Cheilonycha auripennis auripennis ( Lucas, 1857 )

Moravec, Jiří, 2019, Taxonomic and nomenclatorial revision within the Neotropical genera of the subtribe Odontocheilina W. Horn in a new sense- 22. Genus Cheilonycha Lacordaire, 1842 (Coleoptera: Cicindelidae), Zootaxa 4700 (4), pp. 501-534 : 510-519

Cheilonycha auripennis auripennis ( Lucas, 1857) sensu orig.

Cicindela auripennis Lucas, 1857: 31 , pl. 1a, figs 1a, 1b.

Type locality. “ Brésil intérieur” .

Odontochila (Chilonycha) auripennis angustedilatata W. Horn, 1922: 104 , fig. 11, syn. nov.

Type locality. Paraguay .

Cicindela (Euryoda) auripennis: Dokhtouroff (1887: 154) .

Odontochila auripennis: Horn 1910: 203 (in “gruppe Chilonycha ”).

Odontochila (Chilonycha) chalybea auripennis: Horn 1934: 124 .

Odontochila (Chilonycha) chalybea angustedilatata: Horn 1934: 124 .

Cheilonycha auripennis auripennis: Wiesner 1992: 84 .

Cheilonycha auripennis angustedilatata: Wiesner 1992: 84 .

Misapplication. Non Odontochila auripennis: Horn 1910: 203 (as in “gruppe V Chilonycha ”) partim; nec Cheilonycha a. auripennis: Horn 1922: 103 , fig. 10; nec Odontochila (Chilonycha) chalybea auripennis: Horn 1934: 124 ; nec Cheilonycha a. auripennis sensu Wiesner (1992) and sensu auctorum (partim), which is partly Cheilonycha bucephalauripennis sp. nov.

Type material of Cicindela auripennis Lucas. Lectotype (designated here) ♀ in MNHN, bearing a green circu- lar label with “46/45” on its opposite side // “Muséum Paris / Amerique / de Castelnau 1845” [handwritten] // “Lec- totype / Cicindela / auripennis Lucas, 1857 / design. Jiří Moravec 2019 ” [red, printed]. Paralectotype. 1 ♀ in MNHN with same green circular label except for: “84/46” on its opposite side and “Muséum Paris / Amerique / de Castelnau 1845” // “Revision Jiří Moravec 2019: / “ Paralectotype / Cicindela / auripennis Lucas, 1857 ” [red, printed].

Type material of Odontochila (Cheilonycha) auripennis angustedilatata W. Horn syn. nov. Lectotype (designated here) ♂ in SDEI, labelled: “ Paraguay / Dr. Drake 188?” [missing the year, printed, with thin black frame, brownish tarnished] // “Type / W. Horn” [printed] // “Coll. W. Horn / DEI Eberswalde” [printed] // “ Syntypus ” [red, printed] // “f. anguste-di-/ latata / mihi” [large collection label with black frame, handwritten] // “ Lectotype / Odon- tochila ( Chilonycha ) / auripennis / angustedilatata W. Horn, 1922 / design. Jiří Moravec 2017” [red, printed]. Para- lectotypes. 1 ♂, 4 ♀♀ in SDEI with same labels as lectotype, one of them confusedly standing under a large greenish collection-label “auripennis / v.[vidit?] Dokht.[Dokhtouroff]”. 4 ♀♀ in SDEI with same labels as lectotype. 1 ♀ in SDEI: “Rep. Argentina / Prov. Catamarca / 2. 1896 / C. Bruch [leg.] // “Coll. W. Horn / DEI Eberswalde” [printed] // All paralectotypes labelled: “Revision Jiří Moravec 2015: / “ Paralectotype / Odontochila (Chilonycha) / auripennis / angustedilatata W. Horn, 1922 / [red, printed].

