Odontocheila mexicana Laporte de Castelnau, 1834

Moravec, Jiří, Brzoska, David & Huber, Ronald, 2017, Taxonomic and nomenclatorial revision within the Neotropical genera of the subtribe Odontocheilina W. Horn in a new sense — 18. Six Mexican and Central American species related to Odontocheila mexicana Laporte de Castelnau and O. ignita Chaudoir, with a description of O. potosiana sp. nov., Zootaxa 4231 (4), pp. 451-499 : 455-466

Odontocheila mexicana Laporte de Castelnau, 1834

( Figs 1–53 , 208 ).

Odontocheila mexicana Laporte de Castelnau, 1834b: 38 .

Type locality. “Mexique”.

Cicindela quadrina Chevrolat, 1835: 6 (synonymy by Horn 1922). Type locality. “Mexique, Tutepec” (“Tustepec” on the label), but probably Tuxtepec in the state of Oaxaca). Cicindela Nietii Guérin-Méneville, 1844: 254 (synonymy by Fleutiaux 1892). Type locality. “Mexique”.

Odontochila mexicana: Fleutiaux 1892: 125 . Rivalier 1969: 206, fig. 6m, 207, 213, fig. 9mx. Odontochila quadrina: Fleutiaux 1892: 125 ;

Odontochila mexicana Nietoi : Fleutiaux 1892: 125 (emendation of the species-name and as a synonym of O. mexicana ).? Odontochila mexicana quadrina Rivalier: 1969 : 207.

Odontocheila quadrina: Johnson 1996: 37 .

Odontocheila mexicana: Wiesner 1992: 78 .

Nomenclatorial note. The date of the description of O. mexicana has been commonly treated as 1835, thus the same as of the publishing date of the synonymous O. quadrina by Chevrolat (1835). However, according to Sherborne (1928) and also Bousquet (2016), the journal Etudes entomologiques was issued in two parts, and the first one with the description of O. mexicana appeared in 1834—thus see Laporte de Castelnau (1834b). In contrast, according to Sherborne (1930) the description of Cicindela quadrina was published in 1835, and according to Bousquet (2016), the Fascicule 8 with the description of C. quadrina was apparently issued in May 1836 —nevertheless, the date of C. quadrina is cited here as in 1835.

Type material of Odontocheila mexicana Laporte de Castelnau. Lectotype (designated here) ♀ in BMNH, labelled: “Type” [circular label with red border, printed] // “Mexico” [printed] // “ Mexicana / de Lap. / Et. Ent. 1834 / 39[?illegible] / Nietii Guer. R. Z 1844 p. 254 / Mex.” [large circular green label, handwritten] // “Type” [with black border, printed] // “F. Bates Coll. / 1911-248” [printed] // “Lectotype ♀ / Odontocheila / mexicana Lap. / by Erwin ‘96” [printed, partly in red, and handwritten, invalid label] // Lectotype / Odontocheila / mexicana / Laporte de Castelnau, 1835 / design. Jiří Moravec 2012” [red, printed].

Type material of Cicindela quadrina Chevrolat. Lectotype (designated here), ♀ in MNHN, labelled: “♀” [small square green label, handwritten] // “Coll. Chevrolat / Odontocheila quadrina Chv. Coll. Mex. / 1835 / Dejeanii Chev. olim / Mexico Tustepec[Tutepec?] Sallé / Type” [large green label, printed/handwritten] // “ Collection / Chevrolat ” [printed] // “ Type ” [printed] // “Lectotype / Cicindela / quadrina Chevrolat, 1835 / design. Jiří Moravec 2015“ [red, printed] // “ Odontocheila / quadrina ( Chevrolat, 1835) / det. Jiří Moravec 2015” [printed].

Type material of Cicindela Nietii [ nietoi ] Guérin-Méneville. No specimen of exact label data found, but it may be the female in MNHN labelled: “Coll. Chevrolat / Odontocheila / mexicana Cast. Et. p. 38 1844 [sic!] / Princeps Dupnt[Dupont]/Cat. D[Dejean] I / Nieti Guérin R. Z 1844, p 254 / Mexico / Cordova Sallé” [large green label, printed/handwritten].

