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Anomaloglossus saramaka sp. nov. Anomaloglossus sp. “Brownsberg” Grant et al. 2006 Anomaloglossus sp. Fouquet et al. 2007 Anomaloglossus sp. Fouquet et al. 2012 Anomaloglossus baeobatrachus Ouboter and Jairam 2012 Anomaloglossus sp. “Brownsberg” Grant et al. 2017 Anomaloglossus sp. Brownsberg Vacher et al. 2017 Holotype. MNHN _RA_2019.0016 (field n°AF3845), an adult male, collected by A. Fouquet, S. Cally and R. Jairam on 01 February 2016 at Brownsberg, Suriname (4.936527 N 55.194826 W, ~ 500 m elevation; Figs. 2‒3; Ap-pendix 1). Paratypes. Fifteen specimens: MNHN _RA_2019.0018–21, 27, 28, 29 (field n°AF3840–43, 3884, 3889, 3897) and NZCS-A1206 (field n°AF3844) eight adult males and MNHN _RA_2019.0022‒26 (field n° AF3848, 3876, 3877, 3881, 3882) and NZCS-A1207 (field n° AF3847) six females collected with the holotype by A. Fouquet, S. Cally and R. Jairam, and MNHN _RA_2019.0017 (field n°AF3774) an adult male collected by A. Fouquet, S. Cally and R. Jairam on 25 January 2016 at Voltzberg, Suriname (4.68169 N 56.18568 W, ~ 100 m elevation). Etymology. The specific epithet is a noun in apposition and refers to the Saramaka people who live nearby the type locality and are one of the Maroon communities living in Suriname. Definition. (1) Medium-sized Anomaloglossus (average male SVL 19.3 mm [18.6–19.9, n = 10], female SVL 20.3 mm [19.9–21.6, n = 6]) (Table 1); (2) body robust; (3) skin on dorsum finely granular with larger scattered tubercles becoming denser on the posterior half and legs, with a larger tubercle on each eyelid; (4) ventral skin smooth except under thighs where it is finely granular; (5) inconspicuous supratympanic fold; (6) tympanum distinct anteroventrally; (7) snout long and protruding in lateral view; (8) nares oriented ventrolaterally, situated near tip of snout; (9) Finger II smaller than Finger I when fingers adpressed; (10) tip of Finger IV reaching distal edge of distal subarticular tubercle on Finger III when fingers adpressed; (11) distal subarticular tubercle distinct on Finger III and IV, not visible on the other fingers; (12) Finger III swollen dorsally and preaxially in males, extending largely towards dorsal surface of hand forming a cream-coloured gland basally; (13) fringes present on all fingers, particularly developed postaxially on Finger I and prexially on Fingers II and III, in males and females; (14) toes basally webbed, with well-developed fringes on all toes, more developed preaxially on Toes II, III and IV; (15) tarsal keel well-defined, slightly curved; (16) glandular supracarpal pad present in both sexes (larger and more glandular in males); (17) cloacal tubercles present; (18) paracloacal mark present (orangish to light brown in life, cream to grey in preservative); (19) dorsolateral stripe absent; (20) oblique lateral stripe present, solid, undulating (cream but sometimes white anteriorly, yellow at groin level in life, white in preservative), lateral band in life uniformly dark brown/black in males, brown in females, lower flank grey with white speckles, more or less extensively yellowish orange in the axillary and the groin regions; (21) ventrolateral stripe absent; (22) sexual dichromatism in throat colour pattern present, in life light grey in males with sparse minute black melanophores, evenly and entirely yellow in females; (23) sexual dichromatism in ventral colour pattern present, in life abdomen entirely or only posteriorly yellowish orange in males, uniformly bright yellowish orange in females; (24) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (25) median lingual process as long as wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (26) call 0.44–0.69 s in length, consisting of a train of 5–11 notes ranging 0.028–0.036 s in length and spaced by intervals of 0.024–0.030 s with a dominant frequency at 4.55–5.02 kHz (n = 7) (Fig. 4, Table 2); (27) type 4 tadpole (Orton 1953), exotrophic, with a functional mouth with marginal papillae and labial teeth (Fig. 5, Table 3). Morphological comparisons with other Anomaloglossus. The only other species group co-occurring with the Anomaloglossus stepheni group is the A. degranvillei group, represented by A. blanci Fouquet, Vacher, Courtois, Villette, Reizine, Gaucher, Jairam, Ouboter, and Kok, 2018, A. degranvillei (Lescure, 1975), A. dewynteri Fouquet, Vacher, Courtois, Villette, Reizine, Gaucher, Jairam, Ouboter, and Kok, 2018 and A. surinamensis Ouboter and Jairam, 2012. Anomaloglossus saramaka sp. nov. can be distinguished from species of the A. degranvillei group by its basal webbing (moderate in species of the A. degranvillei group), smaller fringes that are more developed on Toes II-IV (well-developed fringes on all toes) and the presence of a solid oblique lateral stripe (absence). Besides these two groups, all the other described Anomaloglossus species exclusively occur in Pantepui (i.e., the Guiana Shield highlands, see Kok et al. 2018) and have moderate to extensive toe webbing (Barrio-Amorós 2006; Barrio-Amorós & Brewer-Carías 2008; Barrio-Amorós et al. 2004, 2010; Fouquet et al. 2015; Kok et al. 2010; La Marca 1997; Meinhardt & Parmalee 1996; Noble 1923; Rivero 1961) except A. meansi Kok, Nicolaï, Lathrop and MacCulloch, 2018, A. kaiei (Kok, Sambhu, Roopsind, Lenglet and Bourne, 2006), A. rufulus (Gorzula, 1990) and A. roraima (La Marca, 1997). However, unlike A. saramaka sp. nov., all these species lack an oblique lateral stripe. Within the Anomaloglossus stepheni group (comparisons follow below), Anomaloglossus saramaka sp. nov. is morphologically most similar to A. baeobatrachus, A. leopardus, A. mitaraka and A. sp. “Bakhuis” (see below) and can easily be confused with these species. Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. baeobatrachus by (1) larger body size (mean = 19.3; range 18.6–19.9 mm in males [n = 10] and mean = 20.3; range 19.9–21.6 mm in females [n = 6] in A. saramaka sp. nov. vs mean = 16.2; range 14.8–17.1 mm in males [n = 16] and mean = 18.4; range 17.3–19.4 mm in females [n = 4] in A. baeobatrachus); (2) venter mostly yellowish orange in males in A. saramaka sp. nov. (presence of a cream/white central patch in A. baeobatrachus); (3) call with lower note rate (mean = 14.6, range 14.0–15.3 notes/s in A. saramaka sp. nov. [n = 7] vs mean = 16.2, range 15.6–16.8 notes/s in A. baeobatrachus [n = 9]) (Table 2). Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. leopardus by (1) narrow, poorly defined dark brown bars on legs (broad and conspicuous in A. leopardus); (2) venter mostly yellowish-orange in males in A. saramaka sp. nov. (presence of a more or less extensive cream/white central patch in A. leopardus); (3) shorter advertisement call (mean = 0.51, range 0.38– 0.69 s in A. saramaka sp. nov. [n = 7] vs mean = 1.52, range 1.08– 2.00 s in A. leopardus [n = 4]); (4) fewer notes in calls (6–11 in A. saramaka sp. nov. [n = 7] vs 14–28 in A. leopardus [n = 4]) (Table 2). Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. mitaraka by (1) venter mostly yellowish orange in males in A. saramaka sp. nov. (presence of a more or less extensive cream/white central patch in A. mitaraka); (2) shorter advertisement call (mean = 0.51, range 0.38– 0.69 s in A. saramaka sp. nov. [n = 7] vs mean = 1.04, range 0.74– 1.23 s in A. mitaraka sp. nov. [n = 6]); (3) higher note rate call (mean = 14.6, range 14.0–15.3 note/s in A. saramaka sp. nov. [n = 7] vs mean = 11.43 note/s, range 10.8–12.3 in A. mitaraka [n = 6]) (Table 2). Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. apiau by (1) solid oblique lateral stripe in A. saramaka sp. nov. (partial, only present as a series of conspicuous white dots extending from groin midway along flank in A. apiau); (2) shorter advertisement call (mean = 0.51, range 0.38– 0.69 s in A. saramaka sp. nov. [n = 7] vs mean = 19.6, range 7.0– 39.4 s in A. apiau [n = 10]). Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. stepheni (Figs. 2‒3, Table 1) by (1) larger body size (mean = 19.3; range 18.6–19.9 mm in males [n = 10] and mean = 20.3; range 19.9–21.6 mm in females [n = 6] in A. saramaka sp. nov. vs mean = 17.2, range 16.5–18.0 mm in males [n = 10] and mean = 17.3, range 17.0–18.0 mm in females [n = 5] [from Martins 1989] in A. stepheni); (2) skin on dorsum finely granular in A. saramaka sp. nov. (evenly tuberculate in A. stepheni); (3) ventral colouration mostly yellowish orange in males in A. saramaka sp. nov. (uniformly cream or white in A. stepheni); (4) longer advertisement call (mean = 0.51, range 0.38– 0.69 s in A. saramaka sp. nov. [n = 7] vs mean= 0.25 s, range 0.18– 0.29 s in A. stepheni [n = 4]); (5) lower note rate call (mean = 14.6, range 14.0–15.3 note/s in A. saramaka sp. nov. [n = 7] vs mean = 21.3, range 20.2–22.6 note/s in A. stepheni [n = 4]) (Table 2). Description of the holotype. An adult male, 18.9 mm SVL; body robust; head wider than long, HL 95 % of HW; HL 33 % of SVL; dorsal skin finely granular, one enlarged tubercle on each eyelid, snout long (SL 52 % of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 42 % of HW; EN 29 % of HL, 75 % of ED. Tympanum distinct anteroventrally; supratympanic fold inconspicuous; choanae small, drop shaped, located anterolaterally. Forelimb slender, skin tuberculate; HAND 24 % of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened; webbing absent on fingers; lateral fringes present on all fingers, particularly developed on postaxial edge of Finger I and on preaxial edges of Finger II and III; Finger III distinctly swollen dorsally and preaxially, swelling largely extending towards dorsal surface of hand; tip of Finger IV reaching distal edge of distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5 times width of digit; width of disc on Finger III 0.6 mm; discs with distinct dorsal scutes; relative lengths of adpressed fingers III> IV> II> I; palmar tubercle large, heart-shaped, 0.8 mm in diameter (larger than disc on Finger III), thenar tubercle elliptic, small (equal to disc on Finger III in maximum diameter), half the size of palmar tubercle, well separated from palmar tubercle; only basal subarticular tubercles on fingers are conspicuous; distal subarticular tubercle of Finger III and IV are almost inconspicuous, basal subarticular tubercles on Fingers I and II the largest, followed by subarticular tubercle and basal subarticular tubercle on Finger III. Hind limb robust, skin tuberculate; TL 48 % of SVL; heels in contact when hind limbs are flexed at right angles to the sagittal plane of body; FL 43 % of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes. Width of disc on Toe IV 0.7 mm. Foot basally webbed; lateral fringes present on all toes, more developed preaxially on Toes II, III and IV. Toe webbing formula I 1+ ‒ 1- II 1+ ‒ 1- III 1+ ‒ 1+ IV 0 ‒ 1+ V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and slightly curved at proximal end. Metatarsal fold strong. Colour of holotype in life. Dorsal colour uniformly dark brown. Oblique lateral stripe solid, continuous, undulating, anteriorly brownish white, cream in the middle and yellow near groin (Fig. 2). Black lateral band below oblique lateral stripe, tapering posteriorly, extending from tip of snout to groin and containing the indistinct dorsal part of tympanum. Upper lip brownish white with a few small white speckles. Lower flank (below black lateral band) grey with white speckles with yellowish orange extensions from the axillary and the groin region. Throat grey covered with minute melanophores, more densely laterally; belly anteriorly greyish white, yellowish orange laterally and posteriorly, ventral surfaces of thighs and arms yellowish orange. Iris with coper metallic pigmentation and pupil ring interrupted dorsally and ventrally by transversal pigmentation (Fig. 2). Upper and lower arm reddish brown dorsally with brown ill-defined spots and white speckles. Bright yellow spot at junction of upper arm with body, and anterior part of upper arm with a black band. Black glandular spot at junction between lower arm and hand, followed by a cream coloured part of the swollen gland extending on Finger III. Dorsal surfaces of thigh, shank and tarsus dark brown with ill-defined dark brown, almost black transverse bars, and ill-defined dark brown blotches. Paracloacal marks light brown, elongate. Toes and digits with small light blueish speckles dorsally and laterally. Palms and soles dark brown. Colour of holotype in preservative. After four years in 70 % ethanol, colours of the specimen faded, and the dorsal colouration now varies from pale brown to grey with a brown interorbital bar; yellowish orange ventral colouration faded to white (Fig. 3). Bluish speckles and orange/reddish marks turned white as well. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. Colouration of limbs varies from brown to reddish brown, females being more reddish. Colour of dorsum varies from light brown to dark brown and some specimens (e.g., NZCS-A1206, field n°AF3844) dis-play diffuse dark brown blotches, more frequently on the interorbital region. Colour of oblique lateral stripe varies from white to orangish. Overall dorsal and lateral colouration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as males carrying tadpoles apparently display overall lighter colours, smoother skin and sharper contrasts, whereas calling males are very dark coloured and highly tuberculate. Ventral colouration of female is entirely yellowish orange while the yellowish orange parts are always limited to the posterior region of throat in males. Vocal sac, slits and minute black melanophores on throat only observed in males as well as swelling of Finger III. Advertisement call. Seven collected specimens calling from the leaf litter were recorded from a distance of about 2 m and at temperatures ranging from 22 to 24°C. Descriptive statistics of call parameters are presented in Table 2. Anomaloglossus saramaka sp. nov. emits trains (call length mean = 0.51, range 0.38– 0.69 s) of 6–11 short notes (note length mean = 0.028 s; range 0.024 –0.031 s; inter-note interval mean = 0.05 s; range 0.04– 0.05 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 4.84 kHz on average (range 4.55–5.02 kHz) with a slight upward modulation (ca. 0.3 kHz) (Fig. 4, Table 2). Larval morphology. The following description is based on six tadpoles at stage 27, 28 and 29 (Fig. 5, Table 3). Tadpoles correspond to a type 4 tadpole of Orton (1953); exotrophic; body skin smooth; TL 12.95–17.19 mm; BL 4.33–5.89 mm, 33–36 % of TL, 146–165 % of BW, 191–238 % of BH; BW 123–149 % of BH (Table 3); body ovoid, snout round in dorsal and lateral view; eyes positioned and directed laterally; ED 0.45–0.60 mm, 53–64 % of IOD; IOD larger than IND; nares frontally positioned and directed laterodorsally; narial opening circular in lateral view; END 0.45–0.56 mm. Spiracular tube sinistral, conical, projecting posterodorsally, its tip located at 60–65 % of BL posteriorly to snout. Lateral-line system inconspicuous. Caudal musculature highest at its base, tapering posteriorly, terminating at tail tip; tail tip rounded; upper fin originating at junction of body and tail, gradually increasing in height to about 3/4 of tail; UTF 40–53 % of TMH; LTF 33–47 % of TMH; MTH 13–15 % of TL (Fig. 5, Table 3). Mouth ventral, oral disc strongly emarginated, width 1.40–1.50 mm. Labial teeth long, in single rows, LTRF 2(2)/3. A-2 consisting of two short rows, separated by a large and deep gap; P-1 not interrupted. Marginal papillae long, of equal size on each labium, tapered, blunt-tipped, in a single row, evenly distributed; median gap on upper labium approximately 2/3 the length of A-1; jaw sheaths large, serrated, lower jaw sheath broadly V-shaped. In life, the entire body is dark grey with abundant golden flecks, particularly on dorsum. Golden flecks become scarce ventrally. Posteriorly, dark grey coloration fades and tail becomes translucent. The tadpole of Anomaloglossus saramaka sp. nov. can be distinguished from those of Anomaloglossus of the stepheni group that are endotrophic (A. stepheni and A. baeobatrachus) by the presence of a functional mouth with marginal papillae and labial teeth. However, it virtually cannot be distinguished from the tadpole of A. mitaraka based on morphological evidence. Distribution and natural history. Anomaloglossus saramaka sp. nov. is a diurnal species inhabiting the leaf litter in primary forest at low to mid elevations (from 100 to 500 m elevation). The species is usually found close to streams next to which it deposits its exotrophic tadpoles into puddles, but some specimens were found more than 50 m away from water bodies. Males call all day long when the weather is rainy but calling activity is otherwise limited to sunset and sunrise. Breeding occurs during the rainy season, from December to July. The rainfall between December and April being highly variable, the reproduction seems to be concentrated in the second half (May-July) of this period. The males respond to intraspecific playbacks with shorter and more rapidly emitted note trills. Males are territorial, their small territories being spaced at least two meters apart.