Other material examined. Historical data (mostly standing as syn. Ch. a. angustedilatata). 1 ♂, 2 ♀♀ in BMNH , 1 ♀ in NMPC : “ Paraguay / Dr. Drake 188[?]” . 1 ♂, 2 ♀♀ in IRSNB : “ Paraguay / Dr. Drake 188[?]”.// “ Ch. auripennis / det. W. Horn 1903”. 14 spms (♂♂, ♀♀) in MNHN , 1 ♀ in CDCL : “ Paraguay / P. Germain / 1881”. 16 spms (♂♂, ♀♀) in MNHN : “ Muséum Paris / Paraguay / Cosset 1900”. 3 spms in SDEI : “ Paraguay / Dr. Drake 188[?]”. // “ Coll. D. Leonhard ” . 1 ♀ in SDEI : “ Cuyaba / Ehlers ” // “ Coll. W. Horn / DEI Eberswalde” // “auripen- nis / Lucas” Other data . 2 ♀♀ in SDEI : “ Paraguay ”. 1 ♂ in SDEI : “ Cheilonycha / auripennis / Paraguay ”. 1 ♂ in MNHN , 2 ♂♂, 2 ♀♀ in NHMW , 1 ♂, 1 ♀ in SDEI : “ Paraguay / Villarica”. 2 ♂♂, 1 ♀ in SDEI : “ Paraguay / Vil- larica / XII.77 ” . 3 ♂♂ in MNHN , 1 ♀ in NHMW , 9 spms in MFNB, 1 ♂ in CCJM , 4 ♂♂, 5 ♀♀ in BMNH , 1 ♀ in NMPC : “ Paraguay ”. 1 ♂, 1 ♀ in NHMW : “ Paraguay / Itapé , 18.I.1932 ” . 2 ♀♀ in CDCL : “ Paraguay / Itapé / Coll Dheurle” . 1 ♂, 1 ♀ in NHMW : “ Cerro Pelado / Paraguay ”. 1 ♀ in MFNB : Matto Grosso ”. 1 ♀ in SDEI : “ Matto Grosso / Vacaria / XII.22 ” // “Lane Coll.” // “ Coll. W. Horn / DEI Eberswalde” . 1 ♂ in RLHC : “ Brazil / M. Grosso / Rio Verde , XI.62 ” . 1 ♂ in RLHC : “ Paraguay: San Pedro / Cororo—Rio Ypane / XII-1 /6-1983 / M. Wasbauer ” 1 ♂ in BMNH : “ Alto Paraguay / Primavera / 22.VII.1956 / E. T. Phillips / MB 1956-752” . 1 ♂ in CDCL : “ Paraguay, depto. Concepcion / Tacuati , XII.1979 J. Wiesner leg.”. Recent data . 4 ♂♂, 4 ♀♀ in CDCL : “ Paraguay, depto. Concepcion / Cororo , 20.XI.1998 / Coll. Dheurle ” . 2 ♂♂, 5 ♀♀ in CDCL : “ Paraguay, Concepcion / Zanja Moroti / 15.X.2012, Walter Gomez leg.” . 12 ♂♂, 9 ♀♀ in CCJM : “ Paraguay, Dpt. Concepcion / Estancia Terrado , 209 m / 23°15’57”S 56°19’05”W / 20.I.2012 leg. Jiří Moravec ” . 12 ♂♂, 10 ♀♀ in CCJM : “ Paraguay, dpt. Canindeyu / RNB Mbaracayu Lagunita / 24°08.1’S, 55°25.6’W, 215 m, / 11.I.2012, leg. Jiří Moravec ” . 6 ♂♂, 4 ♀♀ in CCJM : “ Paraguay, dept. Guaira / Numi env., 187 m. a.s.l. / 25°55’37”S 56°142’25”W / 21.I.2012, leg. Jiří Moravec ” . 1 ♂, 5 ♀♀ in CCJM : “ Paraguay, dpt. Amambay / Rt 5 near Estancia Nelly Victoria / 16.I.2012, leg. J. Moravec ” . 1 ♂ in KCBC : “ Brasil, Mato Grosso / Cuiabá / 9-XII-2011. R. V. Nunes ” .

Specimens cf. Ch. auripennis seen from photographs only (see “Discussion” and “Distribution and biology” below). 4 ♂♂, 2 ♀♀ in DZRJ: “ Brazil, Goiás / Campinaçu, / Vale do Bijuí / mata nativa próximo / a plantio de algodão, Malaise trap, 10.XI.2010, S. Leomar leg”. 2 ♂♂ in DZRJ: ibid., 2.XII.2010, 1 ♂ in DZRJ: ibid., “ 3.I.2011 / S. Leomar leg” .

Differential diagnosis. By its much smaller size (length 10–11.5 mm), the genuine Cheilonycha auripennis , based on Cicindela auripennis Lucas, 1857 , is immediately distinguished from the species currently known under the name Ch. auripennis auripennis (sensu auctorum), which is described below as Cheilonycha bucephalauripennis sp. nov. The type specimens of Cicindela auripennis Lucas possess diagnostic characters corresponding with the taxon described later by Horn (1922) as Odontochila auripennis angustedilatata syn. nov. (see “Remarks” below). The elytra of Ch. a. auripennis are more or less distinctly but notably dilated posteriad towards their apices. The head (measured across the eyes) of Ch. a. auripennis (and of the synonymous Ch. a. angustedilatata) is notably wide due to the conspicuous, distinctly bulged eyes.