Other material examined. Historical data. 1 ♀ in MNHN [Fleutiaux Hist. Coll.]: “ Mexico ” // “ Ex Musaeo / H.W.Bates / 1892” // quadrina / Chevrolat ” . 1 ♂ in MNHN: “8” // “ Muséum Paris / Coll. Chaudoir 1874” // “ Mexicana / Laporte / Nietii Guérin / Mexique ” . 1 ♂ in MNHN: “ quadrina Chvrt. Centrim ? [illegible] / Observ? [illegible]. 1 ♂ in MNHN: “ Mexico / Janson ” . 3 ♂♂, 3 ♀♀ in MNHN: “Muséum Paris / Coll. Chaudoir 1874”. 4 ♂♂, 3 ♀♀ in BMNH: “ Jalisco / Mex.” // “ F. Bates Coll. / 1911–248” . 1 ♂ in BMNH: “ Toxpam / Mexico / Sallé Coll. ” // “ Odontocheila / mexicana / apud Sallé Coll. ” . 1 ♂ in BMNH: “ Chinantla / Mexico / Sallé Coll. ” // “ Odontocheila / mexicana / apud Sallé Coll. ” . 1 ♀ in BMNH: “16277” / Mexico ” . 1 ♂, 1 ♀ in BMNH: “ Truqui [leg.?] / Mexico / (ex Fry)” . 2 ♀♀ in BMNH: “ Mexico / Cordova Sallé Coll. ” // “ Odontocheila / mexicana / apud Sallé Coll. ” . 1 ♂ in BMNH: “ Odontocheila mexicana / C. I. Lap[orte]”. 1 ♀ in BMNH: “ O. mexicana / arm: Cast[elnau]” // “F. Bates Coll. / 1911–248”. 1 ♀ in BMNH: “ Odontocheila / quadrina / C. I. Chvr.[Chevrolat]”. 1 ♀ in MNHN: “ Tuxtla ” // “ Mexico / Sallé Coll. ” // “ H.W. Bates / Biol. Centr. Amer. ” // “ Muséum Paris / Coll. R. Oberthür” // “ quadrina Chevr. / apud Salé ” . 1 ♀ in BMNH: “ Tuxtla ” // “ Mexico / Sallé Coll. ” // “ San André ” // Odontocheila / mexicana Lap. / apud Sallé ” // “ B.C.A. Coll., I(1), / Odontocheila ” . 1 ♀ in MFNB: “ Mexico, Meyer [leg.]” // “ Hist Coll. ( Coleoptera ) / Nr. 3627 / Odontocheila Nieti Guér. / Mexico, Meyer / Zool. Mus . Berlin ” . 1 ♂ in MFNB: “3627” // “ Nieti Guér. / Mexico ” // “ Hist Coll. ( Coleoptera ) / Nr. 3627 / Odontocheila Nieti Guér. / Mexico, Meyer / Zool. Mus . Berlin ” . 1 ♂ in MFNB: “ Hist Coll. ( Coleoptera ) / Nr. 42487 / Odontocheila quadrina Cevr. / Mexico, Coll. Schaum / Zool. Mus . Berlin ” . 1 ♀ in MFNB with same label and: “42487” // “ quadrina / Chevrolat / Mexico ” . 1 ♂ in IRSNB: “ Mexico Sallé”. Other data . 2 ♂♂ in MNHN: Mexico / Jansen ” . 1 ♀ in BMNH, 1 ♂ in SDEI: “ Cordoba / Hôge ” . 1 ♂, 1 ♀ in MFNB, 2 ♂♂ in NMPC: “ Mexico / Hoege ” . 1 ♂, 1 ♀ in MNHN, 1 ♀ in IRSNB: “ Mexique ” . 5 ♂♂, 4 ♀♀ in BMNH, 2 ♂♂, 2 ♀♀ in SDEI: “ Mexico ” . 1 ♂ in SDEI: “ Istma Tehuantepec [partly illegible] / Mexico ” . 2 ♂♂ in SDEI with illegible labels. 1 ♀ in MFNB: “ mexicana / Odont. Castelnau / Mexico ” . 1 ♂, 1 ♀ in IRSNB: “ C. quadrina Chaud. / Mex.” . 1 ♂, 1 ♀ in MFNB: “677”. 1 ♂, 1 ♀ in MFNB: “ Oaxaca ” . 1 ♂ in BMNH, 1 ♂, 3 ♀♀ in MFNB: “Atoyac / Vera Cruz / May H.H.S.”. 1 ♂ in MNHN, 2 ♂♂, 1 ♀ in BMNH: “ Playa Vicente / Mexico / Höge ” . 1 ♀ in CMNH: “ Oaxaca, Mex. / 26.V.1959 / G. y V. Halffter ” // “ Finca San Carlos / Union de lόs ríos / Coatzacoalcos y / Sarabias, 30km / al E de Palomares ” // “ Robert C. Graves / Collection / Carnegie Museum / Acc. No 66.619” . 