenus. Agrilaxia (Costiptera) modesta (Kerremans, 1897) species-group which is characterised by the smooth elytra, well defined central grains in the fine frontal and pronotal ocellation, shorter, pentagonal scutellum which is as wide as long and by the shorter subhumeral carina reaching only midlength of elytral margins and by the anterior margin of the prosternum bearing a peg-like spine (like in Figs 29, 34f); only A. (C) modesta can be included into this species-group. For the remaining species, the A. (C.) occidentalis (Kerremans, 1900) species-group is suggested which is characterised by the longitudinal elytral costae, frontal and pronotal sculpture without central grains, prolonged scutellum which is 1.5 times longer than wide and by the long, well defined subhumeral carina reaching the elytral apex.
According to Bellamy (2008), the genus Agrilaxia comprises 88 species. After removing of 5 species from Agrilaxia to Bilyaxia (see below) and after transferring of 5 species into the subgenus Costiptera subgen. n., 78 species remain in the subgenus Agrilaxia s. str. Quite naturally the subgenus will have to be split into several (or many) species-groups since several very different morphotypes (e.g. Figs 6, 19-25) can be found within the subgenus -some of them have been suggested by Bílý & Brûlé (in press).
Note. The very special morphological character of the subgenus Agrilaxia is the shape of the prosternum (Figs 29, 34a-34f) which is normally flat or slightly convex. In the most species the prosternum is more or less transversely grooved just behind the anterior margin. The "grooving" of the anterior portion of prosternal plate and the relative elevation of the anterior margin of the prosternum which, in the extreme case, forms a sharp, transverse, rolled up ledge which is sometimes transformed into a peglike, medial spine (Figs 29, 34f). The genus was originally described by Hołyński (1989) as a subgenus of Anthaxia and it was upgraded to the genus level by Bílý (2004). According to Bellamy (2008) it contains 18 species but the division to several subgenera is necessary. Three species were transferred to the separate, monotypic genera (Charlesina gen. n., Cobosina gen. n. and Marikia gen. n. -see above), the remaining 15 species is assigned into three subgenera.
Head large, very often somewhat wider than anterior pronotal margin; frons flat or slightly depressed, vertex 1.3-1.8 times as wide as width of eye; sculpture of head consisting of dense, oval or polygonal cells with or without central grains Pronotum flattened, lateral margins subparallel or regularly rounded, posterior margin weakly biarcuate or almost straight, lateroposterior depressions usually well defined; "agriloid" carina short, almost parallel with lateral carina; prescutellar pit missing, basal tubercles small or missing. Scutellum small, subcordiform, usually 1.5 times as long as wide, flat.
Note. Bilyaxia auronotata was described from the Argentinian province Neuquén (Lago Lacar). The lake Lacar lies at the Chilean border at the lowest spot of this part of the Andean range; here is the only "gate" where the Buprestid species can pass through the Andes. B. auronotata was rather frequently collected also in the opposite side in Chile (prov. Villarica).  (Kerremans, 1900), female, 5.5 mm (Brasil, Nova Teutonia).
The distribution of the genus Bilyaxia is similar in distribution to the genera Conognatha Eschscholtz, 1829 and Dactylozodes Chevrolat, 1838, the subgenera of these are also distributed on the opposite sides of the Andes.
Head large, as wide as anterior pronotal margin; clypeus trapezoidal, separated from frons by deep, transverse depression, anterior margin rounded; frons deeply, widely grooved, vertex almost flat, about twice as wide as width of eye; antennae slightly extending beyond midlength of lateral pronotal margins when laid alongside, antennomeres 5-10 widely trapezoidal; sculpture of head consisting of small, oval cells with large, flat central grains which are dense along eyes and very sparse in middle.
Pronotum convex, 2.2 times as wide as long, posterior margin deeply biarcuate; lateral margins slightly S-shaped, posterior angles sharp; "agriloid" carina well defined, reaching posterior third of lateral margins; lateroposterior depressions and basal tubercles weakly defined, prescutellar pit large, deep; sculpture consisting of basal microsculpture and dense, weakly defined polygonal cells without central grains. Scutellum cordiform, somewhat longer than wide, convex.
Ventral surface roughly ocellate, prosternum slightly convex with shallow transverse groove just behind anterior margin which is in this way transformed into fine, transverse, perpendicular ledge (as in Fig. 34e); both anal ventrite and tergite simply rounded, not serrate. Legs moderately long, all tarsi shorter than tibiae; tarsal claws strong, simply hook-shaped.
Male unknown.