Ch. a. chiquitosiana ssp. nov., described below from Bolivian San Jose de Chiquitos, possess similar body size and characters as Ch. a. auripennis including the shape of the head with distinctly bulged eyes, but differs in having the male labrum consistently of a very different shape, possessing notably prolonged tridentate median lobe with anterior teeth in almost same level ( Figs 73–75 ).

Cheilonycha bucephalauripennis sp. nov. (= Ch. a. auripennis sensu auctorum) immediately differs by its notably larger size (13–14.2 mm), somewhat more elongate elytra which are in males usually widest in humeri and moderately narrowed towards apices (in females, rarely in males, mostly subparallel, but never markedly dilated posteriad), notably robust head in contrast to much smaller eyes (well obvious in dorsal view, Figs 97–99 ), thus the robust head when measured across the eyes appears in fact more notably narrower than body ( Figs 85–86 ). For other differences see under that new species below.

Redescription. Body ( Figs 19–21 ) with notably stout elytra, rather variably sized independent of sex, 10.2– 12.4 (LT 10.3) mm long, 3.60–4.60 (HT 4.00) mm wide.

Head ( Figs 22 , 25–29 ) normally shaped with large, conspicuously bulged eyes, but narrower than body, 3.10– 3.60 mm wide, black with violet and green lustre.

Frons, widely and obtusely triangular, moderately or distinctly convex when sloped towards clypeus, black with faint violet or green lustre; surface with fine longitudinally running wrinkles which are more distinct on lateral areas adjacent to supraantennal plates which are elongate-triangular, shiny violet, blue, rarely green; median area fluently passing into vertex over arcuate-convex frons-vertex fold and extremely finely vermicular-rugulose.

Vertex rather dully black with variably expanding violet iridescence and usually with strong green lustre mostly on lateral areas; narrow juxtaorbital edges moderately conically narrowed towards supraantennal plates giving the vertex nearly triangular shape; anteromedian area almost flat or slightly convex, usually with small sublateral impressions on either side, extremely finely vermicular-rugulose (sculpture passing from frons), rugae on median area become more longitudinal when running posteriad, juxtaorbital areas finely parallel-striate, but striae usually irregular and fragmented, posterior and occipital area moderately convex, surface of posteromedian area more distinctly vermicular-rugulose, fine rugae on occipital-median area fragmented into almost granulate sculpture, forming extremely fine, asperate sculpture on temporal area.

Genae chatoyant violaceous with changeable purple, bluish and green iridescences, almost smooth, with a few fine striae on anterior and postgenal areas.

Labrum 4-setose, sexually dimorphic, smooth, metallic blue with changeable violet and green iridescence, convex in middle; male labrum ( Figs 34–38 ) 0.73–0.85 mm long, 1.30–1.55 mm wide, with mostly acute basal teeth, with blunt or mostly entirely effaced lateral teeth, moderately prolonged into indistinct median lobe with mostly right-angled median tooth protruding between blunt anterior teeth; female labrum ( Figs 39–42 ) much longer, 1.00– 1.15 mm long, 1.40–1.65 mm wide, with similar, acute basal teeth, blunt but mostly obvious lateral teeth and tridentate median lobe with distinctly prolonged median tooth between blunt anterior teeth.

Clypeus concolorous with labrum, mostly simply aliform, surface irregularly wrinkled.

Mandibles ( Figs 25–33 ) normally shaped with arcuate lateral margins, rather long, in males with only indistinctly prolonged basal portion; subsymmetrical, each mandible in both sexes with four teeth (and basal molar), inner teeth gradually smaller towards basal molar, fourth tooth in right mandible often notably smaller; coloration metallic black with very faint violet or green iridescence and whitish basolateral stripe which is in males prolonged along the margin anteriad.

Palpi ( Figs 25–29 ). Maxillary palpi entirely metallic black, normally shaped with terminal palpomeres gradually dilated towards rounded apex (in old specimens the apical membrane may be collapsed and consequently the apex may appear as truncate as obvious in Fig 27 ); labial palpi metallic-black except for setose side of penultimate (longest) palpomere which is usually brownish; the longest palpomere of the labial palpi is in both sexes elongate and narrow, only slightly enlarged towards apex.

Antennae ( Figs 19–21 , 25–29 ) rather short, in male passing elytral third, in female even shorter, antennomeres 1–4 metallic black-blue with green and purple iridescences, with sparse, whitish setae, antennomeres 5–11 smoky black-brown with usual micropubescence; scape notably elongate, usually with only one apical seta.