1 ♀ in CMNH: “ Mexico, Oaxaca / Temascal / VI-26-1965 / A. B. Lau ” . 1 ♀ in CMNH: “ Mexico, Veracruz / 2 mi. sw. Fortin de / Las Flores 4000 ft / 15–16-VII-1970 / E. Fischer, P. Sullivan ” // “ Robert D. Ward / Collection ”. Recent data . 1 ♂ in FCCR: “ June 24, 1994 / 2.5 ml. W. of Neuvo San / Juan Chamula Los Margaritas / Chiapas, Mex., W. Johnson ” . 11 ♂♂, 3 ♀♀ RLHC, 1 ♂ in CDCL: “ MX: Oaxaca / 3.2km S. Temascal / Rio Tonto , S. side / 31-VI-1993; 200ft / R.L. Huber ” . 1 ♂ in CDCL: “ Mexico : Oaxaca / 3.2 km S Temazcal / Rio Tonto , S side / 200 ft, 21.VI.1993 ”. 23 ♂♂ , 6 ♀♀ in DBCN: 1 ♂, 2 ♀♀ in CCJM: “Mexico— Oaxaca / Road to Chicali / 3.2 km SE—Temazcal / D. Brzoska 21-VI-1993 ” . 4 ♂♂, 2 ♀♀ in DBCN, 2 ♀♀ in CCJM: “ Mexico—Oaxaca / 3.2 km SE—Temazcal / Rio Tonto / D. Brzoska 16-VI-1995 ” . 31 ♂♂, 22 ♀♀ in DBCN 3 ♂♂, 7 ♀♀ in CCJM: “ Mexico—Oaxaca / Hy. 175—km 51 / 3 km S—Valle Nacional / D. Brzoska 16-VI- 1995 ” . 3 ♂♂ in DBCN: ibid., except for: “ Hy 175, 4 km S- Valle Nacional / D. Brzoska 16-VII-1994 ” . 2 ♂♂ in DBCN: ibid., except for: “ Hy 175, Pto. Eligio, 14 km S- Valle Nacional / D. Brzoska 22-Vi-1993 ” . 1 ♂ in DBCN: ibid., except for: 3 kmS & W Chiltepec, D. Brzoska 21-VI-1993 ”. 3 ♂♂, 6 ♀♀ in DBCN: ibid., except for: “ Hy 182, KM 191 (W- Jalapa) / D. Brzoska 16-VI- 2010 ” . 1 ♂ in DBCN: ibid., except for: “ Hy 175, Pto. Eligio, D. Brzoska 19-VI-2000 ” . 2 ♂♂ in DBCN: ibid., except for: “KM 53, (10 km NE Pto. Eligio ”. 23 ♂♂, 18 ♀♀ in DBCN: “ Mexico—Veracruz / Los Tuxtlas, B.S., 128 m / 18°35.1´N, 95°04.5´W / D. Brzoska 5-6-VI-2010 ”. 7 ♂♂, 4 ♀♀ in CCJM: ibid., except for: “ D. Brzoska 6-7-VI-2009 ”. 1 ♀ RLHC: “Escola, Mexico / Veracruz / 20.VI.1974 / T.W. Taylor ”. 2 ♂♂, 1♀ RLHC: “ Veracruz, Mex. / Los Tuxtlas / Trop. Biol. Sta. / Marlene Palmer” // “ex 3rd instar larva / taken 5.I.1978 / deep woods, seep / black soil + humus”. 2 ♂♂ in JWCW: “ Mexico, Veracruz / 33 km NE Catemaco, 160 m / Las Tuxtlas Biol, Sat. / El Ravine rainforest / 1.VII-1.VIII.1983, S & J. Peck ”. 1 ♀ in WJCM: “ 15 mi East / Teziutlan / Puebla Mex. / VIII-5-6, 60 / H.F. Howden ”. 35 ♂♂, 30 ♀♀ in DBCN, 2 ♂♂, 1 ♀ in CCJM: “ Mexico—Chiapas / P.N. Palenque / Arroyo Otulum / 17°28.9´N; 92°02.8´W / D. Brzoska 5-VI-1998 ”. 2 ♂♂, 4 ♂♂ in CCJM: ibid., except for: “ 17°29.0´N; 92°02.8´W / D. Brzoska 24-V-2009 ”. 4 ♂♂, 3 ♀♀ in DBCN: ibid., except for: “ Hy 186, km 50 (S. Palenque)” . 6 ♂♂, 4 ♀♀ in DBCN: ibid., except for: “ Hy 307, Km. 43.8, (E.- Tziscao ) / D. Brzoska 11-VI-1999 ” . 4 ♂♂, 1 ♀ in DBCN: ibid., except for: “Km 74 1”. 8 ♂♂, 4 ♀♀ in DBCN: ibid., except for: “ Aqua Azul (S. Palenque) / D. Brzoska 6-VI-1998 . 6 ♂♂, 1 ♀ in DBCN, 1 ♀ in CCJM: “ Mexico—Chiapas / Las Margaritas Rd 31 km / SW San Quintin 655m / 16°20.3´N ; 91°29.7´W / D. Brzoska 10- VI-2010 ”.