Anomaloglossus vacheri sp. nov. Anomaloglossus baeobatrachus Ouboter & Jairam 2012 Anomaloglossus sp. Bakhuis Vacher et al. 2017 Holotype. MNHN _RA_2019.0009 (field n��AF3413), an adult male, collected by A. Fouquet, S. Cally and R. Jairam on 30 April 2015 at Bakhuis Mountains, Suriname (4.72462 N 56.7638 W, ~ 200 m elevation; Figs. 2���3; Ap-pendix 1). Paratopotypes. Eight specimens: MNHN _RA_2019.0009���11, 13���15 (field n��AF3412, 3424, 3426, 3430, 3434) and NZCS-A1208���9 (field n��AF3433, 3441) seven adult males and MNHN _RA_2019.0012 (field n�� AF3425) one female collected with the holotype by A. Fouquet, S. Cally and R. Jairam. Etymology. This species is dedicated to Jean-Pierre Vacher, in honour of his strong contribution to the understanding of the diversity and the evolution of the genus Anomaloglossus. Definition. (1) Small-sized Anomaloglossus (average male SVL 17.4 mm [17.0���17.8, n = 8], female SVL 19.7 mm [n = 1]) (Table 1); (2) body robust; (3) skin on dorsum finely granular with larger scattered tubercles becoming denser on the posterior half and legs, with a larger tubercle on each eyelid; (4) ventral skin smooth except under thighs, where it is finely granular; (5) inconspicuous supratympanic fold; (6) tympanum distinct anteroventrally; (7) snout long and protruding in lateral view; (8) nares oriented ventrolaterally, situated near tip of snout; (9) Finger II equal to Finger I when fingers adpressed; (10) tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; (11) distal subarticular tubercle distinct on Finger III and IV, inconspicuous on the other fingers; (12) Finger III swollen dorsally and preaxially in males, extending largely towards dorsal surface of hand; (13) fringes present on all fingers, particularly developed post- and preaxially on Finger II and preaxially on Finger III, almost inconspicuous on Finger IV, in males and females; (14) toes basally webbed, with well-developed fringes on all toes, more developed preaxially on Toes II and III; (15) tarsal keel well-defined, strongly curved; (16) glandular supracarpal pad present in both sexes (larger and more glandular in males); (17) cloacal tubercles present; (18) paracloacal mark present (orangish in life, cream to grey in preservative); (19) dorsolateral stripe absent; (20) oblique lateral stripe present, diffuse (composed of yellow and white speckles), particularly diffuse anteriorly, extending along upper eyelid into a diffuse canthal stripe, lateral band uniformly dark brown/black in males and females in life, lower flank yellowish orange to grey with white speckles; (21) ventrolateral stripe absent; (22) dorsal surface of arms barely blotched in males, orange; (23) sexual dichromatism in throat colour pattern present, in life light grey in males with sparse minute black melanophores (chin often yellow), evenly and entirely yellow in females; (24) sexual dichromatism in ventral colour pattern present, in life abdomen entirely yellowish orange in males, uniformly yellowish orange in females; (25) iris with metallic pigmentation and pupil ring inconspicuously interrupted ventrally by transversal black pigmentation; (26) median lingual process as long as wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (27) call 0.47���0.91 s in length, consisting of a train of 8���18 notes of 0.019���0.021 s in length and spaced by intervals of 0.042 s with a dominant frequency at 5.11���5.35 kHz (n = 5; Table 2); (28) type 4 tadpole (Orton 1953), exotrophic, with a functional mouth with marginal papillae and labial teeth (Fig. 5, Table 3). Morphological comparisons with other Anomaloglossus. Like Anomaloglossus saramaka sp. nov., A. vacheri sp. nov. can be distinguished from species of the A. degranvillei group by its basal webbing (moderate in species of the A. degranvillei group), smaller fringes that are more developed on Toes II-IV (well-developed fringes on all toes) and the presence of a solid oblique lateral stripe (absence) and from all the other described Anomaloglossus species of the Pantepui region by the presence of an oblique lateral stripe. Within the Anomaloglossus stepheni group (comparisons follow below), A. vacheri sp. nov. is morphologically most similar to A. baeobatrachus, A. leopardus, A. mitaraka and A. saramaka sp. nov. and can easily be confused with these species. Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. baeobatrachus by (1) oblique lateral stripe extending along upper eyelid into a canthal stripe in A. vacheri sp. nov. (oblique lateral stripe not extending along upper eyelid into a canthal stripe in A. baeobatrachus); (2) dorsal surface of arms barely blotched and orange in males of A. vacheri sp. nov. (with dark brown blotches and light brown in A. baeobatrachus); (3) ventral coloration yellowish orange in males in A. vacheri sp. nov. (white or only posteriorly yellow in A. baeobatrachus); (4) call with higher note rate (mean = 18.2, range 17.3���19.2 note/s in A. vacheri sp. nov. [n = 8] vs mean = 16.2, range 15.6���16.8 note/s in A. baeobatrachus [n = 9]) (Table 2). Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. leopardus by (1) narrow, poorly defined dark brown bars on legs (broad and conspicuous dark brown in A. leopardus); (2) dorsal surface of arms barely blotched and orange in males of A. vacheri sp. nov. (light brown with dark brown blotches in A. leopardus); (3) oblique lateral stripe extending along upper eyelid into a canthal stripe in A. vacheri sp. nov. (not extending along upper eyelid into a canthal stripe in A. leopardus); (4) advertisement call shorter (mean = 0.63, range 0.47��� 0.91 s in A. vacheri sp. nov. [n = 5] vs mean = 1.52, range 1.08��� 2.00 s in A. leopardus [n = 4]) and higher in frequency (mean = 5.25, range 5.11���5.35 kHz in A. vacheri sp. nov. [n = 5] vs mean = 4.45, range 4.40���4.57 kHz in A. leopardus [n = 4]) (Table 2). Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. mitaraka by (1) smaller male body size (mean = 17.4; range 17.0��� 17.8 mm in males [n = 8] in A. vacheri sp. nov. vs mean = 18.6; range 18.2���19.3 mm in males [n = 7] in A. mitaraka); (2) dorsal surface of arms barely blotched and orange in males of A. vacheri sp. nov. (with dark blotches and light brown in A. mitaraka); (3) oblique lateral stripe extending along upper eyelid into a canthal stripe A. vacheri sp. nov. (not extending along upper eyelid into a canthal stripe in A. mitaraka); (4) advertisement with a higher note rate (mean = 18.2, range 17.3���19.1 note/s in A. vacheri sp. nov. [n = 8] vs mean = 11.4 note/s, range 10.8���12.3 in A. mitaraka [n = 6]) and higher in frequency (mean = 5.25, range 5.11���5.35 kHz in A. vacheri sp. nov. [n = 5] vs mean = 4.43, range 4.12���4.76 kHz in A. mitaraka [n = 6]) (Table 2). Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. saramaka sp. nov. by (1) smaller male body size (mean = 17.4; range 17.0��� 17.8 mm in males [n = 8] in A. vacheri sp. nov. vs mean = 19.3; range 18.6���19.9 mm in males [n = 10] in A. saramaka sp. nov.); (2) dorsal surface of arms barely blotched and orange in males of A. vacheri (with dark blotches and light brown in A. saramaka sp. nov.); (3) oblique lateral stripe extending along upper eyelid into a canthal stripe in A. vacheri sp. nov. (not extending along upper eyelid into a canthal stripe in A. saramaka sp. nov.); (4) advertisement with a higher note rate (mean = 18.2, range 17.3���19.2 note/s in A. vacheri sp. nov. [n = 8] vs mean = 14.5 note/s, range 14.0��� 15.3 in A. saramaka sp. nov. [n = 6]) and higher in frequency (mean = 5.25, range 5.11���5.35 kHz in A. vacheri sp. nov. [n = 5] vs mean = 4.84, range 4.55���5.02 kHz in A. saramaka sp. nov. [n = 6]) (Table 2). Anomaloglossus vacheri sp. nov. can be mainly distinguished from A. apiau by (1) solid oblique lateral stripe in A. vacheri sp. nov. (present as a series of conspicuous white dots extending from groin midway along flank in A. apiau); (2) by its shorter call (mean = 0.63, range 0.47��� 0.91 s in A. vacheri sp. nov. [n = 5] vs mean = 19.6, range 7.0��� 39.4 s in A. apiau [n = 10]). Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. stepheni (Figs. 2���3) by (1) skin on dorsum finely granular in A. vacheri sp. nov. vs evenly tuberculate in A. stepheni; (2) ventral colouration mostly yellowish orange in males in A. vacheri sp. nov. vs uniformly white in A. stepheni; (3) oblique lateral stripe extending along upper eyelid into a canthal stripe in A. vacheri sp. nov. vs oblique lateral stripe not extending along upper eyelid into a canthal stripe in A. stepheni; (4) advertisement call longer (mean = 0.63, range 0.47��� 0.91 s in A. vacheri sp. nov. [n = 5] vs mean= 0.25 s, range 0.18��� 0.29 s in A. stepheni [n = 4]), and higher in frequency (mean = 5.25, range 5.11���5.35 kHz in A. vacheri sp. nov. [n = 5] vs mean = 4.48, range 4.25���4.85 kHz in A. stepheni [n = 4]) (Table 2). Description of the holotype. An adult male, 17.5 mm SVL; body robust; head wider than long, HL 98 % of HW; HL 33 % of SVL; dorsal skin finely tuberculate with larger tubercles posteriorly, one enlarged tubercle on each eyelid, snout long (SL 51 % of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 41 % of HW; EN 30 % of HL, 77 % of ED. Tympanum distinct anteroventrally; supratympanic fold inconspicuous; choanae small, circular, located anterolaterally. Forelimb slender, skin tuberculate; HAND 23 % of SVL; Finger I as long as Finger II when fingers adpressed; fingers large and flattened; webbing absent on fingers; lateral fringes present, particularly developed post and preaxially on Finger II and preaxially on Finger III; Finger III distinctly swollen dorsally and preaxially, swelling largely extending towards dorsal surface of hand; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5 times width of digit; width of disc on Finger III 0.6 mm; discs with distinct dorsal scutes; relative lengths of adpressed fingers III> IV> II> I; palmar tubercle large, heart-shaped, 0.75 mm in diameter (larger than disc on Finger III), thenar tubercle elliptic, small (equal to disc on Finger III in maximum diameter), half the size of palmar tubercle, well separated from palmar tubercle; basal subarticular tubercles on fingers and distal subarticular tubercle of Finger III and IV conspicuous; subarticular tubercles on Fingers I and II the largest, followed by subarticular tubercles and basal subarticular tubercle on Finger III. Hind limb robust, skin tuberculate; TL 48 % of SVL; heels in contact when hind limbs are flexed at right angles to the sagittal plane of body; FL 42 % of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes. Width of disc on Toe IV 0.8 mm. Foot basally webbed; lateral fringes present on all toes, more developed preaxially on Toes II and III. Toe webbing formula I 1+ ��� 1- II 1+ ��� 1- III 1+ ��� 1+ IV 0 ��� 1+ V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end. Metatarsal fold strong. Colour of holotype in life. Dorsal colour light brown with dark brown blotches occurring on the interorbital region and at mid-body forming an hourglass pattern. Blotches of the same colour also occur laterally along the oblique lateral stripe. Oblique lateral stripe thin, diffuse (particularly anteriorly), formed by white and yellow speckles, extending along the upper eyelid into a diffuse canthal stripe (Fig. 2). Dark brown lateral band below oblique lateral stripe, tapering posteriorly, extending from tip of snout to groin and containing the indistinct dorsal part of tympanum, tapering posteriorly. Upper lip grey with a few small white speckles. Lower flank (below dark brown lateral band) grey with white speckles. Throat light grey, chin yellowish orange, throat and chin covered with minute melanophores, more densely laterally; belly bright yellowish orange, ventral surfaces of thighs and arms yellowish orange. Iris with coper metallic pigmentation and pupil ring uninterrupted by transversal pigmentation (Fig. 2). Upper and lower arm light orange dorsally with a few white speckles. Black glandular spot at junction between lower arm and hand, followed by a grey coloured part of the swollen gland extending on Finger III. Dorsal surfaces of thigh, shank and tarsus light brown with ill-defined dark brown transverse bars and ill-defined dark brown blotches. Paracloacal marks light brown, elongate. Toes and digits with small light blueish speckles dorsally and laterally. Palms and soles dark brown. Colour of holotype in preservative. After four years in 70 % ethanol, colours of the specimen faded and the dorsal colouration now varies from pale brown to grey with darker blotches, the yellowish orange ventral colouration faded to white (Fig. 3). Bluish speckles and orange marks turned white as well. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. Colour of oblique lateral stripe varies in its proportion of white and yellow. Overall dorsal and lateral coloration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as calling males are dark coloured and highly tuberculate. Dorsal dark brown hourglass pattern may be absent (MNHN _RA_2019.0013, field n�� AF3426; NZCS-A1208, field n��AF3433). Paracloacal marks sometimes inconspicuous. Throat colouration of female is entirely yellowish orange while the yellowish orange parts are often limited to the chin in males (extent of coloured part of chin varies). Vocal sac, slits and minute black melanophores on throat only observed in males as well as swelling of Finger III. Advertisement call. Five collected specimens calling from the leaf litter were recorded from a distance of about 2 m and at temperatures between 25���26��C and 98% relative humidity. Descriptive statistics of call parameters are presented in Table 2. Anomaloglossus vacheri sp. nov. emits trains (call length mean = 0.63, range 0.47��� 0.91 s) of 8���18 short notes (note length mean = 0.020 s; range 0.019 ���0.021 s; inter-note interval mean = 0.039 s; range 0.036 ���0.042 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 5.25 kHz on average (range 5.11���5.35 kHz) with a slight upward modulation (ca. 0.4 kHz) (Fig. 4, Table 2). Larval morphology. The following description is based on two tadpoles at stage 28 and 30 (Fig. 5). Tadpoles correspond to a type 4 tadpole of Orton (1953); exotrophic; body skin smooth; TL 15.00��� 19.06 mm; BL 5.27���6.46 mm, 34���35 % of TL, 151���160 % of BW, 261���267 % of BH; BW 135���144 % of BH (Table 3); body ovoid, snout round in dorsal and lateral view; eyes positioned and directed laterally; ED 0.49���0.61 mm, 55���58 % of IOD; IOD smaller than IND; nares frontally positioned and directed laterodorsally; narial opening circular in lateral view; END 0.55���0.71 mm. Spiracular tube sinistral, conical, projecting posterodorsally, its tip located at 57���59 % of BL posteriorly to snout. Lateral-line system inconspicuous. Caudal musculature highest at its base, tapering posteriorly, terminating at tail tip; tail tip rounded; upper fin originating at junction of body and tail, gradually increasing in height to about 3/4 of tail; UTF 47���48 % of TMH; LTF 46���49 % of TMH; MTH 13���16 % of TL (Fig. 5, Table 3). Mouth ventral, oral disc strongly emarginated, width 1.27���1.89 mm. Labial teeth long, in single rows, LTRF 2(2)/3. A-2 consisting of two short rows, separated by a large and deep gap; P-1 not interrupted. Marginal papillae long, of equal size on each labium, tapered, blunt-tipped, in a single row, evenly distributed; median gap on upper labium approximately 2/3 the length of A-1; jaw sheaths large, serrated, lower jaw sheath broadly V-shaped. In life, the body is dark grey with abundant golden flecks, particularly on dorsum, except the loreal and postocular regions that are pink. Golden flecks become scarce ventrally. Posteriorly, dark grey coloration fades and tail becomes translucent. The tadpole of Anomaloglossus vacheri sp. nov. can be distinguished from those of Anomaloglossus of the stepheni group that are endotrophic (A. stepheni and A. baeobatrachus) by the presence of a functional mouth with marginal papillae and labial teeth. It can be distinguished from the tadpole of A. mitaraka and A. saramaka sp. nov. by its less dense pigmentation particularly ventrally and by a spiracle located more anteriorly. Distribution and natural history. Anomaloglossus vacheri sp. nov. is a diurnal species inhabiting the leaf litter in primary forest at low elevation (200 m elevaion). The species has only been found in terra firme forest on lateritic crust distant from streams. Males call after rainfall. Breeding apparently occurs during the rainy season, probably extending between December and July, depending on rainfall. The males respond to intraspecific playbacks with shorter and more rapidly emitted note trills. Males are spaced by at least five meters apart. They usually call slightly above the leaf litter, exposed on a branch or a dead leaf. Eggs are probably deposited under or in the fold of a dead leaf. After they hatch the male probably carries the tadpoles to small water bodies formed in the lateritic crust where tadpoles where found (Fig. 5). A single population is known to date, in Bakhuis Mountains, Suriname. The species likely occurs throughout the massif, but it unlikely extends much furthe, Published as part of Fouquet, Antoine, Jairam, Rawien, Ouboter, Paul & Kok, Philippe J. R., 2020, Two new species of Anomaloglossus (Anura: Aromobatidae) of the stepheni group from Suriname, pp. 147-164 in Zootaxa 4820 (1) on pages 156-160, DOI: 10.11646/zootaxa.4820.1.7, http://zenodo.org/record/4397441, {"references":["Ouboter, P. E. & Jairam, R. (2012) Amphibians of Suriname. Brill, Leiden, 376 pp. https: // doi. org / 10.1163 / 9789004208001","Vacher, J. - P., Kok, P. J. R., Rodrigues, M. T., Dias Lima, J., Lorenzini, A., Martinez, Q., Fallet, M., Courtois, E. A., Blanc, M., Gaucher, P., Dewynter, M., Jairam, R., Ouboter, P., Thebaud, C. & Fouquet, A. (2017) Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield. Molecular Phylogenetics and Evolution, 112, 158 - 173. https: // doi. org / 10.1016 / j. ympev. 2017.04.017","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2, 63 - 75. https: // doi. org / 10.2307 / 2411661"]}