Thorax. Pronotum ( Figs 43–46 ) glabrous, slightly but notably wider than long, length 2.10–2.50 mm, width 2.20–2.60 mm, both anterior and posterior sulcus deep, giving the disc almost subcordiform shape; anterior lobe narrower than disc, but markedly wider than posterior lobe, surface finely and irregularly vermicular-rugulose; disc with distinctly convex lateral margins of dorsally visible proepisterna; dorsally visible notopleural sutures running mostly almost parallel with the proepisternal margins and only slightly narrower than the margins; discal surface extremely finely and densely vermicular-wavy to zigzag-wavy striate-rugulose; median well marked, but sometimes almost merging with the surface sculpture; posterior lobe with distinct posterior rim, surface on its wide median area with coarser, but very irregular and wavy rugae, dorsolateral bulges prevailingly smooth; all ventral and lateral thoracic sterna glabrous, metallic black-blue with strong purple-violet or greenish lustre; proepisterna smooth and shiny; mesepisterna smooth with longitudinal furrow which is in females undistinguishable from the furrow in males (female coupling sulci unrecognizable); metepisterna indistinctly wrinkled.

Elytra ( Figs 52–60 ) notably broad, 6.50–7.50 mm long, outer margins mostly notably dilated towards anteapical angles (in females sometimes less distinctly); anteapical angles broadly arcuate, apex in both sexes rounded towards indistinct sutural spine, rarely in males only indistinctly rounded ( Fig. 54 ), microserrulation extremely fine, indistinct, irregular or partly absent; elytral disc conspicuously convex, then rather steeply sloped towards apex; humeral impression rather distinct, discal impression indistinct or absent; anteapical impression deep, usually running along the whole outer anteapical-apical margin and forming rather distinct bulge at the anteapical angle; whole surface covered with mostly isolated and rather spaced punctures which are much coarser on anterior elytral third, very irregular when running within humeral impression and usually also in a short row on elytral base and with only indistinct, 2–3 fovea-like punctures on elytral disc running posteriad; punctures become smaller posteriad, usually, but not always very fine and widely spaced on posterior elytral half or third, while limited apical area usually again with coarser but irregular punctures; sometimes elytral surface rather coarsely punctate throughout ( Figs 58–60 ); the appearance of the punctation is also changeable depending on angle of illumination; elytral surface glabrous, immaculate; coloration mostly prevailingly iridescent green with shiny red-cupreous elytral discal area, or with golden-bronze iridescences, or prevailingly shiny red with shiny green lateral and apical areas; juxtaepipleural area shiny green or green-blue with indistinct, short whitish setae scattered at epipleural margin.

Legs. Coxae metallic black with changeable strong green, purple and blue chatoyant lustre, procoxae rather densely punctate-setose, setae on mesocoxae much sparser, metacoxae with one apical seta and occasional one or two discal setae, their lateral margin fringed with sparse to dense setae (setae easily abraded); trochanters shiny black with strong green, gold-bronze or reddish lustre, glabrous (except for usual, easily abraded apical seta on pro- and mesotrochanters), metatrochanters entirely glabrous; femora metallic black mostly with only faint purpleviolaceous lustre, rarely with blue, or greenish iridescence, profemora notably thick, almost cone-shaped; femoral surface with rows of white, mostly erect and rather spaced setae, stiffer and usually sparser on metafemora; tibiae concolorous with femora, covered with scattered, stiff, whitish to greyish setae (dorsal surface of protibiae glabrous, ventral area of protibiae and apical third of mesotibiae ventrally covered with dense pad of almost rusty setae; tarsi generally concolorous with tibiae, sometimes, usually on protarsi, with violet or green lustre, subclavate towards their apices, first three protarsomeres in male dilated; claws black-brown.

Abdomen metallic black-blue with strong purple-violet, blue or greenish lustre, surface of ventrites glabrous except for irregularly spaced whitish setae at margins of the visible ventrites.

Aedeagus ( Figs 47–49 ) elongate, widest in middle, conically attenuated towards rounded, slightly ventrally directed apex; internal sac ( Figs 50–51 ) well-armed, upper sclerites mostly membranous and indistinctly delineated including central tooth and voluminous upper-ventral piece and basoventral piece; other two basal pieces distinctly sclerotized, consisting of basal-central characteristic piece with acutely triangular base, prolonged into elongateovaliform upper portion which is recognizable only in left lateral view ( Fig. 50 ) while in right lateral view is covered with another sclerite; other diagnostic piece is a strongly sclerotized basoventral spur of almost indefinite shape.

Variability. Only that mentioned in the description above and illustrated in plates.