Differential diagnosis. Odontocheila mexicana is immediately distinguished from all other species treated here by its subglobose, almost smooth and polished pronotal disc (with only few short and shallow rugae along the median line). Similarly coloured O. potosiana sp. nov. differs in having the pronotal disc narrower and clearly finely and shallowly sculptured on its entire surface up to the notopleural sutures; moreover, its labrum is notably longer and apex of its aedeagus is slightly wider.

Redescription. Body ( Figs 1–6 ) markedly shiny metallic coloured, medium-sized, 8.60–10.7 (lectotype 10.6) mm long, 2.60–3.35 (lectotype 3.35) mm wide (females usually larger than males, occasionally smaller, exceptionally much smaller).

Head ( Figs 7–10 ) notably large with pronounced eyes, only slightly narrower than body, 2.55–3.10 mm wide, all head portions glabrous.

Frons steeply sloped towards clypeus, with distinctly convex median area, dorsally bluntly triangular, deep violaceous-blue with gold-bronze and blue-green lustre on lateral areas and reddish-cupreous in middle, sharply delimited from clypeus and confluent with vertex over rounded frons-vertex fold, with rather indistinct lateral edges formed by deep impressions above supraantennal plates which are flat and with strong metallic-violaceous lustre; frons surface smooth at base and with only few shallow wrinkles on lateral areas while the median area on frons-vertex fold is ornamented with mostly transverse-wavy rugae which are usually surrounded by arcuate arranged wavy rugae forming a distinct ornament passing on anteromedian area of vertex.

Vertex with only one juxtaorbital sensory seta (on each side), anteromedian area almost flat, bright reddish cupreous in middle with iridescent greenish, blue, bronze or violaceous lateral areas; frons-vertex fold and limited anteromedian area irregularly transversely wavy and arcuate rugulose (sculpture passing from frons), large juxtaorbital areas and lateral areas distinctly and deeply longitudinal parallel-striate, striae divergent posteriad passing on temples; posterior and occipital area moderately convex, rather distinctly but shallowly irregularly wavy to vermicular rugulose.

Clypeus metallic iridescent green, blue-green, or reddish cupreous with only iridescent green lateral areas, smooth and shiny, sometimes with few indistinct wrinkles.

Genae metallic green, usually with blue or violaceous lustre, almost smooth in middle, with few deep parallel striae on postgenal area passing from temples.

Labrum 4-setose, in both sexes bicolored, ochre-testaceous with black basomedian or whole basal area; male labrum ( Figs 11–14 ) rather short, 0.60–0.70 mm long, 1.05–1.25 mm wide; lateral teeth blunt or subacute, anterolateral teeth mostly rounded, very rarely right-angled or subacute, anterior margin usually moderately convex or shallowly emarginate between subacute anterior teeth, rarely with small, or only indicated median tooth; female labrum ( Figs 15–16 ) much longer, length 1.05–1.20 mm, width 1.00– 1.35 mm, with prominent, acutely tridentate median lobe with protruding median tooth.

Mandibles ( Figs 7–10 ) pale brown to reddish or mahogany-brown except for narrow, ivory-ochre lateral stripe and blackened borders of teeth, normally shaped with arcuate lateral margins, subsymmetrical, each mandible with four teeth (and basal molar), the three inner teeth gradually smaller towards the basal molar.

Palpi ( Figs 7–10 ) with normal (elongate) shape of terminal palpomeres, ivory-yellow to ochre-testaceous except for black-brown to black terminal palpomeres in both labial and maxillary palpi, and with slightly to distinctly brown-darkened to blackened penultimate palpomeres of maxillary palpi; penultimate (longest) palpomere of labial palpus rather narrow with subparallel lateral margins, only gradually dilated towards apex (width up to 0.21 mm).

