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Pogonostoma (Microstenocera) noheli sp. nov. (Figs 1–20) Type locality. Eastern Madagascar: province of Toamasina (= Tamatave), Lakato forest near Moramanga. Pogonostoma (Microstenocera) flavomaculatum sensu Moravec (2007: 398), partim. Non P. (M.) flavomaculatum W. Horn, 1892! Type material. Holotype ♂ in SDEI, labelled: “Madagascar Est / Lakato / Moramanga / 20–23.I.2000, leg. Jan Vybíral” [printed] // “Pogonostoma (Microstenocera) / flavomaculatum / W. Horn, 1892 / det. Jiří Moravec 2003” [printed] // “Holotype / Pogonostoma (Microstenocera) / noheli sp. nov. det. / J. Moravec et Jan Vybíral 2017” [red, printed]. Allotype. 1 ♀ in CJVB: “Madagascar Est / Lakato / (Moramanga) / 20–23.I.2000, leg. Jiří Moravec” [printed] // “Allotype / Pogonostoma (Microstenocera) / noheli sp. nov. det. / J. Moravec et Jan Vybíral 2017” [red, printed]. Paratypes. 2 ♂♂, 1 ♀ in CCJM: “Madagascar Est / Lakato near Moramanga / 19.XII.1998 leg. Jiří Moravec”. 2 ♂♂, 2 ♀♀ in CJVB: ibid., except for leg. Jan Vybíral” // “Paratype / Pogonostoma (Microstenocera) / noheli sp. nov. det. / J. Moravec et Jan Vybíral 2017” [red, printed]. Differential diagnosis. Externally similar to P. (M.) flavomaculatum, but the main diagnostic difference is in the unique shape of the aedeagus of P. (M.) noheli sp. nov. which is in the middle ventrally deeply excavated, and then constricted into elongate thorn-like apex (Figs 14, 16, 18–20). In contrast, P. (M.) flavomaculatum possesses a longer aedeagus with a much wider, gradually conically attenuated apex, usually emarginated dorsally (Figs 37– 38, 40–43). Moreover, the pronotal posterior lobe of P. (M.) noheli sp. nov. is consistently ochre-yellow or testaceous, while in P. (M.) flavomaculatum the posterior lobe is mostly black, rarely partly brownish, or brownishtestaceous, very rarely predominantly testaceous. The pronotal disc in P. (M.) noheli sp. nov. (Figs 5–6) is more narrowed anteriad and notably oviform, particularly so in females, covered with more distinctly transverse rugae, and the labrum (Figs 12–13) is mostly brownish and with clearly outlined central convexity, while the pronotal disc in both sexes of P. (M.) flavomaculatum is almost regularly ellipsoid with much finer and irregular surface sculpture, and the labrum is entirely black and with central convexity almost keel-like raised in middle. Similar coloration of the body and appendages is also found in P. (M.) sawadai Moravec. (2007) described from Ranomafana (south-eastern Madagascar), but it clearly differs from both P. (M.) noheli sp. nov. and P. (M.) flavomaculatum in having its pronotal disc with much coarser surface sculpture consisting of more continuous transverse ridges, and with distinctly elevated notopleural sutures visible in dorsal view along the whole margins of the pronotal disc, and its aedeagus has the conical apical portion notably ventrally directed (Moravec 2007: 403, figs 1448–1450). P. (M.) pusillum (Laporte de Castelnau & Gory, 1835) immediately differs from P. (M.) noheli sp. nov. in having entirely black legs, truncate elytral apices in both sexes and very different shape of the aedeagus (similar to that in P. (M.) flavomaculatum W. Horn, 1892). Description. Body (Figs 1–2) small, length 5.55–7.50 (HT 7.30, AT 7.50) mm, width 1.55–1.90 (HT 1.70, AT 1.90) mm, both ventrally and dorsally shiny pitchy-black, except for ochre-yellow to ochre-testaceous pronotal posterior lobe; setal vesture whitish. Head (Fig. 9) narrower than body, 1.25–1.45 mm wide, with only one orbital sensory seta (on each side), lacking frontoclypeal setae; temples moderately long (2 times shorter than eyes). Frons flat or moderately convex, merging with clypeus in middle and not differentiated from vertex; supraantennal keels consisting of distinctly elevated anterior crest and smaller posterior one; vertex nearly flat on anterior area, convex between posterior half of eyes, with variably shallow or deep posterior impression; surface of the frons, anterior and median area of vertex rather finely scabrous-rugulose to partly areolate (thin intervals forming irregular subreticulate sculpture); posterior area more scabrous-rugulose, occipital area and temples with much more irregular, short and wavy rugae, or partly bumpy; whole dorsal surface densely covered with very short and therefore indistinct, decumbent to erect, whitish ornamental microtrichia which are barely obvious in dorsal view. Genae shiny black, variably very shallowly or markedly rugulose, particularly on postgenal area, finely wrinkled on anterior area, ventral half nearly smooth with sparse whitish micrortichia. Clypeus matt black, rough, distinctly elevated in frontoclypeal area, with sparse, short, whitish setae. Labrum (Fig. 12–13) 4–setose bearing long, yellowish setae, surface brownish or black-brown, rarely black, sparsely covered with short, appressed, white microtrichia; central convexity moderate but clearly outlined by semicircular shallow impression; shape and size similar in both sexes: lateral margins moderately rounded towards mostly blunt, more or less distinct lateral indentations, small, subacute or blunt, right-angled anterolateral teeth, constricted towards narrow median lobe which is short or more anteriad prolonged, shallowly emarginate between two blunt or subacute anterior teeth; male labrum 0.35–0.45 mm long, 0.60–0.85 mm wide; female labrum 0.45– 0.53 mm long, 0.75–0.85 mm wide. Maxillae (Fig. 11). Galea black with brownish apex; lacinia black-brown, elongate and outward-turned, gradually dilated towards simple, subacute apex with sinuous, but never bilobed margin, 0.15–0.70 mm wide; setae ochre-brown to testaceous. Palpi (Fig 9) black-brown to black with rusty to rusty-brown apical area of terminal palpomeres; setae on longest palpomeres very long, ivory-yellow. Mandibles (Figs 9–10) black-brown with reddish-brown to ochre brown to testaceous teeth and ochrebrownish juxtamolar area, separated by distinct edge from large, distinctly outward-convex basolateral portions, normally shaped, lateral margins (with terminal teeth) arcuate and moderately bent in middle, subsymmetrical (terminal tooth of left mandible slightly shorter than that of right one), both left and right mandible in male with second tooth smaller than third one, less markedly so in female. Antennae (Figs 1–2, 9) in male very slightly shorter than body, in female reaching only angles of anteapical convexity, black, indistinctly maculate: scape black or black-brown with ochre-testaceous ventral side, bearing white apical seta and several surface microsetae, pedicel and antennomere 3 black or black-brown; antennomere 4 either entirely black or with more or less paler, brownish or dark testaceous basal half, antennomere 5 with either indistinctly or markedly ochre-testaceous basal half, or entirely testaceous, also antennomere 6 sometimes with pale brownish basal half, remaining antennomeres black-brown or brown; setal vesture usual, antennomeres 3–4 rather densely covered with whitish to ochre microsetae, antennomeres 5–11 with usual micropubescence. Thorax. Pronotum (Figs 5–6) elongate, length 1.45–1.90 mm, width 0.90–1.05 mm; anterior lobe markedly narrower than posterior one, surface irregularly rugulose, nearly glabrous; disc notably oviform, particularly in female, with lateral margins moderately convex and narrowed anteriad; median line thin; notopleural sutures thin, indistinctly elevated, in dorsal view obvious only on posterior discal quarter or third; surface sculpture finely striate-rugulose, consisting of generally transverse rugae, continuous but also partly irregularly anastomosing, particularly on sublateral areas, but always clearly carved; whole discal surface appearing glabrous, but scattered, extremely short, indistinct microtrichia are obvious when the pronotum is observed in its lateral aspect in higher magnification; posterior lobe ochre or ochre-testaceous except for black area adjacent to disc, usually with small black spots, smooth and glabrous; proepisterna shiny and smooth except for shallowly wrinkled juxtanotopleural area, rarely finely wrinkled throughout, glabrous except for sparse microtrichia on ventral area; prosternum usually with testaceous anterior margin, sparsely covered with long and erect, whitish hairlike setae; basal half of mesosternum with scattered hairlike setae; metasternum with sparse microsetae; mesepisterna smooth, shiny, in both sexes with deep anterior-ventral furrow and scarce microtrichia on their ventral area; metepisterna transversely elongate with furrows, glabrous. Elytra (Figs 3–4) elongate, length 3.40–4.30 mm, dorsal surface with moderate basodiscal convexity, deep widely V-shaped discal impression and distinctly convex disc; humeri oblique-arcuate; lateral margins moderately dilated towards rounded angles of anteapical convexity, more distinctly so in female; apices with more or less distinct ochre coloured marginal area, sexually dimorphic: apex in male (Fig. 7) rounded in middle but with wide, shallow sutural emargination towards rounded sutural termination (usually lacking sutural spine); apex in female truncate as obliquely shallowly emarginate towards right-angled or blunt outer tooth, and with deep and steep sutural excision which forms right-angled, subacute, or sharpened inner tooth; elytral surface notably shiny, punctate throughout: punctures deep, isolated, with wide, shiny intervals, much larger, deeper and sparsely anastomosing on elytral base, within the V-shaped discal impression and on lateral areas of anterior elytral half, becoming smaller towards apices, but smallest on posterior half of elytral disc posteriad from the discal impression, particularly so along sutures, in female this area, lateral margins of humeri, basal area adjacent to pronotum and lateral areas of anteapical convexity are often almost effaced and polished; setal vesture white, ornamental setae irregularly arranged, very short, mostly appressed, arising both from the punctures and from micropits of microtexture, rather densely covering shiny intervals, densest on discal area, but on female elytra on the areas with more spaced punctures the setae appearing as only arising from the punctures, because those on the shiny intervals are usually sparser, tightly appressed and therefore barely visible; the ornamental setae are mixed with sparse, long, and erect, whitish hairlike sensory setae which are particularly present on humeral, basolateral and anteapical areas. Abdomen. Ventrites black except for testaceous margin of last ventrite, their surface sparsely or more densely covered with short whitish microtrichia. Legs (Figs 1–2). Coxae black or partly rusty-testaceous; trochanters ochre to brownish-testaceous, metatrochanters sometimes partly blackened; femora black with ochre-yellow apical areas, glabrous or with only few, indistinct, short, whitish microtrichia; tibiae black with ochre-yellow basal areas (“knees” together with tibiae, Fig. 21), protibiae rarely also with paler apical third, rather densely covered with short, whitish or ferrugineous microtrichia which are darker on metatibiae; tarsi black, rarely protarsi and even more rarely mesotarsi with first two tarsomeres brownish-testaceous, densely covered with whitish microtrichia; claws testaceous. Aedeagus of consistent shape (unique within the genus), notably short, 1.35–1.45 mm long, 0.22–0.25 mm wide, in its left lateral aspect (Figs 14, 16, 18–20) nearly straight with dorsal outline deeply excavated in middle, then constricted into narrow apical portion terminated with elongate, moderately ventrally directed and slightly dorsally bent, thorn-like apex which is in dorsal view (Figs 15) and ventral view (Fig. 17) conical and blunt. Variability. The shiny areas of almost or entirely effaced punctures on the elytral disc in female vary. The mostly brown labrum is rarely almost entirely black. While the yellow-testaceous apices of the femora and bases of tibiae (“knees”) are consistent in all specimens, the testaceous areas on antennomeres vary (as mentioned in the description). The male labrum is in some males longer, thus of the same shape as the female labrum. Etymology. Dedicated to Marcel Nohel, one of the friendly colleagues of the second author who is active in the research, specified to the protection of the ecosystems in Madagascar. Biology and distribution. The nine type specimens come only from the eastern evergreen rainforest near the village Lakato southeast of Moramanga, approximately 19°11´S; 48°26´E. The locality is between the forest stations Amboditiana and Ankasoka, 1050–1130 m (Viette 1991). Within our last visit in the January 2002, a great part of the forest was destroyed and other portions were just burning. The area is rather isolated from the Analamazaotra forest where P. (M.) flavomaculatum occurs. Remarks. In the monograph of the genus (Moravec 2007), P. (M.) noheli sp. nov. was previously partly included within the externally very similar P. (M.) flavomaculatum, but due to some differences from the holotype (by monotypy) of P. (M.) flavomaculatum deposited in SDEI, the possibility of an existence of a separate, undescribed species was mentioned in the monograph. For the circumstances leading to the confusion of these two species see “Remarks” under P. (Microstenocera) flavomaculatum below. The recent discovery of the numerous adults of the hitherto very rare P. (M.) flavomaculatum in the Andasibe-Mantadia National Park, has confirmed that these two species possess very different shape of their aedeagi, and that the photos in Moravec (2007, figs 1436–1437) show in fact the aedeagus of P. (M.) noheli sp. nov. The examination of the numerous specimens has enabled the separation and description of this new species.

Pogonostoma (Microstenocera) flavomaculatum W. Horn, 1892 (Figs. 21–44) Pogonostoma pusillum var. flavomaculatum W. Horn, 1892: 372. Type locality. “Madagascar” (Annanarivo on the label of the holotype—see “Biology and distribution” below). Pogonostoma flavomaculatum: Horn 1898: 24. Pogonostoma (Microstenocera) flavomaculatum: Jeannel 1946: 114, 141. Pogonostoma (Microstenocera) flavomaculatum: Rivalier 1970: 327. Pogonostoma (Microstenocera) flavomaculatum: Moravec 2015: 398, partim. Type material. Holotype ♂ (by monotypy) in SDEI, labelled: “Sikora[leg.] / Madagascar / Annanarivo” [printed] / / “P. flavomacule” [handwritten] // “ Type! / Dr W. Horn” [printed] // “Coll. W. Horn / DEI Eberswalde” [printed] // “ Holotype hat / sehr durch Essig- / säure / gelitten” [handwritten] // “flavomacula- / tum / mihi” [additionally attached blue-green collection label, handwritten] // “ Holotypus ” [red, printed] // “Revision Jiří Moravec 2003: / Holotype (by monotypy) / Pogonostoma pusillum / flavomaculatum W.Horn,1892 ” [red, printed] // “Pogonostoma / (Microstenocera) / flavomaculatum W. Horn, 1892 / det. Jiří Moravec 2003” [printed]. Note: the aedeagus of the holotype was destroyed, except for its basal portion glued to a separate small papered board. Other material examined. 1 ♀ in SDEI, 1 ♀ in MRAC: “La Mandraka”. 7 specimens (males and females) in SDEI: “Fôret Anamalazoatra [=Analamazaotra] / -Perinet, Dez. / N. Olsufiew 1930”. 2 ♂♂, 1 ♀ in MRAC: “ Forêt de Fito / ex coll. Dr. Breuning ”. 1 ♀ in NMPC: “ Madagascar ”. Recent data. 1 ♀ in CCJM: “ Madagascar est., 930– 1000 m / Andasibe Perinet / 6–7.II.1993, J. Janák lgt.”. 1 ♀ in CCJM: “ Madagascar / Toamasina distr. / Analamazaotra env. / 3–6.XII.1997 / Jan Stolarczyk lgt.”. 2 ♂♂, 1 ♀ in JWCW, 2 ♂♂, 3 ♀♀ in HSCA: “ Madagascar / Toamasina prov. / Andasibe Nat. Park, Perinet / 19– 31.12.2001, 950 m / 48°27'E. 19°00'S / V. Dolin & Anreeva leg.”. 10 ♂♂, 9 ♀♀ in CJVB, 2 ♂♂, 3 ♀♀ in CCJM: “ Madagascar Centr. / Andasibe Perinet N. P. / Mantadia, 13.I.2017 / Jan & Ondřej Vybíral leg”. 3 ♂♂, 4 ♀♀ in CJVB, 1 ♂, 1 ♀ in CCJM: “ Madagascar Centr. / Analamazaotra Perinet N. P / Circuit Aduant, 12.I.2017 Jan & Ondřej Vybíral leg”. 1 ♂, 1 ♀ in CJVB: “ Madagascar Centr. / Analamazaotra Perinet N. P. / Maromizaha, 13. 1. 2017 Jan & Ondřej Vybíral leg”. 1 ♀ in CJVB: “ Madagascar Centr. / N. P. c. Indri / 28–30.I. 2015, M. Trýzna leg.”. 1 ♂, 1 ♀ in CMTD, 1 ♂, 1 ♀ in CCJM: “ Madagascar / Andasibe – Mantadia N.P. / Analamazaotra forest / 2–14.II.2007 M. Trýzna leg”. Differential diagnosis. Originally described by Horn (1892) as a variety of P. (M.) pusillum with which it shares similar shape of the aedeagus, but P. (M.) flavomaculatum is immediately distinguished by the testaceous apices of femora, bases of tibiae, and sometimes also variably partly, rarely almost entirely testaceous pronotal posterior lobe, as well as the antennomeres 3–5. P. (M.) flavomaculatum is externally very similar to P. (M.) noheli sp. nov., but it is clearly distinguished from the new species by its very different shape of its aedeagus in its lateral view (Figs 37–38, 40–43), almost regularly ellipsoid pronotal disc covered with notably more irregular, much denser and finer rugae, in some adults with more distinct notopleural sutures, and female elytra mostly lacking smooth areas. For other diagnostic differences including the inconsistently coloured posterior pronotal lobe, and also from other similar species, see the “Differential diagnosis” under P. (M.) noheli sp. nov. above, and in the “Variability” below. Redescription. Body (Figs 21–22) small (but generally larger than in P. (M.) noheli sp. nov.), 6.60–8.30 (HT 6.70) mm long, 1.70–2.00 (HT 1.70) mm wide, mostly entirely pitchy black or with the posterior pronotal lobe partly, rarely entirely brownish, or testaceous. Head (Fig. 31) 1.45–1.60 mm wide, shape and surface of frons, vertex, clypeus and genae as in P. (M.) noheli sp. nov., but the surface of frons-vertex area with less distinct areolate sculpture. Labrum shaped and possessing similar variability as in P. (M.) noheli sp. nov., but entirely black and with central convexity almost keel-like-raised as impressed in ether side (with less distinct or missing rounded outlined impression); male labrum (Figs 34–35) length 0.38–0.40 mm, width 0.65–0.80 mm; female labrum (Fig. 36) generally longer, 0.45–0.50 mm long, 0.75–0.85 mm wide. Maxillae (Figs 32–33) as in P. (M.) noheli sp. nov., lacinia up to 0.19 mm wide. Palpi (Fig. 31) generally as in P. (M.) noheli sp. nov. Mandibles (Fig. 31) as in P. (M.) noheli sp. nov., but generally darker. Thorax. Pronotum (Figs 28–30) elongate, length 1.60–2.10 mm, width 0.90–1.20 mm (for that of HT see “Variability” below), anterior lobe narrower than posterior lobe, its surface irregularly rugulose; disc almost regularly ellipsoid, mostly with rather distinctly convex lateral margins and narrow, but distinct notopleural sutures obvious in lateral view along posterior half of the disc; discal surface very finely and irregularly rugulose, rugae short and wavy to vermicular, becoming extremely fine, irregularly anastomosing and fragmented on anterior pronotal third, more transverse only along the median line (sculpture much finer than in P. (M.) noheli sp. nov.); median line mostly indistinct as partly merging with the surface sculpture; discal surface covered with indistinct, extremely short microtrichia obvious only in lateral view; posterior lobe smooth and glabrous, mostly entirely black, rarely partly brownish, reddish brown or brownish-testaceous, very rarely predominantly testaceous; ventral and lateral thoracic sterna as in P. (M.) noheli sp. nov, but the proepisterna larger and always smooth. Elytra (Figs 23–27) elongate, length 4.00– 4.80 mm long, shape and surface impressions and setal vesture as in P. (M.) noheli sp. nov., as well as the punctation in male, but the punctures generally somewhat smaller, and in females the punctures on posterior juxtasutural area mostly less effaced (see the “Variability” below). Legs (Figs 21–22) as in P. (M.) noheli sp. nov., but the yellow to ochre-testaceous areas (“knees”) are sometimes restricted only on the bases of tibiae (Fig. 22) while the apices of femora may be black (either only on meso- and metafemora, or also on profemora). Abdomen as in P. (M.) noheli sp. nov., but all ventrites mostly black in both sexes. Aedeagus almost straight, in its left lateral view (Figs 37–38, 40–43) dorsally shallowly excavated above the middle, the apical portion gradually conically attenuated towards apex, mostly ventrally shallowly emarginate towards blunt or nearly pointed apex; in its ventral view (Fig. 39), as well as in dorsal view, the apex is conical and blunt. Variability. The holotype is rather aberrant specimen and belongs to the smallest specimens of this species. It is in a rather bad shape, now even worse that in the photo (Fig. 21) taken 11 years ago by the first author. Apart from the previously broken aedeagus, now the holotype is missing its right antenna (it is separately glued together with a maxillary palpus on the same papered board with the basal portion of the broken aedeagus), middle right leg and apical portion of left metafemur and missing the metatibia, its abdomen partly crashed and glued together including the right posterior leg. Its pronotum (Fig. 30), has less convex lateral margins of the pronotal disc than in most other specimens, the margins with slightly impressed anterior parts, the shape probably caused during the metamorphoses of the adult. It has black pronotal posterior lobe with only indistinctly reddish-brown narrow area on its posterior margin (as in most other adults including the recently collected specimens), despite the original description by Horn (1892) who mentioned testaceous pronotal base. One of the labels in Fig 44), handwritten by Horn and attached to the holotype says that the holotype is damaged by an action of acetic acid. As noted by the first author (Moravec 2007), the acid barely could have changed the coloration of the pronotum to black, because the legs of the holotype retain their original yellow-testaceous areas (as in other adults). The shape of the aedeagus somewhat varies, the apical portion is in some aedeagi more distinctly (but always shallowly) emarginated and consequently the apex is subacute or nearly pointed. The shape well corresponds with the schematic drawing of the aedeagus (in its reverse right lateral aspect) by Horn (1934a, pl. 2, fig. 46), particularly comparable with Figs 37, 38, 40, 42 here; while the schematic drawing by Rivalier (1970: 326, fig. 23fl) is comparable to Figs 41 and 43 here. However, despite such slight variability, the aedeagi fundamentally differ from the unique and consistent shape of the aedeagi in P. (M.) noheli sp. nov. Also the other historical male specimens from the collection of Walther Horn (SDEI) have the same shape of the aedeagi within the above mentioned variability. Some specimens have the notopleural sutures notably more distinct, obvious in dorsal view along the whole margins of the pronotal disc, while all other characters, including the pattern of the sculpture on the pronotal disc and shape of their aedeagi, correspond with this species. The ochre-testaceous areas on legs are in some adults of P. (M.) flavomaculatum restricted only on the bases of tibiae while the apices of femora are partly or entirely black (Fig. 22). The effaced areas on the elytra also are variable. Biology and distribution. Hitherto rare species missing in most collections including BMNH. The type locality “ Madagascar ” mentioned in the original description by Horn (1892) and “Annanarivo” on the label of the holotype, is ambiguous, because Annanarivo either means an area near Antananarivo, probably including the Angavo massif (Anjozorobe) north-east of the capital, but it also was commonly used by historical insect dealers for a larger area, or as a synonym for Madagascar (see Moravec 2007). Horn (1934b) specified the occurrence as “Analamazaotra”, also mentioned by Jeannel (1946) as the occurrence of P. (M.) flavomaculatum. Most of the confirmed localities of this species lie in the area of the large Analamazaotra forest (preserved and still partly primary eastern evergreen rainforest) in the central-eastern area of the province of Toamasina (Tamatave). Although the non-protected forest remnants are mostly destroyed, the species has a chance to survive in the protected parts of the Andasibe–Mantadia National Park where this hitherto rare species was recently found in numerous adults. It must be noted here that the individual parts of the large national park (155 square kilometre, elevation 800–1260 m, are inconsistently interpreted and cited on the labels. In fact the Mantadia reserve (18°48´S, 48°25´E) is now rather isolated but well preserved rainforest northward of the Analamazaotra forest of the Andasibe-Perinet special reserve (18°57´S, 48°25´E) and the nearby Maromizaha reserve. The examined specimens labelled “Forêt de Fito / ex coll. Dr. Breuning” (MRAC), possess all characters of P. (M.) flavomaculatum including their aedeagi but the locality is in an area near the lake Lac Alaotra, north-east of Toamasina (approximately 17°30´S, 48°30´E), a long way from the confirmed localities. It must be noted here that some of the specimens from Breuning proved to be evidently mislabelled (see Moravec 2010). Rivalier (1970), apart from the occurrence in the environs of Andasibe (Perinet) adopted from Olsoufieff (1934), also mentioned Antsihanaka (also spelled Antsianaka), a record gained from Perrot brothers who mostly collected in the area of the lake Lac Alaotra which also includes the forest Forêt de Fito mentioned above. Occurrence in La Mandraka, the forests station along the road between Antananarivo and Moramanga has not been confirmed with certainty as the specimens examined by the first author in SDEI and MRAC are females, and no other specimen was found in MNHN, although recorded from La Mandraka by Rivalier (1970). The forest of the valley is now nearly destroyed. Remarks. The previous redescription of P. (M.) flavomaculatum by the first author (Moravec 2007) partly included P. (M.) noheli sp. nov. The confusion was primarily caused by the external similarity of these two species which were at the time very rare, and also that the original description by Horn (1892) did not feature the aedeagus. Moreover, the holotype (SDEI) of P. (M.) flavomaculatum is in rather bad state (see “Variability” above) and when at the time examined by the first author, it was found with a broken aedeagus (only a small basal portion remained on the small papered board attached to the holotype). The only illustration of the aedeagus of P. (M.) flavomaculatum was the schematic drawing by Horn (1934a, pl. 2, fig. 46), the same figure adopted by Olsoufieff (1934), and the schematic drawing by Rivalier (1970: 326, fig. 23fl). A colour picture of the habitus of P. (M.) flavomaculatum, showing a partly testaceous posterior pronotal lobe, is in Horn (1910: pl. 6, fig. 6).