Distribution and biology. The nominotypical subspecies (including the synonymous ssp. angustedilatata ) is known from Central Brazil and Paraguay. The type locality of Cicindela auripennis Lucas, 1857 was rather ambiguously stated as “ Brésil intérieur”. One examined female from the Argentinean province of Catamarca was mentioned by Horn (1922) among the type specimens of his ssp. angustedilatata and is therefore treated here as one of the paralectotypes of the synonymous Ch. a. angustedilatata. The same Argentinean locality, obviously based on the same specimen, was recorded by Horn (1910) and also by Bruch (1911), cited also by some subsequent authors. Nevertheless, this species must be very rare in Argentina, because Wiesner & Bandinelli (2014) only cited Bruch (1911) without any other record from this country. Schade (1933) recorded it from Paraguay (Villarica to Itapé), and Sawada & Wiesner (2002) recorded it (as Ch. a. angustedilatata) from the Paraguayan departments of Amambay, Caaguazú, Concepción, Guaira and San Pedro. Horn (1910) mentioned the distribution of this species as “Catama- rca bis Goyaz” (the Goyaz record in fact belonging to Ch. bucephalauripennis sp. nov.). Horn (1924) mentioned also Brazilian Mato Grosso and Vacaria, and Mato Grosso was mentioned also by Naviaux (2002), some of the specimens have been examined (see “Other material examined” above).

Horn (1924) characterized Vacaria (also spelled Vaccaria) as grasslands with small rivers, ravines with bush and scattered forest of stunted trees (cerrado), mostly used as cattle pasture and often with large, cone-shaped termite mounds. He specified the locality Vacaria as situated in Mato Grosso in an area of Rio Vaccaria, 77 km south of Campo Grande on the railway which runs from São Paulo to Corumba (state of Mato Grosso do Sul) on the Para- guay River (the river forms the Brazil-Paraguay border). Later ( Horn 1926b) mentioned “Vacarias” areas as situated in southern Matto Grosso; he mentioned the “Campos de Vacarias (Campos Xeres), in the vicinity of the main river Rio Ivinheima in the southernmost Mato Grosso ”.

Cassola & Pearson (2001) listed this species under the name Ch. auripennis from all the above-mentioned countries and also from Bolivia, but the Bolivian records, including those by Horn (1931) and Mandl (1958) from the province of Sara (department of Santa Cruz), as well as those by Pearson, Guerra & Brzoska (1999) from several localities of the Santa Cruz department, very probably belong to O. a. chiquitosiana ssp. nov. described from the Bolivian San José de Chiquitos (Santa Cruz department) as a new subspecies below.

The exact distribution in Brazil must be revised in collaboration with Brazilian colleagues, because specimens from Brazilian collections are not accessible on loan. Nine specimens from Campinaçu, Vale do Bijuí, deposited in DZRJ (see “Other material examined”, “specimens cf. Ch. auripennis ” above, and “Discussion” below), move the distribution of Ch. auripennis also to the northern parts of the Brazilian state of Goiás, but situated long way from Jataí in the south-western area of the state.

Ch. auripennis belongs to termitophilous tiger beetles (see also Moravec & Brzoska 2014 on Pentacomia lanei W. Horn, 1924 ). The hunting behaviour of Ch auripennis on termite hills (probably of a Cornitermes species) was reported by Wasmann (1895a, b), Berg (1900), Horn (1924), Guerra (1993) and Pearson, Guerra & Brzoska (1999). In Paraguay, we had the same experience (reported by Moravec & Brzoska 2014 under the synonymous Ch. a. an- gustedilatata). In the above-cited locality Lagunita near the Mbaracayu National Park (department of Canindeyu, Paraguay) we observed the adults of Ch. a. auripennis (= “ssp” angustedilatata) together with adults of Brasiella aureola Klug, 1834 , both inhabiting mostly very high termite mounds, usually flying onto neighbouring termite hill or sheltered among surrounding grass when disturbed. They sheltered inside the termite mounds both during the night and the hottest hours of the day. The termitophilous tiger beetles thus can survive during commonly occurring fires as mentioned by the above cited authors and summarized by Erwin & Pearson (2008). In January 2012, we found partly burned termite hills in the Paraguayan department of Guairá, but with some adults (and probably also larvae) survived ( Figs 108–110 ). Nevertheless, Ch. a. auripennis and other termitophilous insectes are highly endangered not only by burning the places, but also because of destruction of the termite hills in the pastures by some farmers, usually using tractors for the extraction; once in January 2012 we witnessed such destructions.