Antennae ( Figs 1–10 ) long, in male markedly surpassing elytral half, in female shorter; scape with only apical seta, deep violaceous or deep purple, sometimes with bluish reflections, pedicel mostly concolorous with the scape, often with paler, reddish base or irregular reddish spots; antennomeres 3–4 ( Figs 7–10 ) variably either almost uniformly metallic-black with purple or violaceous reflections, or with more or less distinctly delineated purple or mahogany-testaceous apical areas of variable intensity, with indistinct, sparse microsetae; antennomeres 5–11 smoky black with usual micropubescence.

Thorax. Pronotum ( Figs 38–45 ) glabrous, notably shiny, metallic iridescent green-blue, often reddish-cupreous or golden on sublateral discal areas, often also with violaceous reflections or purple-violet lateral areas, mostly as long as wide or very slightly wider, length 1.65–2.10 mm, width 1.65–2.15 mm; sulci well pronounced; anterior lobe distinctly wider than posterior lobe, surface of anterior lobe almost smooth and polished with only few median wrinkles which are shallow but light-refracting; disc (including lateral margins of proepisterna) notably subglobose (particularly in females) almost smooth and polished, with only few very shallow and mostly indistinct rugae along the median line, large lateral areas smooth and polished, rarely very shallow, indistinct wrinkles reaching sublateral areas; notopleural sutures indistinct and barely visible from above; medial line shallow but obvious; posterior lobe with distinctly raised and wide dorsolateral bulges, smooth and shiny, or with few shallow wrinkles in middle, with distinct posterior rim; all ventral and lateral sterna glabrous; proepisterna, mesepisterna and metepisterna metallic black with faint green, blue or purple lustre, female mesepisternal coupling sulci indistinct, in form of a sinuous longitudinal furrow which is only somewhat more distinctly carved than the shallower furrow in male, lacking any pit; metepisterna with several wrinkles; prosternum, mesosternum and metasternum metallic black-blue or blackblue with chatoyant, green, golden, purple or violaceous iridescence, smooth and polished.

Elytra ( Figs 19–37 ) elongate, length 5.20–6.50 mm, with rounded humeri in male, mostly subquadrate in female, lateral margins in male subparallel, outer margin in female slightly dilated in middle and usually (variably) with slightly more arched posthumeral epipleural crease, anteapical angles arcuate, then obliquely running towards apices which are rounded towards short but distinct sutural spine in both sexes, but more widely rounded in female; microserrulation indistinct and usually very irregular; elytral dorsal surface almost even, convex on posterior half of elytral disc, humeral impression rather distinct, discal impression shallow, but clearly delimiting moderate basodiscal convexity; anteapical-apical impression rather distinct; whole elytral surface covered with punctures of very variable pattern: punctures larger and mostly isolated on anterolateral area and on basodiscal convexity, with two short rows of 4–6 conspicuously large fovea-like punctures running within and below the humeral impression; punctures on elytral disc either isolated and rarely anastomosing, or very commonly anastomosing in obliquely transverse chains, sometimes forming a cristulate sculpture, even more irregular and denser on juxtasutural area, becoming smaller posteriad and variably even more commonly anastomosing in chains, appearing to form a very irregular sculpture (but their shape also changes depending on angle of illumination); elytral surface glabrous except for several, hairlike sensory setae scattered mostly on basal area, and a few others adjacent to epipleura; elytral coloration notably metallic shiny, with conspicuous, iridescent reddish-cupreous area on elytral disc, mostly sharply delineated from usually wide, blue-green sublateral and apical areas (rarely passing through a narrow area of gold-bronze iridescence); lateral area usually deep violaceous or purple, also on juxtaepipleural area; whitish elytral maculation in male consisting of rather distinct maculae: humeral macula partly but clearly visible from above only in male, in female reduced and mostly invisible from above; lateromedian macula rather large, irregularly rounded or triangular, in female sometimes of a virgule-like shape; anteapical macula rather large in both sexes, slightly elongated.

Abdomen. Ventrites shiny metallic-black with greenish, blue, gold-bronze and violaceous lustre, glabrous, except for usual (easily abraded) hairlike sensory setae on their posterior margins.