Pristimantis bounides sp. nov. Figs 4 & 5 Pristimantis sp. nov. D in Lehr & von May (2017) Common name. English: Hill Dweller Rubber Frog. Spanish: Rana cutín de montaña. Holotype. MUSM 31197 (IWU 182; Figs 4, 5), adult male from the buffer zone of the Pui Pui Protected Forest, Quebrada Tasta, " Runda ", 11° 26' 51.9'' S, 74° 53' 11.9''W, 3463 m a.s.l. (Fig. 2, 16 B), Provincia Satipo, Región Junín, collected on 19 May 2012 by E. Lehr and R. von May. Paratypes. A total of 6 (Figs 6, 7), from Provincia Satipo, Región Junín: 2 males (MUSM 31970, NMP 6 V 75097 [GenBank accession numbers KY962797, KY962790, KY962774]) collected with the holotype; 4 females (MUSM 31198 [GenBank accession numbers KY962794, KY962789, KY962771], 31971, NMP 6 V 75066 [GenBank accession numbers KY962796, KY962773]), 75540 [GenBank accession numbers KY962795, KY962772], from Sector Carrizal, Satipo-Toldopampa Road at km 134 on left side of street coming from Satipo (Fig. 16 A), 11° 29' 03.5'' S, 74° 53' 27.3''W, 3350 m a.s.l., collected on 23 June 2013 by E. Lehr, J. C. Cusi, and J. Moravec. Generic placement. We assign this species to Pristimantis based on molecular evidence (Fig. 3). Diagnosis. A new species of Pristimantis assigned to the Pristimantis danae species Group having the following combination of characters: (1) Skin on dorsum shagreen with scattered tubercles, skin on flanks tuberculate, skin on venter areolate with scattered low tubercles; discoidal fold and thoracic fold present; irregularly shaped, discontinuous dorsolateral folds present; (2) tympanic membrane and tympanic annulus present; (3) snout moderate, rounded in dorsal and in lateral views; (4) upper eyelid with few enlarged conical tubercles; EW shorter than IOD; cranial crests absent; (5) dentigerous processes of vomers present; (6) males with vocal slits and visible subgular vocal sac, nuptial pads absent; (7) Finger I shorter than Finger II; discs of digits slightly expanded, round, having circumferential grooves; (8) fingers with narrow lateral fringes; (9) minute low ulnar and tarsal tubercles present; (10) heel with a small conical tubercle; inner tarsal fold absent; (11) inner metatarsal tubercle ovoid, 1.5 times as large as outer; outer metatarsal tubercle small, rounded; few low supernumerary plantar tubercles; (12) toes with narrow lateral fringes; basal toe webbing present; Toe V longer than Toe III; discs of digits slightly expanded, round, having circumferential grooves, toe discs slightly smaller than those on fingers; (13) in life, dorsal and lateral ground coloration pale grayish brown, orange brown, yellowish brown or reddish brown with dark grayish-brown marmorations, flanks paler than dorsum; canthal and supratympanic stripes grayish brown, not well defined, diffuse; groin pale grayish brown, orange brown, yellowish brown or reddish brown with or without gray flecks; venter pale gray, pale greenish gray or creamish white with or without dark gray mottling; iris pale grayish green with fine black vermiculation, a short vertical black streak from pupil across lower half of iris and a reddish tint horizontally across lower half of iris; (14) SVL in males 18.2–21.0 mm (n = 3), in females 21.6–24.4 mm (n = 4). Comparisons. Pristimantis bounides sp. nov. is readily distinguished from its congeners in Ecuador (181 species), Peru (130 species), and Bolivia (17 species); AmphibiaWeb (2017), by having discontinuous dorsolateral folds, tuberculate flanks, an areolate venter, slightly expanded digits, and a polymorphic dorsal coloration consisting of pale grayish brown, orange brown, yellowish brown or reddish brown with dark grayish brown marmorations. Pristimantis bounides sp. nov. is similar to members of the former Pristimantis orestes species Group (considered to be not monophyletic by Duellman & Lehr 2009, not recognized by Padial et al. 2014). This group contained 17 species at high elevations in southern Ecuador and in Peru (Guayasamin &Artega 2013; Duellman & Lehr 2009), which have narrow digits usually with circumferential grooves (except P. simonsii [Boulenger, 1900]), and all (except P. simonsii) have a tympanum. The following five species are morphologically similar to P. bounides sp. nov.: Two species have discontinuous dorsolateral folds (P. chimu Lehr, 2007; P. cordovae [Lehr & Duellman, 2007b]), whereas three have continuous dorsolateral folds (P. seorsus Lehr, 2007; P. simonsii, P. ventriguttatus Lehr & Köhler, 2007). However, P. bounides sp. nov. lacks cranial crests (present in P. chimu), has males with vocal slits (absent in P. chimu, P. seorsus and in P. simonsii), discs on digits slightly expanded, round (notched in P. cordovae, broadly expanded, truncate in P. ventriguttatus), and tarsal fold absent (present in P. seorsus and in P. ventriguttatus). Furthermore, P. bounides sp. nov. is phylogenetically distant from members of the former P. orestes Group (P. melanogaster [Duellman & Pramuk, 1999], P. orestes [Lynch, 1979], P. simonbolivari [Wiens & Coloma, 1992], P. simonsii, Fig. 3). Pristimantis bounides sp. nov. shares with other members of the Pristimantis danae species Group (Fig. 3), dorsolateral folds and males (unknown for P. albertus) which have vocal slits present (absent in P. attenboroughi) and nuptial pads absent. Furthermore, all but P. attenboroughi and P. puipui sp. nov. have a tympanum and discs with circumferential grooves. Pristimantis bounides sp. nov. is larger than P. albertus (female SVL 21.6–24.4 [n = 4] vs. 19.7–20.7 [n = 2] in P. albertus; Duellman & Hedges 2007) and has the belly colored pale gray, pale greenish gray or creamish white with or without dark gray mottling (cream with brown mottling in P. albertus). Pristimantis bounides sp. nov. has dorsal skin shagreen with scattered tubercle (smooth in P. aniptopalmatus) and ventral skin areolate with scattered low tubercles (smooth in P. aniptopalmatus and P. ornatus). Pristimantis bounides sp. nov. easily can be distinguished from P. stictogaster which has a white belly with prominent dark brown to black, irregularly shaped spots. Pristimantis bounides sp. nov. is most similar to three other species of the Pristimantis danae species Group from upper montane forests and puna of the PPPF and its surroundings (Figs 2, 3): P. attenboroughi, P. humboldti sp. nov., and P. puipui sp. nov. All have discontinuous dorsolateral folds, tuberculate flanks, and an areolate venter. Pristimantis bounides sp. nov. and P. humboldti sp. nov. are sister taxa (Fig. 3), and are of similar size (male SVL 18.2–21.0 mm, n = 3 in P. bounides sp. nov. vs. male SVL 17.2–20.6 mm, n = 3 in P. humboldti sp. nov.). Furthermore, P. bounides sp. nov. and P. humboldti sp. nov. have a tympamum (absent in P. attenboroughi and P. puipui sp. nov.), and have expanded discs with circumferential grooves (absent in P. attenboroughi and P. puipui sp. nov.). Males of P. bounides sp. nov., P. humboldti sp. nov., and P. puipui sp. nov. have vocal slits (absent in P. attenboroughi). However, P. bounides sp. nov. has diffuse canthal and supratympanic stripes (well defined canthal and supratympanic stripes in P. humboldti sp. nov.), interorbital bar absent (present in P. humboldti sp. nov.), scapular without W- or X-shaped marks (W- or X-shaped marks present in P. humboldti sp. nov.), and the venter pale gray, pale greenish gray or creamish white with or without gray mottling (ventral coloration of throat, chest, belly, and anterior ventral surfaces of thighs dark gray and pale gray mottled with white and pale gray spots in P. humboldti sp. nov.). Description of holotype. Head about as long as wide; head length 38% of SVL, head width 37% of SVL; cranial crests absent; snout moderate, rounded in dorsal and lateral views (Figs 4 A, B); eye-nostril distance 89% of eye diameter; nostrils slightly protuberant, directed dorsolateral; canthus rostralis rounded in lateral view, straight in dorsal view; loreal region slightly concave; lips rounded; outer margin of upper eyelid with few enlarged conical tubercles; upper eyelid width 54% of IOD; supratympanic fold short and narrow, extending from posterior margin of upper eyelid curved around posterior margin of tympanum; tympanic membrane present, tympanic annulus weakly defined, its upper and posterior margin concealed by supratympanic fold; two conical, slightly fused postrictal tubercles on each side of head. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomers positioned posterior to level of choanae, vomerine teeth minute, embedded in buccal mucosa, narrowly separated; tongue long, narrow, about three times as long as wide, not notched posteriorly, posterior half one third free; vocal slits slightly curved, located at posterior half of mouth floor between tongue and margin of jaw; subgular vocal sac distinct. Skin on dorsum shagreen with scattered tubercles, skin on flanks tuberculate, irregularly shaped, discontinuous dorsolateral folds extending continuously from posterior level of tympanic area to level of mid body, extending discontinuously (separated into tubercles) to level of hind limb insertion; skin on throat smooth, chest and belly areolate with scattered low tubercles; discoidal fold and thoracic fold weakly defined; cloacal sheath short. Outer ulnar surface of right arm with row of five minute low tubercles, outer ulnar surface of left arm bearing four tubercles; palmar tubercle bifid; thenar tubercle ovoid; subarticular tubercles weakly defined, most prominent on base of fingers, round in ventral view, subconical in lateral view; few, low supernumerary tubercles present, about one third size of subarticular tubercles; fingers with narrow lateral fringes, Finger I shorter than Finger II; nuptial pads absent; discs on digits of fingers slightly expanded, round, having circumferential grooves (Fig. 5 A). Hind limbs short, slender, tibia length 45% of SVL; foot length 48% of SVL; dorsal surfaces of hind limbs tuberculate; inner surface of thighs smooth, posterior surfaces of thighs tuberculate, ventral surfaces of thighs areolate; heels each with a small conical tubercle; outer surface of tarsus with few scattered minute low tubercles; inner tarsal fold absent; inner metatarsal tubercle ovoid, one and a half times the size of round outer metatarsal tubercle; subarticular tubercles weakly defined, most prominent on base of toes, round in ventral view, subconical in lateral view; few low plantar supernumerary tubercles, about one third the size of subarticular tubercles; toes with narrow lateral fringes; basal webbing present; discs slightly expanded, round, less expanded than those on fingers, having circumferential grooves; relative length of toes: 1 Measurements of the holotype (in mm). SVL 18.2; tibia length 8.2; foot length 8.8; head length 6.9; head width 6.8; eye diameter 1.8; tympanum diameter 0.9; inter orbital distance 2.4; upper eyelid width 1.3; internarial distance 2.1; eye–nostril distance 1.6. Characters Pristimantis bounides sp. nov. Pristimantis humboldti sp. nov. Males (n = 3) Females (n = 4) Males (n = 3) Females (n = 6) SVL 18.1–21.0 (20.0 ± 1.3) 21.6–24.4 (22.9 ± 1.3) 17.2–20.6 (18.9 ± 1.4) 19.7–25.7 (22.5 ± 1.9) TL 8.2–9.2 (8.8 ± 0.4) 9.9–10.8 (10.3 ± 0.4) 7.8–8.7 (8.3 ± 0.4) 9.2–10.6 (10.1 ± 0.4) FL 8.8–9.9 (9.2 ± 0.5) 9.9–12.0 (10.8 ± 0.9) 8.3–9.0 (8.9 ± 0.5) 9.5–11.2 (10.4 ± 0.5) HL 6.9–7.9 (7.5 ± 0.4) 7.6–8.9 (8.4 ± 0.5) 6.8–8.5 (7.4 ± 0.8) 7.5–8.7 (8.2 ± 0.4) HW 6.8–8.0 (7.6 ± 0.5) 7.7–8.5 (8.0 ± 0.3) 6.1–7.5 (6.9 ± 0.6) 6.9–8.7 (7.8 ± 0.5) ED 1.8–2.2 (2.0 ± 0.2) 2.1–2.5 (2.3 ± 0.2) 2.1–2.2 (2.1 ± 0.0) 2.2–2.6 (2.4 ± 0.2) TY 0.9–1.0 (0.9 ± 0.0) 1.0–1.1 (1.1 ± 0.1) 0.6–0.9 (0.8 ± 0.1) 0.8–1.0 (0.9 ± 0.1) IOD 2.4–2.7 (2.5 ± 0.1) 2.7–3.0 (2.8 ± 0.1) 2.3–2.7 (2.4 ± 0.2) 2.5–3.3 (2.9 ± 0.2) EW 1.3–2.0 (1.7 ± 0.3) 1.4–1.7 (1.6 ± 0.1) 1.3–1.7 (1.5 ± 0.2) 1.4–2.0 (1.7 ± 0.2) IND 2.0–2.3 (2.1 ± 0.1) 2.2–2.5 (2.3 ± 0.1) 1.9–2.3 (2.1 ± 0.2) 2.0–2.6 (2.4 ± 0.2) E–N 1.4–1.8 (1.6 ± 0.2) 1.6–1.9 (1.8 ± 0.1) 1.4–1.7 (1.6 ± 0.1) 1.5–2.2 (1.8 ± 0.2) TL/SVL 0.43–0.45 0.44–0.46 0.42–0.45 0.40–0.49 FL/SVL 0.42–0.48 0.46–0.50 0.46–0.48 0.41–0.49 HL/SVL 0.38 0.35–0.37 0.37–0.41 0.34–0.38 HW/SVL 0.37–0.38 0.33–0.36 0.35–0.38 0.33–0.36 HW/HL 1.00 0.9–1.0 0.9–1.0 0.92–1.0 E–N/ED 0.64–0.95 0.72–0.79 0.64–0.81 0.58–0.88 TY/ED 0.45–0.50 0.40–0.52 0.27–0.43 0.35–0.42 EW/IOD 0.54–0.76 0.50–0.63 0.48–0.74 0.50–0.71 Coloration of the holotype in life (Fig. 4). Dorsal and lateral ground coloration pale yellowish brown with dark grayish-brown marmorations dorsally, flanks paler than dorsum. Canthal and supratympanic stripes grayish brown, not well defined, diffuse. Tympanum creamish white. Interorbital bar narrow grayish brown. Groin pale yellowish brown, anterior surfaces of thighs with a reddish tint. Throat and vocal sac pale yellowish brown, anterior half of chest pale grayish brown, posterior half of chest and belly pale creamish white, thighs pale grayish brown. Iris pale grayish green with fine black vermiculation, a short vertical black streak from pupil across lower half of iris and a reddish tint horizontally across lower half of iris. Coloration of the holotype in preservative. Dorsal and lateral ground coloration pale tan with grayish-brown marmorations dorsally, flanks paler than dorsum. Canthal and supratympanic stripes grayish brown, not well defined, diffuse. Narrow grayish brown interorbital bar. Groin pale tan, anterior surfaces of thighs pale tan, posterior surfaces of thighs pale grayish brown. Throat, chest, and belly pale tan with small gray dots, and narrow gray midventral line across chest and belly. Lower arms pale tan with narrow gray midventral line, tarsi and ventral surfaces of thighs pale grayish brown and tan mottled. Iris pale gray with fine black vermiculation. Variation. All paratypes (Figs 6, 7) are similar to the holotype regarding morphology and proportions (Tables 3, 4). Besides differences in SVL, notable morphological variation includes prominence of dorsolateral folds. Two females (MUSM 31971, NMP 6V 75540; Figs 7 E, H) have prominent dorsolateral folds of pale grayish coloration on dark grayish brown dorsum. Most striking is the coloration polymorphism which resembles the color range of plants the frogs inhabit (Fig. 16 C). The dorsal and lateral ground coloration is pale grayish brown (MUSM 31971, NMP 6V 75540; Figs 7 D, G), orange brown (NMP 6V 75097; Fig. 6 B), pale yellowish brown (MUSM 31198; Fig. 7 B) or reddish brown (MUSM 31970; Fig. 6 E) with dark grayish-brown marmorations. The dorsum and flanks have reddish brown flecks in one male (MUSM 31970; Fig. 6 E), pale yellowish brown in one female (MUSM 31198; Fig. 7 B), and orange in one female (NMP 6V 75540; Fig. 7 E). Flanks are paler than dorsum. Males have the throat and vocal sac pale grayish orange (MUSM 31970; Fig. 6 F) or salmon ((NMP 6V 75097; Fig. 6 C). The venter is pale gray (NMP 6V 75540, Fig. 7 F; MUSM 31970, Fig. 6 F; MUSM 31971, Fig. 7 I; NMP 6V 75066), pale greenish gray (NMP 6V 75097; Fig. 6 C) or creamish white (MUSM 31198; Fig. 7 C) with or without dark gray mottling. Etymology. The species epithet bounides is derived from the Greek noun “bounos” and means "dweller in the hills". The name refers to the habitat of montane forests where the new species was found. Distribution, natural history, and threat status. Pristimantis bounides sp. nov. is known from two locations in the humid montane forest ecoregion (Brack 1986, Figs 2, 16 A, B): the type locality in the Tasta valley, "Runda" at 3463 m a.s.l (7.5 km [airline] from the border of PPPF), and the Satipo-Toldopampa Road at km 134 on left side of street coming from Satipo at 3350 m a.s.l. (4.2 km [airline] from the type locality). The vegetation at the type locality consists of Peruvian feather grass (Stipa ichu), large layers of mosses, small bushes, trees, and forest patches (Fig. 16 B). The vegetation is disturbed by cattle breeding and deforestation. The male holotype was found calling from the center of Peruvian feather grass in the early afternoon on 19 May 2012 during light rain. No sympatric anuran species were found at the type locality. At km 134 of the Satipo-Toldopampa Road, calling males were heard from the moss layer in the early afternoon on 23 June 2013. On the same day a female P. bounides sp. nov. (NMP 6V 75540; Fig. 16 C) guarding 20 eggs was found there inside moss. The eggs were pale cream colored and had an average diameter of 3.4 ± 0.1 mm (3.2–3.7 mm, n = 20). Sympatric anurans include Phrynopus sp. A (IWU 285) and Gastrotheca griswoldi Shreve, 1941 (MUSM 31972). We propose that Pristimantis bounides sp. nov. should be classified as "Data Deficient" according to the IUCN Red List criteria and categories (IUCN Standards and Petitions Subcommittee 2016) based on the low number of specimens known. Threats include agriculture and cattle breeding in both valleys, more acute in the Toldopampa valley than in the Tasta valley.