Arndt et al. (1996) and Erwin & Pearson (2008) mentioned that according to Guerra (1993), larvae of Ch. auripennis make their burrows in the surface of the termite mounds. However, in our experience it was difficult to recognize the burrows of this species according to their shapes among the numerous holes on the surface of the mounds.

Discussion. Lucas (1857: 31, pl. 1a, fig. 1) did not indicate if he based the original description of Cicindela auripennis on only one or more specimens. The drawing of the labrum ( Lucas 1857, fig. 1b) is largely scholastic, but due to the prolonged median lobe with protruding median tooth it evidently figures female sex, which corresponds also with the original description: “ le lèvre supérieure avancée, lisse présente dans son milieu une dent bien marquée, recourbée, et trois autres beaucoup plus petites de chaque côté”. The colour picture of the body (fig. 1 in the same plate by Lucas) very probably figures the same female.

Two females in MNHN labelled “Amerique du Sud / de Castelnau 1847”, corresponding with the original paper by Lucas (1857) and coming from the “Voyages de Castelnau” realized by Laporte de Castelnau in 1843–1847, are evidently the syntypes of this species. Apart from the above-mentioned label, they bear a circular green labels with “46/45” and 84/46” as usual attached to specimens from the “Voyages de Castelnau”. The female labelled “46/45” perfectly corresponds with the original description by Lucas (1857), also in its very small body length (10.3 mm) and coloration: “Longueur 10 milim.; ” and “La tête, le thorax sont d’une belle couleur bleue légèrement violacée, assez brillante, avec des élytres d’un beau vert doré”. The coloration and the shape of the head with markedly bulged eyes correspond with the original description by Lucas: “le yeux: ceux-ci sont très gros, saillants ...... Le thorax plus ètroit que la tête. Les élytres son moins allongèes et un peu plus large que dans la C. chylybea ; eles sont convexes et arrondies à leur extrémité”. These diagnostic characters clearly differ from specimens of Cheilonycha bucephalauripennis sp. nov. (= Ch. auripennis sensu auctorum) described below. Consequently, the female in MNHN, coming from the Voyages de Castelnau corresponding with the paper by Lucas (1857) and labelled “46/45”, is designated here as the lectotype of Ch. auripennis ( Lucas, 1857) , to fix identity of the taxon.

The examined specimens (SDEI) from Brazilian provinces of Cuiabá (= Cuyaba) in the state of Mato Grosso and those labelled “ Mato Grosso ”, as well as specimens from Brazilian Vacaria, proved to be conspecific with Ch. a. auripennis (= “ssp” angustedilatata syn. nov.). The other Brazilian specimens, those from Jataí (=Jatahy) in the state of Goiás (= Goyaz), deposited in MNHN, SDEI, BMNH, IRSNB, NMPC, are notably much larger and differ in a complex of diagnostic characters (see “Differential diagnosis” above and under Ch. bucephalauripennis sp. nov. below).

When Horn (1922) described his “ Odontochila ” auripennis angustedilatata, he did not examine the true type specimens of Cicindela auripennis Lucas, 1857 (see below); he discussed the differences of the specimens from Goiás, but merely on the level of his speculations. In the discussion Horn recalled that an exceptionally small male (sic!) from Goiás might correspond with a “type” by Lucas, which Horn, but only according to his memory (sic!), saw in 1903 (without mentioning of the depository of the alleged “type”). He also considered the type locality “Bré- sil intérieur” ambiguous and supposed that Castelnau travelled via Goiás (without any evidences and irrelevantly, because the male specimen mentioned by Horn was caught much later by Pujol as also other specimens from Jataí (state of Goiás). In the same paper, Horn (1922) also speculated whether Lucas (1857) described and illustrated male or female sex and noted that the gender of the type was ambiguous. However, as mentioned above, the description and illustrations by Lucas (1857, pl. 1 a, figs 1a, 1b) almost certainly feature female sex, including the prolonged median lobe of the labrum (fig. 1b). The most important is the shape of the elytra illustrated by Lucas (1857, pl. 1 a, fig. 1), which are clearly dilated posteriad, corresponding with the lectotype and paralectotype (MNHN) of the true Ch. auripennis Lucas from the Voyage de Castelnau, as well as with the type specimens of the synonymous Ch. a. angustedilatata. Therefore, the description of the latter by Horn (1922), who apart from other errors overlooked the illustration by Lucas, was superfluous, because of Horn’s misinterpretation. It is also noteworthy that the specimens from Brazilian Vacaria, Mato Grosso and Cuyaba were considered by Horn as “ Chilonycha ” auripennis (Lucas) , despite all their characters identical with his ssp. angustedilatata (as revealed within this present revision).