Anomaloglossus saramaka sp. nov. Anomaloglossus sp. ���Brownsberg��� Grant et al. 2006 Anomaloglossus sp. Fouquet et al. 2007 Anomaloglossus sp. Fouquet et al. 2012 Anomaloglossus baeobatrachus Ouboter and Jairam 2012 Anomaloglossus sp. ���Brownsberg��� Grant et al. 2017 Anomaloglossus sp. Brownsberg Vacher et al. 2017 Holotype. MNHN _RA_2019.0016 (field n��AF3845), an adult male, collected by A. Fouquet, S. Cally and R. Jairam on 01 February 2016 at Brownsberg, Suriname (4.936527 N 55.194826 W, ~ 500 m elevation; Figs. 2���3; Ap-pendix 1). Paratypes. Fifteen specimens: MNHN _RA_2019.0018���21, 27, 28, 29 (field n��AF3840���43, 3884, 3889, 3897) and NZCS-A1206 (field n��AF3844) eight adult males and MNHN _RA_2019.0022���26 (field n�� AF3848, 3876, 3877, 3881, 3882) and NZCS-A1207 (field n�� AF3847) six females collected with the holotype by A. Fouquet, S. Cally and R. Jairam, and MNHN _RA_2019.0017 (field n��AF3774) an adult male collected by A. Fouquet, S. Cally and R. Jairam on 25 January 2016 at Voltzberg, Suriname (4.68169 N 56.18568 W, ~ 100 m elevation). Etymology. The specific epithet is a noun in apposition and refers to the Saramaka people who live nearby the type locality and are one of the Maroon communities living in Suriname. Definition. (1) Medium-sized Anomaloglossus (average male SVL 19.3 mm [18.6���19.9, n = 10], female SVL 20.3 mm [19.9���21.6, n = 6]) (Table 1); (2) body robust; (3) skin on dorsum finely granular with larger scattered tubercles becoming denser on the posterior half and legs, with a larger tubercle on each eyelid; (4) ventral skin smooth except under thighs where it is finely granular; (5) inconspicuous supratympanic fold; (6) tympanum distinct anteroventrally; (7) snout long and protruding in lateral view; (8) nares oriented ventrolaterally, situated near tip of snout; (9) Finger II smaller than Finger I when fingers adpressed; (10) tip of Finger IV reaching distal edge of distal subarticular tubercle on Finger III when fingers adpressed; (11) distal subarticular tubercle distinct on Finger III and IV, not visible on the other fingers; (12) Finger III swollen dorsally and preaxially in males, extending largely towards dorsal surface of hand forming a cream-coloured gland basally; (13) fringes present on all fingers, particularly developed postaxially on Finger I and prexially on Fingers II and III, in males and females; (14) toes basally webbed, with well-developed fringes on all toes, more developed preaxially on Toes II, III and IV; (15) tarsal keel well-defined, slightly curved; (16) glandular supracarpal pad present in both sexes (larger and more glandular in males); (17) cloacal tubercles present; (18) paracloacal mark present (orangish to light brown in life, cream to grey in preservative); (19) dorsolateral stripe absent; (20) oblique lateral stripe present, solid, undulating (cream but sometimes white anteriorly, yellow at groin level in life, white in preservative), lateral band in life uniformly dark brown/black in males, brown in females, lower flank grey with white speckles, more or less extensively yellowish orange in the axillary and the groin regions; (21) ventrolateral stripe absent; (22) sexual dichromatism in throat colour pattern present, in life light grey in males with sparse minute black melanophores, evenly and entirely yellow in females; (23) sexual dichromatism in ventral colour pattern present, in life abdomen entirely or only posteriorly yellowish orange in males, uniformly bright yellowish orange in females; (24) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (25) median lingual process as long as wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (26) call 0.44���0.69 s in length, consisting of a train of 5���11 notes ranging 0.028���0.036 s in length and spaced by intervals of 0.024���0.030 s with a dominant frequency at 4.55���5.02 kHz (n = 7) (Fig. 4, Table 2); (27) type 4 tadpole (Orton 1953), exotrophic, with a functional mouth with marginal papillae and labial teeth (Fig. 5, Table 3). Morphological comparisons with other Anomaloglossus. The only other species group co-occurring with the Anomaloglossus stepheni group is the A. degranvillei group, represented by A. blanci Fouquet, Vacher, Courtois, Villette, Reizine, Gaucher, Jairam, Ouboter, and Kok, 2018, A. degranvillei (Lescure, 1975), A. dewynteri Fouquet, Vacher, Courtois, Villette, Reizine, Gaucher, Jairam, Ouboter, and Kok, 2018 and A. surinamensis Ouboter and Jairam, 2012. Anomaloglossus saramaka sp. nov. can be distinguished from species of the A. degranvillei group by its basal webbing (moderate in species of the A. degranvillei group), smaller fringes that are more developed on Toes II-IV (well-developed fringes on all toes) and the presence of a solid oblique lateral stripe (absence). Besides these two groups, all the other described Anomaloglossus species exclusively occur in Pantepui (i.e., the Guiana Shield highlands, see Kok et al. 2018) and have moderate to extensive toe webbing (Barrio-Amor��s 2006; Barrio-Amor��s & Brewer-Car��as 2008; Barrio-Amor��s et al. 2004, 2010; Fouquet et al. 2015; Kok et al. 2010; La Marca 1997; Meinhardt & Parmalee 1996; Noble 1923; Rivero 1961) except A. meansi Kok, Nicola��, Lathrop and MacCulloch, 2018, A. kaiei (Kok, Sambhu, Roopsind, Lenglet and Bourne, 2006), A. rufulus (Gorzula, 1990) and A. roraima (La Marca, 1997). However, unlike A. saramaka sp. nov., all these species lack an oblique lateral stripe. Within the Anomaloglossus stepheni group (comparisons follow below), Anomaloglossus saramaka sp. nov. is morphologically most similar to A. baeobatrachus, A. leopardus, A. mitaraka and A. sp. ���Bakhuis��� (see below) and can easily be confused with these species. Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. baeobatrachus by (1) larger body size (mean = 19.3; range 18.6���19.9 mm in males [n = 10] and mean = 20.3; range 19.9���21.6 mm in females [n = 6] in A. saramaka sp. nov. vs mean = 16.2; range 14.8���17.1 mm in males [n = 16] and mean = 18.4; range 17.3���19.4 mm in females [n = 4] in A. baeobatrachus); (2) venter mostly yellowish orange in males in A. saramaka sp. nov. (presence of a cream/white central patch in A. baeobatrachus); (3) call with lower note rate (mean = 14.6, range 14.0���15.3 notes/s in A. saramaka sp. nov. [n = 7] vs mean = 16.2, range 15.6���16.8 notes/s in A. baeobatrachus [n = 9]) (Table 2). Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. leopardus by (1) narrow, poorly defined dark brown bars on legs (broad and conspicuous in A. leopardus); (2) venter mostly yellowish-orange in males in A. saramaka sp. nov. (presence of a more or less extensive cream/white central patch in A. leopardus); (3) shorter advertisement call (mean = 0.51, range 0.38��� 0.69 s in A. saramaka sp. nov. [n = 7] vs mean = 1.52, range 1.08��� 2.00 s in A. leopardus [n = 4]); (4) fewer notes in calls (6���11 in A. saramaka sp. nov. [n = 7] vs 14���28 in A. leopardus [n = 4]) (Table 2). Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. mitaraka by (1) venter mostly yellowish orange in males in A. saramaka sp. nov. (presence of a more or less extensive cream/white central patch in A. mitaraka); (2) shorter advertisement call (mean = 0.51, range 0.38��� 0.69 s in A. saramaka sp. nov. [n = 7] vs mean = 1.04, range 0.74��� 1.23 s in A. mitaraka sp. nov. [n = 6]); (3) higher note rate call (mean = 14.6, range 14.0���15.3 note/s in A. saramaka sp. nov. [n = 7] vs mean = 11.43 note/s, range 10.8���12.3 in A. mitaraka [n = 6]) (Table 2). Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. apiau by (1) solid oblique lateral stripe in A. saramaka sp. nov. (partial, only present as a series of conspicuous white dots extending from groin midway along flank in A. apiau); (2) shorter advertisement call (mean = 0.51, range 0.38��� 0.69 s in A. saramaka sp. nov. [n = 7] vs mean = 19.6, range 7.0��� 39.4 s in A. apiau [n = 10]). Anomaloglossus saramaka sp. nov. can mainly be distinguished from A. stepheni (Figs. 2���3, Table 1) by (1) larger body size (mean = 19.3; range 18.6���19.9 mm in males [n = 10] and mean = 20.3; range 19.9���21.6 mm in females [n = 6] in A. saramaka sp. nov. vs mean = 17.2, range 16.5���18.0 mm in males [n = 10] and mean = 17.3, range 17.0���18.0 mm in females [n = 5] [from Martins 1989] in A. stepheni); (2) skin on dorsum finely granular in A. saramaka sp. nov. (evenly tuberculate in A. stepheni); (3) ventral colouration mostly yellowish orange in males in A. saramaka sp. nov. (uniformly cream or white in A. stepheni); (4) longer advertisement call (mean = 0.51, range 0.38��� 0.69 s in A. saramaka sp. nov. [n = 7] vs mean= 0.25 s, range 0.18��� 0.29 s in A. stepheni [n = 4]); (5) lower note rate call (mean = 14.6, range 14.0���15.3 note/s in A. saramaka sp. nov. [n = 7] vs mean = 21.3, range 20.2���22.6 note/s in A. stepheni [n = 4]) (Table 2). Description of the holotype. An adult male, 18.9 mm SVL; body robust; head wider than long, HL 95 % of HW; HL 33 % of SVL; dorsal skin finely granular, one enlarged tubercle on each eyelid, snout long (SL 52 % of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 42 % of HW; EN 29 % of HL, 75 % of ED. Tympanum distinct anteroventrally; supratympanic fold inconspicuous; choanae small, drop shaped, located anterolaterally. Forelimb slender, skin tuberculate; HAND 24 % of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened; webbing absent on fingers; lateral fringes present on all fingers, particularly developed on postaxial edge of Finger I and on preaxial edges of Finger II and III; Finger III distinctly swollen dorsally and preaxially, swelling largely extending towards dorsal surface of hand; tip of Finger IV reaching distal edge of distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5 times width of digit; width of disc on Finger III 0.6 mm; discs with distinct dorsal scutes; relative lengths of adpressed fingers III> IV> II> I; palmar tubercle large, heart-shaped, 0.8 mm in diameter (larger than disc on Finger III), thenar tubercle elliptic, small (equal to disc on Finger III in maximum diameter), half the size of palmar tubercle, well separated from palmar tubercle; only basal subarticular tubercles on fingers are conspicuous; distal subarticular tubercle of Finger III and IV are almost inconspicuous, basal subarticular tubercles on Fingers I and II the largest, followed by subarticular tubercle and basal subarticular tubercle on Finger III. Hind limb robust, skin tuberculate; TL 48 % of SVL; heels in contact when hind limbs are flexed at right angles to the sagittal plane of body; FL 43 % of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes. Width of disc on Toe IV 0.7 mm. Foot basally webbed; lateral fringes present on all toes, more developed preaxially on Toes II, III and IV. Toe webbing formula I 1+ ��� 1- II 1+ ��� 1- III 1+ ��� 1+ IV 0 ��� 1+ V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and slightly curved at proximal end. Metatarsal fold strong. Colour of holotype in life. Dorsal colour uniformly dark brown. Oblique lateral stripe solid, continuous, undulating, anteriorly brownish white, cream in the middle and yellow near groin (Fig. 2). Black lateral band below oblique lateral stripe, tapering posteriorly, extending from tip of snout to groin and containing the indistinct dorsal part of tympanum. Upper lip brownish white with a few small white speckles. Lower flank (below black lateral band) grey with white speckles with yellowish orange extensions from the axillary and the groin region. Throat grey covered with minute melanophores, more densely laterally; belly anteriorly greyish white, yellowish orange laterally and posteriorly, ventral surfaces of thighs and arms yellowish orange. Iris with coper metallic pigmentation and pupil ring interrupted dorsally and ventrally by transversal pigmentation (Fig. 2). Upper and lower arm reddish brown dorsally with brown ill-defined spots and white speckles. Bright yellow spot at junction of upper arm with body, and anterior part of upper arm with a black band. Black glandular spot at junction between lower arm and hand, followed by a cream coloured part of the swollen gland extending on Finger III. Dorsal surfaces of thigh, shank and tarsus dark brown with ill-defined dark brown, almost black transverse bars, and ill-defined dark brown blotches. Paracloacal marks light brown, elongate. Toes and digits with small light blueish speckles dorsally and laterally. Palms and soles dark brown. Colour of holotype in preservative. After four years in 70 % ethanol, colours of the specimen faded, and the dorsal colouration now varies from pale brown to grey with a brown interorbital bar; yellowish orange ventral colouration faded to white (Fig. 3). Bluish speckles and orange/reddish marks turned white as well. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. Colouration of limbs varies from brown to reddish brown, females being more reddish. Colour of dorsum varies from light brown to dark brown and some specimens (e.g., NZCS-A1206, field n��AF3844) dis-play diffuse dark brown blotches, more frequently on the interorbital region. Colour of oblique lateral stripe varies from white to orangish. Overall dorsal and lateral colouration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as males carrying tadpoles apparently display overall lighter colours, smoother skin and sharper contrasts, whereas calling males are very dark coloured and highly tuberculate. Ventral colouration of female is entirely yellowish orange while the yellowish orange parts are always limited to the posterior region of throat in males. Vocal sac, slits and minute black melanophores on throat only observed in males as well as swelling of Finger III. Advertisement call. Seven collected specimens calling from the leaf litter were recorded from a distance of about 2 m and at temperatures ranging from 22 to 24��C. Descriptive statistics of call parameters are presented in Table 2. Anomaloglossus saramaka sp. nov. emits trains (call length mean = 0.51, range 0.38��� 0.69 s) of 6���11 short notes (note length mean = 0.028 s; range 0.024 ���0.031 s; inter-note interval mean = 0.05 s; range 0.04��� 0.05 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 4.84 kHz on average (range 4.55���5.02 kHz) with a slight upward modulation (ca. 0.3 kHz) (Fig. 4, Table 2). Larval morphology. The following description is based on six tadpoles at stage 27, 28 and 29 (Fig. 5, Table 3). Tadpoles correspond to a type 4 tadpole of Orton (1953); exotrophic; body skin smooth; TL 12.95���17.19 mm; BL 4.33���5.89 mm, 33���36 % of TL, 146���165 % of BW, 191���238 % of BH; BW 123���149 % of BH (Table 3); body ovoid, snout round in dorsal and lateral view; eyes positioned and directed laterally; ED 0.45���0.60 mm, 53���64 % of IOD; IOD larger than IND; nares frontally positioned and directed laterodorsally; narial opening circular in lateral view; END 0.45���0.56 mm. Spiracular tube sinistral, conical, projecting posterodorsally, its tip located at 60���65 % of BL posteriorly to snout. Lateral-line system inconspicuous. Caudal musculature highest at its base, tapering posteriorly, terminating at tail tip; tail tip rounded; upper fin originating at junction of body and tail, gradually increasing in height to about 3/4 of tail; UTF 40���53 % of TMH; LTF 33���47 % of TMH; MTH 13���15 % of TL (Fig. 5, Table 3). Mouth ventral, oral disc strongly emarginated, width 1.40���1.50 mm. Labial teeth long, in single rows, LTRF 2(2)/3. A-2 consisting of two short rows, separated by a large and deep gap; P-1 not interrupted. Marginal papillae long, of equal size on each labium, tapered, blunt-tipped, in a single row, evenly distributed; median gap on upper labium approximately 2/3 the length of A-1; jaw sheaths large, serrated, lower jaw sheath broadly V-shaped. In life, the entire body is dark grey with abundant golden flecks, particularly on dorsum. Golden flecks become scarce ventrally. Posteriorly, dark grey coloration fades and tail becomes translucent. The tadpole of Anomaloglossus saramaka sp. nov. can be distinguished from those of Anomaloglossus of the stepheni group that are endotrophic (A. stepheni and A. baeobatrachus) by the presence of a functional mouth with marginal papillae and labial teeth. However, it virtually cannot be distinguished from the tadpole of A. mitaraka based on morphological evidence. Distribution and natural history. Anomaloglossus saramaka sp. nov. is a diurnal species inhabiting the leaf litter in primary forest at low to mid elevations (from 100 to 500 m elevation). The species is usually found close to streams next to which it deposits its exotrophic tadpoles into puddles, but some specimens were found more than 50 m away from water bodies. Males call all day long when the weather is rainy but calling activity is otherwise limited to sunset and sunrise. Breeding occurs during the rainy season, from December to July. The rainfall between December and April being highly variable, the reproduction seems to be concentrated in the second half (May-July) of this period. The males respond to intraspecific playbacks with shorter and more rapidly emitted note trills. Males are territorial, their small territories being spaced at least two meters apart., Published as part of Fouquet, Antoine, Jairam, Rawien, Ouboter, Paul & Kok, Philippe J. R., 2020, Two new species of Anomaloglossus (Anura: Aromobatidae) of the stepheni group from Suriname, pp. 147-164 in Zootaxa 4820 (1) on pages 150-153, DOI: 10.11646/zootaxa.4820.1.7, http://zenodo.org/record/4397441, {"references":["Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262. https: // doi. org / 10.1206 / 0003 - 0090 (2006) 299 [1: PSODFA] 2.0. CO; 2","Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmell, N. J. (2007) Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses. PLoS One, 2, e 1109. https: // doi. org / 10.1371 / journal. pone. 0001109","Fouquet, A., Noonan, B. P., Rodrigues, M. T., Pech, N., Gilles, A. & Gemmell, N. J. (2012) Multiple quaternary refugia in the eastern Guiana Shield revealed by comparative phylogeography of 12 frog species. Systematic Biology, 61, 461 - 489. https: // doi. org / 10.1093 / sysbio / syr 130","Ouboter, P. E. & Jairam, R. (2012) Amphibians of Suriname. Brill, Leiden, 376 pp. https: // doi. org / 10.1163 / 9789004208001","Vacher, J. - P., Kok, P. J. R., Rodrigues, M. T., Dias Lima, J., Lorenzini, A., Martinez, Q., Fallet, M., Courtois, E. A., Blanc, M., Gaucher, P., Dewynter, M., Jairam, R., Ouboter, P., Thebaud, C. & Fouquet, A. (2017) Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield. Molecular Phylogenetics and Evolution, 112, 158 - 173. https: // doi. org / 10.1016 / j. ympev. 