Odontocheila exilis Bates, 1884 stat. restit (Figs 177���207, 208). Odontocheila exilis Bates, 1884: 260. Type locality. Panama: Chiriqui province, Volcan de Chiriqui. Odontochila exilis: Fleutiaux 1892: 123; Odontochila ignita exilis: Horn 1922: 102, 103; 1926: 120. Odontocheila ignita exilis: Wiesner 1992: 78. Type material. Lectotype (designated here) ♂ in BMNH, labelled: ���Type��� [circular with red frame, printed] // ���Bugaba / Panama / Champion��� // ���B. C. A. Coll., I (1). / Odontocheila / exilis ��� [printed] // ��� Odontocheila / exilis / Bates��� [handwritten] // ���Lectotype ♂ / Odontocheila / exilis Bts. / by Erwin 76��� [printed/handwritten] // ���The lectotype label / by Erwin is invalid / (unpublished)��� [printed]; ���Lectotype / Odontocheila exilis Bates, 1884 / design. Jiř�� Moravec 2012��� [red, printed]. Paralectotypes. 1 ♂ in BMNH with same second and third label. 2 ♂♂, 1 ♀ in MNHN: ���Bugaba / Panama / Champion��� [printed] // ���H. W. Bates / Biol. Cent. Amer��� [printed] // ���Mus��um Paris / 1952 / Coll. R. Oberth��r��� [greenish, printed] // ��� Odontocheila / exilis / Bates��� [handwritten]. 2 ♂♂ in MNHN [Fleutiaux Hist. Coll.] with same label data. 1 ♂, 1 ♀ in BMNH: ���Bugaba, / 800���1,500 ft. / Champion��� // ���Co- / type��� [circular with yellow border, printed] // ���B. C. A. Coll., I (1). / Odontocheila / exilis ��� [printed] ���F. Bates Coll. / 1911-248��� [printed]. 4 ♂♂, 2 ♀♀ in BMNH: ���Bugaba, / 800���1,500 ft. / Champion��� // ���B. C. A. Coll., I (1)��� [printed]. 1 ♂ in BMNH: ���Type��� [circular with red border, printed] // ���V. de Chiriqui / 2���3000 ft / Champion��� [printed] // ���sp. figured��� // ���B. C. A., Coll., I (1). / Odontocheila / exilis ��� [printed] // ��� Odontocheila / exilis / Bates��� [handwritten]. 1 ♀ in SDEI: ���Bugaba, / 800���1,500 ft. / Champion��� // ���Mus. Berolin��� [handwritten] / / ���Type! / Coll. W. Horn / DEI Eberswalde��� [printed] // ���f. / exilis / Bates��� [greenish with black border, handwritten] // ��� Odontocheila ignita exilis / Bates Type (DEI-Eberswalde / borrowed by D. L. Pearson / 23 Oct.1996 (drawer # 58)��� [printed]. 2 ♂♂, in MFNB with same first label. 1 ♀ in MFNB with same firs label and: ���73895��� // ��� Odontocheila / exilis Bates ���. 1 ♂ in IRSNB: ���Coll. R. I. Bc. N.B / Panama:��� [dark violet, printed] // ���Bugaba 800���1500 ft / Champion���[small, glued onto the large label, printed] // ��� Odontocheila / exilis, Bates / Biol. Centr. Amer. / 18484, 2: 260, 1��� // ���Paratype��� [sic!, orange with black border, printed]. All paralectotypes labelled: ���Revision Jiř�� Moravec 2012 (2013 or 2014 respectively): ���Paralectotype, Odontocheila exilis Bates, 1884! [red, printed]. Other material examined. Historical data. 1 ♂ in MNHN: ��� Turrialba [sic!] / Costa Rica ���// ��� Mus��um Paris / Ex Coll. M. Maindron / Coll. G. Babault 1830���. Other data. 2 ♂♂, 2 ♀♀ in FCCR: ���Progreso Panama / Chiriqui Prov. / 16���23.IV.1923 / F.M. Gaige ���. 1 ♂ in MNHN: ��� Peninsula / Costa Rica ���. Recent data. 1 ♂ in KCBC: ��� Panama Chiriqui Prov. / Bunca Peninsula / Reserve El Chirogo, 700 m / 8��18.05��N, 83��58.18��W / 10���11.VII.2009, L. Sekerka & D.Windsor lgt.���. 2 ♂♂ in DBCN: ��� Panama���Cocl�� / 7.2km NNE���El Cop�� / P.N. Omar Torrijos / D. Brzoska 20-V-1995 ���. 2 ♂♂, 2 ♀♀ in NMPC: ��� Panama, Cocl�� prov., El Cop�� / P.N. Omar Torrijos, 300���850 m / 0.8��40.083��N, 80��35.551��W / 4���7.V.2015, individual collection / L. Sekerka & k. ��tajnerov�� lgt.���. 1 ♂ DDCP: ��� Panama, Cocl�� province, El Cop��, 5.X.2003 / leg. Daniel Duran ���. 1 ♂ in DBCN: ��� Panama Bocas Del Toro / Almirante Rd. km 14 / 09��000.1��N, 80��15.9��W / D. Brzoska, 10-VI-1994 ���. 9 ♂♂ in DBCN, 1 ♂ in CCJM: ��� Costa Rica: Puntarenas / Tiskita Lodge���Punta Banco / D. Brzoska / 15-VI-1997 ���. 41 ♂♂, 4 ♀♀ in DBCN: ��� Costa Rica: Puntarenas / Manuel Antonio N.P. / D. Brzoska 24-VI-1990 ���. 24 ♂♂, 4 ♀♀ in DBCN: ibid., except for: ��� 20-V-1993 ���. 1 ♀ in CMNH: ��� Costa Rica, Puntarenas / Prov., Golfito / 21���26-VII-1981 / H.W. Weems, Jr, G.B. Edwards / forest edge���. 2 ♂♂, 2 ♀♀ in DBCN: ��� Costa Rica: Puntarenas / NE- Golfito Airport / D. Brzoska 26-V- 1989 ���. 1 ♀ in CCJM: ��� Costa Rica, Puntarenas / Ref. Nac. de Fauna / Silvestre Golfito / D. Brzoska, 28.V.1993 ���. 1 ♂ in DBCN, 1 ♀ in CCJM: ��� Costa Rica, Puntarenas / 7 km NE / Potrero Grande / D. Brzoska 13-VIII-1992 ���. 3 ♂♂, 2 ♀♀ in DBCN, 1 ♀ in CCJM: ��� Costa Rica: Puntarenas / 21.3 km S+ W P. Jimenez / Osa Peninsula / D. Brzoska 16-V-1996 ���. 44 ♂♂, 3 ♀♀ in DBCN: ��� Costa Rica: Puntarenas / Osa Peninsula / Morenco B.S. / D. Brzoska 23-VI-1990 ���. 32 ♂♂, 11 ♀♀ in DBCN: Costa Rica: Puntarenas / P.N. Piedras Blancas / Esquinas Lodge / D. Brzoska 15-V-1996 ���. 28 ♂♂, 7 ♀♀ in DBCN: ibid., except for: ��� 17-VI-1997 ���. 1 ♂ in CCJM: ��� Costa Rica, Puntarenas Prov. / (Golfito) La Gamba, 220m / Tropical Research Station / 8��42.46��N;83��12.9��W; Ocelot Trail / 18.V.2013, leg. Jiř�� Moravec���. 12 ♂♂, 4 ♀♀ in DBCN, 2 ♂♂, 2 ♀♀ in CCJM, 1 ♂, 1 ♀ in CJWB: ��� Costa Rica: Puntarenas / Wilson Botanical Garden / Gamboa Annex (Forest Trail) / D. Brzoska 13-VI-1997. 1 ♂, 1 ♀ in CCJM: ��� Costa Rica, Puntarenas Prov. / Wilson Botanical Garden, 1420m / (Las Cruces) 8��47��N;82��57��W / 13.V.2013 leg. Jiř�� Moravec���. 1 ♂ in RLHC: ��� Costa Rica: Osa Peninsula / Corcovado Nat. Park. / 22.VIII.1977 / D.L. Pearson ���. 1 ♂ in RLHC: ��� Costa Rica Pts. / nr Palmar Sur / & Hy to Cerro / 18.VIII.1965 / D.F. Veirs ���. 1 ♀ in RLHC: ��� Costa Rica: / Puntarenas Prov. / Rincon de Osa / pitfall in forest / 14���26.VII.1969 / T. Schuh, J. Crane ���. Differential diagnosis. O. exilis has been commonly considered to be a subspecies of O. ignita. However, although the external characters of these two species are similar, namely the dense and commonly anastomosing elytral punctures, O. exilis can be distinguished from O. ignita by its generally larger body size, longer labrum and almost subacute elytral apices in both sexes; moreover, females of O. exilis have reticulate-punctate sculpture on the elytral discal area. Males are immediately distinguished from other species by the different apex of their aedeagi, which is ventrally rounded, but curved dorsad, with dorsal margin blunt or more or less sharpened (cusped); (Figs 197���207), in contrast to the capitate apex of the aedeagus in O. ignita (Figs 161���176). O. exilis is in collections rarely confused with O. iodopleura with which it shares similar, dense and irregular elytral punctation and rather small whitish maculae, but the elytral punctures in O. exilis are even denser, and in females forming almost reticulate sculpture. The pronotal disc in O. exilis is more iridescent-coloured and notably narrower than the mostly dully coloured and wider, almost subglobose pronotal disc in O. iodopleura. Males of O. iodopleura are immediately distinguished by the very different apex of their aedeagi (Figs 105���120). For differences of other species treated in this paper see under the taxa above. Odontocheila camposi W. Horn, 1925 which occurs in Ecuador, possesses somewhat similar apex of its aedeagus, but the apex is longer and more convex ventrally, of a somewhat transiting shape to the aedeagus of O. margineguttata (Dejean, 1825). Moreover, O. camposi, is immediately distinguished from O. exilis by its pronotum with coarser and markedly zigzag-wavy rugae on the pronotal disc. Rivalier (1969), due to the somewhat similar shape of the aedeagi, and because O. camposi was unknown to him, tentatively considered specimens from Ecuadorian Chimbo to be ��� O. margineguttata pavida Erichson, 1844 ���. The specimens from Chimbo, arranged in MNHN under ��� O. m. pavida ���, and examined by the first author, proved to be conspecific with O. camposi. Unfortunately, some females of O. exilis are barely recognizable from females of O. salvini Bates, 1874. Both species are partially sympatric in Panama. Although O. exilis does not occur in south-eastern Panamanian provinces, identification problems may arise with individually caught females in central Panama where both species occur. Females of O. salvini may be distinguished merely by their generally larger size, almost entirely black mandibles (pale lateral stripe is very reduced) and almost entirely metallic black legs, but although the coloration of femora may vary in both species, trochanters in O. salvini are darkened, meso- and metatrochanters black. However, old specimens have all their leg segments faded to testaceous, such for instance one of the specimens standing in the historical Fleutiaux collection (MNHN) as ��� O. secedens Steinheil ���, which, except for its aedeagus characteristic of O. salvini, strongly resembles O. exilis. A better, more constant difference is in the shape of the elytral apices which are in females of O. salvini rounded, and rugae on the pronotum denser and zigzag wavy. Males of O. salvini are immediately recognizable by a very different and much narrower apex of their aedeagi, and their mandibles with conspicuously arcuately emarginate inner subapical margin of terminal teeth. Redescription. Body (Figs 177���180) of very variable size independent of sex, 8.10���11.2 (LT 9.60) mm long, 2.60���3.50 (LT 2.90) mm wide. Head (Fig.182) with notably pronounced eyes, notably large, as wide as body or slightly smaller, 2.70���3.40 mm wide, all head portions glabrous. Frons, vertex, genae and clypeus basically as in O. ignita with similar pattern and size of the striae and rugae on the surface of vertex, but vertex almost flat in middle. Labrum 4-setose; distinctly bicoloured; male labrum (Figs 183���186) basically shaped as in O. ignita, but notably longer, length 0.65���0.80 mm, width 1.10���1.25 mm, and its anterior margin is either shallowly emarginate in middle, or with small, or only indicated median tooth); female labrum (Fig. 187) 1.20���1.35 mm long, 1.25���1.50 mm wide, of similar shape as in females of other species of this complex, with prominent, acutely tridentate median lobe with protruding median tooth. Mandibles (Fig.182) variably brownish to metallic-black (faded to brown in old specimens), with distinct or diffusing, ivory-yellow lateral stripe (more extended in male), normally shaped and with teeth as in preceding species. Palpi (Fig. 182) shaped as in O. ignita, ivory-yellow to ochre-testaceous, terminal palpomeres in both labial and maxillary palpi metallic black; penultimate palpomeres of maxillary palpi variably (independent of sex) yellow-ochre to brownish-darkened, or black. Antennae (Figs 177���180, 182) notably long, in male surpassing elytral half, in female shorter, but almost reaching the elytral half; scape with only apical seta; coloration of antennomeres 1���4 (Fig. 182) as in O. iodopleura and O. ignita. Thorax. Pronotum (Figs 193���196) glabrous, coloured and shaped as in O. ignita, but even more distinctly longer than wide particularly in male, length 1.80���2.35 mm, width 1.70���2.10 mm; sulci well pronounced; anterior lobe slightly wider than posterior lobe and slightly narrower than disc; surface generally as in O. ignita; disc normally shaped (even narrower than in O. ignita) with only moderately convex or subparallel lateral margins of dorsally visible proepisterna; dorsally visible notopleural sutures are narrower and mutually subparallel, in female mostly slightly attenuated posteriad; medial line usually distinct, or almost merging with the surface sculpture; surface of the disc and posterior lobe sculptured as in O. ignita; all ventral and lateral sterna generally as in O. ignita. Elytra (Figs 188���192) elongate, length 5.60���7.60 mm, in male with rounded humeri which are subquadrate in female; lateral margins subparallel, outer margin only very slightly dilated in female, anteapical angles arcuate, then obliquely running towards apices which are subacute towards small sutural spine; dorsal elytral surface and pattern of commonly anastomosing punctures as in O. ignita, with cristulate pattern on elytral disc in male, but in female (rarely also in male) the thin intervals of the punctures form almost regularly punctate-reticulate sculpture; elytral coloration as in O. ignita, the green-blue lateral stripe is sometimes less distinct; whitish elytral maculation in both sexes as in O. ignita, but the humeral macula in male mostly smaller and in female very reduced, never visible from above or entirely missing. Abdomen as in O. ignita. Legs as in O. ignita, metatrochanters in some females blackened, and femora generally darker, particularly in female almost black (faded in old specimens). Aedeagus (Figs 197���207) voluminous in middle, 3.30���3.50 mm long, 0.90���0.95 mm wide, apex conspicuously shaped, ventrally rounded, curved dorsad and the dorsal margin blunt or more or less sharpened (cusped); internal sac (Fig. 200) as in the preceding species and other species of this complex. Variability. Apart from the insignificant variability in brightness of the body coloration comparable to that in O. ignita, the subacute elytral apex is slightly variable in shape, but always differs from the widely rounded apex in the elytra of O. ignita and O. salvini. The dense elytral sculpture with thin intervals is in females, rarely in males, rather punctate-reticulate than cristulate. The apex of the aedeagi somewhat varies in shape, as demonstrated in Figs 197���207; but only one aedeagus (Fig. 203) possessing somewhat intermediate shape between O. exilis and O. ignita has been found among the numerous aedeagi examined within this revision (see also under O. ignita above). Coloration of legs in old specimens is markedly faded. Biology and distribution. The type locality of O. exilis in the Panamanian province of Chiriqui, the Volcan de Chiriqui, is now named Volc��n Bar�� (3 474 m) and is situated in the Volc��n Bar�� National Park. This species occurs in southeastern Costa Rica, in most of the province of Puntarenas including the Osa Peninsula, spreading towards the Panamanian border and to the neighbouring Panamanian provinces of Chiriqui and Bocas Del Toro. Individual specimens of O. exilis come from the central Panamanian province of Cocl�� where it is sympatric with O. salvini, which, in contrast to O. exilis, is spread through the southeastern Panamanian provinces, including the province of Darien, to Colombia and Venezuela. One historical male specimen (missing its aedeagus) in MNHN (ex collections Maindron and Babault), labelled ���Turrialba, Costa Rica ��� (see ���Other Material examined��� above), possesses its external characters as those in O. exilis, but the specimen was obviously mislabelled (it has passed through different collections and the locality is additionally added in handwriting onto the same, printed label with ���Heine���). The occurrence of O. exilis in Turrialba in the Costa Rican province of Cartago, where only O. iodopleura commonly occurs, was not confirmed within this revision. As also mentioned under O. ignita above, the records of O. exilis (and O. ignita respectively) from Colombia and Venezuela were based either on a few confused or mislabelled specimens (see under O. ignita above), or belong in fact to O. salvini, while some other records from Costa Rica (outside of the province of Puntarenas) belong in fact to O. iodopleura. Werner (1993: 24, plate 12, fig. 90) published a colour photograph of ��� O. ignita exilis ��� from Pandora in the Costa Rican province of Lim��n, but either the label data or the specimens were confused with those of O. iodopleura, which is common in the province of Lim��n (the identity of the species is impossible to recognize from the small figure by Werner). Mandl (1961) listed this species (under O. ignita exilis) from El Salvador (���San Salvador���), but his specimens were obviously misidentified and belong probably to O. ignita. They were not found in the NHMW or the other collection listed in the acronyms in the ���Material and methods��� here. As no other specimens of O. ignita and O. exilis with locality labels outside of their confirmed occurrence have been found in collections except for the only few specimens mentioned under the taxa above, they are considered here mislabelled. It is noteworthy that the occurrence of mislabelled specimens (some old ones and mostly those spread by insect dealers) has been confirmed in collections also during previous revisions of tiger beetles from other regions (see Moravec 2002, 2007 and 2010). Remarks. The name Odontocheila exilis was not mentioned by Rivalier (1969) in his brief revision, as he probably confused it with O. ignita (see under O. ignita above). As mentioned in the ���Introductory��� above, O. exilis was considered as a subspecies of O. ignita by Horn (1922, 1926) and following authors including Mandl (1961), Boyd (1982), Wiesner (1992), Werner (1993), Johnson (1996), Cassola & Pearson (2001) and most recently by Erwin & Pearson (2008). However, particularly because of the different shape of the apex of its aedeagus, it is treated here as a separate species, thus restored to its original species status. A schematic, but rather appropriate shape (in reverse view) of the aedeagus of O. exilis was for the first time illustrated by Horn (1929, figs 24���25)., Published as part of Moravec, Ji����, 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., pp. 451-499 in Zootaxa 4231 (4) on pages 489-492, DOI: 10.11646/zootaxa.4231.4.1, http://zenodo.org/record/292697, {"references":["Bates, H. W. (1884) Coleoptera, Carabidae, Cicindelinae. Vol. 1. Part 1. Supplement. In: Godman, F. D. & Salvin, O. (Eds.), Biologia Centrali American. Taylor & Francis, London, pp. 256 - 316.","Fleutiaux, E. (1892) Catalogue systematique des Cicindelidae. Imprimerie H. Vaillant-Carmanne, Liege, 186 pp.","Horn, W. (1922) Studien uber neue und alte Cicindeliden (Col.), (Neubeschreibungen, Synonymie, Faunistik). Zoologische Mededeelingen, 7, 1 - 112.","Wiesner, J. (1992) Verzeichnis der Sandlaufkafer der Welt. Checklist of the tiger beetles of the world (Coleoptera, Cicindelidae). Verlag Erna Bauer, Keltern, 364 pp.","Rivalier, E. (1969) Demembrement du genre Odontochila (Col. Cicindelidae) et revision des principales especes. - Annales de la Societe entomologique de France, Nouvelle Serie, 5 (1), 195 - 237.","Werner, K. (1993) Cicindelidae Regionis Nearcticae (bis zum Panama Kanal und einschließlich der karibischen Inseln), Collyrini Cicindela. Die Kafer der Welt, 18, 1 - 163. [Sciences Naurelles, Venette, France]","Mandl, K. (1961) Eine Cicindeliden-Ausbeute aus San Salvador. Koleopterologische Rundschau, 39, 26 - 28.","Moravec, J. (2002) Tiger beetles of Madagascar 2. A monograph of the genus Physodeutera (Coleoptera: Cicindelidae). Nakladatelstvi Kabourek, Zlin, 290 pp.","Moravec, J. (2007) Tiger beetles of Madagascar 1. A monograph of the genus Pogonostoma (Coleoptera: Cicindelidae). Nakladatelstvi Kabourek, Zlin, 499 pp.","Moravec, J. (2010) Tiger beetles of the Madagascan Region (Madagascar, Seychelles, Comoros, Mascarenes, and other islands. Taxonomic revision of the 17 genera occurring in the region (Coleopterea: Cicindelidae). Biosfericka rezervace Dolni Morava, o. p. s., Lednice na Morave, 429 pp.","Horn, W. (1926) Carabidae, Cicindelinae (Pars 86). In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. Junk, Berlin, pp. 1 - 345.","Boyd, H. P. (1982) Checklist of Cicindelidae, the tiger beetles, annotated checklist of Cicindelidae (Coleoptera) of North and Central America and the West Indies. Plexus Publishing, Inc., New Jersey, VIII + 32 pp.","Johnson, W. (1996) A new species of Odontocheila from Honduras with notes on other Central American species (Coleoptera: Cicindelidae). Cicindela, 28 (3 / 4), 37 - 44.","Cassola, F. & Pearson, D. L. (2001) Neotropical Tiger Beetles (Coleoptera: Cicindelidae): Checklist and Biogeography. Biota Colombiana, 2 (1), 3 - 24.","Erwin, T. L. & Pearson, D. L. (2008) A treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions, and ways of the life. Vol. II. Carabidae - Nebriformes 2 - Cicindelitae. Pensoft Series Faunistica, 84. Pensoft Publishers, Sofia, 366 pp., 33 pls.","Horn, W. (1929) Sur deux especes nouvelles d'Odontochila neotropiques et quelques autres especes rapprochees. Revista Chilena de Historia Natural, 33, 154 - 158."]}