Legs. Coxae metallic-green with blue or violaceous iridescence; pro- and mesocoxae densely white setose, metacoxae with only one apical and one discal seta (easily abraded) and densely setose lateral margin; trochanters glabrous (except for usual apical seta), yellow-ochre; femora on dorsal area brownish-testaceous, usually darker in female (fading in old specimens), often with mahogany lustre, with mostly distinctly delineated paler subapical area, basal and almost whole ventral area yellow-ochre to ochre-testaceous; pro- and mesofemora covered with rows of white to greyish, rather long, erect ore semierect setae which are densest on profemora and much sparser on metafemora; tibiae dark brown in male usually blackened on their apical third, rarely with testaceous apical half, usually darker in female, black-brown (in old specimens considerably faded to testaceous), covered with scattered white semierect setae and rusty thorn-like setae; tibial basal third to half covered by dense pad of greyish setae; metatibiae with only sparse semierect and stiffer short white setae and rusty thorns; tarsi virtually concolorous with tibiae, often with metallic-blue, greenish and purple lustre, mostly on protarsi, first three tarsomeres in male dilated and with usual pad of dense, greyish-white setae.

Aedeagus ( Figs 46–53 ) as in most other species of the genus moderately voluminous in middle, 3.20–3.40 mm long, 0.80–0.90 mm wide, apical portion gradually attenuated, towards rather narrow, elongate-spatulate, above rounded to subtruncate apex; apical orifice with a small, convex protrusion (usually with protruding flagellum); structure of internal sack ( Fig. 50 ) characteristic of the genus, with large voluminous, reniform, usually above folded central-ventral piece, but in this species complex connected with moderately bulbous base of convoluted flagellum, its flagelliform part usually protruding from the dorsoapical orifice; other sclerites comprise: thin, upper dorsal arciform piece with short, thin basal appendage, and large, elongate basodorsal piece.

Variability. Besides the variability stressed in the “Redescription” above, the pronotal disc is variably reddish or green-blue, particularly in middle; its smooth surface possesses only a few, very indistinct and shallow wrinkles along the median line, rarely a few, very shallow (light-refracting) wrinkles reach sublateral areas ( Figs 38, 40, 42 ), but never reach lateral areas.

Some adults are more vividly reddish-cupreous coloured on the elytral disc. The shiny coloration of the pronotum varies from reddish, golden, to blue-green. Type specimens both of O. mexicana and the synonymous O. quadrina have shiny, almost smooth and notably polished surface of the subglobose pronotal disc ( Figs 38–45 ) with only indistinct, shallow wrinkles along the median line. In some adults, mostly males, the few, short and shallow, more distinct wrinkles along the median line may appear in photographs as strongly light-refracting, but the lateral areas are always polished.

Because of the variable pattern of the elytral punctation, O. quadrina , was considered to be a separate species (see also in “Remarks” below). However, the isolated and only occasionally anastomosing elytral punctures in some adults including the type of O. mexicana commonly vary not only in populations, but also in syntopic adults, as demonstrated in Figs (19–37). Moreover, the pattern of the elytral punctation considerably changes depending on the angle of illumination: the isolated punctures, particularly on posterior elytral half, appear as anastomosing in chains when illuminated from a different angle; thus photos of the elytra taken under different angle of illuminations may show very different pattern of the punctation as demonstrated in Figs 22–23 and 24–25 .

The size and shape of the white elytral maculae also vary and can hardly be differentiating characters; the lateromedian macula is in some females of a virgule-like shape ( Figs 31–32 ), and the humeral macula in syntopic females, independent of the locality, is either reduced, barely visible from above, or entirely missing. Rivalier (1969) hesitated to consider O. quadrina a subspecies of O. mexicana , and he somewhat misleadingly noted that O. quadrina may differ in having smaller white maculae. Nevertheless, the female lectotype of the synonymous O. quadrina in MNHN from the collection Chevrolat (which was not obviously examined by Rivalier) have the whitish elytral maculae quite large ( Figs 6 , 26 ), comparable with those of the lectotype of O. mexicana ( Figs 1 , 24– 25 ). Some adults from Palenque (state of Chiapas) have large whitish maculae and coarser and more anastomosing elytral punctation as in the female lectotype of the synonymous O. quadrina , but the pattern of the elytral punctation varies not only in populations but also in syntopic adults as demonstrated in Figs 19–37 , with contrasting pattern also in the syntopic adults from Palenque ( Figs 33–37 ).

The coloration of the antennomeres also vary in syntopic adults independent of the locality, pattern of elytral punctation and other characters. Most adults from Chiapas and the female lectotype of the synonymous O. quadrina have their antennomeres 3–4 with lustrous purple to mahogany-testaceous apical areas ( Figs 7, 9 ), while most adults from the Valle National and Temazcal in the state of Oaxaca have these antennomeres mostly almost unicoloured ( Fig. 10 ). Nevertheless, some adults from Oaxaca also have the antennomeres 3–4 with contrasting purple apical areas, or possess an intermediate coloration with only diffusely purple to mahogany-testaceous apical areas ( Fig. 8 ). Most specimens from the state of Veracruz have the antennomeres 3–4 with deep purple apical areas. Such variability in antennal coloration is common also in some other species of Odontocheila .