2017.04.017","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2, 63 - 75. https: // doi. org / 10.2307 / 2411661","Fouquet, A., Vacher, J. - P., Courtois, E. A., Villette, B., Reizine, H., Gaucher, P., Jairam, R., Ouboter, P. & Kok, P. J. R. (2018) On the brink of extinction: two new species of Anomaloglossus from French Guiana and amended definition of Anomaloglossus degranvillei and A. surinamensis (Anura: Aromobatidae). Zootaxa, 4379 (1), 001 - 023. https: // doi. org / 10.11646 / zootaxa. 4379.1.1","Lescure, J. (1975) Contribution l'etude des amphibiens de Guyane Francaise. III. Une nouvelle espece de Colostethus (Dendrobatidae): Colostethus degranvillei n. sp. Bulletin du Museum National d'Histoire Naturelle. Paris, Serie 3, Zoologie, 293, 413 - 420.","Kok, P. J. R., Nicolai ¨, M. P. J., Lathrop, A. & MacCulloch, R. D. (2018) Anomaloglossus meansi sp. n., a new Pantepui species of the Anomaloglossus beebei group (Anura, Aromobatidae). ZooKeys, 759, 99 - 116. https: // doi. org / 10.3897 / zookeys. 759.24742","Barrio-Amoros, C. L. (2006) A new dendrobatid frog (Anura: Dendrobatidae: Colostethus) from Aprada tepui, southern Venezuela. Zootaxa, 1110 (1), 59 - 68. https: // doi. org / 10.11646 / zootaxa. 1110.1.6","Barrio-Amoros, C. L. & Brewer-Carias, C. (2008) Herpetological results of the 2002 expedition to Sarisarinama, a tepui in Venezuelan Guayana, with the description of five new species. Zootaxa, 1942 (1), 1 - 68. https: // doi. org / 10.11646 / zootaxa. 1942.1.1","Barrio-Amoros, C. L., Fuentes-Ramos, O. & Rivas-Fuenmayor, G. A. (2004) Two new species of Colostethus (Anura: Dendrobatidae) from the Venezuelan Guayana. Salamandra, 40, 183 - 200.","Barrio-Amoros, C. L., Santos, J. C. & Jovanovic, O. (2010) A new dendrobatid frog (Anura: Dendrobatidae: Anomaloglossus) from the Orinoquian rainforest, southern Venezuela. Zootaxa, 2413 (1), 37 - 50. https: // doi. org / 10.11646 / zootaxa. 2413.1.3","Fouquet, A., Souza, S. M., Nunes, P. M. S., Kok, P. J. R. Curcio, F. F., de Carvalho, C. M., Grant, T. & Rodrigues, M. T. (2015) Two new endangered species of Anomaloglossus (Anura: Aromobatidae) from Roraima State, northern Brazil. Zootaxa, 3926 (2), 191 - 210. https: // doi. org / 10.11646 / zootaxa. 3926.2.2","Kok, P. J. R., MacCulloch, R. D., Lathrop, A., Willaert, B. & Bossuyt, F. (2010) A new species of Anomaloglossus (Anura: Aromobatidae) from the Pakaraima Mountains of Guyana. Zootaxa, 2660, 18 - 32. https: // doi. org / 10.11646 / zootaxa. 2660.1.2","La Marca, E. (1997 1996) Ranas del genero Colostethus (Amphibia: Anura: Dendrobatidae) de la Guayana Venezolana con la descripcion de siete especies nuevas. Publicaciones de la Asociacioin de Amigos de Donana, 9, 1 - 64.","Noble, G. K. (1923) New batrachians from the Tropical Research Station British Guiana. Zoologica, 3, 289 - 299.","Rivero, J. A. (1961) Salientia of Venezuela. Bulletin of the Museum of Comparative Zoology. Cambridge, Massachusetts, 126, 1 - 207.","Kok, P. J. R., Sambhu, H., Roopsind, I., Lenglet, G. L. & Bourne, G. R. (2006) A new species of Colostethus (Anura: Dendrobatidae) with maternal care from Kaieteur National Park, Guyana. Zootaxa, 1238 (1), 35 - 61. https: // doi. org / 10.11646 / zootaxa. 1238.1.3","Gorzula, S. J. (1990 \" 1988 \") Una nueva especie de Dendrobates (Amphibia: Dendrobatidae) del Macizo del Chimantai, Estado Boliivar, Venezuela. Memoria Sociedad de Ciencias Naturales La Salle, 48, 143 - 149.","Martins, M. (1989) Nova especie de Colostethus da Amazonia central (Amphibia: Dendrobatidae). Revista Brasileira de Biologia, 49, 1009 - 1012.","Dezecache, C., Faure, E., Gond, V., Salles, J. M., Vieilledent, G. & Herault, B. (2017) Gold-rush in a forested El Dorado: deforestation leakages and the need for regional cooperation. Environmental Research Letters, 12, 034013. https: // doi. org / 10.1088 / 1748 - 9326 / aa 6082"]}

Anomaloglossus vacheri sp. nov. Anomaloglossus baeobatrachus Ouboter & Jairam 2012 Anomaloglossus sp. Bakhuis Vacher et al. 2017 Holotype. MNHN _RA_2019.0009 (field n°AF3413), an adult male, collected by A. Fouquet, S. Cally and R. Jairam on 30 April 2015 at Bakhuis Mountains, Suriname (4.72462 N 56.7638 W, ~ 200 m elevation; Figs. 2‒3; Ap-pendix 1). Paratopotypes. Eight specimens: MNHN _RA_2019.0009‒11, 13‒15 (field n°AF3412, 3424, 3426, 3430, 3434) and NZCS-A1208‒9 (field n°AF3433, 3441) seven adult males and MNHN _RA_2019.0012 (field n° AF3425) one female collected with the holotype by A. Fouquet, S. Cally and R. Jairam. Etymology. This species is dedicated to Jean-Pierre Vacher, in honour of his strong contribution to the understanding of the diversity and the evolution of the genus Anomaloglossus. Definition. (1) Small-sized Anomaloglossus (average male SVL 17.4 mm [17.0–17.8, n = 8], female SVL 19.7 mm [n = 1]) (Table 1); (2) body robust; (3) skin on dorsum finely granular with larger scattered tubercles becoming denser on the posterior half and legs, with a larger tubercle on each eyelid; (4) ventral skin smooth except under thighs, where it is finely granular; (5) inconspicuous supratympanic fold; (6) tympanum distinct anteroventrally; (7) snout long and protruding in lateral view; (8) nares oriented ventrolaterally, situated near tip of snout; (9) Finger II equal to Finger I when fingers adpressed; (10) tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; (11) distal subarticular tubercle distinct on Finger III and IV, inconspicuous on the other fingers; (12) Finger III swollen dorsally and preaxially in males, extending largely towards dorsal surface of hand; (13) fringes present on all fingers, particularly developed post- and preaxially on Finger II and preaxially on Finger III, almost inconspicuous on Finger IV, in males and females; (14) toes basally webbed, with well-developed fringes on all toes, more developed preaxially on Toes II and III; (15) tarsal keel well-defined, strongly curved; (16) glandular supracarpal pad present in both sexes (larger and more glandular in males); (17) cloacal tubercles present; (18) paracloacal mark present (orangish in life, cream to grey in preservative); (19) dorsolateral stripe absent; (20) oblique lateral stripe present, diffuse (composed of yellow and white speckles), particularly diffuse anteriorly, extending along upper eyelid into a diffuse canthal stripe, lateral band uniformly dark brown/black in males and females in life, lower flank yellowish orange to grey with white speckles; (21) ventrolateral stripe absent; (22) dorsal surface of arms barely blotched in males, orange; (23) sexual dichromatism in throat colour pattern present, in life light grey in males with sparse minute black melanophores (chin often yellow), evenly and entirely yellow in females; (24) sexual dichromatism in ventral colour pattern present, in life abdomen entirely yellowish orange in males, uniformly yellowish orange in females; (25) iris with metallic pigmentation and pupil ring inconspicuously interrupted ventrally by transversal black pigmentation; (26) median lingual process as long as wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (27) call 0.47–0.91 s in length, consisting of a train of 8–18 notes of 0.019–0.021 s in length and spaced by intervals of 0.042 s with a dominant frequency at 5.11–5.35 kHz (n = 5; Table 2); (28) type 4 tadpole (Orton 1953), exotrophic, with a functional mouth with marginal papillae and labial teeth (Fig. 5, Table 3). Morphological comparisons with other Anomaloglossus. Like Anomaloglossus saramaka sp. nov., A. vacheri sp. nov. can be distinguished from species of the A. degranvillei group by its basal webbing (moderate in species of the A. degranvillei group), smaller fringes that are more developed on Toes II-IV (well-developed fringes on all toes) and the presence of a solid oblique lateral stripe (absence) and from all the other described Anomaloglossus species of the Pantepui region by the presence of an oblique lateral stripe. Within the Anomaloglossus stepheni group (comparisons follow below), A. vacheri sp. nov. is morphologically most similar to A. baeobatrachus, A. leopardus, A. mitaraka and A. saramaka sp. nov. and can easily be confused with these species. Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. baeobatrachus by (1) oblique lateral stripe extending along upper eyelid into a canthal stripe in A. vacheri sp. nov. (oblique lateral stripe not extending along upper eyelid into a canthal stripe in A. baeobatrachus); (2) dorsal surface of arms barely blotched and orange in males of A. vacheri sp. nov. (with dark brown blotches and light brown in A. baeobatrachus); (3) ventral coloration yellowish orange in males in A. vacheri sp. nov. (white or only posteriorly yellow in A. baeobatrachus); (4) call with higher note rate (mean = 18.2, range 17.3–19.2 note/s in A. vacheri sp. nov. [n = 8] vs mean = 16.2, range 15.6–16.8 note/s in A. baeobatrachus [n = 9]) (Table 2). Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. leopardus by (1) narrow, poorly defined dark brown bars on legs (broad and conspicuous dark brown in A. leopardus); (2) dorsal surface of arms barely blotched and orange in males of A. vacheri sp. nov. (light brown with dark brown blotches in A. leopardus); (3) oblique lateral stripe extending along upper eyelid into a canthal stripe in A. vacheri sp. nov. (not extending along upper eyelid into a canthal stripe in A. leopardus); (4) advertisement call shorter (mean = 0.63, range 0.47– 0.91 s in A. vacheri sp. nov. [n = 5] vs mean = 1.52, range 1.08– 2.00 s in A. leopardus [n = 4]) and higher in frequency (mean = 5.25, range 5.11–5.35 kHz in A. vacheri sp. nov. [n = 5] vs mean = 4.45, range 4.40–4.57 kHz in A. leopardus [n = 4]) (Table 2). Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. mitaraka by (1) smaller male body size (mean = 17.4; range 17.0– 17.8 mm in males [n = 8] in A. vacheri sp. nov. vs mean = 18.6; range 18.2–19.3 mm in males [n = 7] in A. mitaraka); (2) dorsal surface of arms barely blotched and orange in males of A. vacheri sp. nov. (with dark blotches and light brown in A. mitaraka); (3) oblique lateral stripe extending along upper eyelid into a canthal stripe A. vacheri sp. nov. (not extending along upper eyelid into a canthal stripe in A. mitaraka); (4) advertisement with a higher note rate (mean = 18.2, range 17.3–19.1 note/s in A. vacheri sp. nov. [n = 8] vs mean = 11.4 note/s, range 10.8–12.3 in A. mitaraka [n = 6]) and higher in frequency (mean = 5.25, range 5.11–5.35 kHz in A. vacheri sp. nov. [n = 5] vs mean = 4.43, range 4.12–4.76 kHz in A. mitaraka [n = 6]) (Table 2). Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. saramaka sp. nov. by (1) smaller male body size (mean = 17.4; range 17.0– 17.8 mm in males [n = 8] in A. vacheri sp. nov. vs mean = 19.3; range 18.6–19.9 mm in males [n = 10] in A. saramaka sp. nov.); (2) dorsal surface of arms barely blotched and orange in males of A. vacheri (with dark blotches and light brown in A. saramaka sp. nov.); (3) oblique lateral stripe extending along upper eyelid into a canthal stripe in A. vacheri sp. nov. (not extending along upper eyelid into a canthal stripe in A. saramaka sp. nov.); (4) advertisement with a higher note rate (mean = 18.2, range 17.3–19.2 note/s in A. vacheri sp. nov. [n = 8] vs mean = 14.5 note/s, range 14.0– 15.3 in A. saramaka sp. nov. [n = 6]) and higher in frequency (mean = 5.25, range 5.11–5.35 kHz in A. vacheri sp. nov. [n = 5] vs mean = 4.84, range 4.55–5.02 kHz in A. saramaka sp. nov. [n = 6]) (Table 2). Anomaloglossus vacheri sp. nov. can be mainly distinguished from A. apiau by (1) solid oblique lateral stripe in A. vacheri sp. nov. (present as a series of conspicuous white dots extending from groin midway along flank in A. apiau); (2) by its shorter call (mean = 0.63, range 0.47– 0.91 s in A. vacheri sp. nov. [n = 5] vs mean = 19.6, range 7.0– 39.4 s in A. apiau [n = 10]). Anomaloglossus vacheri sp. nov. can mainly be distinguished from A. stepheni (Figs. 2‒3) by (1) skin on dorsum finely granular in A. vacheri sp. nov. vs evenly tuberculate in A. stepheni; (2) ventral colouration mostly yellowish orange in males in A. vacheri sp. nov. vs uniformly white in A. stepheni; (3) oblique lateral stripe extending along upper eyelid into a canthal stripe in A. vacheri sp. nov. vs oblique lateral stripe not extending along upper eyelid into a canthal stripe in A. stepheni; (4) advertisement call longer (mean = 0.63, range 0.47– 0.91 s in A. vacheri sp. nov. [n = 5] vs mean= 0.25 s, range 0.18– 0.29 s in A. stepheni [n = 4]), and higher in frequency (mean = 5.25, range 5.11–5.35 kHz in A. vacheri sp. nov. [n = 5] vs mean = 4.48, range 4.25–4.85 kHz in A. stepheni [n = 4]) (Table 2). Description of the holotype. An adult male, 17.5 mm SVL; body robust; head wider than long, HL 98 % of HW; HL 33 % of SVL; dorsal skin finely tuberculate with larger tubercles posteriorly, one enlarged tubercle on each eyelid, snout long (SL 51 % of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 41 % of HW; EN 30 % of HL, 77 % of ED. Tympanum distinct anteroventrally; supratympanic fold inconspicuous; choanae small, circular, located anterolaterally. Forelimb slender, skin tuberculate; HAND 23 % of SVL; Finger I as long as Finger II when fingers adpressed; fingers large and flattened; webbing absent on fingers; lateral fringes present, particularly developed post and preaxially on Finger II and preaxially on Finger III; Finger III distinctly swollen dorsally and preaxially, swelling largely extending towards dorsal surface of hand; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5 times width of digit; width of disc on Finger III 0.6 mm; discs with distinct dorsal scutes; relative lengths of adpressed fingers III> IV> II> I; palmar tubercle large, heart-shaped, 0.75 mm in diameter (larger than disc on Finger III), thenar tubercle elliptic, small (equal to disc on Finger III in maximum diameter), half the size of palmar tubercle, well separated from palmar tubercle; basal subarticular tubercles on fingers and distal subarticular tubercle of Finger III and IV conspicuous; subarticular tubercles on Fingers I and II the largest, followed by subarticular tubercles and basal subarticular tubercle on Finger III. Hind limb robust, skin tuberculate; TL 48 % of SVL; heels in contact when hind limbs are flexed at right angles to the sagittal plane of body; FL 42 % of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes. Width of disc on Toe IV 0.8 mm. Foot basally webbed; lateral fringes present on all toes, more developed preaxially on Toes II and III. Toe webbing formula I 1+ ‒ 1- II 1+ ‒ 1- III 1+ ‒ 1+ IV 0 ‒ 1+ V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end. Metatarsal fold strong. Colour of holotype in life. Dorsal colour light brown with dark brown blotches occurring on the interorbital region and at mid-body forming an hourglass pattern. Blotches of the same colour also occur laterally along the oblique lateral stripe. Oblique lateral stripe thin, diffuse (particularly anteriorly), formed by white and yellow speckles, extending along the upper eyelid into a diffuse canthal stripe (Fig. 2). Dark brown lateral band below oblique lateral stripe, tapering posteriorly, extending from tip of snout to groin and containing the indistinct dorsal part of tympanum, tapering posteriorly. Upper lip grey with a few small white speckles. Lower flank (below dark brown lateral band) grey with white speckles. Throat light grey, chin yellowish orange, throat and chin covered with minute melanophores, more densely laterally; belly bright yellowish orange, ventral surfaces of thighs and arms yellowish orange. Iris with coper metallic pigmentation and pupil ring uninterrupted by transversal pigmentation (Fig. 2). Upper and lower arm light orange dorsally with a few white speckles. Black glandular spot at junction between lower arm and hand, followed by a grey coloured part of the swollen gland extending on Finger III. Dorsal surfaces of thigh, shank and tarsus light brown with ill-defined dark brown transverse bars and ill-defined dark brown blotches. Paracloacal marks light brown, elongate. Toes and digits with small light blueish speckles dorsally and laterally. Palms and soles dark brown. Colour of holotype in preservative. After four years in 70 % ethanol, colours of the specimen faded and the dorsal colouration now varies from pale brown to grey with darker blotches, the yellowish orange ventral colouration faded to white (Fig. 3). Bluish speckles and orange marks turned white as well. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. Colour of oblique lateral stripe varies in its proportion of white and yellow. Overall dorsal and lateral coloration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as calling males are dark coloured and highly tuberculate. Dorsal dark brown hourglass pattern may be absent (MNHN _RA_2019.0013, field n° AF3426; NZCS-A1208, field n°AF3433). Paracloacal marks sometimes inconspicuous. Throat colouration of female is entirely yellowish orange while the yellowish orange parts are often limited to the chin in males (extent of coloured part of chin varies). Vocal sac, slits and minute black melanophores on throat only observed in males as well as swelling of Finger III. Advertisement call. Five collected specimens calling from the leaf litter were recorded from a distance of about 2 m and at temperatures between 25‒26°C and 98% relative humidity. Descriptive statistics of call parameters are presented in Table 2. Anomaloglossus vacheri sp. nov. emits trains (call length mean = 0.63, range 0.47– 0.91 s) of 8–18 short notes (note length mean = 0.020 s; range 0.019 –0.021 s; inter-note interval mean = 0.039 s; range 0.036 –0.042 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 5.25 kHz on average (range 5.11–5.35 kHz) with a slight upward modulation (ca. 0.4 kHz) (Fig. 4, Table 2). Larval morphology. The following description is based on two tadpoles at stage 28 and 30 (Fig. 5). Tadpoles correspond to a type 4 tadpole of Orton (1953); exotrophic; body skin smooth; TL 15.00– 19.06 mm; BL 5.27–6.46 mm, 34–35 % of TL, 151–160 % of BW, 261–267 % of BH; BW 135–144 % of BH (Table 3); body ovoid, snout round in dorsal and lateral view; eyes positioned and directed laterally; ED 0.49–0.61 mm, 55–58 % of IOD; IOD smaller than IND; nares frontally positioned and directed laterodorsally; narial opening circular in lateral view; END 0.55–0.71 mm. Spiracular tube sinistral, conical, projecting posterodorsally, its tip located at 57–59 % of BL posteriorly to snout. Lateral-line system inconspicuous. Caudal musculature highest at its base, tapering posteriorly, terminating at tail tip; tail tip rounded; upper fin originating at junction of body and tail, gradually increasing in height to about 3/4 of tail; UTF 47–48 % of TMH; LTF 46–49 % of TMH; MTH 13–16 % of TL (Fig. 5, Table 3). Mouth ventral, oral disc strongly emarginated, width 1.27–1.89 mm. Labial teeth long, in single rows, LTRF 2(2)/3. A-2 consisting of two short rows, separated by a large and deep gap; P-1 not interrupted. Marginal papillae long, of equal size on each labium, tapered, blunt-tipped, in a single row, evenly distributed; median gap on upper labium approximately 2/3 the length of A-1; jaw sheaths large, serrated, lower jaw sheath broadly V-shaped. In life, the body is dark grey with abundant golden flecks, particularly on dorsum, except the loreal and postocular regions that are pink. Golden flecks become scarce ventrally. Posteriorly, dark grey coloration fades and tail becomes translucent. The tadpole of Anomaloglossus vacheri sp. nov. can be distinguished from those of Anomaloglossus of the stepheni group that are endotrophic (A. stepheni and A. baeobatrachus) by the presence of a functional mouth with marginal papillae and labial teeth. It can be distinguished from the tadpole of A. mitaraka and A. saramaka sp. nov. by its less dense pigmentation particularly ventrally and by a spiracle located more anteriorly. Distribution and natural history. Anomaloglossus vacheri sp. nov. is a diurnal species inhabiting the leaf litter in primary forest at low elevation (200 m elevaion). The species has only been found in terra firme forest on lateritic crust distant from streams. Males call after rainfall. Breeding apparently occurs during the rainy season, probably extending between December and July, depending on rainfall. The males respond to intraspecific playbacks with shorter and more rapidly emitted note trills. Males are spaced by at least five meters apart. They usually call slightly above the leaf litter, exposed on a branch or a dead leaf. Eggs are probably deposited under or in the fold of a dead leaf. After they hatch the male probably carries the tadpoles to small water bodies formed in the lateritic crust where tadpoles where found (Fig. 5). A single population is known to date, in Bakhuis Mountains, Suriname. The species likely occurs throughout the massif, but it unlikely extends much furthe

Anomaloglossus mitaraka sp. nov. Anomaloglossus sp. “Mitaraka” Vacher et al. 2017 Holotype. MNHN2018.64 (field n° AF2814), an adult male, collected by A. Fouquet and M. Dewynter, 24 February 2015, Mitaraka, French Guiana, 2.23577° N 54.44928° W, ~ 150 m elevation (Figs. 2–3). Paratopotypes. Nine specimens: MNHN 2018.65–70 (field n° AF 2732, 2750-1, 2754, 2808, 2878) six adult males and MNHN 2018.71–73 (field n° AF 2724, 2731, 2824) three females collected with the holotype by A. Fouquet and M. Dewynter. Etymology. The specific epithet is a noun in apposition and refers to the type locality (Mitaraka, French Guiana). Definition. (1) Medium-sized Anomaloglossus (average male SVL 18.6 mm [18.2–19.3, n = 7], female SVL 20.1 mm [19.2–21.0, n = 3]) (Table 1); (2) body robust; (3) skin on dorsum with irregularly scattered tubercles becoming denser on the posterior half and legs, with a larger tubercle on each eyelid, ventral skin smooth; (4) inconspicuous supratympanic fold; (5) tympanum distinct anteroventrally; (6) snout short and protruding in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II equal to Finger I when fingers adpressed; (9) tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle distinct on Finger III, absent on the other fingers; (11) Finger III swollen dorsally and preaxially, extending largely towards dorsal surface of hand in males; (12) fringes present on all fingers particularly developed preaxially on Finger II in males and females; (13) toes basally webbed, with well-developed fringes on all toes, more developed preaxially on Toe II, and pre- and postaxially on toes III and IV (sensu Grant et al. 2006; keel-like lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved; (15) black arm gland at the junction with wrist in males (sensu Grant & Castro 1998, see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark present (orangish in life, cream in preservative); (18) pale dorsolateral stripe present, solid (white to orangish anteriorly, yellow posteriorly in life, white in preservative), flanks and dorsum uniformly dark brown in males, reddish brown with dark brown blotches in females in life; (19) ventrolateral stripe absent; (20) sexual dichromatism in throat color pattern present in life, sometimes anteriorly yellow in males with sparse black melanophores, evenly and entirely yellow in females; (21) sexual dichromatism in ventral color pattern present, abdomen mostly cream, posteriorly and laterally yellow in males, uniformly bright yellow in females in life; (22) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (23) median lingual process as long as wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (24) a 0.74– 1.23 s length call consisting of a train of 8–16 notes of 0.028– 0.036 s in length and spaced by intervals of 0.051– 0.068 s and of dominant frequency at 4.13–4.76 kHz (n = 6); (25) tadpole of type 4 (Orton 1953), exotrophic, with a functional mouth with marginal papillae and labial teeth (Fig. 4; Table 2). Morphological comparisons with other Anomaloglossus. The only other species group co-occurring with the Anomaloglossus stepheni group is the A. degranvillei group, represented by A. blanci, A. degranvillei, A. dewynteri and A. surinamensis, which are readily distinguishable by their moderate webbing (basal in A. mitaraka) and well-developed fringes on all toes (smaller and more developed on Toe II, III, IV) and the absence of a dorsolateral stripe (present). Outside these two groups all the other described Anomaloglossus species exclusively occur in Pantepui (i.e., the Guiana Shield highlands, see Kok et al. 2018) and have moderate to extensive toe webbing except A. meansi Kok, Nicolaï, Lathrop & MacCulloch, 2018, A. kaiei (Kok, Sambhu, Roopsind, Lenglet & Bourne, 2006), A. rufulus (Gorzula, 1990) and A. roraima (La Marca, 1997). However, unlike A. mitaraka, A. meansi lacks a conspicuous dorsolateral stripe, Finger III in males of A. kaei is not distinctly swollen, A. rufulus has as an extensively spotted ventral face, and A. roraima lacks well-developed fringes. Within the Anomaloglossus stepheni group, A. mitaraka can be distinguished from its probable sister species (Fouquet et al. 2019) A. leopardus (Figs. 2–3) by (1) narrower dark brown bars on legs vs large and more conspicuous dark transverse bars on legs in A. leopardus; (3) ventral coloration with orange restricted to the posterior part in males in A. mitaraka vs. generally entirely yellow in A. leopardus; (4) dorsolateral line thin in A. mitaraka vs. thick and well defined in A. leopardus; (5) advertisement call with a slower note rate (mean = 11.43, range 10.8–12.3 note/s in A. mitaraka [n = 6] vs. mean = 13.4, range 12.9–13.6 notes/s in A. leopardus [n = 4]) (Table 2). Anomaloglossus mitaraka can be distinguished from A. stepheni (Figs. 2–3, Table 1) by (1) a larger body size (mean = 18.6; range 18.2–19.3 mm in males [n = 7] and mean = 20.1; range 19.2–21.0 mm in females [n = 3] in A. mitaraka vs. mean = 17.15, range 16.5–18.0 mm in males [n = 10] and mean = 17.3, range 17.0–18.0 mm in females [n = 5] [from Martins 1989] in A. stepheni); (2) skin of dorsum with scattered tubercules in A. mitaraka vs. evenly tuberculate in A. stepheni; (3) orange ventral coloration posteriorly and laterally in males of A. mitaraka vs. entirely translucent white in A. stepheni; (4) dorsolateral line thin but well defined and continuous in A. mitaraka vs. thick and ill-defined dorsally in A. stepheni; (5) advertisement call much longer (mean = 1.04 s, range 0.74– 1.23 s in A. mitaraka [n = 6] vs. mean= 0.25 s, range 0.18– 0.29 s in A. stepheni [n = 4]), emitted at a slower pace (call rate mean = 0.43, range 0.32–0.52 calls/s in A. mitaraka [n = 6] vs. mean = 1.40, range 1.10–1.89 calls/s in A. stepheni [n = 4]) (Table 2). Anomaloglossus mitaraka can be distinguished from A. baeobatrachus by (1) a larger body size (mean = 18.6; range 18.2–19.3 mm in males [n = 7] and mean = 20.1; range 19.2–21.0 mm in females [n = 3] in A. mitaraka vs. mean = 16.2; range 14.8–17.1 mm in males [n = 16] and mean = 18.4; range 17.3–19.4 mm in females [n = 4] in A. baeobatrachus); (2) ventral coloration posteriorly and laterally in males in A. mitaraka vs. yellow and less extensive in A. baeobatrachus; (3) dorsolateral line thin but well defined and solid in A. mitaraka vs. narrower and interrupted in A. baeobatrachus; (4) call with lower note rate (mean = 11.43 notes/s, range 10.83–12.28 in A. mitaraka [n = 6] vs. mean = 16.10, range 15.62– 16.85 s in A. baeobatrachus [n = 9]) lower dominant frequency (mean= 4.44, range 4.13–4.76 kHz in A. mitaraka [n = 6] vs. mean = 5.39, range 4.96–5.59 kHz in A. baeobatrachus [n = 9]) (Table 2). Description of the holotype. An adult male, 18.6 mm SVL; body robust; head wider than long, HL 94 % of HW; HL 33 % of SVL; dorsal skin tuberculate, one enlarged tubercle on each eyelid, snout long (SL 52 % of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 39 % of HW; EN 29 % of HL, 75 % of ED. Tympanum distinct anteroventrally; supratympanic fold inconspicuous; choanae small, circular, located anterolaterally (Figs. 2–3). Forelimb slender, skin tuberculate; metacarpal ridge present; HAND 24 % of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened; webbings absent on fingers; lateral fringes present on preaxial edges of Finger II; Finger III distinctly swollen dorsally and preaxially; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5 times width of digit; width of disc on Finger III 0.6 mm; discs with distinct dorsal scutes. Relative lengths of adpressed fingers III> IV> I> II; palmar tubercle large, heart-shaped, 0.8 mm in diameter (larger than Finger’s III disc), thenar tubercle elliptic, small (equal to Finger III’s disc in maximum diameter), elliptic, half the size of palmar tubercle, well separated from palmar tubercle. Only basal subarticular tubercles on Fingers are conspicuous; subarticular tubercles on fingers I and II the largest, followed by Finger IV’s subarticular tubercles and basal subarticular tubercle on Finger III. Hind limb robust, skin tuberculate; TL 47 % of SVL; heels in contact when hind limbs are flexed at right angles to the sagittal plane of body; FL 42 % of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes. Width of disc on Toe IV 0.7 mm. Foot basally webbed; lateral fringes present on all toes. Toe webbing formula I 1 + –1 - II 1 + –1 - III 1 + –1 + IV 0–1 + V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end. Metatarsal fold strong. Color of holotype in life. Dorsal color uniformly dark brown. Dorsolateral stripe thin, continuous, bluish white (yellow near groin) (Fig. 2). Dark brown lateral band extending from tip of snout to the groin and containing the indistinct dorsal part of tympanum, tapering posteriorly. Upper lip with small white blotches. Lower flanks pale white anteriorly and orange posteriorly with small white blotches. Throat white covered with melanophores, more densely anterolaterally; belly anteriorly white, white laterally, orange posteriorly, ventral surfaces of thighs and arms orange. Iris with coper metallic pigmentation and pupil ring interrupted dorsally and ventrally by transversal pigmentation (Fig. 2). Upper and lower arm pale red dorsally, light brown with small dark brown spots elsewhere. Dorsal surfaces of thigh, shank and tarsus brown with dark brown transverse bars and ill-defined blotches. Paracloacal marks orangish, elongated anteroposteriorly. Toes and digits with small light blue dots dorsally and laterally. Palms and soles dark brown. Color of holotype in preservative. After three years in 70 % ethanol, some colors of the specimen faded and the dorsal coloration now varies from brown to grey with interorbital, mid-dorsal and sacral dark brown blotches and the orangish ventral coloration disappeared (Fig. 3). Bluish freckles and orange and reddish marks turned cream. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. Coloration of limbs varies from brown to reddish brown, females being paler and with more conspicuous dark dorsal blotches. Colour of dorsolateral line varies from white to orangish. Overall dorsal and lateral coloration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as males carrying tadpoles apparently display overall lighter colors, smoother skin and sharper contrasts, whereas calling males are very dark and highly tuberculate. Ventral coloration of female is entirely orangish while the orangish parts are limited in various extent to the posterior region and throat in males. Vocal sac, slits and small dark melanophores on throat only observed in males as well as swelling on the Finger III. Advertisement call. Six specimens (one uncollected) calling from the leaf litter were recorded from a distance of about 1 m and at temperatures ranging from 23 to 26°C. Descriptive statistics of call parameters are presented in Table 2. Anomaloglossus mitaraka emits trains (call length mean = 1.04 s; range 0.74– 1.23 s) of short notes (note length mean = 0.030 s; range 0.028– 0.036 s; inter-note interval mean = 0.060 s; range 0.051– 0.068 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 4.44 kHz on average (range 4.13–4.76 kHz) with a slight upward modulation (ca. 0.2 kHz) (Fig. 4, Table 2). Larval morphology. The following description is based on three tadpoles at stage 26, 28 and 30 (Fig. 6). Tadpoles correspond to a Type 4 tadpole of Orton (1953); exotrophic; body skin smooth; TL 12.21–19.43 mm; BL 4.41–7.14 mm, 35–37 % of TL, 164–181 % of BW, 221–256 % of BH; BW 131–143 % of BH (Table 3); body ovoid, snout round in dorsal and lateral view; eyes positioned and directed laterally; ED 0.48–0.63 mm, 45–98 % of IOD; IOD equal to IND; nares frontally positioned and directed laterodorsally; narial opening reniform in lateral view; END 0.34–0.73 mm. Spiracular tube sinistral, conical, projecting posterodorsally, its tip located at 60–63 % of BL posteriorly to snout. Lateral-line system inconspicuous. Caudal musculature highest at its base, tapering posteriorly, terminating at tail tip; tail tip rounded; upper fin originating at junction of body and tail, gradually increasing in height to about 3/4 of tail; UTF 28–53 % of TMH; LTF 38–55 % of TMH; MTH 14–17 % of TL (Fig. 6, Table 3). Mouth ventral, oral disc strongly emarginated, width 1.90 mm. Labial teeth long, in single rows, LTRF 2(2)/3. A-2 consisting of two short rows, separated by a large and deep gap; P-1 not interrupted. Marginal papillae long, of equal size on each labium, tapered, blunt-tipped, in a single row, evenly distributed; median gap on upper labium approximately 2/3 the length of A-1; jaw sheaths large, serrated, lower jaw sheath broadly V-shaped. In life, the entire body is dark grey with abundant golden flecks, particularly on dorsum. Golden flecks become scarce ventrally. Posteriorly, dark grey coloration fades and tail becomes translucent. Tadpoles of Anomaloglossus mitaraka can be distinguished from those of any other described Anomaloglossus of the stepheni group (A. stepheni and A. baeobatrachus - this study) by the presence of a functional mouth with marginal papillae and labial teeth. Distribution and natural history. Anomaloglossus mitaraka is a diurnal species inhabiting the leaf litter in primary forest at low to mid elevations (from 150 to 500 m a.s.l.). The species is usually found close to streams next to which it deposits its exotrophic tadpoles into puddles, but some specimens can be found more than 50 m from water bodies. Males call all day long when the weather is rainy. Breeding occurs during the rainy season, between January and May. The males respond to intraspecific playbacks with shorter and more rapidly emitted note trills. Males are territorial, their small territories being spaced at least a few meters apart. Males usually call slightly above the leaf litter, exposed on a branch or a dead leaf. Eggs are deposited in the fold of a dead leaf (a single observation). This clutch was raised (Fig. 6) and some tadpoles were genotyped. Populations have been documented in southwestern French Guiana and adjacent Suriname (Mitaraka massif, Pic Coudreau, Haute Marwini, upper Tapanahony; Fig. 1). It also likely occurs in adjacent Brazil and may occupy similar habitats in other regions of Suriname.