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 Gu��rin-M��neville, 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. Jiř�� 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?] Sall�� / Type��� [large green label, printed/handwritten] // ��� Collection / Chevrolat ��� [printed] // ��� Type ��� [printed] // ���Lectotype / Cicindela / quadrina Chevrolat, 1835 / design. Jiř�� Moravec 2015��� [red, printed] // ��� Odontocheila / quadrina (Chevrolat, 1835) / det. Jiř�� Moravec 2015��� [printed]. Type material of Cicindela Nietii [nietoi] Gu��rin-M��neville. 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 Gu��rin R. Z 1844, p 254 / Mexico / Cordova Sall����� [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��� // ��� Mus��um Paris / Coll. Chaudoir 1874��� // ��� Mexicana / Laporte / Nietii Gu��rin / Mexique ���. 1 ♂ in MNHN: ��� quadrina Chvrt. Centrim? [illegible] / Observ? [illegible]. 1 ♂ in MNHN: ��� Mexico / Janson ���. 3 ♂♂, 3 ♀♀ in MNHN: ���Mus��um Paris / Coll. Chaudoir 1874���. 4 ♂♂, 3 ♀♀ in BMNH: ��� Jalisco / Mex.��� // ��� F. Bates Coll. / 1911���248���. 1 ♂ in BMNH: ��� Toxpam / Mexico / Sall�� Coll. ��� // ��� Odontocheila / mexicana / apud Sall�� Coll. ���. 1 ♂ in BMNH: ��� Chinantla / Mexico / Sall�� Coll. ��� // ��� Odontocheila / mexicana / apud Sall�� Coll. ���. 1 ♀ in BMNH: ���16277��� / Mexico ���. 1 ♂, 1 ♀ in BMNH: ��� Truqui [leg.?] / Mexico / (ex Fry)���. 2 ♀♀ in BMNH: ��� Mexico / Cordova Sall�� Coll. ��� // ��� Odontocheila / mexicana / apud Sall�� 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 / Sall�� Coll. ��� // ��� H.W. Bates / Biol. Centr. Amer. ��� // ��� Mus��um Paris / Coll. R. Oberth��r��� // ��� quadrina Chevr. / apud Sal�� ���. 1 ♀ in BMNH: ��� Tuxtla ��� // ��� Mexico / Sall�� Coll. ��� // ��� San Andr�� ��� // Odontocheila / mexicana Lap. / apud Sall�� ��� // ��� B.C.A. Coll., I(1), / Odontocheila ���. 1 ♀ in MFNB: ��� Mexico, Meyer [leg.]��� // ��� Hist Coll. (Coleoptera) / Nr. 3627 / Odontocheila Nieti Gu��r. / Mexico, Meyer / Zool. Mus. Berlin ���. 1 ♂ in MFNB: ���3627��� // ��� Nieti Gu��r. / Mexico ��� // ��� Hist Coll. (Coleoptera) / Nr. 3627 / Odontocheila Nieti Gu��r. / 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 Sall�����. Other data. 2 ♂♂ in MNHN: Mexico / Jansen ���. 1 ♀ in BMNH, 1 ♂ in SDEI: ��� Cordoba / H��ge ���. 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 / H��ge ���. 1 ♀ in CMNH: ��� Oaxaca, Mex. / 26.V.1959 / G. y V. Halffter ��� // ��� Finca San Carlos / Union de l��s r��os / 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, Published as part of Moravec, Ji����, 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., pp. 451-499 in Zootaxa 4231 (4) on pages 455-466, DOI: 10.11646/zootaxa.4231.4.1, http://zenodo.org/record/292697, {"references":["Chevrolat, L. A. A. (1835) Coleopteres du Mexique. 8 e Fascicule. G. Silbermann, Strasbourg, 65 pp.","Horn, W. (1922) Studien uber neue und alte Cicindeliden (Col.), (Neubeschreibungen, Synonymie, Faunistik). Zoologische Mededeelingen, 7, 1 - 112.","Guerin-Meneville, M. (1844) Description de Coleopteres nouveaux decouveris par M. A. NIETO dans l'interieur du Mexique. Revue Zoologique pa la Societe Cuvierienne, 7, 253 - 255.","Fleutiaux, E. (1892) Catalogue systematique des Cicindelidae. Imprimerie H. Vaillant-Carmanne, Liege, 186 pp.","Rivalier, E. (1969) Demembrement du genre Odontochila (Col. Cicindelidae) et revision des principales especes. - Annales de la Societe entomologique de France, Nouvelle Serie, 5 (1), 195 - 237.","Johnson, W. (1996) A new species of Odontocheila from Honduras with notes on other Central American species (Coleoptera: Cicindelidae). Cicindela, 28 (3 / 4), 37 - 44.","Wiesner, J. (1992) Verzeichnis der Sandlaufkafer der Welt. Checklist of the tiger beetles of the world (Coleoptera, Cicindelidae). Verlag Erna Bauer, Keltern, 364 pp.","Sherborne, C. D. (1928) Part 16. In: Index Animalium. Sectio Secunda. B. Quaritch, Ltd., London, pp 3971 - 4194.","Bousquet, Y. (2016) Literatura Coleopterologica (1758 - 1900): a guide to selected books related to the taxonomy of Coleoptera with publication dates and notes. ZooKeys, 583, 1 - 776.","Sherborne, C. D. (1930) Part 22. In: Index Animalium. Sectio Secunda. B. Quaritch, Ltd., London, pp. 5349 - 5702.","Horn, W. (1910) Coleoptera Adephaga, Fam. Carabidae, Subfam. Cicindelinae. In: Wytsman, P. (Ed.), Genera Insectorum, 82, 105 - 208, pls. 1 - 15.","Boyd, H. P. (1982) Checklist of Cicindelidae, the tiger beetles, annotated checklist of Cicindelidae (Coleoptera) of North and Central America and the West Indies. Plexus Publishing, Inc., New Jersey, VIII + 32 pp.","Erwin, T. L. & Pearson, D. L. (2008) A treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions, and ways of the life. Vol. II. Carabidae - Nebriformes 2 - Cicindelitae. Pensoft Series Faunistica, 84. Pensoft Publishers, Sofia, 366 pp., 33 pls.","Palmer, M. (1981) Notes on the biology and behavior of Odontochila mexicana. Cicindela, 13 (3 / 4), 29 - 36.","Werner, K. (1993) Cicindelidae Regionis Nearcticae (bis zum Panama Kanal und einschließlich der karibischen Inseln), Collyrini Cicindela. Die Kafer der Welt, 18, 1 - 163. [Sciences Naurelles, Venette, France]","Cambefort, Y. (2006) Des coleopteres, des collections et des homes. Museum national d'Histoire naturelle, Paris, 375 pp.","Horn, W. & Kahle, I. (1935) Uber entomologischen Sammlungen (Ein Beitrag zur Geschichte der Entomologie). Entomologische Beihefte aus Berlin-Dahlem, 2, 1 - 160.","Dejean, P. M. F. A. (1837) Catalogue des Coleopteres de la collection de M. le Comte Dejean. Troisieme edition, revue, corrigee et augmentee. Mequignon-Marvis, Paris, 501 pp.","Horn, W. (1905) Systematischer Index der Cicindeliden. Deutsche Entomologische Zeitschrift, 2, 1 - 56.","Horn, W., Becker, R. & Hoge, C. F. (1897) Die mexicanischen Cicindeliden. Deutsche Entomologische Zeitschrift, 1, 161 - 186.","Cassola, F. & Pearson, D. L. (2001) Neotropical Tiger Beetles (Coleoptera: Cicindelidae): Checklist and Biogeography. Biota Colombiana, 2 (1), 3 - 24.","Horn, W. (1926) Carabidae, Cicindelinae (Pars 86). In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. Junk, Berlin, pp. 1 - 345."]}

Pristimantis humboldti sp. nov. Figs 8 & 9 Pristimantis sp. nov. C in Lehr and von May (2017) Common name. English: Humboldt’s Rubber Frog. Spanish: Rana cutín de Humboldt. Holotype. MUSM 31190 (IWU 168, Figs 8, 9), adult male from the Pui Pui Protected Forest, Quebrada Tarhuish, left bank of Antuyo River, " Shiusha ", upper cloud forest, 11° 21' 21.4''S, 74° 56' 10.8''W, 3318 m a.s.l. (Figs 2, 16 D, E), Provincia Satipo, Región Junín, collected on 14 May 2012 by E. Lehr and R. von May. Paratypes. A total of 8 (Figs 10, 11), all collected with the holotype: 6 females (MUSM 31189, 31191, 31192, 31194 [GenBank accession numbers KY962798, KY962791, KY962775], NMP 6V 75095, 75538 [GenBank accession numbers KY962799, KY962792, KY962776]), 2 males (NMP 6V 75096, 75539). Generic placement. We assign this species to Pristimantis based on molecular evidence (Fig. 3). Diagnosis. A new species of Pristimantis assigned to the Pristimantis danae species Group having the following combination of characters: (1) Skin on dorsum shagreen with scattered tubercles, skin on flanks tuberculate, skin on venter areolate; discoidal fold and thoracic fold present; irregularly shaped, discontinuous dorsolateral folds present; (2) tympanic membrane and tympanic annulus present; (3) snout moderate, rounded in dorsal and in lateral views; (4) upper eyelid with few enlarged conical tubercles; EW shorter than IOD; cranial crests absent; (5) dentigerous processes of vomers present; (6) males with vocal slits and visible subgular vocal sac, nuptial pads absent; (7) Finger I shorter than Finger II; discs of digits expanded, rounded, having circumferential grooves; (8) fingers with narrow lateral fringes; (9) small conical ulnar and tarsal tubercles present; (10) heel with a small conical tubercle; inner tarsal fold absent; (11) inner metatarsal tubercle ovoid, twice as large as outer; outer metatarsal tubercle small, rounded; few low supernumerary plantar tubercles; (12) toes with narrow lateral fringes; basal toe webbing absent; Toe V longer than Toe III; discs of digits expanded, rounded, having circumferential grooves, toe discs slightly smaller than those on fingers; (13) in life, dorsal and lateral ground coloration orange brown with brownish-olive blotches, orange brown with grayish-browm blotches and flecks, reddish brown with grayish-browm blotches or grayish brown with orange brown blotches; interorbital bar brownish olive or grayish brown, often followed by grayish-brown or brownish-olive marks in shape of a W or X or a combination of both on scapular region; dark grayish-brown canthal and supratympanic stripes present; groin blackish brown with white or pale gray spots and usually with an orange tint; throat, chest, belly, anterior and ventral surfaces of thighs, tibia and axilla dark gray and pale gray mottled with white and pale gray spots of different sizes and density; iris pale golden with fine black vermiculation and reddish tint horizontally across lower half of iris; (14) SVL in males 17.2–20.6 mm (n = 3), in females 19.7–25.7 mm (n = 6). Comparisons. Pristimantis humboldti sp. nov. is readily distinguished from its congeners in Ecuador (181 species), Peru (130 species), and Bolivia (17 species); AmphibiaWeb (2017), by having discontinuous dorsolateral folds, tuberculate flanks, and an areolate venter which is dark gray and pale gray mottled with white and pale gray spots. Pristimantis humboldti sp. nov. and P. phalaroinguinis (Lehr & Duellman, 2007b) from elevations between 1800 and 2600 m a.s.l. in northern Peru have the groin dark brown with white spots, but P. humboldti sp. nov. is much smaller (SVL in males 25.0– 31.4 mm in P. phalaroinguinis, Lehr & Duellman 2007b), has venter areolate (smooth), dorsolateral folds present (absent), males with vocal slits (absent), and throat, chest, belly, ventral surfaces of tibia, and axilla dark gray and pale gray mottled with white and pale gray spots (white spots restricted to groin). Pristimantis humboldti sp. nov. and P. ventriguttatus from 1800 m a.s.l. in northern Peru have dorsum shagreen, dorsolateral folds, areolate venter with pale spots, and have males of similar size (male SVL 17.8–21.8 mm, n = 3 in P. ventriguttatus, Lehr & Köhler 2007), but have digits moderately expanded (broadly expanded), and inner tarsal fold absent (present). Pristimantis humboldti sp. nov. and P. phoxocephalus (Lynch, 1979) from elevations between 1850 and 2770 m in southern Ecuador and northern Peru (Duellman & Lehr 2009) have a polymorphic dorsal coloration (dorsum gray to red or brown with few or no darker markings in P. phoxocephalus vs. dorsum orange brown, reddish brown or grayish brown with brownish olive, grayish brown or orange brown blotches and flecks in P. humboldti sp. nov.). However, P. humboldti sp. nov. has smaller males (male SVL 22.3– 29.9 mm [n = 29] in P. phoxocephalus, Lynch 1979), has dorsolateral folds (absent in P. phoxocephalus), males without nuptial pads (present), and venter dark gray and pale gray mottled with white and pale gray spots (venter cream). Pristimantis humboldti sp. nov. shares with other members of the Pristimantis danae species Group (Fig. 3), dorsolateral folds and males (unknown for P. albertus) which have vocal slits present (absent in P. attenboroughi) and nuptial pads absent. Furthermore, all but P. attenboroughi and P. puipui sp. nov. have a tympanum and discs with circumferential grooves. Pristimantis humboldti sp. nov. is larger than P. albertus (female SVL 19.7–25.7 [n = 6] vs. 19.7–20.7 [n = 2] in P. albertus; Duellman & Hedges 2007) and has the belly colored dark gray and pale gray mottled with white and pale gray spots of different sizes and density (cream with brown mottling in P. albertus). Pristimantis humboldti sp. nov. has dorsal skin shagreen with scattered tubercle (smooth in P. aniptopalmatus) and ventral skin areolate (smooth in P. aniptopalmatus and P. ornatus). Pristimantis humboldti sp. nov. easily can be distinguished from P. stictogaster which has a white belly with prominent dark brown to black, irregularly shaped spots. Pristimantis humboldti sp. nov. is most similar to three other species of the Pristimantis danae species Group from the upper montane forests and puna of the PPPF and its surroundings (Figs 2, 3): P. attenboroughi, P. bounides sp. nov., and P. puipui sp. nov. All have discontinuous dorsolateral folds, tuberculate flanks, and an areolate venter. Pristimantis humboldti sp. nov. and P. bounides sp. nov. are sister taxa (Fig. 3), and are of similar size (male SVL 17.2–20.6 mm, n = 3 in P. humboldti sp. nov. vs. 18.2–21.0 mm, n = 3 in P. bounides sp. nov.). Furthermore, P. humboldti sp. nov. and P. bounides sp. nov. have a tympamum (absent in P. attenboroughi and P. puipui sp. nov.), have expanded discs with circumferential grooves (absent in P. attenboroughi and P. puipui sp. nov.), and male P. humboldti sp. nov., P. bounides sp. nov., and P. puipui sp. nov. have vocal slits (absent in P. attenboroughi). However, P. humboldti sp. nov. has well defined canthal and supratympanic stripes (diffuse canthal and supratympanic stripes in P. bounides sp. nov.), interorbital bar present (interorbital bar absent in P. bounides sp. nov.), scapular with W- or X-shaped marks (scapular without W- or X-shaped marks in P. bounides sp. nov.), has ventral coloration of throat, chest, belly, and anterior ventral surfaces of thighs dark gray and pale gray mottled with white and pale gray spots (venter pale gray, pale greenish gray or creamish white with or without gray mottling in P. bounides sp. nov.), and has ventral coloration of throat, chest, belly, and anterior ventral surfaces of thighs dark gray and pale gray mottled with white and pale gray spots (venter pale gray, pale greenish gray or creamish white with or without gray mottling in P. bounides sp. nov.). Description of holotype. Head slightly shorter than wide; head length 37% of SVL; head width 38% of SVL; cranial crests absent; snout short, rounded in dorsal view, rounded in lateral view (Figs 8 A, B); eye-nostril distance 76% of eye diameter; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis short, rounded in lateral view, weakly concave in dorsal view; loreal region slightly concave; lips rounded; upper eyelids with enlarged conical tubercles (three on right, four on left upper eyelid); upper eyelid width 74% of IOD; supratympanic fold short and broad, extending from posterior margin of upper eyelid curved around posterior margin of tympanum; tympanic membrane present, tympanic annulus distinct, its upper and posterior margin concealed by supratympanic fold; conical postrictal tubercles present (two right, one left). Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomers positioned posterior to level of choanae, vomerine teeth minute, embedded in buccal mucosa, widely separated; tongue short, oval, about 1.5 times as long as wide, not notched posteriorly, posterior half free; vocal slits slightly curved, located at posterior half of mouth floor between tongue and margin of jaw; subgular vocal sac distinct. Skin on dorsum shagreen with low scattered tubercles, skin on flanks weakly tuberculate, irregularly shaped, discontinuous dorsolateral folds extending from posterior level of arm insertion with a distinct conical tubercle at its anterior end to level of hind limb insertion; skin on throat smooth, skin on chest and belly areolate; discoidal and thoracic folds weakly defined; cloacal sheath short. Outer ulnar surface of right arm with three minute low tubercles, that of left arm with two minute low tubercles; palmar tubercle bifid; thenar tubercle ovoid; subarticular tubercles distinct, most prominent on base of fingers, round in ventral view, subconical in lateral view; few, low supernumerary tubercles present, about one third the size of subarticular tubercles; fingers with narrow lacking lateral fringes, Finger I shorter than Finger II; nuptial pads absent; discs on digits of fingers expanded, round, having circumferential grooves (Fig. 9 A). Hind limbs moderate, slender; tibia length 46% of SVL; foot length 47% of SVL; dorsal surfaces of hind limbs tuberculate; inner surface of thighs smooth, posterior surfaces of thighs tuberculate, ventral surfaces of thighs areolate; heels with minute conical tubercles; outer surface of tarsus with few scattered minute low tubercles; inner tarsal fold absent; inner metatarsal tubercle ovoid, three times the size of round outer metatarsal tubercle; subarticular tubercles weakly defined, more distinct on base of toes, round in ventral view, subconical in lateral view; few low plantar supernumerary tubercles, about one third the size of subarticular tubercles; toes with narrow lateral fringes; basal webbing present; discs expanded, round, less expanded than those on fingers, having circumferential grooves; relative length of toes: 1 Measurements of the holotype (in mm). SVL 19.0; tibia length 8.8; foot length 9.0; head length 7.0; head width 7.2; eye diameter 2.1; TY 0.9; inter orbital distance 2.3; upper eyelid width 1.7; internarial distance 2.0; eye– nostril distance 1.6. Coloration of the holotype in life (Fig. 8). Dorsal and lateral ground coloration orange brown with brownish olive flecks. Brownish-olive interorbital bar and W-shaped mark on scapular region. Dark grayish-brown canthal and supratympanic stripes outlined by a narrow pale brownish yellow margin. Labial bars on upper lip brownish olive, diffuse. Groin grayish brown with white and pale gray spots and an orange tint. Throat, chest, belly and anterior ventral surfaces of thighs dark gray, pale gray, and orange brown mottled. Anterior and posterior surfaces of thighs, tibia, dorsal surfaces of thighs and axilla mottled with white and pale gray spots. Iris pale golden with fine black vermiculation and reddish tint horizontally across lower half of iris. Coloration of the holotype in preservative. Dorsal and lateral ground coloration pale tan and pale gray mottled. Dark brown interorbital bar and W-shaped mark on scapular region. Dark grayish-brown canthal and supratympanic stripes outlined by a narrow pale tan margin. Labial bars on upper lip pale grayish brown, diffuse. Groin grayish brown with white spots. Throat, chest, belly and anterior ventral surfaces of thighs dark gray, and tan brown mottled. Anterior and posterior surfaces of thighs, tibia, dorsal surfaces of tarsi and axilla with white and pale tan spots. Iris pale gray with fine black vermiculation. Variation. All paratypes (Figs 10, 11) are similar to the holotype regarding morphology and proportions (Tables 3, 4). The dorsal and lateral ground coloration is orange brown with brownish-olive blotches (NMP 6V 75096; Figs 10 A, B, D), orange brown with grayish-brown blotches and flecks (MUSM 31192; Figs 11 I–K), reddish brown with grayish-brown blotches (MUSM 31194, NMP 6V 75095, 75538, 75539; Figs 10 E, 11E, F) or grayish brown with orange-brown blotches (MUSM 31189; Figs 11 A–C). The interorbital bar is brownish olive or grayish brown, often followed by grayish-brown or brownish-olive mark in shape of a W (MUSM 31194) or X (NMP 6V 75096; Fig. 10 B) or a combination of both (MUSM 31189, 31191; Fig. 11 B) on scapular region. Dark grayish-brown canthal and supratympanic stripes present, outlined by a narrow pale gray margin (e.g., MUSM 31189, NMP 6V 75096; Figs. 10 A, 11A) or without such an outline (e.g., NMP 6V 75095, 75538; Fig. 11 E). Labial bars on upper lip are brownish olive, pale or dark grayish brown with or without pale tan outline, mostly diffuse (e.g., NMP 6V 75095, 75539; Figs 10 E, 11E), sometimes distinct (MUSM 31189, 31191, NMP 6V 75096; Figs 10 A, 11A) or absent (MUSM 31194). The groin is blackish brown with white or pale gray spots and usually with an orange tint (e.g., MUSM 31189, NMP 6V 75096; Figs 10 A, 11C). The throat, chest, belly and anterior ventral surfaces of thighs are dark gray and pale gray mottled with white and pale gray spots of different sizes and density (NMP 6V 75095, 75538, 75539; Figs 10 C, F, 11D, H, L), a pattern which seems to resemble lichens growing on a dark rock. Two specimens have anterior throat with a dark gray blotch (MUSM 31194, NMP 6V 75096; Fig. 10 C) and three specimens have a narrow dark grayish-brown midventral line across throat, chest, and belly (MUSM 31189, 31192, NMP 6V 75095; Figs 11 D, H, L). Etymology. We dedicate this species to Alexander von Humboldt (1769–1859), German geographer, explorer, naturalist, in recognition of his exploration of the New World, especially the Andes, and whose legacy continues to inspire generations. Distribution, natural history, and threat status. The type locality at 3318 m a.s.l. on the border of the PPPF (Figs 2, 16 D, E) is in the Quebrada Tarhuish on the left bank "Shiusha" of the Antuyo River and belongs to the upper montane forest (Brack 1986). The vegetation consists of small bushes, small trees, forest patches, Peruvian feather grass, fern, and moss layers. Pristimantis humboldti sp. nov. was found inside Peruvian feather grass bunches (MUSM 31189) in the early afternoon and at night (1900–2100 hrs) on low vegetation up to 1 m from ground (MUSM 31190, 31191, 31194, NMP 6 V 75095, 75096) and on ground (MUSM 31192) inside a forest patch. Males called during rain in late morning and afternoon, and at night. Sympatric anurans are Gastrotheca griswoldi (MUSM 31193), and Pristimantis attenboroughi (MUSM 31186, 31187, NMP 6 V 75522, 75523). Four specimens had ectoparasitic mite (Trombiculidae) infestations (MUSM 31192, 31194, NMP 6 V 75096, 75539; Fig. 10) ranging from one (MUSM 31192) to 17 (NMP 6 V 75096) mite capsules distributed on the flanks and posterior surfaces of thighs. We propose that P. humboldti sp. nov. should be classified as "Data Deficient" according to the IUCN Red List criteria and categories (IUCN Standards and Petitions Subcommittee 2016). Given that P. humboldti sp. nov. occurs inside the PPPF, the long-term protection of this species should be supported by this nationally recognized protected area. However, other factors such as fungal infections, climate change, and pollution continue to threaten many Andean amphibians even inside protected areas (Catenazzi & von May 2014).