Biology and distribution (map Fig. 208 ) O. mexicana has been confirmed during this revision from Mexico. The examined specimens listed above come from the states of Veracruz, Oaxaca, Chiapas, Puebla and possibly Guerrero. The records from Honduras by Horn (1910) and from Belize by Boyd (1982) have not been confirmed during this revision, but a colour photograph of a male of O. mexicana from San Antonio, Belize, is in Erwin & Pearson (2008: Plate 24).

As in most other species of Odontocheila , the adults inhabit trails through forests. The detailed behaviour of adults, their mating, ovipositing, larval tunnels and information of the larvae, as well as description of the biotopes in the tropical rainforest at Los Tuxtlas (Veracruz), in a field station of the University of Mexico was published by Palmer (1981). The two males and one female (RLHC), listed in the “Other Material examined” above, were reared by Marlene Palmer from 3rd instar larvae taken in deep woods with humid black soil and humus.

While the type locality of O. mexicana and the synonymous Cicindela nietoi is Mexico (without a specific locality), the type locality of Cicindela quadrina was mentioned by Chevrolat (1835) as Tutepec: “ Trouvée en terre chande, á Tutepec, par le jeune Sallé, prés des chemins, pendant le mois de juin ”. Tutepec is located in the state of Guerrero, about 110 km (by air) southeast of Chilpancingo de los Bravos, the capital of the region, and approximately 100 km (by air) northeast of Acapulco. However, as there is “Tustepec” written on the label ( Fig. 18 ) of the female lectotype in BMNH, the locality can alternatively be Tuxtepec, a region and the city in the state of Oaxaca. This is supported by the numerous adults caught in the Valle de National thus in the San Juan Bautista Tuxtepec area, as well as the adults caught along the road from Chiltepec to Temazcal. The locality Playa Vicente also is near Tuxtepec. This species was reported from Tuxtlas (Veracruz) also by Werner (1993), who accompanied it by a colour photograph.

Remarks. When Laporte de Castelnau (1834b) described Odontocheila mexicana , he did not mention the sex, nor number of specimens. He only noted that the beetle came from Mexico and from the collection of Audinet de Serville. Although Laporte did not describe the shape of the labrum, he described the elytra with only two whitish maculae on each elytron. Therefore, we should consider that the description was based on one or more females, because the whitish humeral macula is in this species in all examined males clearly visible from above, but barely visible in females. We only may suppose that the type or more syntypes were distributed by Laporte (François Luis Nompar de Caumont, Comte de Castelnau) to other museums, because the most probable genuine type is the female in BMNH designated here as the lectotype. It is well known that specimens including types were commonly distributed (sold or exchanged) by most of the historical authors ( Cambefort 2006). The large collection of Laporte was partly housed in the National Museum in Melbourne (see Horn & Kahle 1935), but also dispersed through sale to other museum and private collections. In 1842, a large part of his enormous collection was donated to the Smithsonian Institute in Washington, where it was destroyed during the large fire in 1865, but owing to the previous distributions, some specimens were preserved in other collections (see Evenhuis 1997).

The problem is rather complicated by the synonymous taxon validly described as Cicindela Nietii Guérin- Meneville, 1844, the species name later validly emended as “ nietoi ” by Fleutiaux (1892). These two names are written on the labels both of the lectotype in BMNH and one historical specimen in MNHN from the collection of Chaudoir, which hardly could have been a syntype because of its male sex. Another female labelled by these two names and additional name “ princeps ”, cannot be a syntype of O. mexicana as well, particularly because of the locality “ Mexico Cordova”, and coming from the collection of Sallé as written on the label (see under the “ Type material”). This female is very probably a type of C. nietoi , also because José Apolinario Nieto lived his whole life in the state of Veracruz between the cites of Cordoba and Orizaba, which may agree with the locality “Cordova” (= Cordoba) on the label of this female.

Cicindela princeps and C. dejeanii listed by Dejean (1837) without descriptions, and treated as synonyms of O. quadrina by Fleutiaux (1892) and Horn (1905, 1910), are unavailable names (nomina nuda).