Nine specimens (see “Other material examined, specimens cf. Ch. auripennis ” and “Distribution and biology” above) from northern area of the Brazilian state of Goiás, seen only from photographs taken in DZRJ by André Silva Roza (DZRJ), possess their characters (A. S. Roza pers. com) such as small body (length 9.60–10.8 mm), head with markedly bulged eyes and posteriad-dilated elytra, shared with Ch. a. auripennis , only the green coloration of the head and pronotum is shared with Ch. bucephalauripennis sp. nov. described below. Due to their small size and above-mentioned characters, the specimens in DZRJ are certainly conspecific with Ch. auripennis , but without examination of the specimens it is difficult to decide if they represent another subspecies of the species by Lucas. In any event, they fundamentally differ from Ch. bucephalauripennis sp. nov.

The erroneous concept of Ch. auripennis was followed by all subsequent authors.

It must be noted that some authors starting by Horn (1910) cited the redescription of Cheilonycha auripennis by Dokthouroff (1887) as if he described a new (homonymous) species (under the name combination Cicindela (Eu- ryoda) auripennis ) identical with Ch. a. angustedilatata. Consequently, some of the syntypes of ssp. angustedilatata are confusedly standing in SDEI under a large greenish collection-label “auripennis / v. Dokht.[Dokhtouroff]”, obvi- ously arranged there by DÖBLER (see DÖBLER 1973). Naturally, no such homonymous name by Dokthouroff exists, because Dokthouroff (1887: 154) underlined his redescription as “ Cicindela (Euryoda) auripennis Luc. ”, explicitly citing Lucas as the author of this taxon. Dokhtouroff thus only redescribed C. auripenni s Lucas, and this present revision confirmed that he had this taxon in the correct concept. Erwin & Pearson (2008) cited Dokhtouroff correctly under Ch. a. auripennis , but erroneously as if the author of a homonym, and by a mistake these authors have Lacordaire as the author of Ch. auripennis instead of Lucas.

Horn (1934: 384, fig. 32 and 1938: pl. 87, figs 21–22) illustrated the aedeagi of these taxa (in his concept) as somewhat differing in their apical parts. The same small differences in the aedeagus apex of the lectotype of “ ssp. angustedilatata ( Fig. 47 ) have been observed within the revision presented here, but the shape varies as revealed after more aedeagi of syntopic males have been examined (as obvious from Figs 47–49 , 65–68 , 104–105 here) and the shape depends also on the state of the aedeagi (compare Figs 67, 68 ). Nevertheless, as stressed in the “Differential diagnosis” under Ch. chalybea above, the internal sacs within the aedeagi of Ch. a. auripennis and Ch. a. chiquitosiana differ from the internal sac in Ch. chalybea .

It is noteworthy that Horn (1934) treated both Ch. auripennis in his sense (here Ch. bucephalauripennis sp. nov.) and his Ch. a. angustedilatata as only subspecies of Ch. chalybea , but his concept was not accepted by subsequent authors.

Because of the shape of the head and elytra, the previous colour picture of Ch. auripennis by Horn (1910, pl. 13, fig. 4) almost certainly shows the true Ch. auripennis (= Ch. a. angustedilatata).

Figures

FIGURE 108. Habitat of Cheilonycha a. auripennis (Lucas), Paraguay, Guaira. termite mound on an area with cut down trees and partly burned.

FIGURE 109. Habitat of Cheilonycha a. auripennis (Lucas), Paraguay, Guaira. ditto, with one imago on the mound surface.

FIGURE 110. Habitat of Cheilonycha a. auripennis (Lucas), Paraguay, Guaira. the same but more heavily burned area with termite mounds.

FIGURES 71–84. Cheilonycha a. chiquitosiana ssp. nov., all from Bolivia, San Chose de Chiquitos. 71–72––head: 71–– ♂, HT (UASC); 72––♀, PT (CCJM); 73–78––labrum: 73––♂, HT; 74––♂, PT (DBCN); 75––♂, PT (CCJM ex COSJ); 76––♀, AT; 77–78––♀, PT (CCJM and DBCN); 79–80—pronotum: 79––♂, HT; 80––♀, AT (DBCN); 81–84––elytron: 81––♂, HT; 82––♀, PT (CCJM); 83––♀, PT (DBCN); 84––♂, PT (DBCN). Bars = 1 mm.