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): 451-499, DOI: https://doi.org/10.11646/zootaxa.4231.4.1

Pristimantis puipui sp. nov. Figs 12 & 13 Pristimantis sp. nov. E in Lehr & von May (2017) Common name. English: Pui Pui Rubber Frog. Spanish: Rana cutín de Pui Pui. Holotype. MUSM 31982 (IWU 303; Figs 12, 13), adult male from the Pui Pui Protected Forest, Laguna Sinchon, puna, 11° 16' 56.3''S, 75° 03' 11.7''W, 3890 m a.s.l. (Figs 16 F, G), Provincia Satipo, Región Junín, collected on 30 June 2013 by E. Lehr, J.C. Cusi, and J. Moravec. Paratypes. A total of 6 (Figs 14, 15), all collected with the holotype: 4 females (NMP 6V 75056, 75541, 75542 [GenBank accession numbers KY962800, KY962777], MUSM 31981), 2 males (MUSM 31983, NMP 6V 75057). Generic placement. We assign this species to Pristimantis based on molecular evidence (Fig. 2). Diagnosis. A new species of Pristimantis assigned to the Pristimantis danae species Group having the following combination of characters: (1) Skin on dorsum shagreen with low scattered tubercles, skin on flanks tuberculate, skin on venter areolate; discoidal fold absent, thoracic fold present; irregularly shaped, discontinuous dorsolateral folds present; (2) tympanic membrane and tympanic annulus absent; (3) snout short, rounded in dorsal and in lateral views; (4) upper eyelid without enlarged conical tubercles; EW shorter than IOD; cranial crests absent; (5) dentigerous processes of vomers present; (6) males with vocal slits, subgular vocal sac not discernible, nuptial pads absent; (7) Finger I shorter than Finger II; tips of digits narrow, rounded, lacking circumferential grooves; (8) fingers without lateral fringes; (9) small conical ulnar and tarsal tubercles present; (10) heel without tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle ovoid, half the size of outer; outer metatarsal tubercle prominent, rounded; supernumerary plantar tubercles absent; (12) toes without lateral fringes; basal toe webbing absent; Toe V longer than Toe III; tips of digits narrow, rounded, lacking circumferential grooves, toe discs slightly smaller than those on fingers; (13) in life, dorsum and flanks pale orange brown, reddish brown or grayish brown with or without grayish-brown mottling, flanks paler than dorsum; canthal and supratympanic stripes absent; groin colored as flanks; venter pale cream and pale gray mottled; iris pale grayish green with fine black vermiculation and dark brown horizontal streak across pupil and lower half of iris; (14) SVL in males 16.1– 17.1 mm (n = 3), in females 20.6–22.4 mm (n = 4). Comparisons. Pristimantis puipui sp. nov. shares having narrow digits without circumferential grooves and the absence of a tympanic annulus and tympanic membrane with the Andean genera Phrynopus Peters, 1873 (except for Phrynopus auriculatus Duellman & Hedges, 2008, and P. peruanus Peters, 1873, 32 species between 2200 and 4400 m a.s.l. in central and northern Peru; Duellman & Lehr 2009, Frost 2017) and Bryophryne Hedges, Duellman & Heinicke, 2008 (except for B. flammiventris Lehr & Catenazzi, 2010 and B. gymnotis, Lehr & Catenazzi, 2009, 9 species between 2900 and 4120 m a.s.l. in southern Peru; Duellman & Lehr 2009, Frost 2017). Pristimantis puipui sp. nov. is readily distinguished from its congeners in Ecuador (181 species), Peru (130 species), and Bolivia (17 species); AmphibiaWeb (2017), by having tips of digits narrow without circumferential grooves, by lacking a tympanic annulus and tympanic membrane, and by having irregularly shaped, discontinuous dorsolateral folds. In Peru 19 species of Pristimantis lack a tympanum. These are P. academicus Lehr, Moravec & Gagliardi Urrutia, 2010, P. altamazonicus (Barbour & Dunn, 1921), P. ashaninka, P. attenboroughi, P. colodactylus (Lynch, 1979), P. coronatus Lehr & Duellman, 2007a, P. croceoinguinis (Lynch, 1968), P. cruciocularis (Lehr, Lundberg, Aguilar & von May, 2006), P. flavobracatus (Lehr, Lundberg, Aguilar & von May, 2006), P. imitatrix (Duellman, 1978b), P. lirellus (Dwyer, 1995), P. leucorrhinus Boano, Mazzotti & Sindaco, 2008, P. martiae (Lynch, 1974), P. minutulus Duellman & Hedges, 2007, P. rhabdocnemus (Duellman & Hedges, 2005), P. simonsii, P. tantanti (Lehr, Torres-Gastello & Suárez-Segovia, 2007), P. ventrimarmoratus (Boulenger, 1912), and P. vilcabambae Lehr, 2007. Of these, only P. attenboroughi and P. simonsii (member of the former Pristimantis orestes species Group, not recognized by Padial et al. 2014) share with P. puipui sp. nov. the characteristics of having narrow digits without circumferential grooves. Besides digit morphology and absence of a tympanum, P. puipui sp. nov. shares with P. attenboroughi, from the upper montane forests and puna of the PPPF and its buffer zone, a similar SVL (16.1–17.1 mm vs. 14.6–19.2 mm in males of P. attenboroughi, Lehr & von May 2017), tuberculate flanks, irregularly shaped, and discontinuous dorsolateral folds. However, males of P. puipui sp. nov. have vocal slits (absent in P. attenboroughi), and dorsum and flanks pale orange brown, reddish brown or grayish brown (pale or dark gray, reddish brown or brownish olive with dark gray scattered flecks), and venter pale cream and pale gray mottled (dark gray, pale gray, grayish brown or pale grayish green with or without dark gray mottling). Pristimantis puipui sp. nov. is smaller than P. simonsii from northern Peru (SVL in males 19.3–25.9 mm in P. simonsii, Lynch 1975), lacks an inner tarsal fold (present), and has males with vocal slits (absent) which lack nuptial pads (present). Pristimantis puipui sp. nov. shares with other members of the Pristimantis danae species Group (Fig. 3), dorsolateral folds and males (unknown for P. albertus) which have vocal slits present (absent in P. attenboroughi) and nuptial pads absent. However, P. puipui sp. nov. and P. attenboroughi are the only members of the Pristimantis danae species Group that lack both a tympanum and discs with circumferential grooves. Description of holotype. Head about as long as wide; head length and head width 35% of SVL; cranial crests absent; snout short, rounded in dorsal and in lateral views (Figs 12 B, C); eye-nostril distance 56% of eye diameter; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis short, rounded in lateral view, weakly concave in dorsal view; loreal region concave; lips rounded; upper eyelid with few small tubercles; upper eyelid width 57% of IOD; supratympanic fold broad, extending from posterior margin of upper eyelid slightly diagonally to insertion of arm; tympanic membrane and annulus absent; postrictal tubercles fused into a low ridge on each side of head. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomers positioned posterior to level of choanae, vomerine teeth minute, embedded in buccal mucosa, narrowly separated; tongue long, oval, about two times as long as wide, not notched posteriorly, posterior third free; vocal slits short, slightly curved, located at posterior fifth of mouth floor between tongue and margin of jaw; subgular vocal sac not discernible. Skin on dorsum shagreen with low scattered tubercles, skin on flanks tuberculate, irregularly shaped, discontinuous dorsolateral folds extending from level of arm insertion to anterior level of hind limb insertion; skin on throat smooth, skin on chest, and belly areolate; discoidal fold absent, thoracic fold present; cloacal sheath short. Outer ulnar surfaces each with a row of three minute low tubercles; palmar tubercle bifid; thenar tubercle ovoid; subarticular tubercles weakly defined, more distinct on base of fingers, round in ventral and lateral views; supernumerary tubercles absent; fingers short and stout lacking lateral fringes, Finger I shorter than Finger II; tips of digits of fingers narrow, round, lacking circumferential grooves (Fig. 13 A). Hind limbs short, slender, tibia length 40% of SVL; foot length 39% of SVL; dorsal surfaces of hind limbs tuberculate; inner surface of thighs smooth, posterior surfaces of thighs tuberculate, ventral surfaces of thighs areolate; heels lacking conical tubercles; outer surface of tarsus with few scattered minute low tubercles; inner tarsal fold absent; inner metatarsal tubercle ovoid, one and a half times smaller than round outer metatarsal tubercle; subarticular tubercles weakly defined, round in ventral and lateral views; plantar supernumerary tubercles absent; toes without lateral fringes; basal webbing absent; tips of digits narrow, round, slightly smaller than those on fingers, lacking circumferential grooves; relative length of toes: 1 Measurements of the holotype (in mm): SVL 16.1; tibia length 6.5; foot length 6.2; head length 5.7; head width 5.6; eye diameter 1.8; inter orbital distance 2.3; upper eyelid width 1.3; internarial distance 1.7; eye–nostril distance 1.0. Coloration of the holotype in life (Fig. 12). Dorsal and lateral ground coloration pale orange brown, flanks slightly paler than dorsum. Canthal, supratympanic stripes, and labial bars absent, but rim of canthus rostralis and outer edge of upper eyelid pale gray. Groin colored as flanks. Anterior surfaces of thighs pale grayish brown, posterior surfaces of thighs colored as flanks. Throat pale cream, chest, belly, and thighs pale creamish white and gray mottled. Pale grayish-brown line ventrally across arms and chest, and midventrally across throat, chest, and belly. Iris pale grayish green with fine black vermiculation and dark brown horizontal streak across pupil and lower half of iris. Coloration of the holotype in preservative. Dorsum pale tan and brown mottled, flanks pale gray and brown mottled. Throat pale cream with tiny brown spots, chest and belly pale cream and gray mottled, posterior surfaces of thighs pale cream and brown mottled. Fine dotted brown narrow line ventrally across arms and chest and midventrally across throat, chest, and belly. Iris gray with black vermiculation. Variation. All paratypes (Figs 14, 15) are similar to the holotype regarding morphology and proportions (Tables 3, 4). Coloration in life is weakly polymorphic. Dorsum and flanks are pale orange brown (MUSM 31981, NMP 6V 75542; Figs 15 D, E), reddish brown (MUSM 31983, NMP 6V 75056, 75057; Figs 14 A, B, D, E, 15G, H) or grayish brown (NMP 6V 75541; Figs 15 A, B) with grayish-brown mottling. Flanks are paler than dorsum (e.g., MUSM 31983; Fig. 14 A). Canthal and supratympanic stripes absent, but one specimen (MUSM 31983; Fig. 14 A) has a dark brown blotch on canthus rostralis and a dark brown line along upper lip. Three specimens have the rim of the canthus rostralis pale gray (MUSM 31981, NMP 6V 75057, 75542; Figs 14 D, E, 15D, E) and upper eyelids gray and brown mottled. Four specimens have a pale grayish-brown line ventrally across arms and chest, and midventrally across throat, chest, and belly as distinct as in the holotype (MUSM 31981, NMP 6V 75541; Figs 15 C) or weakly defined (MUSM 31983, NMP 6V 75056, 75542; Figs 15 F, 14C), and one specimen (NMP 6V 75057; Fig. 14 F) only has a weak midventral line. Etymology. The species epithet " puipui " is derived from the Quechua name Pui Pui meaning eyes of water in reference to the many lagunas in the Pui Pui Protected Forest and is used as noun in apposition. The type locality of P. puipui sp. nov. is in proximity of Laguna Sinchon which marks the approximate center of the Pui Pui Protected Forest. Distribution, natural history, and threat status. Pristimantis puipui sp. nov. is only known from the type locality at Laguna Sinchon inside the PPPF at 3880 m a.s.l, where it co-occurs with Phrynopus sp. A (MUSM 31984, 31985). Specimens of Pristimantis puipui sp. nov. were found during early afternoon inside moss (Figs 16 F, G). Two females (Fig. 16 H) guarding eggs were found on 30 June 2013. The eggs of the two clutches were pale cream colored and had an average diameter of 4.0 ± 0.2 mm (3.6–4.3 mm, n = 13, MUSM 31981) and 3.7 ± 0.1 mm (3.5–3.9 mm, n = 20, NMP 6 V 75542). We propose that P. puipui sp. nov. should be classified as "Data Deficient" according to the IUCN Red List criteria and categories (IUCN Standards and Petitions Subcommittee 2016).

ODONTOcheila micrOPTera nom. nov. (Figs 30–58) Odontochila euryoides W. Horn, 1922: 100, 101, fig. 9, primary junior homonym of Odontocheila bennigseni euryoides W. Horn, 1906: 85 (currently a subspecies of Euryarthron bennigseni (W. Horn, 1897) – see “Remarks” below. Type locality. “Brasilia”. Odontocheila ? euryoides: Wiesner 1992: 80. Odontocheila euryoides: Lorenz 1998a: 35. Type material. Holotype [originally of O. euryoides] (by monotypy), ♂ in SDEI, labelled: “Brasilia / ex cab Dieck.” [handwritten] // “Coll. Ehlers / V. de Poll” [printed]; “ Type! / Dr. W. Horn” [printed] // “coll. W. Horn / DEI Eberswalde” [printed] // “ Holotypus ” [red, printed] // “ euryoides, mihi” [large green-blue collection label with black frame, handwritten] // “ Odontocheila euryoides W. Horn Type (DEI –Eberswalde) / borrowed by D. L. Pearson / 23.Oct.1996 (drawer # 60)”[printed] // “Revision Jiří Moravec 2012: / Holotype (by monotypy) / Odontochila / euryoides W. Horn, 1922 ” [red, printed] // “ Odontocheila / euryoides W. Horn, 1922 / det Jiří Moravec 2012” [printed]. The holotype is accompanied with separately mounted thoracic wing, labelled by Horn: “Unterflügel / von / Od. euryoides / m.” [handwritten]. Other material examined. Historical data. 1 ♀ in MFNB: “Hist. Coll. (Coleoptera), Nr. 3642, Odontocheila nitidicollis / Dej. / Bahia, Freireiss / Zool. Mus. Berlin”. 1 ♀ in BMNH: “Estancia San Noria / Rio San Javier, / Santa Fe, Argentine / G. E. Bryant / 12.I.1912 ”; “ Odontochila / nitidicollis / Dej. / Dr. W. Horn det, 1925”; “Pres. by / Imp. Bur. Ent. / Brit. Mus. / 1925–292”. All specimens labelled: “ Odontocheila / euryoides W. Horn, 1922), det Jiří Moravec 2012 (2014 respectively)” Recent data. 2 ♂♂, 10 ♀♀ in DBCN, 3 ♂♂, 1 ♀ in CCJM: “Uruguay / Hyw. 5, KM. 395 (N.- Tacuarembo) 179 m, -31.684, -55.926° / D. Brzoska 15-I-2017 ”. 2 ♂♂, 4 ♀♀ in JCDG: ibid., except for “leg. Johnnie Chong”. 4 ♂♂, 2 ♀♀ in DBCN: ibid. except for: “D. Brzoska 16-I-2017 ”. 8 ♂♂, 5 ♀♀ in JCDG: ibid., except for: “leg. Johnnie Chong”. 3 ♂♂ in DBCN: ibid., except for: “Hyw. 26, KM. 220 (W.- Tacuarembo) 164 m., -31.763, -56.077°, D. Brzoska 16-I- 017°”. 5 ♂♂, 2 ♀♀ in JCDG: ibid., except for “leg. Johnnie Chong”. 1 ♂, 1♀ in DBCN ibid., except for: “246 (W.- Tacuarembo) 171 m, 31.711, -55.866”. 2 ♂♂ in JCDG: ibid., except for: “leg. Johnnie Chong”. Differential diagnosis. Odontocheila microptera nom. nov. shares the notably nitid, reddish-cupreous pronotal surface, immaculate elytra and coloration of other body portions with Odontocheila nitidicollis. However, O. microptera nom. nov. can be immediately distinguished by its uniquely mutually ovoid elytra in both sexes (Figs 30–36). Its male labrum is distinctly bicolored with large ivory-yellow anterior area (Figs 47–48) and has 4-dentate anterior margin which is truncate or shallowly emarginated in middle (lacking median tooth) with the anterolateral and anterior teeth in the same level, giving the male labrum almost transverse shape, in contrast to the predominantly black male labrum with 5-dentate anterior margin in O. nitidicollis. In contrast to the entirely black female labrum in O. nitidicollis, the female labrum in O. microptera nom. nov. is either entirely black or bicoloured and may resemble the bicoloured labrum in females of the species-group related to O. rutilans, including O. fulgens (but see annotation in the “Variability” below). Moreover, the terminal palpomeres of maxillary palpi are in O. microptera nom. nov. slightly but notably narrower (Figs 38–40, 42) than the almost spatulate-dilated terminal palpomeres in males of O. nitidicollis (Figs 10–12, 19), and also the antennal scape (Figs 20–21) is in O. nitidicollis slightly more dilated (but in both species never distinctly spatulate-dilated). Because of its uniquely atrophic thoracic wings which are firmly connate with the soft thoracic tissue (Fig. 49), O. microptera nom. nov., is the only flightless species of Odontocheila; the total length of the atrophic wings (Figs 50–51) does not surpass the elytral third, in contrast to the normally developed, long wings in O. nitidicollis (Fig. 22). The aedeagus apex (Figs 55–57) in O. microptera nom. nov. is a rather thick, dorsally directed hook, but the hook is on its ventral side rounded, thus more similar to the aedeagus apex of O. rutilans. Nevertheless, as in others of the species-group related to O. rutilans and O. fulgens (Klug, 1834), the shape in syntopic adults may vary, and in one male of O. microptera nom. nov. from Tacuarembo, Uruguay the aedeagus apex (Fig. 56) is somewhat similar to that in some of the aedeagi in O. nitidicollis. In turn, the aedeagus apex of the male of O. nitidicollis from Paraguayan Estancia Terrado (Fig. 28), as well as that from Bolivian Alta Vista (Fig. 29), is almost of the same shape as in most males of O. microptera nom. nov. Redescription. Body (Figs 30–32) medium-sized (of a variable size independent of sex) 9.40–11.4 (HT 9.50) mm long, 2.90–3.80 (HT 3.20) mm wide, dark coppery or more vividly reddish-cupreous except for shiny reddishcupreous pronotum or also head, with faint green-blue lateral iridescence. Head (Figs 10–12) shaped and coloured as in O. nitidicollis, width 2.80–3.35 mm. Frons, vertex genae and clypeus coloured and with surface sculptures as in O. nitidicollis, but vertex usually less convex in middle, sometimes almost flat or with shallow anteromedian impression. Labrum 4-setose, often strongly sexually dimorphic in coloration; male labrum (Figs 47–48) predominantly ivory-white (ochre-yellow in old specimens) except for narrow, black basolateral area, rather short, length 0.60– 0.75 mm, width 1.10–1.40 mm, basolateral and anterolateral teeth subacute or acute, anterior margin 4-dentate with anterolateral and anterior teeth in same level giving the labrum nearly transverse shape, truncate or shallowly emarginate in middle, lacking median tooth; female labrum (Figs 44–46) variably either entirely metallic black or with testaceous anteromedian or whole anterior area, but always with black basal area expanded to the middle, much longer, length 1.15–1.25 mm, width 1.40–1.50 mm, its tridentate median lobe with protruding median tooth of rather variable length. Mandibles (Figs 37–41) shaped and coloured as in O. nitidicollis, but often paler (faded to brownish-testaceous in old specimens, such as in HT), and with similar variability in shape of inner teeth, but the third tooth is more often and also in left mandible notably smaller than the second and fourth (Fig. 41). Palpi (Figs 38–40, 42) as in O. nitidicollis, but terminal palpomeres of maxillary in both sexes notably less distinctly dilated, width 0.15–0.17 mm. Antennae (Figs 30–32, 37–39) shaped and coloured as in O. nitidicollis, but notably shorter, in male reaching only elytral third, in female only elytral shoulders, coloration in old specimens usually faded (antennomeres 3–4 in HT faded to pale mahogany with only apices metallic-black); scape moderately dilated, width up to 0. 28 mm. Thorax. Pronotum (Figs 52–54) coloured and with surface sculpture as in O. nitidicollis, but the transverse rugae on pronotal disc are sometimes, particularly in male (also in HT) denser and more wavy; in male slightly longer than wide, length 2.00– 2.30 mm, width 1.90-2.15 mm, in female as long as wide or slightly wider, 2.20– 2.30 mm long, 2.20–2.40 mm wide. Elytra (Figs 33–36) mutually notably ovoid, their outer margin markedly arched, length 5.70–7.00 mm, with widely arcuate humeri; anteapical angles widely arched towards apices which are in both sexes mostly almost acute with only small emargination towards distinct sutural spine; elytral surface punctate, but coloration slightly less variable and generally darker; whitish elytral maculation entirely absent. Abdomen as in O. nitidicollis. Legs as in O. nitidicollis, the black coloration in old specimens (HT) usually faded. Aedeagus (Figs 55–57) moderately voluminous in middle, 3.10–3.50 mm long, 0.70–0.85 mm wide, with simply hooked apex which is ventrally rounded, rarely subquadrate (Fig. 56); internal sac as in O. nitidicollis and most other species of the species-group related to O. rutilans with large, normally shaped voluminous base of the flagellum (partly visible in Figs 55, 56). Variability. Apart from that emphasized in the “Differential diagnosis” and the “Redescription” above, the most important is the variable coloration of the female labrum which is either entirely black, or distinctly bicoloured, thus similar to the bicoloured labrum in most females of O. fulgens. Although O. fulgens immediately differs by its whitish elytral maculation, it is important to rectify here the error by Rivalier (1969) who stated that one of the distinguishing characters of O. fulgens is metallic black female labrum, and such misleading error was adopted by Pearson et al. (1999). In fact, the labrum of the female holotype (MFNB) of O. fulgens is distinctly bicolored, as also are the labra of majority of the other examined females of O. fulgens from Argentina and Paraguay, except for females from Rio Salado, Argentina (IRSNB), where the female labrum is prevailingly metallic black with only testaceous median area which is often very narrow. Etymology. The species name microptera = having small or rudimentary wings, is derived from ancient Greek. Biology and distribution. This species was hitherto known (under the name O. euryoides) only from the holotype labelled “Brasilia”. Horn (1922, 1910, 1926) mentioned only Brazil as its occurrence. Only two other historical specimens were identified in collections within this revision of the genus by the first author: one of them the female (MFNB) from Bahia, Brazil, the other the female (BMNH) from the Estancia San Noria, Santa Fe, Argentina, both cited above. Although the female (BMNH) from Argentina was previously labelled by Horn as O. nitidicollis, he later (Horn 1928) mentioned a possible occurrence of this species (under O. euryoides) also in Argentina. Therefore, the most surprising is that this flightless species was recently discovered by the third author and his collecting companion Johnnie Chong, in north-central Uruguay, in the three adjoining localities north of Tacuarembo (cited above), in a good number of 55 adults of both sexes. Nevertheless, it remains a great enigma how this flightless species could penetrate from the Atlantic Rainforest of north- and southeastern Brazil to the very distant country, as well as to Argentina. The flightless adults run and hide in the vegetation and due to their atrophic thoracic wings, this species was obviously forced out of the vegetative cover. It is one of the few Odotocheila that can survive in Uruguay where probably most of the native forest has been removed and replaced with pine and Eucalyptus, as also in the area of the locality of O. microptera. The biotope is partly similar to that of O. nitidicollis, which is, however, a very good flyer, and therefore may occupy somewhat more various biotopes. Unlike most species of Odontocheila which usually occur along forest trails and more shaded areas, these two species inhabit more open and grassy areas, but O. microptera nom. nov. was mostly found in heavier grassy places. At one of the three Uruguayan localities with pine trees (Fig. 58), the adults were found among grass, but not under the trees; in the other two places they were found in heavy grassy areas along the shoulders of highways where no trees were present. It also may be significant that no other tiger beetle species was found in the habitats together with O. microptera nom. nov. Remarks. Horn (1922) described Odontocheila euryoides despite the existence of Odontocheila bennigseni euryoides described previously by himself (Horn 1906) from Tanzania, and he kept this African taxon in the genus Odontocheila also later (Horn 1910, 1926) in his wide concept both of the subtribe Odontocheilina (as Odontochilina) and of the genus Odontocheila (as Odontochila) which in his sense also comprised some African and Asian taxa. Odontocheila bennigseni W. Horn, 1897 was first transferred to the African genus Euryarthron Guerin, 1849 by Rivalier (1957), but without mentioning its subspecies euryoides that was mentioned as a member of Euryarthron for the first time by Wiesner (1992). Thus both O. bennigseni euryoides W. Horn, 1906 and its junior primary homonym O. euryoides W. Horn, 1922 were in use congeneric until 1992. The homonymy was overlooked also by Lorenz (1998a, b, 2005a. b). According to Article 23.9.5 (ICZN 1999), when a species-group name in use is a junior primary homonym of another species-group name also in use, but the names apply to taxa not considered congeneric after 1899, the author must not automatically replace the junior homonym. However, in this case both homonyms were considered congeneric after 1899, so the junior name is permanently invalid (Article 57.2). Since there is no available synonym for Odontocheila euryoides W. Horn. 1922, the name must be replaced by a new substitute name (Article 60.3).