Anomaloglossus leopardus Ouboter & Jairam, 2012 Figures 2–3 Amended definition. We here provide an amended definition of A. leopardus based on the examination of 11 additional specimens (MNHN2018.53–63) collected in 2014 by two of us (AF, JPV) at Apalagadi, Suriname, the species type locality, as well as by the re-examination of the type series deposited in the NZCS (NZCS A249–51) (Paramaribo, Suriname). (1) Medium-sized Anomaloglossus (mean = 18.8 mm [18.4–19.2, n = 7] in males, mean = 21.1 mm [20.5–21.5, n = 4] in females) (Table 1); (2) body robust; (3) skin slightly tuberculate on dorsum (more tuberculated posteriorly) and legs, with a larger tubercle on each eyelid, ventral skin smooth; (4) diffuse supratympanic fold; (5) tympanic annulus distinct anteroventrally; (6) snout short and protruding in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II slightly shorter than Finger I when fingers adpressed; (9) tip of Finger IV hardly reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle distinct on Finger III and absent on the other fingers; (11) Finger III swollen dorsally and preaxially in males extending largely on dorsal surface of hand, with well-developed fringes in females; (12) fringes on all fingers particularly developed preaxially on Fingers II, in both sexes; (13) toes poorly webbed (Toe webbing formula I 2 - –1 - II 1 + –1 - III 1 + –1 + IV 0–1 + V MNHN2018.58, AF2035), with well-developed fringes on all toes but more developed preaxially on Toe II, and pre- and postaxially on Toes III and IV (sensu Grant et al. 2006; keellike lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved; (15) black arm gland at the junction of forearm with wrist in males (sensu Grant & Castro 1998, see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark present (orangish in life, cream in preservative); (18) dorsolateral stripe present, thick and continuous (cream and yellowish posteriorly in life, white in preservative), flanks and dorsum dark brown in males, paler with dark brown blotches in females; (19) ventrolateral stripe absent; (20) sexual dichromatism in throat color pattern present, in life sometimes anteriorly yellow with uniformly distributed sparse black melanophores in males, evenly yellow in females; (21) sexual dichromatism in ventral color pattern present, abdomen entirely or only posteriorly and laterally orange, centrally cream in males in life, abdomen uniformly orange in females in life; (22) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (23) median lingual process as long as wide, tapered, bluntly pointed, smooth (nonpapillate), reclined in pit; (24) call 1.08– 2.00 s long consisting of a train of 14–28 notes 0.024– 0.029 s long and spaced by intervals of 0.051– 0.055 s, dominant frequency at 4.20–4.61 kHz (n = 9) (Fig. 4; Table 2). Distribution and natural history. Anomaloglossus leopardus is only known from two streams at its type locality Apalagadi above 400 m elevation, and from the southern border of Suriname with Brazil (2.4788N, 55.63161W). The species also likely occurs between these two localities and maybe throughout the massif extending northwest in Suriname and southeast in Brazil. The tadpole is unknown, but since the species is associated with streams and has a more extensive webbing than the endotrophic A. baeobatrachus, and because all closely-related species (except A. baeobatrachus) are associated to streams and have exotrophic tadpoles transported to the water, we assume this is likely also the case for A. leopardus. It co-occurs with A. stepheni, which is endotrophic, throughout its range.

Anomaloglossus mitaraka sp. nov. Anomaloglossus sp. “Mitaraka” Vacher et al. 2017 Holotype. MNHN2018.64 (field n° AF2814), an adult male, collected by A. Fouquet and M. Dewynter, 24 February 2015, Mitaraka, French Guiana, 2.23577° N 54.44928° W, ~ 150 m elevation (Figs. 2–3). Paratopotypes. Nine specimens: MNHN 2018.65–70 (field n° AF 2732, 2750-1, 2754, 2808, 2878) six adult males and MNHN 2018.71–73 (field n° AF 2724, 2731, 2824) three females collected with the holotype by A. Fouquet and M. Dewynter. Etymology. The specific epithet is a noun in apposition and refers to the type locality (Mitaraka, French Guiana). Definition. (1) Medium-sized Anomaloglossus (average male SVL 18.6 mm [18.2–19.3, n = 7], female SVL 20.1 mm [19.2–21.0, n = 3]) (Table 1); (2) body robust; (3) skin on dorsum with irregularly scattered tubercles becoming denser on the posterior half and legs, with a larger tubercle on each eyelid, ventral skin smooth; (4) inconspicuous supratympanic fold; (5) tympanum distinct anteroventrally; (6) snout short and protruding in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II equal to Finger I when fingers adpressed; (9) tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle distinct on Finger III, absent on the other fingers; (11) Finger III swollen dorsally and preaxially, extending largely towards dorsal surface of hand in males; (12) fringes present on all fingers particularly developed preaxially on Finger II in males and females; (13) toes basally webbed, with well-developed fringes on all toes, more developed preaxially on Toe II, and pre- and postaxially on toes III and IV (sensu Grant et al. 2006; keel-like lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved; (15) black arm gland at the junction with wrist in males (sensu Grant & Castro 1998, see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark present (orangish in life, cream in preservative); (18) pale dorsolateral stripe present, solid (white to orangish anteriorly, yellow posteriorly in life, white in preservative), flanks and dorsum uniformly dark brown in males, reddish brown with dark brown blotches in females in life; (19) ventrolateral stripe absent; (20) sexual dichromatism in throat color pattern present in life, sometimes anteriorly yellow in males with sparse black melanophores, evenly and entirely yellow in females; (21) sexual dichromatism in ventral color pattern present, abdomen mostly cream, posteriorly and laterally yellow in males, uniformly bright yellow in females in life; (22) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (23) median lingual process as long as wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (24) a 0.74– 1.23 s length call consisting of a train of 8–16 notes of 0.028– 0.036 s in length and spaced by intervals of 0.051– 0.068 s and of dominant frequency at 4.13–4.76 kHz (n = 6); (25) tadpole of type 4 (Orton 1953), exotrophic, with a functional mouth with marginal papillae and labial teeth (Fig. 4; Table 2). Morphological comparisons with other Anomaloglossus. The only other species group co-occurring with the Anomaloglossus stepheni group is the A. degranvillei group, represented by A. blanci, A. degranvillei, A. dewynteri and A. surinamensis, which are readily distinguishable by their moderate webbing (basal in A. mitaraka) and well-developed fringes on all toes (smaller and more developed on Toe II, III, IV) and the absence of a dorsolateral stripe (present). Outside these two groups all the other described Anomaloglossus species exclusively occur in Pantepui (i.e., the Guiana Shield highlands, see Kok et al. 2018) and have moderate to extensive toe webbing except A. meansi Kok, Nicolaï, Lathrop & MacCulloch, 2018, A. kaiei (Kok, Sambhu, Roopsind, Lenglet & Bourne, 2006), A. rufulus (Gorzula, 1990) and A. roraima (La Marca, 1997). However, unlike A. mitaraka, A. meansi lacks a conspicuous dorsolateral stripe, Finger III in males of A. kaei is not distinctly swollen, A. rufulus has as an extensively spotted ventral face, and A. roraima lacks well-developed fringes. Within the Anomaloglossus stepheni group, A. mitaraka can be distinguished from its probable sister species (Fouquet et al. 2019) A. leopardus (Figs. 2–3) by (1) narrower dark brown bars on legs vs large and more conspicuous dark transverse bars on legs in A. leopardus; (3) ventral coloration with orange restricted to the posterior part in males in A. mitaraka vs. generally entirely yellow in A. leopardus; (4) dorsolateral line thin in A. mitaraka vs. thick and well defined in A. leopardus; (5) advertisement call with a slower note rate (mean = 11.43, range 10.8–12.3 note/s in A. mitaraka [n = 6] vs. mean = 13.4, range 12.9–13.6 notes/s in A. leopardus [n = 4]) (Table 2). Anomaloglossus mitaraka can be distinguished from A. stepheni (Figs. 2–3, Table 1) by (1) a larger body size (mean = 18.6; range 18.2–19.3 mm in males [n = 7] and mean = 20.1; range 19.2–21.0 mm in females [n = 3] in A. mitaraka vs. mean = 17.15, range 16.5–18.0 mm in males [n = 10] and mean = 17.3, range 17.0–18.0 mm in females [n = 5] [from Martins 1989] in A. stepheni); (2) skin of dorsum with scattered tubercules in A. mitaraka vs. evenly tuberculate in A. stepheni; (3) orange ventral coloration posteriorly and laterally in males of A. mitaraka vs. entirely translucent white in A. stepheni; (4) dorsolateral line thin but well defined and continuous in A. mitaraka vs. thick and ill-defined dorsally in A. stepheni; (5) advertisement call much longer (mean = 1.04 s, range 0.74– 1.23 s in A. mitaraka [n = 6] vs. mean= 0.25 s, range 0.18– 0.29 s in A. stepheni [n = 4]), emitted at a slower pace (call rate mean = 0.43, range 0.32–0.52 calls/s in A. mitaraka [n = 6] vs. mean = 1.40, range 1.10–1.89 calls/s in A. stepheni [n = 4]) (Table 2). Anomaloglossus mitaraka can be distinguished from A. baeobatrachus by (1) a larger body size (mean = 18.6; range 18.2–19.3 mm in males [n = 7] and mean = 20.1; range 19.2–21.0 mm in females [n = 3] in A. mitaraka vs. mean = 16.2; range 14.8–17.1 mm in males [n = 16] and mean = 18.4; range 17.3–19.4 mm in females [n = 4] in A. baeobatrachus); (2) ventral coloration posteriorly and laterally in males in A. mitaraka vs. yellow and less extensive in A. baeobatrachus; (3) dorsolateral line thin but well defined and solid in A. mitaraka vs. narrower and interrupted in A. baeobatrachus; (4) call with lower note rate (mean = 11.43 notes/s, range 10.83–12.28 in A. mitaraka [n = 6] vs. mean = 16.10, range 15.62– 16.85 s in A. baeobatrachus [n = 9]) lower dominant frequency (mean= 4.44, range 4.13–4.76 kHz in A. mitaraka [n = 6] vs. mean = 5.39, range 4.96–5.59 kHz in A. baeobatrachus [n = 9]) (Table 2). Description of the holotype. An adult male, 18.6 mm SVL; body robust; head wider than long, HL 94 % of HW; HL 33 % of SVL; dorsal skin tuberculate, one enlarged tubercle on each eyelid, snout long (SL 52 % of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 39 % of HW; EN 29 % of HL, 75 % of ED. Tympanum distinct anteroventrally; supratympanic fold inconspicuous; choanae small, circular, located anterolaterally (Figs. 2–3). Forelimb slender, skin tuberculate; metacarpal ridge present; HAND 24 % of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened; webbings absent on fingers; lateral fringes present on preaxial edges of Finger II; Finger III distinctly swollen dorsally and preaxially; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5 times width of digit; width of disc on Finger III 0.6 mm; discs with distinct dorsal scutes. Relative lengths of adpressed fingers III> IV> I> II; palmar tubercle large, heart-shaped, 0.8 mm in diameter (larger than Finger’s III disc), thenar tubercle elliptic, small (equal to Finger III’s disc in maximum diameter), elliptic, half the size of palmar tubercle, well separated from palmar tubercle. Only basal subarticular tubercles on Fingers are conspicuous; subarticular tubercles on fingers I and II the largest, followed by Finger IV’s subarticular tubercles and basal subarticular tubercle on Finger III. Hind limb robust, skin tuberculate; TL 47 % of SVL; heels in contact when hind limbs are flexed at right angles to the sagittal plane of body; FL 42 % of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes. Width of disc on Toe IV 0.7 mm. Foot basally webbed; lateral fringes present on all toes. Toe webbing formula I 1 + –1 - II 1 + –1 - III 1 + –1 + IV 0–1 + V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end. Metatarsal fold strong. Color of holotype in life. Dorsal color uniformly dark brown. Dorsolateral stripe thin, continuous, bluish white (yellow near groin) (Fig. 2). Dark brown lateral band extending from tip of snout to the groin and containing the indistinct dorsal part of tympanum, tapering posteriorly. Upper lip with small white blotches. Lower flanks pale white anteriorly and orange posteriorly with small white blotches. Throat white covered with melanophores, more densely anterolaterally; belly anteriorly white, white laterally, orange posteriorly, ventral surfaces of thighs and arms orange. Iris with coper metallic pigmentation and pupil ring interrupted dorsally and ventrally by transversal pigmentation (Fig. 2). Upper and lower arm pale red dorsally, light brown with small dark brown spots elsewhere. Dorsal surfaces of thigh, shank and tarsus brown with dark brown transverse bars and ill-defined blotches. Paracloacal marks orangish, elongated anteroposteriorly. Toes and digits with small light blue dots dorsally and laterally. Palms and soles dark brown. Color of holotype in preservative. After three years in 70 % ethanol, some colors of the specimen faded and the dorsal coloration now varies from brown to grey with interorbital, mid-dorsal and sacral dark brown blotches and the orangish ventral coloration disappeared (Fig. 3). Bluish freckles and orange and reddish marks turned cream. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. Coloration of limbs varies from brown to reddish brown, females being paler and with more conspicuous dark dorsal blotches. Colour of dorsolateral line varies from white to orangish. Overall dorsal and lateral coloration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as males carrying tadpoles apparently display overall lighter colors, smoother skin and sharper contrasts, whereas calling males are very dark and highly tuberculate. Ventral coloration of female is entirely orangish while the orangish parts are limited in various extent to the posterior region and throat in males. Vocal sac, slits and small dark melanophores on throat only observed in males as well as swelling on the Finger III. Advertisement call. Six specimens (one uncollected) calling from the leaf litter were recorded from a distance of about 1 m and at temperatures ranging from 23 to 26°C. Descriptive statistics of call parameters are presented in Table 2. Anomaloglossus mitaraka emits trains (call length mean = 1.04 s; range 0.74– 1.23 s) of short notes (note length mean = 0.030 s; range 0.028– 0.036 s; inter-note interval mean = 0.060 s; range 0.051– 0.068 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 4.44 kHz on average (range 4.13–4.76 kHz) with a slight upward modulation (ca. 0.2 kHz) (Fig. 4, Table 2). Larval morphology. The following description is based on three tadpoles at stage 26, 28 and 30 (Fig. 6). Tadpoles correspond to a Type 4 tadpole of Orton (1953); exotrophic; body skin smooth; TL 12.21–19.43 mm; BL 4.41–7.14 mm, 35–37 % of TL, 164–181 % of BW, 221–256 % of BH; BW 131–143 % of BH (Table 3); body ovoid, snout round in dorsal and lateral view; eyes positioned and directed laterally; ED 0.48–0.63 mm, 45–98 % of IOD; IOD equal to IND; nares frontally positioned and directed laterodorsally; narial opening reniform in lateral view; END 0.34–0.73 mm. Spiracular tube sinistral, conical, projecting posterodorsally, its tip located at 60–63 % of BL posteriorly to snout. Lateral-line system inconspicuous. Caudal musculature highest at its base, tapering posteriorly, terminating at tail tip; tail tip rounded; upper fin originating at junction of body and tail, gradually increasing in height to about 3/4 of tail; UTF 28–53 % of TMH; LTF 38–55 % of TMH; MTH 14–17 % of TL (Fig. 6, Table 3). Mouth ventral, oral disc strongly emarginated, width 1.90 mm. Labial teeth long, in single rows, LTRF 2(2)/3. A-2 consisting of two short rows, separated by a large and deep gap; P-1 not interrupted. Marginal papillae long, of equal size on each labium, tapered, blunt-tipped, in a single row, evenly distributed; median gap on upper labium approximately 2/3 the length of A-1; jaw sheaths large, serrated, lower jaw sheath broadly V-shaped. In life, the entire body is dark grey with abundant golden flecks, particularly on dorsum. Golden flecks become scarce ventrally. Posteriorly, dark grey coloration fades and tail becomes translucent. Tadpoles of Anomaloglossus mitaraka can be distinguished from those of any other described Anomaloglossus of the stepheni group (A. stepheni and A. baeobatrachus - this study) by the presence of a functional mouth with marginal papillae and labial teeth. Distribution and natural history. Anomaloglossus mitaraka is a diurnal species inhabiting the leaf litter in primary forest at low to mid elevations (from 150 to 500 m a.s.l.). The species is usually found close to streams next to which it deposits its exotrophic tadpoles into puddles, but some specimens can be found more than 50 m from water bodies. Males call all day long when the weather is rainy. Breeding occurs during the rainy season, between January and May. The males respond to intraspecific playbacks with shorter and more rapidly emitted note trills. Males are territorial, their small territories being spaced at least a few meters apart. Males usually call slightly above the leaf litter, exposed on a branch or a dead leaf. Eggs are deposited in the fold of a dead leaf (a single observation). This clutch was raised (Fig. 6) and some tadpoles were genotyped. Populations have been documented in southwestern French Guiana and adjacent Suriname (Mitaraka massif, Pic Coudreau, Haute Marwini, upper Tapanahony; Fig. 1). It also likely occurs in adjacent Brazil and may occupy similar habitats in other regions of Suriname., Published as part of Fouquet, Antoine, Vacher, Jean-Pierre, Courtois, Elodie A., Deschamps, Chloé, Ouboter, Paul, Jairam, Rawien, Gaucher, Philippe, Dubois, Alain & Kok, Philippe J. R., 2019, A new species of Anomaloglossus (Anura: Aromobatidae) of the stepheni group with the redescription of A. baeobatrachus (Boistel and de Massary, 1999), and an amended definition of A. leopardus Ouboter and Jairam, 2012, pp. 439-460 in Zootaxa 4576 (3) on pages 447-454, DOI: 10.11646/zootaxa.4576.3.2, http://zenodo.org/record/3715704, {"references":["Vacher, J. - P., Kok, P. J. R., Rodrigues, M. T., Dias Lima, J., Lorenzini, A., Martinez, Q., Fallet, M., Courtois, E. A., Blanc, M., Gaucher, P., Dewynter, M., Jairam, R., Ouboter, P., Thebaud, C. & Fouquet, A. (2017) Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield. Molecular Phylogenetics and Evolution, 112, 158 - 173. https: // doi. org / 10.1016 / j. ympev. 