Etymology. The subgenus Paraguayetta subgen. n. (feminine) is named after the country of the origin (Paraguay).
Differential diagnosis. The subgenus Paraguayetta subgen. n. differs from other subgenera of Bilyaxia by the large, robust body, anteriorly rounded clypeus, S-shaped lateral, pronotal margins, weakly defined lateroposterior pronotal depressions, large and deep prescutellar pit, ledge-shaped anterior prosternal margin (as in Fig. 34e) and by the simple, not serrate anal ventrite.
Diagnosis. Small to medium sized (3.3-4.8 mm), slender, subcylindrical species. Head relatively large, slightly wider or as wide as anterior pronotal margin; clypeus wide, trapezoidal, anterior margin straight or weakly emarginate; frons flat or convex, rarely weakly grooved; vertex flat or weakly convex, 1.8-2.2 times as wide as width of eye; eyes large, reniform, often slightly projecting beyond outline of head; sculpture of head consisting of small, fine, oval or polygonal cells.
Pronotum subcylindrical or lateral margins regularly rounded, 1.8-2.0 times as wide as long (exceptionally lateral margins S-shaped: B. (T.) hayeki (Cobos, 1972)); lateroposterior depressions wide, well defined; "agriloid" carina fine but well defined, usually reaching posterior third of lateral margins; prescutellar pit missing, basal tubercles weakly defined or missing; pronotal sculpture consisting of fine, poorly defined, polygonal cells without central grains and usually also by very fine basal microsculpture. Scutellum small to very small, cordiform, longer than wide.
Elytra regularly convex, smooth or with more or less defined, longitudinal costae, 2.2-2.4 times as long as wide; humeral swellings small but well defined, basal transverse depressions usually wide, deep, rarely shallow but always reaching scutellum; elytral epipleura narrowing posteriorly, reaching elytral apex; each elytron narrowly rounded, sometimes slightly caudiform; subhumeral carina well defined, usually reaching apical portion of elytra, sometimes shortened, scarcely reaching elytral midlength.
Etymology. The subgenus Tomasia subgen. n. (feminine) is named after my friend and colleague Tomás Moore Rodríguez (Santiago, Chile), the well-known specialist in the taxonomy of Chilean Buprestidae.
Note. Bilyaxia (T.) cinctipennis (Fig. 15) somewhat contradicts the definition of the subgenus by its colouration (black and yellow pattern) and by the widely rounded elytral apices. In 2010 I found in Paraguay (Prov. Presidente Hayes) a specimen with entirely yellow elytra.
Note. The nominotypical subspecies possesses rather strong sexual dichromatism (Figs 17, 18). The female is rather similar to P. bimaculata litigiosa and only the different form of the male genitalia (Figs 45-48 in Cobos, 1956) justifies treating the subspecies litigiosa as a valid taxon.
The genus was originally described by Théry (1934) as a subgenus of Tetragonoschema Thompson, 1857 which was accepted also by Bellamy (2008). The subgenus was subsequently upgraded to a genus by Bílý (2012). The genus Anilaroides contains only two very similar species distributed in Brasil and revised by Bílý (2012).  Thomson, 1857 (by monotypy; currently junior subjective synonym of T. quadratum (Buquet, 1841)).
The genus contains 17 species distributed from Mexico to Patagonia. The genus was quite recently revised by Bílý (2012) and divided into two subgenera.
Tetragonoschema (Tetragonoschema) s. str. Fig. 27 Remarks. The nominotypical subgenus contains 14 black, bronze or multicolorous, shortened (sometimes nearly as wide as long ) species; elytra usually very short, flat, conspicuously uneven with one common, medial depression and with deep lateral and preapical depressions; elytral epipleura wide, well-developed, reaching or nearly reaching elytral suture; frons usually deeply impressed with projecting prominences above antennal insertions, rarely frons convex; pronotum weakly convex, transverse, usually with more or less distinct lateroposterior depressions; antennae and tarsi black; aedeagus spindle-shaped or elongate, parameres very often with lateral hooks or spines. The subgenus is distributed from Mexico to central Argentina. Fig. 28 Type species. Tetragonoschema patagonicum Obenberger, 1922 by original designation.
Note. Tetragonoschema (P.) patagonicum (Fig. 28) is the southernmost distributed species of the genus Tetragonoschema (Patagonia, Santa Cruz). The province of Santa Cruz is situated south of latitude 46 where the climatic conditions are rather extreme and T. (P.) patagonicum is most probably one of the southernmost distributed species of the Neotropical Buprestids.