Pristimantis boucephalus sp. nov. urn:lsid:zoobank.org:act: BCF78234-F3F2-4A81- AE 80-1B4 BE 26B1E1F Figs 3–7, Tables 1 – 4 Pristimantis sp. 4 – Angulo et al. 2016: 4, figs 76–77. Suggested English name: Bigheaded Rubber Frog. Suggested Spanish name: Rana cutín cabezona. Fig. 2 (opposite page). The Bayesian phylogenetic tree of selected South American Pristimantis; for taxon sampling design see Material and methods. Nodes were collapsed if they appeared in less than 50% of the post burn-in tree samples. Support values of posterior probabilities (pp) given only for discussed lineages (stars indicate full support, pp = 1.00). Taxa in red occur in the Cordillera Yanachaga region and represent six divergent lineages/species groups (uncorrected p -distance 13–17%). The new species, P. boucephalus sp. nov., is a member of a speciose clade, an as yet unnamed species group, comprising both montane taxa from the Andes and Guiana Shield, and lowland taxa from the intervening Amazon Basin. Species-group names follow Padial et al. (2014). DNA sequences of taxa retrieved from GenBank are labelled by standard GenBank accession numbers, all other codes stand for our new material. Etymology The species epithet boucephalus is derived from the Greek prefix “ bou -“ meaning large, huge, or great, and the Greek noun “ cephalo ” meaning head. The name refers to the disproportionally large head of the new species. Material examined Holotype PERU: ♂, adult (Figs 3–5), Yanachaga-Chemillén National Park (Sector San Daniel), Distrito de Huancabamba, Provincia de Oxapampa, Región Pasco, Quebrada Yanachaga, 10°22′46.3 S, 75°27′43.0 W, 2950 m a.s.l., Edgar Lehr, Juan C. Cusi and Jiři Moravec leg., 19 Jan. 2012 at 20h30 (MUSM 31102, GenBank 16S rRNA barcode: KY006091). Paratypes (n = 4) PERU: 2 ♂♂ (MUSM 24477–78), 1 subadult ♀ (MUSM 24479) and 1 juvenile (MUSM 24474), all from the Yanachaga-Chemillén National Park (based on field numbers, detailed collecting data missing), collected during the 1990s, Javier Icochea leg., see Fig. 6. Generic placement We assign this species to Pristimantis based on its general morphological similarity to other members of the genus and our molecular data (Fig. 2). Definition A new species of Pristimantis, not assigned to any species group, having the following combination of characters: (1) Skin on dorsum smooth with few low scattered tubercles, skin on venter areolate with low scattered tubercles; discoidal and thoracic folds absent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus absent; (3) snout short, rounded in dorsal and lateral views; (4) upper eyelid with one enlarged conical tubercle at its center and one enlarged conical tubercle at its posterior end; EW slightly shorter than IOD; cranial crests absent; (5) dentigerous processes of vomers absent; (6) males with vocal slits, nuptial pads absent; (7) Finger I shorter than Finger II; discs of digits expanded, rounded; (8) fingers with lateral fringes; (9) small conical ulnar and tarsal tubercles present; (10) heel with a conical tubercle; inner tarsal fold absent; (11) inner metatarsal tubercle ovoid, 2.5 times as large as outer; outer metatarsal tubercle small, rounded; numerous low, supernumerary plantar tubercles; (12) toes with lateral fringes; basal toe webbing present; Toe V longer than Toe III; toe discs slightly smaller than those on fingers; (13) in life, dorsal ground coloration greenish gray with reddishbrown blotches and scattered brown flecks surrounded by black; canthal and supratympanic stripes absent; groin and anterior surfaces of thighs greenish yellow with black blotches; venter gray with pale reddish and greenish brown mottling and scattered dark gray flecks; iris reddish copper with fine black vermiculation and black narrow vertical streak from pupil across lower half of iris; (14) SVL in adult males 13.4–14.5 mm (n = 3), in adult females unknown (12.5 mm in single subadult female). Differential diagnosis Pristimantis boucephalus sp. nov. is readily distinguished from its congeners in Ecuador and Peru by its minute SVL, short snout, absence of a tympanum, large head, groin and anterior surfaces of thighs greenish yellow with black blotches, and reddish-copper iris. It is currently one of the smallest of the 130 species of Pristimantis from Peru (AmphibiaWeb 2017) and the 199 species of Pristimantis from Ecuador (Ron et al. 2017, see Table 2). Pristimantis boucephalus sp. nov. can be distinguished from other minute Andean species of Pristimantis from Ecuador (P. andinognomus, P. minimus, P. trachyblepharis) and Peru (P. caeruleonotus, P. coronatus, P. minutulus, P. trachyblepharis) as follows (characters of P. boucephalus in parenthesis, if not otherwise stated): Pristimantis minutulus Duellman & Hedges, 2007 from elevations of 250–1200 m a.s.l. in central Peru has larger males (SVL 13.1–17.6 mm [n = 7] vs 13.4–14.5 mm [n = 3] in P. boucephalus sp. nov.; Duellman & Hedges 2007). Pristimantis minutulus and P. boucephalus sp. nov. lack dorsolateral folds, discoidal folds, a tympanum, and dentigerous processes of vomers and both have a smooth dorsum and areolate venter, but P. minutulus has the snout moderately long (short), males without vocal slits (present), groin with large yellow spot (greenish yellow with black blotches), and iris reddish gray (reddish copper with fine black vermiculation). Pristimantis caeruleonotus Lehr, Aguilar, Siu-Ting & Jordán, 2007 from elevations of 2500–2900 m a.s.l. of humid montane forests in northern Peru has males of similar size (SVL 11.6–14.6 mm [n = 4] vs 13.4–14.5 mm [n = 3] in P. boucephalus sp. nov.; Lehr et al. 2007). Both P. caeruleonotus and P. boucephalus sp. nov. have fingers and toes with lateral fringes, heel with conical tubercle, and males that have vocal slits, but lack nuptial pads. However, P. caeruleonotus has a distinct tympanum (absent), dorsolateral folds (absent), small dentigerous processes of vomers (absent), a discoidal fold (absent), prominent dorsolateral folds ending in a sacral tubercle (absent), and a long snout with terminal papillae (short snout without papillae). Furthermore, P. caeruleonotus has the groin blackish brown with yellow and pale blue blotches (greenish yellow with black blotches). Pristimantis coronatus Lehr & Duellman, 2007 is known from a single female specimen of 15.3 mm SVL from 2850 m a.s.l. in northern Peru (Lehr & Duellman 2007). It shares with P. boucephalus sp. nov. the absence of dorsolateral folds and a tympanum, and the presence of conical tubercles on the upper eyelid and fingers and toes with lateral fringes. However, P. coronatus has dentigerous processes of vomers (absent), groin red and black (greenish yellow with black blotches), and the iris dark brown (reddish copper). Pristimantis trachyblepharis (Boulenger, 1918) from elevations of 100–1250 m a.s.l. in southern Ecuador and northern Peru and P. boucephalus sp. nov. have smooth dorsum, areolate venter, dorsolateral folds absent, and males that lack nuptial pads. However, P. trachyblepharis tends to have larger males (SVL 12.1–15.8 mm [n = 20] vs 13.4–14.5 mm [n = 3] in P. boucephalus sp. nov.), tympanum present (absent), males without vocal slits (present), dentigerous processes of vomers (absent), and fingers and toes without lateral fringes (present) (Lynch & Duellman 1980). Pristimantis andinognomus Lehr & Coloma, 2008 from 2450–2800 m a.s.l. in the Cordillera Oriental in southern Ecuador (Zamora Chinchipe) has males of similar size (SVL 10.0– 14.5 mm [n = 20] vs 13.4–14.5 mm [n = 3] in P. boucephalus sp. nov.), males that have vocal slits but lack nuptial pads, fingers and toes with lateral fringes, and supernumerary plantar tubercles (Lehr & Coloma 2008). However, Pristimantis andinognomus has short dorsolateral folds that end in a conical sacral tubercle (absent), tympanic membrane and tympanic annulus present (absent), a short snout with terminal tubercle (terminal tubercle absent), minute dentigerous processes of vomers (absent), flanks with tubercles coalescing into short ridges (ridges absent), heel with two enlarged tubercles (one heel tubercle), and inner tarsal fold present (absent). Pristimantis minimus Terán-Valdez & Guayasamin, 2010 from the Cordillera del Condor in Ecuador at elevations between 1250 and 1685 m a.s.l. has smaller males on average (SVL 9.5–13.7 mm [n = 46] vs 13.4–14.5 mm [n = 3] in P. boucephalus sp. nov.; Terán-Valdez & Guayasamin 2010). Pristimantis minimus and P. boucephalus sp. nov. lack dorsolateral, discoidal and thoracic folds, have a short snout, and males have vocal slits present and nuptial pads absent. However, P. minimus has dentigerous processes of vomers (absent), fingers and toes without lateral fringes (present), heel without tubercles (heel with a conical tubercle), upper eyelid without enlarged tubercles (enlarged upper eyelid tubercles present), and has red iris with black reticulations (iris reddish copper with fine black vermiculation and narrow black vertical streak from pupil across lower half of iris). Pristimantis boucephalus sp. nov. can be distinguished from other species of Pristimantis from Ecuador and Peru that are on average larger but have males of similar size as follows: Pristimantis carvalhoi (Lutz, 1952) from the Amazonian lowlands and P. boucephalus sp. nov. both have males of similar size (SVL 13.4–14.8 mm vs 13.4–14.5 mm [n = 3] in P. boucephalus sp. nov.; Lynch 1980), no dorsolateral folds, and a groin with yellow coloration. However, P. carvalhoi has a long snout (short), tympanic annulus present (absent), prominent dentigerous processes of vomers (absent), vocal slits absent (present), and fingers and toes without lateral fringes (present). Pristimantis imitatrix (Duellman, 1978) from the Amazonian lowlands has slightly smaller males (SVL 13.0–14.0 mm [n = 9] vs 13.4–14.5 mm [n = 3] in P. boucephalus sp. nov.; Duellman 2005) and shares with P. boucephalus sp. nov. the absence of a tympanum, dorsolateral folds, dentigerous processes of vomers, and nuptial pads, and both species have fingers and toes with lateral fringes. However, P. imitatrix lacks vocal slits (present), has the groin mottled black and white (greenish yellow with black blotches) and the iris is bronze (reddish copper). Pristimantis lirellus (Dwyer, 1995) from elevations of 470–1200 m a.s.l. in northern Peru and P. boucephalus sp. nov. both have males with vocal slits, and fingers and toes with lateral fringes, and both species lack a tympanum, dorsolateral folds, and nuptial pads, but P. lirellus has low longitudinal dermal ridges (absent), prominent dentigerous processes of vomers (absent), groin with single pale yellow to orange-yellow spot (greenish yellow with black blotches), and iris bronze to reddish brown (reddish copper). Pristimantis martiae (Lynch, 1974) from the Amazonian lowlands up to 1330 m a.s.l. and P. boucephalus sp. nov. lack dorsolateral folds, a tympanum, and nuptial pads, and both species have males with vocal slits, fingers and toes with lateral fringes. However, P. martiae has larger males (SVL 11.8–16.8 mm vs 13.4–14.5 mm [n = 3], Lynch 1980), small dentigerous processes of vomers (absent), a groin that is dull cream, pale orange or tan with brown to black bars or mottling (greenish yellow with black blotches), and bronze iris (reddish copper). Pristimantis croceoinguinis (Lynch, 1968) from the Amazonian lowland forests in Ecuador and Peru has larger males (13.9–18.2 mm vs 13.4–14.5 mm [n = 3], Lynch 1968) which lack vocal slits (present). Pristimantis llosintuta (Köhler & Lötters, 1999) from 2000–2200 m a.s.l. in Bolivia has a tympanum (absent) and much larger males (SVL 20.3–21.2 mm [n = 7] vs 13.4–14.5 mm [n = 3], Köhler & Lötters 1999). Pristimantis cruciocularis (Lehr, Lundberg, Aguilar & von May, 2006) from the YCNP and its surroundings and P. boucephalus sp. nov. are of similar size (SVL 11.4–15.4 [n = 7] vs 13.4–14.5 mm [n = 3]; Lehr et al. 2006) and share the absence of a tympanic annulus and membrane, but male P. cruciocularis lack vocal slits (present). Furthermore, P. cruciocularis has the iris with a cruciform mark (absent) and a groin that is orange-red (greenish yellow with black blotches). Pristimantis sp. Pui Pui and P. boucephalus sp. nov. share the absence of a tympanic annulus and membrane, but P. sp. Pui Pui is much larger (SVL up to 31.3 mm vs 13.4–14.5 mm [n = 3]), and has a gray groin (greenish yellow with black blotches). Description of the holotype Head broader than body, as long as wide; head length 41.1% of SVL; head width 41.1% of SVL; cranial crests absent; snout short, rounded in dorsal view, rounded in lateral view (Figs 3, 5 A–B); eye-nostril distance 50% of eye diameter; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis short, broadly rounded in lateral view, weakly concave in dorsal view; loreal region slightly concave; lips rounded; upper eyelid each with an enlarged conical tubercle at its center and one enlarged conical tubercle at its posterior end; upper eyelid width 57.1% of IOD; few small tubercles on scapular region (see photos in life Fig. 3 A–B); supratympanic fold short and broad, extending from posterior margin of upper eyelid slightly curved to insertion of arm; tympanic membrane and annulus absent; small low postrictal tubercles present bilaterally. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomers absent; tongue oval, tongue width is about three-fourths of the tongue length, posterior half free. Skin on dorsum and flanks smooth, with low scattered tubercles (denser on flanks than on dorsum), dorsolateral folds absent; occipital and scapular region with sinusoidal ridge; skin on throat, chest, and belly areolate, with scattered low tubercles; discoidal and thoracic folds absent; cloacal sheath short. Outer ulnar surface with minute low tubercles; palmar tubercle partially divided distally; thenar tubercle ovoid; subarticular tubercles well defined, most prominent on base of fingers, round in ventral view, subconical in lateral view; supernumerary tubercles distinct, ovoid, approximately half the size of subarticular tubercles; fingers with lateral fringes; Finger I shorter than Finger II; discs on digits of fingers expanded, round (Fig. 5C). Hind limbs moderately long, slender, tibia length 54.6% of SVL; foot length 48.9% of SVL; upper surfaces of hind limbs smooth, with low scattered tubercles; inner surface of thighs smooth, posterior and ventral surfaces of thighs weakly areolate; heels each with a prominent conical tubercle; outer surface of tarsus with scattered minute low tubercles; inner tarsal fold absent; inner metatarsal tubercle ovoid, two and a half times the size of round outer metatarsal tubercle; subarticular tubercles well defined, round in ventral view, subconical in lateral view; plantar supernumerary tubercles distinct, about half the size of subarticular tubercles; toes with lateral fringes; basal webbing present; discs expanded, round, less expanded than those on fingers; relative length of toes: 1Coloration of the holotype in life (Fig. 3) The dorsal ground coloration is greenish gray with reddish-brown and scattered brown flecks surrounded by black; a reddish-brown blotch mid-dorsally behind the eyes; sinusoidal ridge on occipital and scapular region dark brown; upper lip with two irregularly shaped grayish-brown bars bordered by black on each side of head below eye, bars separated by an irregular pale tan blotch; narrow brown bar laterally in area of canthus rostralis, but not forming a canthal stripe, supratympanic stripe grayish brown, bordered by black; flanks pale greenish and pale reddish brown with scattered dark brown flecks; groin and anterior surfaces of thighs greenish yellow with black blotches and black stripes that extend dorsally to posterior surfaces of thighs; throat, chest, belly and thighs gray with pale reddish and greenish brown mottling and scattered dark gray flecks; palmar and plantar surfaces gray, fingers I and II cream, fingers III and IV gray except for cream discs, toes I and II cream, toes III–V gray with reddish-brown lateral fringes; iris reddish copper with fine black vermiculation and narrow black vertical streak from pupil across lower half of iris. Coloration of the holotype in preservative (Fig. 4) Dorsal coloration grayish tan with scattered brown flecks; brown blotch middorsally behind eyes; sinusoidal ridge on occipital and scapular region dark brown; upper lip with two irregularly shaped grayish-brown bars on each side of head below eye, bars separated by an irregular pale gray blotch; narrow brown bar laterally in area of canthus rostralis, but not forming a canthal stripe, supratympanic stripe grayish brown; flanks tan with scattered brown flecks; groin and anterior surfaces of thighs tan with black blotches and black stripes that extend dorsally to posterior surfaces of thighs; throat, chest, belly and thighs grayish tan with dark gray flecks; palmar and plantar surfaces pale gray, fingers I and II tan, fingers III and IV gray except for tan discs, toes I and II tan, toes III–V gray; iris gray with fine black vermiculation and black narrow vertical streak from pupil across lower half of iris. Variation All paratypes are similar to the holotype regarding morphology (Tables 3–4) and coloration pattern (Fig. 6). They can easily be assigned to the species by having a small size with a disproportionally large head with short snout. The juvenile specimen (MUSM 24474) has two prominent tarsal and two prominent ulnar tubercles on each hind limb. One male specimen (MUSM 24477) has black bars dorsally, laterally and ventrally on the tibia and a brown blotch midventrally on the throat. One specimen (MUSM 24479) is considered a subadult female because of its small size, presence of ovaries, but absence of ovarian eggs. Distribution, natural history, and threat status Pristimantis boucephalus sp. nov. is currently only known from the YCNP. The type locality (Figs 1, 7) is located on a mountain crest at 2950 m a.s.l. covered with primary cloud forest. The holotype was found at night, sitting on a leaf at 1 m above the ground. Syntopic anurans included Pristimantis cf. aniptopalmatus (MUSM 31103–05), P. cf. mendax (MUSM 31101, 31107), and Phrynop, Published as part of Lehr, Edgar, Moravec, Jiří, Cusi, Juan Carlos & Gvoždík, Václav, 2017, A new minute species of Pristimantis (Amphibia: Anura: Craugastoridae) with a large head from the Yanachaga-Chemillén National Park in central Peru, with comments on the phylogenetic diversity of Pristimantis occurring in the Cordillera Yanachaga, pp. 1-22 in European Journal of Taxonomy 325 on pages 7-15, DOI: 10.5852/ejt.2017.325, http://zenodo.org/record/3829345, {"references":["Angulo A., von May R., Icochea J., Chaparro J. C., Hedges B., Lehr E., Lopez D. G., Lundberg M. & Moravec J. 2016. Parque Nacional Yanachaga-Chemillen, Pasco, Peru. Anfibios del bosque de nubes y su zona de amortiguamiento. Field Guides 738: 1 - 6.","Padial J. M., Grant T. & Frost D. R. 2014. Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa 3825: 1 - 132. https: // doi. org / 10.11646 / zootaxa. 3825.1.1","Ron S. R., Guayasamin J. M., Yanez-Munoz M. H., Merino-Viteri A., Ortiz D. A. & Nicolalde D. A. 2017. AmphibiaWebEcuador. Version 2017.0. Museo de Zoologia, Pontificia Universidad Catolica del Ecuador. Available from http: // zoologia. puce. edu. ec / Vertebrados / anfibios [accessed 13 May 2017].","Duellman W. E. & Hedges S. B. 2007. Three new species of Pristimantis (Lissamphibia, Anura) from montane forests of the Cordillera Yanachaga in Central Peru. Phyllomedusa 6: 119 - 135. https: // doi. org / 10.11606 / issn. 2316 - 9079. v 6 i 2 p 119 - 135","Lehr E., Aguilar C., Siu-Ting K. & Jordan J. C. 2007. Three new species of Pristimantis (Anura: Leptodactylidae) from the Cordillera de Huancabamba in northern Peru. Herpetologica 63: 519 - 536. https: // doi. org / 10.1655 / 0018 - 0831 (2007) 63 % 5 B 519: TNSOPA % 5 D 2.0. CO; 2","Lehr E. & Duellman W. E. 2007. A diminutive new species of Pristimantis (Amphibia: Anura: Leptodactylidae) from northern Peru. Salamandra 43: 165 - 171.","Boulenger G. A. 1918. Descriptions of new South-American batrachians. Annals and Magazines of Natural History, Series 9 2: 427 - 433. https: // doi. org / 10.1080 / 00222931808673782","Lynch J. D. & Duellman W. E. 1980. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Miscellaneous Publication of the Museum of Natural History, University of Kansas 69: 1 - 86.","Lehr E. & Coloma L. A. 2008. A minute new Ecuadorian Andean frog (Anura: Strabomantidae, Pristimantis). Herpetologica 64: 354 - 367. https: // doi. org / 10.1655 / 07 - 089.1","Teran-Valdez A. & Guayasamin J. M. 2010. The smallest terrestrial vertebrate of Ecuador: A new frog of the genus Pristimantis (Amphibia: Strabomantidae) from the Cordillera del Condor. Zootaxa 2447: 53 - 68.","Duellman W. E. 1978. Three new species of Eleutherodactylus from Amazonian Peru (Amphibia: Anura: Leptodactylidae). Herpetologica 34: 264 - 270.","Lynch J. D. 1968. Two new frogs of the genus Eleutherodactylus from eastern Ecuador (Amphibia: Leptodactylidae). Journal of Herpetology 2: 129 - 135. https: // doi. org / 10.2307 / 1563112","Kohler J. & Lotters S. 1999. New species of the Eleutherodactylus unistrigatus group (Amphibia: Anura: Leptodactylidae) from montane rain forest of Bolivia. Copeia 1999: 422 - 427. https: // doi. org / 10.2307 / 1447487","Lehr E., Lundberg M., Aguilar C. & von May R. 2006. New species of Eleutherodactylus (Anura: Leptodactylidae) from the eastern Andes of central Peru with comments on central Peruvian Eleutherodactylus. Herpetological Monographs 20: 105 - 128.","IUCN Standards and Petitions Subcommittee. 2016. Guidelines for using the IUCN Red List categories and criteria. Version 12. Prepared by the Standards and Petitions Subcommittee. Available from http: // www. iucnredlist. org / documents / RedListGuidelines. pdf [accessed 27 Apr. 2017]."]}