The original description of Cicindela nietoi (as C. Nietii) by Guérin-Méneville (1844) as well as of Cicindela quadrina by Chevrolat (1835), clearly described female gender, because of the protruding median tooth of the labrum.

Odontocheila quadrina , was considered by Fleutiaux (1892), Horn et al (1897), Horn (1905, 1910,), and even recently by Johnson (1996) and Cassola & Pearson (2001) as a separate species. Rivalier (1969) considered it either as merely d’une simple forme individuelle, or possibly a subspecies of O. mexicana . This present revision has confirmed that the characters considered to be differentiating are variable (see the “Variability” above). Consequently, O. quadrina is here considered a junior synonym of O. mexicana in accordance with Horn (1922). The reasons for the synonymy presented by Horn (1922), particularly the variability in the elytral punctation, perfectly agree with the results demonstrated here. O. quadrina was listed as the synonym also by Horn (1926), followed by most subsequent authors such as Boyd (1982), Wiesner (1992), Werner (1993) and Erwin & Pearson (2008).

Figures

FIGURE 208. Map of distribution.

FIGURES 1 – 6. Odontocheila mexicana Laporte de Castelnau, habitus (all from Mexico). 1 — ♀, 10.6 mm, LT (BMNH); 2 — ♂, 8.6 mm, Valle National (CCJM ex DBCN); 3 — ♂, 9.3 mm, Los Tuxtlas (CCJM ex DBCN); 4 — ♂, 9.4 mm, Palenque (CCJM ex DBCN); 5 — ♂, 9.1 mm, “ quadrina, Mexique ” ex Sallé (MNHN); 6 — ♀, 9.3 mm, “ Tustepec [Tutepec?] ”, LT of syn. O. quadrina (MNHN).

FIGURES 7 – 18. Odontocheila mexicana Laporte de Castelnau. 7 – 10 — head: 7 –– ♂, Palenque (CCJM ex DBCN); 8 — ♂, Valle National (CCJM); 9 — ♀, 9.3 mm, “ Tustepec [Tutepec?] ”, LT of syn. O. quadrina (MNHN); 10 — ♀, Valle National (CCJM); 11 – 16 — labrum: 11 — ♂, Palenque (CCJM); 12 — ♂, Valle National (CCJM); 13 — ♂, Los Tuxtlas (CCJM); 14 — ♂, Temazcal (DBCN); 15 — ♀, LT (BMNH); 16 — ♀, LT of syn. O. quadrina (MNHN). 17 — original labels, LT of O. mexicana (BMNH); 18 — original labels, LT of syn. O. quadrina (MNHN). Bars = 1 mm.

FIGURES 19 – 27. Odontocheila mexicana Laporte de Castelnau, left elytron. 19 – 20 — ♂, Los Tuxtlas (CCJM ex DBCN); 21 — ♀, Fortin Las Flores (CMNH); 22 – 23 — ditto, detail of elytral sculpture under different angle of illumination; 24 – 25 — ♀, LT (BMNH) the same elytron under two slightly different angles of illumination; 26 — ♀, LT of syn. O. quadrina (MNHN); 27 — ♂, “ quadrina, Mexique ” ex Sallé (MNHN). Bar = 1 mm.

FIGURES 28 – 37. Odontocheila mexicana Laporte de Castelnau, left elytron. 28 — ♂, Valle National (CCJM ex DBCN); 29 – 30 — ♀, ibid., CCJM ex DBCN); 31 — ♀, Temazcal (DBCN); 32 — ♀, ex Chaudoir 1874 (MNHN); 33 — ♀, Palenque (CCJM ex DBCN); 34 – 37 — ♂, ibid., (CCJM ex DBCN). Bar = 1 mm.

FIGURES 38 – 45. Odontocheila mexicana, pronotum. 38 — ♀, LT (BMNH); 39 — ♀, LT of syn. O. quadrina (MNHN); 40 — ♀, ex Chaudoir (MNHN); 41 — ♀, Valle National (CCJM); 42 — ♂, ibid., (CCJM). 43 — ♀, Temazcal (DBCN); 44 – 45 — ♂, Palenque, (CCJM). Bar = 1 mm.

FIGURES 46 – 53. Odontocheila mexicana Laporte de Castelnau, aedeagus. 46 — Valle National (CCJM ex DBCN); 47 — Palenque (CCJM); 48 – 49 — Los Tuxtlas (CCJM ex DBCN); 50 — ditto, internal sac; 51 — apex, Los Tuxtlas (CCJM ex DBCN); 52 — apex, Temazcal (DBCN); 53 — apex, Palenque (CCJM). Bar = 1 mm.