FIGURES 93–103. Cheilonycha bucephalauripennis sp. nov., all from Brazil, Jataí (Jatahy on labels). 93–95––head: 93––♂, HT (MNHN); 94––♂, PT (MNHN); 95––♂, PT (SDEI); 96––ditto, mandibles; 97–99––head in dorsal view: 97––♂, PT (SDEI); 98––♂, HT (MNHN); 99––♂, PT (MNHN); 100–103––labrum: 100––♂, HT; 101––♂, PT (MNHN); 102––♂, PT (SDEI); 103––♀, AT (MNHN). Bars = 1 mm.

FIGURES 85–92. Cheilonycha bucephalauripennis sp. nov., all from Brazil, Jataí (Jatahy on labels). 85–86––habitus: 85––♂, 14 mm, HT (MNHN); 86––♂, 13 mm, PT (MNHN); 87–88––pronotum: 87––♂, HT; 88––♀, AT (MNHN); 89––labels of HT; 90–92––elytron: 90––♂, HT; 91––♀, AT; 92––♂, PT (MNHN). Bars = 1 mm.

FIGURES 19–24. Cheilonycha a. auripennis (Lucas). 19–21––habitus: 19––♀, 10.3 mm, “Brésil intérieur”, LT (MNHN); 20––♂, 11 mm, Paraguay, LT of syn. C. a. angustedilatata W. Horn (SDEI); 21––♂, 11.2 mm, Paraguay, Mbaracayu (CCJM); 22––head in dorsal view, ibid. (CCJM); 23–24––labels: 23––LT (MNHN); 24––LT of syn. C. a. angustedilatata W. Horn (SDEI).

FIGURES 25–42. Cheilonycha a. auripennis (Lucas). 25–29––head: 25––♀, “Brésil intérieur”, LT (MNHN); 26––♀, Para- guay, Mbaracayu (CCJM); 27––♂, Paraguay, LT of syn. C. a. angustedilatata W. Horn (SDEI); 28–29––♂, Mbaracayu (CCJM); 30–33––left mandible, ibid. (CCJM): 30–32––♂; 33––♀; 34–42––labrum: 34––♂, LT of syn. C. a. angustedilatata W. Horn (SDEI); 35–37––♂, Mbaracayu (CCJM); 38––♂, Brazil, Cuyaba (SDEI); 39––♀, LT (MNHN); 40–– anomalous labrum, ♀, PLT of syn. C. a. angustedilatata W. Horn (SDEI); 41––♀, Mbaracayu (CCJM) 42––♀, Argentina, Catamarca, PLT of syn. C. a. angustedilatata (SDEI). Bars = 1 mm.

FIGURES 43–51. Cheilonycha a. auripennis (Lucas). 43–46––pronotum: 43––♀, “Brésil intérieur”, LT (MNHN); 44––♂, Brazil, Cuyaba (SDEI); 45––♂, Mbaracayu (CCJM); 46––♀, ibid. (CCJM); 47–51––male aedeagus: 47––Paraguay, LT of syn. C. a. angustedilatata W. Horn (SDEI); 48–49––Paraguay, Mbaracayu (CCJM); 50–51––ditto, internal sac in left and right lateral view. Bars = 1 mm.

FIGURES 52–61. Cheilonycha a. auripennis (Lucas). 52–60––elytron: 52––♀, “Brésil intérieur”, LT (MNHN); 53––♂, Para- guay, LT of syn. C. a. angustedilatata W. Horn (SDEI); 54––♀, ibid., PLT of syn. C. a. angustedilatata W. Horn (SDEI); 55––♀, Argentina, Catamarca, PLT of syn. C. a. angustedilatata W. Horn (SDEI): 56––♂, Brazil, Cuyaba (SDEI); 57––♀, Brazil, Vac- aria (SDEI); 58–59––♂, Mbaracayu (CCJM); 60––♀, ibid. (CCJM); 61––thoracic wing, ♂, Paraguay, Colonia Azotel (CCJM). Bars = 1 mm.

FIGURES 62–70. Cheilonycha a. chiquitosiana ssp. nov., all from Bolivia, San Chose de Chiquitos. 62–63––habitus: 62–– ♂, 11.4 mm, HT (UASC); 63––♂, 11.5 mm, PT (CCJM ex COSJ); 64––head in dorsal view, ♂, PT (DBCN); 65–70––aedeagus or its apex: 65––HT; 66––PT (DBCN); 67––PT (CCJM) 68—ditto, dry; 69–70–– ditto, internal sac in left and right lateral views. Bars = 1 mm.

FIGURES 104–107. Cheilonycha bucephalauripennis sp. nov., all from Brazil, Jataí (Jatahy on labels), male aedeagus or its apex. 104––HT (MNHN); 105–– PT (SDEI); 106–107––ditto, internal sac in left and right lateral view. Bars = 1 mm.