Opisthencentrus dentipennis (Germar, 1843) Oxygonia dentipennis Germar, 1843: 3. Type locality. Brazil. Opisthencentrus dentipennis: Horn 1893: 196. Type material. Lectotype (designated here), ♀ deposited in MFNB, labelled: “ 42560 ” [printed] // “ Oxygonia / dentipennis / Germ. / Brasil ” [greenish label, handwritten] // “ Lectotype / Oxygonia / dentipennis Germar, 1843 / design. Jiří Moravec 2014 ” [red, printed] // “ Opisthencentrus / dentipennis Germar, 1843 / det. Jiří Moravec 2014 ”. Other material examined. Following five specimens in SDEI: 1 ♂, 1 ♀: “Angra dos Reis / Rio de Janeiro / XII. 1913 ” [head of a male glued on the pronotum of the female!]. 1 ♀: “Coll. Weber / Brasilia ” // “Coll W. Horn / DEI Eberswalde” // “ dentipennis / Germ”. 1 ♂: “ Oxygonia / dentipen- / nis / Germ” // “ Brasilia / Ex Cab. Thomson” // “Coll. V. de Poll” // “Coll W. Horn / DEI Eberswalde”. 1 ♂: “ Brasilia / Coll. Baden Ruge” // “ Opisthencentrus / dentipennis / Germ. / Dr. W. Horn det. 1936 ”. Redescription. Body (Figs 1–3) small, 7.90 –9.00 (LT 8.90) mm long, 2.40–2.65 (LT 2.60) mm wide, mostly (also in LT) metallic black-copper, sometimes with faint, rarely more intense greenish or blue lustre, elytral surface of rather matt appearance. Head (Figs 4–5) with large eyes, but narrower than body, 2.00– 2.30 mm wide, black-copper with faint greenish or blue lustre; all head portions glabrous. Frons moderately sloped towards clypeus, cupreous with strong green lustre, almost smooth, with several fine longitudinal striae in middle; supraantennal plates irregularly triangular, cupreous with strong green lustre. Vertex cupreous with greenish lustre (also in LT), green coloration in one male prevailing; surface irregularly finely vermicular-rugulose in middle, lateral areas with coarser and parallel striae which are partly fragmented on anterior area of juxtaorbital plates; posteriad-divergent parallel rugae passing onto temples; occipital area irregularly rugulose to finely tuberculate. Genae glabrous, finely parallel-striate. Labrum in male (Figs 6–7) ochre-testaceous, transverse, sinuous anterior margin only moderately prolonged anteriad towards blunt, only indicated median tooth; female labrum (Figs 8–9) brownish-darkened, sinuous anterior margin more distinctly prolonged anteriad, forming wide, blunt median tooth. Clypeus comparatively large, bright iridescent green or with reddish-cupreous areas, surface glabrous, finely irregularly wrinkled. Palpi (Figs 4–5) in male yellow, terminal palpomeres with black-brown darkened apical area, normally shaped (moderately and gradually dilated), palpi in female usually darker, ochre to testaceous, whole terminal palpomeres blackened. Mandibles (Figs 4–5) almost symmetrical, lateral margins arcuate, each mandible with four teeth (and basal molar), the inner teeth become gradually smaller towards the basal molar; coloration in male reddish-brown with wide, pale ochre basolateral areas; in female much darker, almost black-brown except for testaceous basolateral areas. Antennae filiform, rather slim and short, in male barely reaching elytral third, in female somewhat shorter; scape and pedicel and mostly also antennomeres 3-4 in male testaceous, occasionally partly black-brown darkened and with indistinct metallic green lustre particularly on scape, generally darker in female; remaining antennomeres brownish, gradually smoky-blackened. Thorax. Pronotum (Figs 10–12) black-copper with faint green lustre (also in LT) rarely with stronger green or blue-green lustre, oblong in shape, slightly longer than wide, 1.50–1.65 mm long, 1.30–1.50 mm wide; anterior lobe almost as wide as the posterior one, surface rather coarsely but sparsely wavy-rugose; lateral margins of disc moderately convex to almost parallel, mostly slightly dilated posteriad, formed by distinct notopleural sutures which are clearly visible from above; surface of disc rather densely but distinctly irregularly transversely striaterugulose; median line thin but usually distinct; posterior lobe with distinct dorsolateral bulges, surface very irregularly rugulose; lateral and ventral thoracic sterna glabrous, metallic black with blue, green and gold-bronze reflections; female mesepisternal coupling sulci in form of longitudinal furrow impressed in its dorsal area to deep, rather wide pit with indistinctly defined margins. Elytra (Figs 15–18) elongate, humeri rounded to subquadrate, lateral margins subparallel, anteapical angles arcuate, then sinuous towards conspicuously long, slightly bent or straight lateral-median spine (the length of the spine varies in both sexes), then truncate in middle towards shorter but distinct sutural spine; elytral surface almost even, lacking deep impressions, matt, densely irregularly punctate throughout; more isolated punctures are only on anterior area, then punctures become irregularly anastomosing with bumpy to tuberculate intervals (Fig. 19); elytral maculation consisting of three distinct ivory-white maculae: mostly rounded humeral macula clearly visible from above, rounded sublateral-median macula, and anteapical macula which is prolonged towards the lateralmedian spine, but not covering the truncate elytral apex; surface glabrous except for usual, a few hairlike sensory setae; elytral coloration black-copper (also in LT) to cupreous, sometimes with faint, rarely more intense blue or cyaneous tinge. Legs with metallic-green coxae, pro- and mesocoxae setose; lateral margins of metacoxae widely fringed with dense setae; trochanters testaceous with apical seta on pro- and mesotrochanters; femora brownish in male, blackbrown in female, in both sexes with yellow to testaceous basal area which in male extends to basal half, and limited testaceous subapical area; femoral apices with two distinct spines (Fig. 13); whitish semierect setae on femora rather sparse, more dense on profemora, sparser on metafemora; tibiae in male brownish with testaceous basal half, much darker in female, covered with sparse, short but rather stiff semierect setae, pro- and mesotibiae with usual denser greyish setose pad on their apical third; tarsi thin and long, brown in male, almost black in female, often with faint metallic lustre; protarsomeres sexually dimorphic: first three protarsomeres in male moderately, but distinctly dilated, with usual whitish pad. Abdomen metallic-black with blue reflections, ventrites glabrous, except for usual hairlike sensory setae at their margins. Aedeagus (Fig. 20) elongate, 2.60 mm long, 0.40 mm wide, apical half conically attenuated towards elongate, moderately ventrally bent, subclavate-rounded apex; internal sac (Fig. 21) with conspicuous, longitudinal, distinctly serrate dorsal sclerite; other sclerites comprise thick spike-like arciform piece, thin, longitudinal, doubled central piece, distinct ventral spur with thin, rather short, acute projection and stiffening rib, and three basal sclerites: dorsally placed small ellipsoidal piece, and elongate, almost transverse sclerite covered with a horseshoeshaped piece. Variability. The original diagnosis given by Germar (1843) was: “ O. supra obscure aenea, subtus viridiaurea, nitida, capite thoraceque transversim strigosis, elytris punctatis, apice truncatis exctus spinosis, maculis tribus albis, pedibus fuscis, femoribus basi testaceis”, thus describing the dorsal surface as obscure copper. In the original description Germar also described the dorsal coloration of the elytra as “ densius et profundis punctata, aeneo-nigra, opaca ” (= densely and deeply punctate, black-copper, dull), which well corresponds with the lectotype in MFNB (Figs 1 and 16) and other specimens examined here; all have the same matt elytral appearance due to the irregularly and densely arranged punctures with bumpy intervals (fig 19). Only one female (SDEI) has the elytra with deep blue tinge (Fig. 17), and it was probably the same female illustrated in colour aquarelle by Horn (1910: T. 13, fig. 6) —the illustration was adopted by Erwin and Pearson (2008: Pl. 26). The male published by Huber (2012) has bluish tinge on its elytra which are of the same matt appearance as in all other specimens examined (R. Huber, pers. com). Biology and distribution. Opisthencentrus dentipennis is a very rare tiger beetle, known only from seven specimens, all collected in a comparatively large area of the Atlantic Rainforest in Brazil. (Horn 1926) also mentioned the locality Alto da Serra, Sao Paulo, but no specimens labelled as such were found in the collection of Walther Horn (now in SDEI), nor in other collections. Reserva Biológica do Alto da Serra de Paranapiacaba is located at Paranapiacaba district (municipality of Santo André, São Paulo Metropolitan Region), near the Industrial Center of Cubatão, which is one of the most polluted industrial area in the world; interestingly, the preserve still includes remnants of the Atlantic rainforest and its associated biodiversity. Apart from the male lectotype in MFNB, only five historical specimens have been found in SDEI (two males and three females), two of them from the locality Angra dos Reis which lies in the southern coastal part of the Brazilian state of Rio de Janeiro. The hilly area of the Atlantic Rainforest also comprises several forested islands. The most recent record for this species was published by Huber (2012) with a colour imagine of the habitus of a single male with the following label data: “ Brazil: Rio de Janeiro / Rio Macacú” // On RJ 116, Km 65 / 22 ° 22.786 'S / 42 ° 33.945 'W, elev. 933m / 17 -iii- 1996 ”, collected during an intensive survey by the University of Minnesota team of Holzenthal, Rochetti and Oliveir; the specimen is deposited in RLHC. The Guapi-Macacú River Basin is located within the Brazilian state of Rio de Janeiro, northeast of Guanabara Bay and the city of Rio de Janeiro. The Cachoeiras de Macacu is a lowland area of Atlantic Rainforest, close to the Serra dos Órgãos Mountain Range. It was severely deforested in most of its territory, but there are still some sites of well preserved forest and its valuable biodiversity(André Sylva Roza, pers. com). Little is known of the biology of this very rare tiger beetle. Huber (2012) mentioned that the biotope of the male from Macacú was described by the collectors as: “forested slopes at upper reaches of Macacú basin; wet, rich with epiphytes, steep slopes”. Erwin and Pearson (2008) mentioned that the only recently collected specimen was captured in a flight interruption trap in forested vegetation along a mountain stream, referring to “Huber, pers. com.”, so, the record was probably the same as published by Huber (2012). These authors (Erwin and Pearson 2008: Pl. 26) published a photograph of a “forested coastal mountain stream landscape in which members of the genus may dwell (Salto Morato Nature Reserve, Mata Atlântica, Paraná, Brazil, ca 100 m. altitude)”. While the Salto Morato Nature Reserve in Guaraquecaba, Paraná, is a well preserved area of the Atlantic Rainforest, its phytogeographic profile may be different from the other localities where Opisthencentrus dentipennis has been recorded; thus considering that Erwin and Pearson (2008) did not provide evidence of a specimen collected in this locality, there is no proof of the occurrence of this rare tiger beetle in the Paraná area. As discussed by Roza and Mermudes (2015), the Atlantic Rainforest is one of the most threatened biomes of the world, with only 11−12 % of its original cover. Remarks. Opisthencentrus dentipennis was described by Germar (1843) probably from only one female (MFNB), but as no number of specimens was given in the original description, the female is here designated as a lectotype to assure stability of the taxon.

Pentacomia (Mesochila) procera (Chaudoir, 1860) Odontochila procera Chaudoir, 1860: 324, 325. Type locality. Brazil: Petrópolis (state of Rio de Janeiro). Odontochila Chaudoiri Dokhtouroff, 1887: 157, 158 (synonymy by HORN 1892). Type locality. Brazil: Petrópolis (state of Rio de Janeiro). Pentacomia (Mesochila) procera: Rivalier 1969: 224 (222, fig. 15 pr, fig. 16 pr, 224, fig. 18 pr.). Misapplication. Non Odontochila procera sensu Horn (1929: 157 fig. 20, 158), which is Pentacomia (Mesochila) proceroides sp. nov. Type material of Odontochila procera Chaudoir. Lectotype (designated here) ♂ in MNHN, labelled: “ procera / Chaud. / Brésil / Petropol. / 53. Sahlb. j.” [brownish-tarnished, with black frame, handwritten] // “Muséum Paris / Coll. Chaudoir 1874 ” [greenish with black border, printed] // “ 1521 / Rivalier” [handwritten, referring to aedeagus mounted separately by Rivalier] // “ Lectotype / Odontochila / procera Chaudoir, 1860 / design. Jiří Moravec 2014 ” [red label, printed] // “ Pentacomia (Mesochila) / procera (Chaudoir, 1860) / det. Jiří Moravec 2014 ” [printed]. Paralectotype. 1 ♀ in MNHN: “Muséum Paris / Coll. Chaudoir 1874 ” [greenish with black border, printed] // “ AEruginosa / Reiche” (sic!) [additionally attached handwritten label]. Type material of the synonymous Odontochila chaudoiri Dokhtouroff. Holotype (by monotypy), labelled: “ Chaudoiri. Sahlb.” [ochre-tarnished, handwritten] / “ Type / Dokhthurow” [printed] // “ Holotypus ” [red, printed] // “Coll. W. Horn / DEI Eberswalde” [printed] // “(Chaudoiri / Dokht.)” [green with black frame, handwritten] // “ Pentacomia (Mesochila) / procera chaudoiri Dokthouroff / Type (DEI Eberswalde) / borrowed by D. L. Pearson / 23 Oct. 1996 (Drawer # 56)” [printed] // “ Pentacomia (Mesochila) / procera (Chaudoir, 1860) / det. Jiří Moravec 2014 ” [printed]. Other material examined. 1 ♂ in MNHN: “Rio Jan.” // “Ex Musaeo / H. W. Bates / 1892 ” // “Muséum Paris, 1952 / Coll. R. Oberthür”. 1 ♂ in MNHN: “Muséum Paris / Coll. Chaudoir, 1874 ” // “ 1522 / Rivalier”. 1 ♂ in MNHN: “ Brésil ” // “Muséum Paris, 1952 / Coll. R. Oberthür” // “Ex Musaeo / Mniszech”. 1 ♀ in MNHN: “ Brasil ” // “Odont. / procera ” // “Muséum Paris”. 1 ♂ in BMNH: “Fry / Rio Jan.” // “ 9353 ” // “Fry Coll. / 1905 – 100 ”. 2 ♀♀ in BMNH [standing as “ conformis ”]: “Petropolis” // F. Bates Coll. / 1911 – 248 ” // “ procera Chd. / v. W. Horn”. 1 ♂ in BMNH: “Fry / Rio Jan.[eiro]” // Fry Coll. / 1905 – 100 ”. 2 ♀♀ in BMNH: “ Brasilia ” // “ Odontocheila / Conformis / Dejean”. 1 ♂ in BMNH: “Petropolis”. 1 ♀ in BMNH: “St. Paul[o]”. 1 ♀ in BMNH: “ 485 ” // “Fry / Rio Jan.”. 1 ♀ in MFNB: “ Brasilia ”. 1 ♀ in MFNB: “Rio Janeiro”. 1 ♀ in MFNB: “ 3626 ” // “attenuata [sic!] / N. / Bona S....[illegible] / “ Chaudoiri / Doct. / det. Horn”. 1 ♂ in SDEI: “ Brasilien / Sao Paulo”. 1 ♂ in SDEI: “ Brasil ”. 1 ♂ in SDEI: “Speyer / Brasilia ”. 1 ♀ in RLHC: “ Brazil, Sao Paulo / Cubatão River / 4.X. 1960 / V.N. Alin / forest”. 1 ♂ in CEIOC: “ Brazil / Petrópolis, Itambraty / J. Zikan, 27.XI. 1908 ”. 1 ♂ in DZRJ: “ Brasil, Rio de Janeiro / Teresópolis, Parque Nacional / da Serra dos Órgãos / PVE, Ponto 6 A / 22 ° 28 '11,5"S, 43 °00'06,0"W, 877m / XII. 2014 Ricardo Monteiro Col. (Malaise Trap)”. 1 ♂ in MZH: “Petrop.” // "F. Sahlb." // " Odontochila / chaudoiri / Sahlb.". // " GAC 20273 / Brazil, Rio de Janeiro / Petrópolis / 22.49 S, 43.18 W / II. 1850 / Ferdinand Sahlberg leg." [syntopic with HT of O. chaudoiri Dokhtouroff]. Differential diagnosis. P. (M.) procera is distinguished from the externally similar P. (M.) conformis by its generally larger and slightly more elongated body, particularly so in female, pronotum with almost parallel lateral margins, elytra (Figs 58–62) with more parallel margins and finer punctation, white lateromedian macula mostly longitudinal or rounded and mostly more distant from the epipleuron, and immediately by the shape of its labrum, which is in male of P. (M.) procera (Figs 44-48) with subtruncate to semicircular anterior margin of the median lobe between rounded but prominent anterolateral teeth (in contrast to mostly truncate anterior lobe between flat, right-angled anterolateral teeth in the male labrum of P. (M.) conformis (Figs 9-12)); female labrum of P. (M.) procera (Figs 49–51) also significantly differs in having the prominent anterolateral teeth (of the same shape as in male), while they are blunt and flattened in female of P. (M.) conformis (Figs 8, 13). Males can be immediately distinguished by the shape of the aedeagi: in P. (M.) procera with distinctly hooked apex (Figs 67 –70, 72– 75), while in P. (M.) conformis the apex is always rounded, sometimes dorsally emarginated; a significant difference is also in the structure of the internal sac in the aedeagus of P. (M.) procera (Figs 74–75) from that in P. (M.) conformis (Figs 31-36). P. (M.) proceroides sp. nov., also has its labrum in both sexes with prominent anterolateral teeth, and a similar structure of the internal sac as in P. (M.) procera, but the median lobe of the male labrum in the new species is always truncate, and the shape of the aedeagus distinctly differs (see under the new species below). Redescription. Body (Figs. 37–42) medium-sized to large, length 11.7–13.6 (LT 11.7) mm (exceptionally only 10.2 mm long), width 3.10–3.70 (LT 3.50) mm, females mostly larger than males; body generally almost uniform, notably elongate, particularly so in female, similarly coloured as in P. (M.) conformis, but mostly with brighter and prevailing iridescent-green lustre, and darker appendages. Head (Figs 43, 85) conspicuously large with wide eyes, but slightly narrower than body, width 2.80–3.50 mm; shape, surface sculpture and coloration of frons, vertex, genae and clypeus as in P. (M.) conformis. Labrum primarily 4 -setose, usually one or two lateral setae absent (either broken or rarely not developed), sexually dimorphic in shape; male labrum (Figs 44–48) ochre-yellow to brownish-testaceous with indistinctly black-brown darkened basal, sometimes with brown areas or patches, rather long, length 1.05–1.10 mm, width 1.30–1.45 mm, lateral margins moderately arcuate with indistinctly indicated or mostly entirely effaced basolateral teeth; anterolateral teeth always well pronounced, rounded, but prominent, anteriad-prolonged, but not surpassing the anterior margin of medial lobe; the rather variably shaped median lobe has right-angled or rounded lateral margins (rarely indicating blunt anterolateral teeth), its anterior margin subtruncate, rarely indicating indistinct median tooth (exceptionally the median tooth developed as in the aberrant male (Fig. 66)), or the anterior margin is moderately to more distinctly prolonged anteriad, of almost semicircular shape (as in LT, Fig. 45); female labrum (Figs 49–51) almost as long as wide, length 1.60–1.80 mm, width 1.60–1.75 mm, similarly shaped as in male, but median lobe with projecting, mostly subacute median tooth; coloration variable as in male, usually brownishtestaceous, rarely almost entirely blackened. Mandibles (Figs 43, 65, closed ones Figs. 47, 49), shape and coloration as in P. (M.) conformis, but each mandible notably wider. Palpi (Figs 43, 47, 51, 65), shape and coloration as in P. (M.) conformis, but the terminal palpomeres mostly almost entirely black, exceptionally pale testaceous (Fig. 65). Antennae as in P. (M.) conformis, but generally much darker. Thorax. Pronotum (Figs 52–57), glabrous, iridescent reddish-cupreous on median area, with iridescent-green lateral areas, the cupreous median area rarely prevailing, slightly or more distinctly longer than wide, length 2.10– 2.50 mm, width 1.80–2.10 mm, sulci well pronounced; anterior lobe as wide as posterior lobe, or very slightly wider, and almost as wide as the disc, its anterior margin usually almost transverse, or only moderately prolonged anteriad; surface of anterior lobe coarsely and very irregularly rugulose; disc with mostly notably parallel to subparallel, or only slightly convex lateral margins (including margins of proepisterna), notopleural sutures rather distinct, clearly visible from above, almost subparallel with the proepisternal lateral margins, or even narrowed in middle; medial line distinct; discal surface sculpture as in P. (M.) conformis, but notably coarser; posterior lobe basically as in P. (M.) conformis, but the posterior margin less emarginated; all lateral and ventral thoracic sterna generally as in P. (M.) conformis. Elytra (Figs 58–64) elongate, particularly markedly elongate in female, length 6.90–8.40 mm (exceptionally 6.20mm), in both sexes with rounded humeri; elytral margins as in P. (M.) conformis, but more subparallel in male and markedly parallel in female; shape of anteapical angles and apices as in P. (M.) conformis; elytral dorsal surface even, except for rather distinct humeral and apical impressions, while discal impression and basodiscal convexity are only indicated, the even surface is particularly notable in female; elytral coloration as in P. (M.) conformis, but the cupreous or green lustre generally much brighter (depending on angle of illumination, the chatoyant reddish-cupreous coloration may change to iridescent green); elytral surface glabrous except for the usual, a few and often very indistinct hairlike sensory setae as in P. (M.) conformis and other species; whole elytral surface rather densely and regularly punctate, pattern of the punctation similar as in P. (M.) conformis, but generally finer and more isolated (the sculpture optically changeable depending on angle of illumination); particularly much finer and almost uniform punctures on female elytra; whitish elytral maculation consisting of three maculae as in P. (M.) conformis, but the sublateral-median macula rounded, or irregularly rhombiform, or slightly longitudinal- prolonged (never mesad-prolonged). Legs as in P. (M.) conformis, but mostly much darker. Abdomen generally as in P. (M.) conformis. Aedeagus (Figs 67–73), elongate, ventral side almost straight, while dorsal outline dilated in middle, apical half attenuated towards distinctly hooked apex; internal sac (Figs 74–75) containing ventral spur of characteristic, wing-like dilated base and short, thin projection; other sclerites are: small basodorsal spike, dorsal ovaliform piece (its shape better visible in right lateral view (Fig. 75), combined with longitudinal dorsal tooth (never distinctly protruding from dorsoapical orifice), and central, partly membranous piece with sclerotized hook, combined ventrally with barely defined longitudinal pieces. Note. The aedeagus (Fig. 67) of the lectotype of P. (M.) procera (MNHN) shows only its outer membrane, because the internal sac was extracted by Rivalier and mounted together with the empty aedeagus between two glasses, numbered as “ 1521 / Rivalier”. The internal sac has been destroyed by such wrong mounting treatment using a brown glue, which in time dried out and destroyed the shape of the internal sac (as in many type specimens of other tiger beetles –see Moravec 2010, 2014). Variability. The male (Fig. 39) from Rio de Janeiro (BMNH) has anomalously wide pronotum (resembling that of P. (M.) conformis), but its aedeagus (Fig. 68) has the distinctly hooked apex as in other males of P. (M.) procera. The aberrant male (Fig. 40) from Sao Paulo (SDEI) notably differs in its much smaller size and much paler legs, labrum with more pronounced median tooth (Fig. 66), the lateromedian macula closer to the epipleuron (Fig 61), paler palpi (Fig. 65) and somewhat less distinctly hooked aedeagus (Fig. 72). The differences are considered here a variability of the probably juvenile male, rather than to consider this aberrant male to be another undescribed species. Distribution, ecology and biology. The type locality of P. (M.) procera is situated in the large area near the municipality of Petrópolis in the Brazilian state of Rio de Janeiro, 70 km from the state capital. The Petrópolis area covers the valley of the Quitandinha and Piabanha rivers with forested hills of the Serra dos Órgãos National Park near Teresópolis with still well preserved biotopes of the Atlantic Rainforest. The “ Mata Atlantica” included also the forested area of Sao Paulo. The specimen (SDEI) labelled “Sao Paulo” comes probably from the area of today’s Reserva Biológica do Alto da Serra de Paranapiacaba, located at Paranapiacaba district (municipality of Santo André, São Paulo Metropolitan Region). The female (RLHC) comes from Rio Cubatão in the same metropolitan region. P. (M.) procera is obviously sympatric with P. (M.) conformis in the state of Rio de Janeiro, as two specimens in MNHN (ex H. W. Bates) bear identical handwritten labels “Rio Jan.”, but there is no proof of also syntopic occurrence of these two species in the large area. In addition, several other species of Mesochila occur in the same area, as already mentioned by Horn (1910, 1926) and Zikán (1929). Rivalier (1969), obviously in error, mentioned the occurrence of P. (M.) procera also in the states of Bahia and Matto Grosso, but no specimen with such labels has been found in MNHN or in other collections within the present revision. Little is known of the habitat and behaviour of adults and larvae. The behaviour of adults described by Erwin & Pearson (2008) are without a reference to any exact locality and experience, but they refer to Zikán (1929). Zikán (1929) described and illustrated larval tunnels and development of adults from their immature stages during rearing of P. (M.) procera, but in fact mostly of P. (M.) proceroides sp. nov. (see under this new species below). Remarks. Chaudoir (1860) based the original description of this species on two specimens of both sexes: one male caught by Sahlberg in Petrópolis (the lectotype designated here to assure stability of this taxon), and one female from “interior Brésil ” received from a collection Laferté. Chaudoir (1860) described the coloration of the labrum of the female syntype as “ piceo ”, which means pitchy-black. Unfortunately, Chaudoir never labelled type specimens of his taxa by type labels. The only female that corresponds by its black labrum with the female syntype, is that from the Chaudoir collection (MNHN), and is therefore considered here to be the paralectotype. It bears the additional label “ aeruginosa / Reiche”, evidently attached to the female subsequently, probably by Reiche, and it has no impact to the individuality of the syntype. It is noteworthy that the unavailable name Cicindela aeruginosa was listed without any description by Dejean (1831: 208) under Cicindela smaragdula Dejean, 1825 with only a note that he received the specimen under the name “ AEruginosa ” from Schönherr; the name is therefore a nomen nudum. Examination of the holotype (Figs 42, 49, 57, 64) of Odontochila chaudoiri Dokhtouroff, 1887 has confirmed the synonymy with P. (M.) procera by Horn (1892). Fleutiaux (1892), evidently without examination of the holotype in SDEI, listed inappropriately O. chaudoiri as a synonym of Odontochila virens Audouin & Brullé, 1839, which is in fact a junior synonym of Pentacomia (Mesochila) smaragdula (Dejean, 1825), as stated by Horn (1892) and confirmed within the present revision.

Moravec, Jiří (2016): Taxonomic and nomenclatorial revision within the Neotropical genera of the subtribe Odontocheilina W. Horn in a new sense— 15. The genus Opisthencentrus W. Horn (Coleoptera: Cicindelidae). Zootaxa 4097 (3): 332-340, DOI: http://doi.org/10.11646/zootaxa.4097.3.2

Moravec, Jiří (2016): Taxonomic and nomenclatorial revision within the Neotropical genera of the subtribe Odontocheilina W. Horn in a new sense— 16. Pentacomia (Mesochila) procera (Chaudoir), P. (M.) conformis (Dejean), and P. (M.) proceroides sp. nov. (Coleoptera: Cicindelidae). Zootaxa 4127 (2): 276-300, DOI: http://doi.org/10.11646/zootaxa.4127.2.3

Pentacomia (Mesochila) proceroides sp. nov. Type locality. Brazil: “Fazenda Jerusalem” near Alegre, state of Espirito Santo. Misinterpretation. Odontochila procera sensu Horn (1929: 157 fig. 20, 158), and sensu Zikán (1929) partim. Non Odontochila procera Chaudoir, 1860 (= Pentacomia (Mesochila) procera (Chaudoir, 1860). Type material. Holotype ♂ in SDEI, labelled: “ Brazil / Espirito Santo / Faz. Jerusalem / 8.- 11.- 1912 ” / J. F. Zikan” [printed, the date handwritten] // “♂” [handwritten] // “Coll. W. Horn / DEI Eberswalde” [printed]. Allotype. 1 ♀ in SDEI with same labels as holotype except for: “ 3.- 11.- 1911 ” // “♀” //. Paratypes. 2 ♂♂ in SDEI, with same labels except for: “ 4.- 11.- 1912 ”. 1 ♂ in SDEI: “Zikan[leg.], Castello / bei Palmital / Espir. Sant.” [handwritten] // “ 11 / 5 / 1905 ” [handwritten] // “Coll. W. Horn / DEI Eberswalde” [printed]. 2 ♂♂ in SDEI: “ Brazil / Villa de Alegre, Espi / rito Santo / J. F. Zikán” [with black border, handwritten/printed] // “ 30./ 11. / 1911 ”. 1 ♀ in MNHN: “ Brazil / Espirito Santo / Faz Jerusalem / J. F. Zikan” [printed] // “Muséum Paris, Coll. Chaudoir, 1874 ” [green, printed]. All type specimens labelled: “ Holotype (or Allotype or Paratype respectively) / Pentacomia (Mesochila) / proceroides sp. nov. / det. Jiří Moravec 2016 ” [red, printed]. Following paratypes examined only from photos of the habitus, labrum and aedeagi, taken and sent by André Silva Roza (Federal University, Rio de Janeiro): 4 ♂♂ in CEIOC with the same locality labels and collector as in the holotype, except for: “ 30.II.1911, 3.XI.1911, 3.XII.1911, 11.XI. 1914 ”. 1 ♂, 1 ♀ in CEIOC: “ Brazil / Palmital / Espirito Santo / Faz. Castello / J. Zikan 11.I. 1905.” Other material examined. 1 ♂ in CMNH: “Minas (Brazil)” // “J. F. Zikán” // “Field Mus. / (F. Psota Coll.)” / / “Robert D. Ward / Collection” // “ Pentacomia / (Mesochila) / conformis Dejean / Det. R. Ward 1977 ” // “ Mesochila / proceroides Moravec, 2016 / det. Jiří Moravec 2016 ”. Differential diagnosis. P. (M.) proceroides sp. nov. superficially resembles P. (M.) procera because of its similar, but generally much smaller body, similar shape of the pronotum and elongate, almost parallel-side elytra with fine elytral punctation; its labrum (Figs 84–90) in both sexes also has the same rounded and prominent anterolateral teeth, but the median lobe of the male labrum (Figs 84–88) is truncate to subtruncate, never distinctly arcuate or semicircular as in most males of P. (M.) procera; males of this new species are reliably distinguished from those of P. (M.) procera by the very different shape of their aedeagi (Figs 98–103) which have their apical portion directed ventrally, and with rounded, sometimes dorsally emarginated apex (never distinctly hooked as in the almost straight aedeagi of P. (M.) procera). P. (M.) conformis clearly differs in having its labrum in both sexes with flattened anterolateral teeth (never prolonged anteriad), and different shape of its aedeagus with characteristic, unique structure of the internal sac (see under that species above). Description. Body (Fig. 76–79) medium-sized to large, length 10.2–12.90 (HT 10.3, AT 12.8) mm, width 2.90–3.50 (HT 3.00, AT 3.50) mm, females mostly larger than males; body shape similar to that in P. (M.) procera, notably elongate, particularly so in female, but iridescent-green coloration mostly prevailing. Head (Figs 80–82) conspicuously large with wide eyes, as wide as the body, width 2.90–3.45 mm; shape, surface sculpture and coloration of frons, vertex, genae and clypeus as in P. (M.) procera, but with prevailing iridescent-green lustre. Labrum primarily 4 -setose, usually one or two lateral setae absent (either broken or rarely not developed), shape sexually dimorphic; male labrum (Figs 84–88) ochre-yellow to brownish-testaceous with indistinctly blackbrown darkened basal or also basolateral areas, sometimes with brown patches, length 0.85 –1.00 mm, width 1.30– 1.45 mm, shape basically similar to that in P. (M.) procera, in having prominent anteriad-prolonged anterolateral teeth, but the teeth are usually in the same level as the median lobe, or slightly surpassing its anterior margin which is mostly truncate or subtruncate, very rarely indistinctly anteriad-prolonged; female labrum (Figs 90–91) almost as long as wide, in AT 1.55 mm long, 1.50 mm wide, similar to that in P. (M.) procera, but in examined specimens never blackened. Mandibles ((Figs 82–83), closed ones (Figs 80 –81, 89), shape and coloration as in P. (M.) procera, but the inner teeth gradually smaller towards the basal molar (the fourth tooth in examined specimens well developed). Palpi (Figs 80 –83, 90), shape as in P. (M.) procera, but coloration darker, often also penultimate palpomeres of maxillary palpi black (apart from the black terminal palpomeres). Antennae as in P. (M.) procera. Thorax. Pronotum (Figs 91–92), glabrous, iridescent-green with mostly indistinct reddish-cupreous median area, notably longer than wide, length 2.05–2.45 mm (HT, AT), width 1.65 –2.00 mm (HT, AT), sulci well pronounced; anterior lobe slightly wide that the posterior, almost as wide as the disc, with anterior margin only moderately prolonged anteriad; surface very irregularly rugulose; disc with parallel to subparallel, or only slightly convex lateral margins (including those of clearly visible proepisterna), notopleural sutures rather distinct, clearly visible from above and distant from the lateral margins of the dorsally visible proepisterna and slightly narrowed in middle; medial line distinct; discal surface finely but striate-rugulose, transverse-striate in middle, striae converging towards the median line, sublateral areas covered with finer and irregularly wavy rugae; lateral juxtanotopleural areas covered with shorter and shallower rugae which do not surpass notopleural sutures; posterior lobe notably high, surface very finely and irregularly rugulose, large dorsolateral bulges smooth and shiny; all lateral and ventral thoracic sterna generally as in P. (M.) conformis and P. (M.) procera, but with stronger green lustre; female mesepisternal coupling sulci indistinct, in form of the longitudinal furrow which is only slightly deeper than in male, somewhat sinuous and with indistinct median impression (lacking any deep pit). Elytra (Figs 93-97) in both sexes notably elongate, length 6.30 –8.00 mm, in both sexes with rounded humeri; elytral margins as in P. (M.) procera, but even more subparallel in both sexes; shape of anteapical angles and apices as in P. (M.) conformis and P. (M.) procera; elytral dorsal surface as in P. (M.) procera almost even; elytral coloration as in P. (M.) procera but prevailingly iridescent green, rarely reddish-cupreous, but usually changing to green depending on angle of illumination; whole elytral surface densely and almost regularly punctate, the pattern of the punctation even finer than in P. (M.) procera, notably finer and almost uniform punctures particularly on female elytra; elytral surface glabrous except for the usual, a few and often very indistinct hairlike sensory setae (as in other species); whitish elytral maculation consisting in male of three maculae as in P. (M.) conformis and P. (M.) procera, but the humeral macula is absent in female, and sublateral-median macula is generally smaller, particularly so in female, mostly rounded, or only very indistinctly longitudinal, or mesad prolonged. Legs as in P. (M.) procera, but trochanters in female much darker, particularly metatrochanters black, and femora with more intense mahogany lustre. Abdomen generally as in P. (M.) conformis and P. (M.) procera, but ventrites with prevailing deep blue coloration. Aedeagus (Figs 98–103, anomalous one Fig. 104) widest in middle, with notably ventrally bent apical half which is conically (or rarely more abruptly) attenuated towards rounded apex which is only indistinctly dorsally emarginated; internal sac (Figs 105–110) structured as in P. (M.) procera, with similar ventral spur with wing-like dilated base and rather short filiform projection and other sclerites, of which the dorsal tooth partly penetrates the dorsal orifice, but the upper-central, partly membranous large piece is wide, compacter and with more sclerotized hooks. Variability. Despite the almost constant external characters and consistent structure of the internal sac, the aedeagi somewhat vary in the shape of the ventrally bent apical half. Nevertheless, the rounded apex is almost consistent in shape with only more or less distinct dorsal emargination. The differences are obviously caused by the state of the membrane of the dorsoapical orifice, which is usually collapsed and may change the shape. Moreover, the aedeagi, cleared and mounted in order to show their internal sacs, become usually widened and straightened by the clearing procedure. The aedeagus (Fig. 90) was inappropriately treated (probably using acetic acid) when mounted by Horn in his collection (SDEI). Teratology. The anomalously developed aedeagus (Fig. 104) is with forked apical half, thus showing an additional apical portion of the penis, while the primary portion of the penis, including its apex, is normally developed. Distribution, ecology and biology. The type locality of this new species, Fazenda Jerusalem, one of the farms of the collector J. F. Zikán, is situated near the municipal area of Alegre in the southeastern Brazilian state Espirito Santo. The paratypes (SDEI and CEIOC), labelled “Villa Alegre” and “Castello del Palmital”, also come from the same area. The only male (CMNH), labelled “Minas”, comes from the state of Minas Gerais, probably from an area adjoining the states Espirito Santo and Rio de Janeiro. The biotopes of the Atlantic Rainforest in the areas of the type locality are now only partially preserved (André Silva Roza, pers. com.) The behaviour of adults (under “ Odontochila procera ”) was partially described by Zikán (1929) during rearing in captivity, mostly female ovipositing and developing of larva and pupa to metamorphosis; he also illustrated larval tunnels. As Zikán (1929) mentioned an occurrence of “ Odontochila ” procera together with “ Odontochila ” cyaneomarginata (= Pentacomia (Mesochila) cyaneomarginata (W. Horn, 1900)), also in mountains near Palmital and Fazenda Jerusalem near Alegre (the type locality of P. (M.) proceroides sp. nov.), he evidently treated under the name “ Odontochila procera ”, at least partially, P. (M.) proceroides sp. nov. from its type locality. He briefly mentioned the occurrence of these species as “in jungle”, and that adults seldom flew onto ground. Remarks. As mentioned under P. (M.) procera above, Horn (1929: 157 fig. 20, 158) confused P. (M.) proceroides sp. nov. with P. (M.) procera. Zikán (1929) also treated this new species partly under the name P. (M.) procera (see “Biology and distribution” above). The initials of the baptismal names of Joseph Francisco Zikán, the collector of this new species and entomologist who specialized in immature stages of tiger beetles, are often variably and wrongly cited for his publication (Zikán 1929).

Genus Opisthencentrus W. Horn, 1893 Opisthencentrus W. Horn, 1893: 196. Type species. Oxygonia dentipennis Germar, 1843, by original designation (and monotypy). Differential diagnosis. Monobasic genus immediately distinguished from all other genera of the subtribe Odontocheilina by the conspicuous, long lateral-median spine of the elytral apex (Figs 15–18), and by the femoral apices possessing two conspicuous spines (Fig. 13). The aedeagus and internal sac are similar to that in Pentacomia Bates, 1872, as the internal sac contains a ventral spur with a short projection (among other sclerites). However, the internal sac in Opisthencentrus contains characteristic distinctly serrate dorsal sclerite (Fig. 21), unknown in other genera. The spiny femoral apices in Opisthencentrus are similar to those in the genus Oxygonia Mannerheim, 1837. However, the diagnostic characters which differentiate Opisthencentrus from Oxygonia comprise particularly the shape of the elytra and their matte appearance (Fig. 19) due to the irregular punctures with bumpy to tuberculate intervals (corresponding with the pattern of elytral sculptures illustrated by Moravec (2007: 42, figs 128-129), the dimorphic shape of the labrum and only four teeth in each mandible (plus basal molar), as well as the different shape of the aedeagus and internal sac lacking a flagellum. In contrast, all species of Oxygonia can be immediately distinguished from Opisthencentrus by their shiny body surface due to much more regularly spaced, or partly effaced elytral punctation, mandibles with six to seven teeth (plus basal molar), elytral apex with only a sutural spine, and aedeagus with the internal sac possessing in all species of Oxygonia a flagellum. Moreover, the terminal protarsomere 5 in Oxygonia is inserted dorsally into the protarsomere 4 as emphasized by Horn (1905) and illustrated by Horn (1915) and Kippenhan (1997).

in English The motivation for this thesis is to analyze Bitcoin as Bitcoin is still a new phenomenon and requires an economical, legal, and legislative evaluation. In this thesis we seek an answer to the question what is Bitcoin. This thesis is divided into three main blocks, within them we use analytical, descriptive, and comparative methods. In the first part, we aim mostly on the economic side of Bitcoin, comparing Bitcoin to money in general sense. In the second part, we argue whether Bitcoin shall be considered a thing in a legal sense under the Czech law. In the third part, we describe the current Bitcoin's regulatory situation and suggest a drafting of Bitcoin specific regulation. The conclusion of this thesis is that Bitcoin is a digital medium of exchange that cannot be considered money or currency. To the extent, Bitcoin might be interpreted in ways where it does not satisfy the legal definition of a thing in legal sense, but for regulatory purposes it shall be, at least for now, considered a thing in legal sense. We also suggest that Bitcoin shall be systematically regulated by imposing various duties on persons who's scope of business is relating to the Bitcoin.

The rigorous thesis deals with involvement of young people in nature conservation within the Brontosaurus Movement, both children under 15 years of age and teenagers. The work is primarily based on documentation of the Communist Party of Czechoslovakia, Socialist Union of Youth, Security Services Archive and on private archives of former activists of the Brontosaurus Movement and magazine Mladý svět (Young World). Structured interviews with the former Brontosaurus Movement activists are the most important source of knowledge. One of the main goals of the work is to map the history of the Brontosaurus Movement and the extent to which the movement activities were influenced by Party and Union bodies. The work was concurrently confronted with a wide variety of personal testimonies. The work should contribute to development of civil society and environmental education. Powered by TCPDF (www.tcpdf.org)