2017.04.017","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262. https: // doi. org / 10.1206 / 0003 - 0090 (2006) 299 [1: PSODFA] 2.0. CO; 2","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147. https: // doi. org / 10.1206 / 308.1","Grant, T. & Castro, F. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology, 32, 378 - 392. https: // doi. org / 10.2307 / 1565452","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2, 63 - 75. https: // doi. org / 10.2307 / 2411661","Kok, P. J. R., Nicolai, M. P. J., Lathrop, A. & MacCulloch, R. D. (2018) Anomaloglossus meansi sp. n., a new Pantepui species of the Anomaloglossus beebei group (Anura, Aromobatidae). ZooKeys, 759, 99 - 116. https: // doi. org / 10.3897 / zookeys. 759.24742","Kok, P. J. R., Sambhu, H., Roopsind, I., Lenglet, G. L. & Bourne, G. R. (2006) A new species of Colostethus (Anura: Dendrobatidae) with maternal care from Kaieteur National Park, Guyana. Zootaxa, 1238, 35 - 61. http: // dx. doi. org / 10.11646 / zootaxa. 1238.1.3","Gorzula, S. J. (1990) Una nueva especie de Dendrobates (Amphibia: Dendrobatidae) del Macizo del Chimanta, Estado Bolivar, Venezuela. Memoria Sociedad de Ciencias Naturales La Salle, 48, 143 - 149.","La Marca, E. (1997) Ranas del genero Colostethus (Amphibia: Anura: Dendrobatidae) de la Guayana Venezolana con la descripcion de siete especies nuevas. Publicaciones de la Asociacion de Amigos de Donana, 9, 1 - 64.","Fouquet, A., Ferrier, B., Salmona, J., Tirera, S., Vacher, J. - P., Courtois, E. A., Gaucher, P., Dias Lima, J., Nunes, P. M. S., de Souza, S. M., Rodrigues, M. T., Noonan, B. & de Thoisy, B. (2019) Phenotypic and life-history diversification in Amazonian frogs despite past introgressions. Molecular Phylogenetics and Evolution, 130, 169 - 180. https: // doi. org / 10.1016 / j. ympev. 2018.09.010","Martins, M. (1989) Nova especie de Colostethus da Amazonia central (Amphibia: Dendrobatidae). Revista Brasileira de Biologia, 49, 1009 - 1012. https: // doi. org / 10.1643 / 0045 - 8511 (2007) 7 [114: ANASOT] 2.0. CO; 2"]}

Anomaloglossus baeobatrachus (Boistel & de Massary, 1999) Figs. 2–5 Neotype. MNHN2018.31 (field n o AF2589), an adult male, collected by Jean-Pierre Vacher, 21 February 2015, Saint Eugène, French Guiana, 4.82167° N 53.06766° W, ~ 60 m elevation (Figs. 2, 3). Additional specimens examined. Twenty-one specimens: MNHN 2018.34–36 (field n° AF 0207–8, 0214) two adult males and one female, collected by A. Fouquet and Régis Gallais, 0 8 April 2007, Angoulème, French Guiana, 5.383333° N 53.650000° W, ~ 20 m elevation. MNHN 2018.37 (field n° AF 0244) an adult female, collected by A. Fouquet, 10 May 2007, Nouragues field station, French Guiana, 4.083333° N 52.683333° W, ~ 100 m elevation. MNHN 2018.38 (field n° AF 0821) an adult male, collected by A. Fouquet, 31 December 2012, Chutes Voltaire, French Guiana, 5.031222° N 54.087778° W, ~ 60 m elevation. MNHN 2018.39–40 (field n° AF 1839, 1841) two adult males, collected by A. Fouquet and Jean-Pierre Vacher, 24 March 2014, RN2 corridor 3, French Guiana, 4.10583° N 52.05096° W, ~ 50 m elevation. MNHN 2018.41–44 (field n° AF 2098-9, 2101-2) four adult males, collected by A. Lorenzini, 0 1 April 2014, RN2 corridor 5, French Guiana, 4.03297° N 51.99078° W, ~ 50 m elevation. MNHN 2018.32–33 (field n° AF 2590-1) two adult males with same data as neotype. MNHN 2018.45–48,50 (field n° AF 2667, 2704-6, 2709) four adult males and MNHN 2018.49 (field n° AF 2707) and adult female, collected by Jean-Pierre Vacher, 18 February 2015, Alikéné, French Guiana, 3.21153° N 52.39981° W, ~ 280 m elevation. MNHN 2018.51–52 (field n° AF 2942, 2963) an adult female and an adult male, collected by Antoine Fouquet and Pedro S. Nunes, 15 March 2015, Mémora, French Guiana, 3.31774° N 52.19329° W, ~ 50 m elevation. Definition. (1) A small-sized Anomaloglossus (average male SVL 16.2 mm [14.8–17.1, n = 16], female SVL 18.4 mm [17.3–19.4, n = 4]) (Table 1); (2) body robust; (3) skin tuberculate on dorsum (particularly the posterior half) and legs, with a large tubercle on each eyelid, ventral skin smooth; (4) diffuse supratympanic fold; (5) tympanic annulus distinct anteroventrally; (6) snout short and protruding in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II same length as Finger I when fingers adpressed; (9) tip of Finger IV reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle distinct on Finger III but absent on the other fingers; (11) Finger III swollen dorsally and preaxially extending largely on dorsal surface of hand in males, with well-developed lateral fringe in females; (12) Fingers II with preaxial fringes in both sexes; (13) toes basally webbed, with fringes on all toes more developed preaxially on Toe II and pre- and postaxially on Toes III and IV (sensu Grant et al. 2006; keel-like lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved; (15) black arm gland at the junction of distal forearm with wrist in males (sensu Grant & Castro 1998, see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark present (orangish in life, cream in preservative); (18) dorsolateral stripe present, discontinuous (bluish to yellowish particularly posteriorly in life, white in preservative), flanks and dorsum dark brown in males paler in females; (19) ventrolateral stripe absent; (20) sexual dichromatism in throat color present, in life sometimes anteriorly and laterally yellow in males, evenly yellow in females, with sparse black melanophores; (21) sexual dichromatism in ventral color present, in life abdomen mostly cream with flanks often yellow in males, abdomen entirely and uniformly bright yellow in females in life; (22) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (23) median lingual process as long as wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (24) call 0.84– 1.33 s long, consisting of a train of 13–28 pulsed notes 0.018– 0.021 s long spaced by intervals of 0.043– 0.047 s, dominant frequency at 4.960–5.591 kHz (n = 9); (25) tadpole of type 2 (Orton 1953), endotrophic, with bare mouth and prominent trophic reserves (Fig. 4; Table 2). Morphological comparisons with other Anomaloglossus. The only other species group co-occurring with the Anomaloglossus stepheni group is the A. degranvillei group, currently represented by A. blanci Fouquet, Vacher, Courtois, Villette, Reizine, Gaucher, Jairam, Ouboter & Kok, 2018, A. degranvillei (Lescure, 1975), A. dewynteri Fouquet, Vacher, Courtois, Villette, Reizine, Gaucher, Jairam, Ouboter & Kok, 2018, and A. surinamensis Ouboter & Jairam, 2012, which are readily distinguishable by their moderate webbing (basal in A. baeobatrachus) and well-developed fringes on all toes (smaller and more developed on Toe II, III, IV) and the absence of a dorsolateral stripe (present). Outside these two groups all the other described Anomaloglossus species exclusively occur in Pantepui (i.e., the Guiana Shield highlands, see Kok et al. 2018) and have moderate to extensive toe webbing except A. meansi Kok, Nicolaï, Lathrop & MacCulloch, 2018, A. kaiei (Kok, Sambhu, Roopsind, Lenglet & Bourne, 2006), A. rufulus (Gorzula, 1990) and A. roraima (La Marca, 1997). However, unlike A. baeobatrachus, these last four species lack developed fringes on toes. Within the Anomaloglossus stepheni group, A. baeobatrachus can be distinguished from A. stepheni (Figs. 2– 3, Table 1) by (1) skin of dorsum irregularly tuberculate in A. baeobatrachus vs. evenly tuberculate in A. stepheni; (2) ventral coloration with yellow shades, at least posteriorly in males in A. baeobatrachus vs. entirely translucent white in A. stepheni; (3) dorsolateral line thin and interrupted in A. baeobatrachus vs. thick and ill-defined margins dorsally in A. stepheni; (4) call much longer (mean = 1.12, range 0.84– 1.33 s) in A. baeobatrachus [n = 9] vs. mean = 0.25, range 0.18– 0.29 s in A. stepheni [n = 4]) emitted at a slower pace (call rate mean = 0.33, range 0.18–0.48 calls/s in A. baeobatrachus [n = 11] vs. mean = 1.40, range 1.10–1.89 calls/s in A. stepheni [n = 4]), with a higher dominant frequency (mean = 5.39, range 4.96–5.59 kHz in A. baeobatrachus [n = 6] vs. mean = 4.48, range 4.30– 4.85 kHz in A. stepheni [n = 4]). Anomaloglossus baeobatrachus can be distinguished from A. leopardus (Figs. 2–3, Table 1) by (1) a smaller body size (mean = 16.2; range 14.8–17.1 mm in males [n = 16] and mean = 18.4; range 17.3–19.4 mm in females [n = 4] in A. baeobatrachus vs. mean = 18.8, range 18.5–19.2 mm in males [n = 10] and mean = 21.1, range 20.5– 21.5 mm in females [n = 4] in A. leopardus); (2) dorsal coloration uniformly dark brown with inconspicuous dark transverse bars on legs in A. baeobatrachus vs dorsal coloration with large dark blotches, particularly visible in preservative and conspicuous dark transverse bars on legs in A. leopardus; (3) ventral coloration with yellow restricted to the posterior part in males in A. baeobatrachus vs. generally entirely yellow in A. leopardus; (4) dorsolateral line thin and discontinuous in A. baeobatrachus vs. thick and continuous in A. leopardus; (5) call with a faster note rate (mean = 16.1, range 15.7–16.8 note/s in A. baeobatrachus [n = 9] vs. mean = 13.4, range 12.9– 13.6 notes/s in A. leopardus [n = 4]) with a higher dominant frequency (mean = 5.39, range 4.96–5.59 kHz in A. baeobatrachus [n = 6] vs. mean=4.49, range 4.40–4.57 kHz in A. leopardus [n = 4], Table 2). Description of the neotype. An adult male, 15.9 mm SVL; body robust; head wider than long, HL 94 % of HW; HL 32 % of SVL; dorsal skin irregularly tuberculate, one enlarged tubercle on each eyelid, snout long (SL 55 % of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 44 % of HW; EN 31 % of HL, 70 % of ED. Tympanum distinct anteroventrally; supratympanic fold diffuse; choanae small, circular, located anterolaterally to palatine (Fig. 3). Forelimb slender, skin dorsally tuberculate; metacarpal ridge present; HAND 24 % of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened without webbing, lateral fringes present on preaxial edges of Fingers II; Finger III distinctly swollen dorsally and preaxially; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5X width of digit; width of disc on Finger III 0.5 mm; discs with distinct dorsal scutes. Relative lengths of adpressed fingers III> IV> I> II; palmar tubercle large, heart-shaped, 0.7 mm in diameter (larger than Finger III disc), thenar tubercle small (equal to Finger III disc), elliptic, half the size of palmar tubercle, well separated from palmar tubercle. Only basal subarticular tubercles on Fingers I, II and IV are conspicuous; Subarticular tubercle of Finger I larger than Finger II and IV subarticular tubercles. Hind limb robust, skin dorsally tuberculate; TL 47 % of SVL; heels not in contact when hind limbs are flexed at right angle to sagittal plane of body; FL 42 % of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes; disc on Toe I only slightly larger than width of digit. Width of disc on Toe IV 0.7 mm. Feet poorly webbed; lateral fringes present on all toes. Toe webbing formula I 1 + –1 - II 1 + –1 - III 1 ½ –1 ½ IV 0–1 + V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end. Metatarsal fold conspicuous and strongly curved. Color of neotype in life. Dorsal color uniformly dark brown. Discontinuous pale dorsolateral stripes consisting of irregular bluish, white, orangish and yellow (near groin) small blotches (Fig. 2). Dark brown lateral band extending from tip of snout to the groin and containing the indistinct dorsal part of tympanum. Upper lip with small iridescent light blue blotches. Lower flanks pale yellow with small iridescent white blotches. Throat anteriorly yellow, centrally white and posteriorly pinkish, covered with melanophores more densely anterolaterally; belly centrally white and laterally and posteriorly yellow, ventral surface of thighs and arms pale orange. Iris with copper metallic pigmentation and pupil ring interrupted dorsally and ventrally by transversal dark pigmentation (Fig. 2). Upper arm pale orange dorsally with a conspicuous proximal white spot, anteriorly and posteriorly light brown with small bluish spots. Lower arm light brown with ill-defined dark brown blotches and small bluish spots. Dorsal surfaces of thigh, shank and tarsus dark brown with ill-defined darker blotches. Paracloacal marks orangish, elongated anteroposteriorly. Toes and digits with small light blue dots. Palms and soles dark brown. Color of neotype in preservative. After three years in 70 % ethanol, some colors of the specimen faded and the dorsal coloration now varies from brown to grey and the yellow ventral coloration disappeared. All bluish freckles and orange and reddish marks turned cream (Fig. 3). Variation. Measurements (range, mean, and standard deviation) are provided in Table 1. Adult dorsal coloration varies from brown to reddish brown in males, females being paler and with more conspicuous darker dorsal blotches. The dorsolateral line varies from almost continuous to continuous; bluish flecks may be numerous or absent. Additionally, overall coloration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as males carrying tadpoles apparently display overall lighter colors, smoother skin and sharper contrasts while calling males are very dark and highly tuberculate. Ventral coloration of female is entirely yellow while the yellow parts are limited in various extent to the posterior region and throat in males. Vocal sac, slits and small dark melanophores on throat only observed in males as well as swelling on the Finger III. Advertisement call. Nine specimens (two not collected) calling from the leaf litter were recorded from a distance of about 1 m and at temperatures ranging from 23 to 26°C. Descriptive statistics of call parameters are presented in Table 2. Anomaloglossus baeobatrachus emits trains (call length mean = 1.12 s; range 0.84– 1.33 s) of short notes (note length mean = 0.019 s; range 0.018– 0.022 s; inter-note interval mean = 0.045 s; range 0.043– 0.046 s) (Fig. 4). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 5.39 kHz on average (range 4.96–5.59 kHz) with a slight upward modulation (ca. 0.2 kHz) (Fig. 4, Table 2). Larval morphology. The following description is based on four tadpoles at stage 30, 32, 33 and 34 (Fig. 5). Measurements are presented in Table 3. Tadpoles correspond to Type 2 tadpoles of Orton (1953) although an external spiracle is absent; tadpoles endotrophic; body skin smooth; TL 10.86–12.92 mm; BL 3.97–4.62 mm, 36– 37 % of TL, 141–159 % of BW, 212–241 % of BH; BW 137–171 % of BH (Table 2); body inflated from trophic reserves until stage 33 when it becomes ovoid; round snout in dorsal and lateral view; eyes positioned and directed laterally (Fig. 5); ED 0.46–0.68 mm, 65–81 % of IOD; IOD more than IND; nares positioned frontally and directed laterodorsally; narial opening reniform in lateral view; END 0.2 8– 0.40 mm. Spiracular tube absent. Lateral-line system inconspicuous. Caudal musculature highest at its base, tapering posteriorly, terminating at tail tip; tail tip rounded; upper fin originating at junction of body and tail, gradually increasing in height to about 3/4 of tail; UTF 39–50 % of TMH; LTF 26–42 % of TMH; MTH 15–18 % of TL. Mouth small and bare (without marginal papillae, labial teeth nor sheaths), directed ventrally, oral disc width 0.5 mm (from edges of jaws) (Fig. 5, Table 3). In life, until stage 32 the entire body is light grey except the tail fins that are translucent and the abdomen that is yellowish. From stage 33 the body becomes dark grey with golden speckles, and the tail and fin become translucent posteriorly (Fig. 5). Tadpoles of Anomaloglossus baeobatrachus can be distinguished from those of any other Anomaloglossus of the stepheni group (except A. apiau and A. stepheni) by the absence of functional mouth and the prominent trophic reserves. Only two other Anomaloglossus with an endotrophic tadpole have been described, A. stepheni (Juncá et al. 1994) and A. degranvillei (Lescure 1984); although the latter may in fact correspond to A. surinamensis or A. blanci (Fouquet et al. 2018). Tadpoles of A. baeobatrachus can be distinguished from those of Anomaloglossus stepheni by the absence of a spiracle (present in A. stepheni), a mouth located anteriorly to the eyes (at the level of the eyes in A. stepheni), eyes located dorsolaterally (laterally in A. stepheni), a proportionally shorter tail – BL 36– 37 % of TL (vs. 33 % in A. stepheni) (Juncá et al. 1994). They can be distinguish from tadpoles of A. degranvillei by the presence of an oral disc (absent in A. degranvillei), presence of a vent tube (absent in A. degranvillei), eyes located dorsolaterally (laterally in A. degranvillei) and by their development, which is not completed on the back of the male parent. Distribution and natural history. Populations of Anomaloglossus baeobatrachus have been documented throughout French Guiana and the state of Amapá in Brazil (Fig. 1). However, southern FG and Amapá populations form genetic clusters distinct from the ones from northern FG, where lies the type locality (Fouquet et al. 2019). These three clusters remain indistinguishable based on call and morphology and display similar larval development modes (Fouquet et al. 2019). The species is absent west of the Maroni River where it is replaced by A. stepheni, a species that seems to have a similar niche. A similar situation may occur on the eastern margin of the Jari River (state of Pará, Brazil) but too few surveys are available to confirm that hypothesis. Several still undescribed species with exotrophic larvae have been confused with A. baeobatrachus (Vacher et al. 2017) in Suriname (Ouboter & Jairam 2012) and in Pará state (Avila-Pires et al. 2011). Populations with exotrophic larvae have also been documented in French Guiana and Amapá but are assumed to have undergone historical admixture and their status remains ambiguous (Fouquet et al. 2019). Anomaloglossus baeobatrachus is a diurnal species inhabiting the leaf litter in primary and secondary terrafirme forest at low to mid elevations (from 10 to 800 m elevation). It can be found alongside or far from any water bodies. Males call all day long when the weather is rainy, with two apparent peaks of daily activity, a minor one in the morning (08:00–11:00h) and a major one at the end of the day (15:00–18:00h). The species seems to be more abundant on slopes and plateaus. Males aggregate in groups of two to several dozen individuals. Breeding occurs mainly at the beginning of the rainy season (December to April). Males respond to playback of intraspecific advertisement calls with shorter and more rapidly emitted trills and also emit a different vocalization during courtship. They are territorial and defend, sometimes engaging in physical contact, territories of approximately 13 m ² (Deschamps 2005), which are separated by a few meters only. Males usually call slightly above the leaf litter, exposed on a branch or a leaf. Males generally use the same calling and oviposition sites throughout the season (Deschamps 2005). Eggs are usually deposited in the fold of a dead leaf. Breeding behaviours are very similar to those observed in A. stepheni in central Amazonia (Juncá 1998), although these species are not each other’s sister group (Vacher et al. 2017). When a female approaches a calling, Published as part of Fouquet, Antoine, Vacher, Jean-Pierre, Courtois, Elodie A., Deschamps, Chloé, Ouboter, Paul, Jairam, Rawien, Gaucher, Philippe, Dubois, Alain & Kok, Philippe J. R., 2019, A new species of Anomaloglossus (Anura: Aromobatidae) of the stepheni group with the redescription of A. baeobatrachus (Boistel and de Massary, 1999), and an amended definition of A. leopardus Ouboter and Jairam, 2012, pp. 439-460 in Zootaxa 4576 (3) on pages 443-446, DOI: 10.11646/zootaxa.4576.3.2, http://zenodo.org/record/3715704, {"references":["Boistel, R. & de Massary, J. - C. (1999) Les amphibiens veneneux de la famille des dendrobatides. Le Courrier de la Nature, 176, 34 - 39.","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262. https: // doi. org / 10.1206 / 0003 - 0090 (2006) 299 [1: PSODFA] 2.0. CO; 2","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147. https: // doi. org / 10.1206 / 308.1","Grant, T. & Castro, F. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology, 32, 378 - 392. https: // doi. org / 10.2307 / 1565452","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2, 63 - 75. https: // doi. org / 10.2307 / 2411661","Fouquet, A., Vacher, J. - P., Courtois, E. A., Villette, B., Reizine, H., Gaucher, P., Jairam, R., Ouboter, P. & Kok, P. J. R. (2018) On the brink of extinction: two new species of Anomaloglossus from French Guiana and amended definition of Anomaloglossus degranvillei and A. surinamensis (Anura: Aromobatidae). Zootaxa, 4379 (1), 001 - 023. https: // doi. org / 10.11646 / zootaxa. 4379.1.1","Lescure, J. (1975) Contribution a l'etude des amphibiens de Guyane Francaise. III. Une nouvelle espece de Colostethus (Dendrobatidae): Colostethus degranvillei n. sp. Bulletin du Museum National d'Histoire Naturelle. Paris. Serie 3, Zoologie, 293, 413 - 420.","Ouboter, P. E. & Jairam, R. (2012) Amphibians of Suriname. Brill, Leiden, 376 pp.","Kok, P. J. R., Nicolai, M. P. J., Lathrop, A. & MacCulloch, R. D. (2018) Anomaloglossus meansi sp. n., a new Pantepui species of the Anomaloglossus beebei group (Anura, Aromobatidae). ZooKeys, 759, 99 - 116. https: // doi. org / 10.3897 / zookeys. 759.24742","Kok, P. J. R., Sambhu, H., Roopsind, I., Lenglet, G. L. & Bourne, G. R. (2006) A new species of Colostethus (Anura: Dendrobatidae) with maternal care from Kaieteur National Park, Guyana. Zootaxa, 1238, 35 - 61. http: // dx. doi. org / 10.11646 / zootaxa. 1238.1.3","Gorzula, S. J. (1990) Una nueva especie de Dendrobates (Amphibia: Dendrobatidae) del Macizo del Chimanta, Estado Bolivar, Venezuela. Memoria Sociedad de Ciencias Naturales La Salle, 48, 143 - 149.","La Marca, E. (1997) Ranas del genero Colostethus (Amphibia: Anura: Dendrobatidae) de la Guayana Venezolana con la descripcion de siete especies nuevas. Publicaciones de la Asociacion de Amigos de Donana, 9, 1 - 64.","Junca, F. A., Altig, R. & Gascon, C. (1994) Breeding biology of Colostethus stepheni, a dendrobatid frog with a nontransported nidicolous tadpole. Copeia, 747 - 750. https: // doi. org / 10.2307 / 1447191","Lescure, J. (1984) Larvas de Dendrobatidae. II Reunion Iberoamer. Cons. Zool. Uert., 37 - 45.","Fouquet, A., Ferrier, B., Salmona, J., Tirera, S., Vacher, J. - P., Courtois, E. A., Gaucher, P., Dias Lima, J., Nunes, P. M. S., de Souza, S. M., Rodrigues, M. T., Noonan, B. & de Thoisy, B. (2019) Phenotypic and life-history diversification in Amazonian frogs despite past introgressions. Molecular Phylogenetics and Evolution, 130, 169 - 180. https: // doi. org / 10.1016 / j. ympev. 2018.09.010","Vacher, J. - P., Kok, P. J. R., Rodrigues, M. T., Dias Lima, J., Lorenzini, A., Martinez, Q., Fallet, M., Courtois, E. A., Blanc, M., Gaucher, P., Dewynter, M., Jairam, R., Ouboter, P., Thebaud, C. & Fouquet, A. (2017) Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield. Molecular Phylogenetics and Evolution, 112, 158 - 173. https: // doi. org / 10.1016 / j. ympev. 2017.04.017","Deschamps, C. (2005) Etude sur le comportement reproducteur et territorial d'un dendrobatide: Colosthetus baeobatrachus. 40 pp.","Junca, F. A. (1998) Reproductive biology of Colostethus stepheni and Colostethus marchesianus (Dendrobatidae), with the description of a new anuran mating behavior. Herpetologica, 54, 377 - 387."]}

Anomaloglossus leopardus Ouboter & Jairam, 2012 Figures 2���3 Amended definition. We here provide an amended definition of A. leopardus based on the examination of 11 additional specimens (MNHN2018.53���63) collected in 2014 by two of us (AF, JPV) at Apalagadi, Suriname, the species type locality, as well as by the re-examination of the type series deposited in the NZCS (NZCS A249���51) (Paramaribo, Suriname). (1) Medium-sized Anomaloglossus (mean = 18.8 mm [18.4���19.2, n = 7] in males, mean = 21.1 mm [20.5���21.5, n = 4] in females) (Table 1); (2) body robust; (3) skin slightly tuberculate on dorsum (more tuberculated posteriorly) and legs, with a larger tubercle on each eyelid, ventral skin smooth; (4) diffuse supratympanic fold; (5) tympanic annulus distinct anteroventrally; (6) snout short and protruding in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II slightly shorter than Finger I when fingers adpressed; (9) tip of Finger IV hardly reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle distinct on Finger III and absent on the other fingers; (11) Finger III swollen dorsally and preaxially in males extending largely on dorsal surface of hand, with well-developed fringes in females; (12) fringes on all fingers particularly developed preaxially on Fingers II, in both sexes; (13) toes poorly webbed (Toe webbing formula I 2 - ���1 - II 1 + ���1 - III 1 + ���1 + IV 0���1 + V MNHN2018.58, AF2035), with well-developed fringes on all toes but more developed preaxially on Toe II, and pre- and postaxially on Toes III and IV (sensu Grant et al. 2006; keellike lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved; (15) black arm gland at the junction of forearm with wrist in males (sensu Grant & Castro 1998, see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark present (orangish in life, cream in preservative); (18) dorsolateral stripe present, thick and continuous (cream and yellowish posteriorly in life, white in preservative), flanks and dorsum dark brown in males, paler with dark brown blotches in females; (19) ventrolateral stripe absent; (20) sexual dichromatism in throat color pattern present, in life sometimes anteriorly yellow with uniformly distributed sparse black melanophores in males, evenly yellow in females; (21) sexual dichromatism in ventral color pattern present, abdomen entirely or only posteriorly and laterally orange, centrally cream in males in life, abdomen uniformly orange in females in life; (22) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (23) median lingual process as long as wide, tapered, bluntly pointed, smooth (nonpapillate), reclined in pit; (24) call 1.08��� 2.00 s long consisting of a train of 14���28 notes 0.024��� 0.029 s long and spaced by intervals of 0.051��� 0.055 s, dominant frequency at 4.20���4.61 kHz (n = 9) (Fig. 4; Table 2). Distribution and natural history. Anomaloglossus leopardus is only known from two streams at its type locality Apalagadi above 400 m elevation, and from the southern border of Suriname with Brazil (2.4788N, 55.63161W). The species also likely occurs between these two localities and maybe throughout the massif extending northwest in Suriname and southeast in Brazil. The tadpole is unknown, but since the species is associated with streams and has a more extensive webbing than the endotrophic A. baeobatrachus, and because all closely-related species (except A. baeobatrachus) are associated to streams and have exotrophic tadpoles transported to the water, we assume this is likely also the case for A. leopardus. It co-occurs with A. stepheni, which is endotrophic, throughout its range., Published as part of Fouquet, Antoine, Vacher, Jean-Pierre, Courtois, Elodie A., Deschamps, Chlo��, Ouboter, Paul, Jairam, Rawien, Gaucher, Philippe, Dubois, Alain & Kok, Philippe J. R., 2019, A new species of Anomaloglossus (Anura: Aromobatidae) of the stepheni group with the redescription of A. baeobatrachus (Boistel and de Massary, 1999), and an amended definition of A. leopardus Ouboter and Jairam, 2012, pp. 439-460 in Zootaxa 4576 (3) on pages 446-447, DOI: 10.11646/zootaxa.4576.3.2, http://zenodo.org/record/3715704, {"references":["Ouboter, P. E. & Jairam, R. (2012) Amphibians of Suriname. Brill, Leiden, 376 pp.","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262. https: // doi. org / 10.1206 / 0003 - 0090 (2006) 299 [1: PSODFA] 2.0. CO; 2","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147. https: // doi. org / 10.1206 / 308.1","Grant, T. & Castro, F. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology, 32, 378 - 392. https: // doi. org / 10.2307 / 1565452"]}

Anomaloglossus dewynteri sp. nov. Anomaloglossus sp. “ Itoupé” Vacher et al. 2017 Holotype. MNHN2017.0111 (field no. AF3644/APA-973-23-2), an adult male, collected by Antoine Fouquet, 11 January 2016, near base camp on Mont Itoupé, French Guiana, 3.0230 N 53.0955 W, elevation ca. ~ 600 m (Figure 7). Paratopotypes. Three specimens: MNHN2017.0112 (field no. AF0574; not measured because badly preserved), an adult male, collected by Maël Dewynter, 0 5 April 2010, near base camp on Mont Itoupé, French Guiana, 3.0230 N 53.0955 W, ~ 600 m elevation. MNHN2017.0113–4 (field no. AF3686/APA-973-23-1, AF3645/ APA-973-23-3), two adult males, collected by Antoine Fouquet, with the holotype. Etymology. This species is dedicated to our friend Maël Dewynter for his invaluable contribution to the herpetology of French Guiana. Adult definition and diagnosis. We assigned the new species to the genus Anomaloglossus based on previous studies (Fouquet et al. 2012; Vacher et al. 2017) and the presence of a median lingual process. The new species belongs to the A. degranvillei clade according to Vacher et al. (2017). (1) Medium-sized Anomaloglossus (average male SVL= 19.9 mm [19.4–20.4, n=3], female unknown) (Table 1); (2) body robust; (3) skin tuberculate on dorsum (particularly the posterior half) and legs, with a larger tubercle on each eyelid, ventral skin smooth; (4) conspicuous dark brown to black glandulous supratympanic fold flanked anteroventrally by a discontinuous stripe (orange in life, cream in preservative) running from the ventroposterior edge of the eye and extending onto the upper arm; (5) tympanum indistinct; (6) snout short and acute in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II shorter than Finger I when fingers adpressed; (9) tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle on Finger III and IV indistinct; (11) Finger III not distinctly swollen in males; (12) fingers with fringes particularly developed on preaxial edges of Fingers II and III; (13) toes moderately webbed, with well-developed fringes (sensu Grant et al. 2006; keel-like lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved, slightly tuberclelike; (15) no black arm gland in males (sensu Grant & Castro 1998. see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark inconspicuous; (18) dorsolateral stripe absent or inconspicuous, dorsal part of flanks darker than dorsum; (19) ventrolateral stripe sometimes present as a broken, poorly defined pale brown blotch, ventral part of flanks with flecks white or bluish (in life); (20) sexual dichromatism in throat colour pattern unknown (although likely because present in other species of the clade), throat solid black in reproductive males; (21) abdomen and ventral side of legs dark grey with small ill-defined white dots; (22) iris with metallic pigmentation and pupil ring interrupted ventrally by transversal black pigmentation; (23) median lingual process longer than wide, tapered, bluntly pointed, smooth (nonpapillate), reclined in pit; (24) single note call of 0.221– 0.237 s length and dominant frequency at 3.45–3.59 kHz (n=3) (Table 2; Figure 3). Morphological comparisons with other lowland Anomaloglossus. The only other species group cooccurring with the Anomaloglossus degranvillei species group is the A. stepheni species group, represented by A. baeobatrachus in French Guiana, which is readily distinguishable by conspicuous dorsolateral stripes, swollen third fingers in males and absence of webbing and fringes on toes. Within the Anomaloglossus degranvillei group, A. dewynteri can be distinguished from A. surinamensis (Figure 4) by (1) a shorter snout, acute in lateral view (vs. rounded in A. surinamensis); (2) a larger body size in males (X¯ = 19.9 mm, range 19.4–20.4 mm in A. dewynteri [n=3] vs. X¯ =14.75, range 14.0– 15.3 mm in A. surinamensis [n=8]); (3) more developed fringes on fingers and toes; (4) presence of a large tubercle on the top of the eyelid (inconspicuous in A. surinamensis); (5) belly dark grey with small white spots (vs. cream with irregular grey blotches less abundant posteriorly in A. surinamensis); (6) ventral surface of thigh dark grey (vs. cream with dark spots on the edges in A. surinamensis); (7) larger MLP (> 0.5 mm vs A. surinamensis; (8) call characterized by much longer pulsed notes (pulsed note X¯ =0.228, range 0.221– 0.237 s [n=3] in A. dewynteri vs. tonal note X¯ =0.032, range 0.028– 0.037 s in A. surinamensis [n=20]) emitted between longer intervals (X¯ =0.76, range 0.60– 0.88 s in A. dewynteri [n=3] vs. X¯ =0.573, range 0.372– 0.825 s in A. surinamensis [n=20]) and with lower dominant frequency (X¯ =3.52, range 3.45–3.59 kHz in A. dewynteri [n=4] vs. X¯ =4.89 kHZ, range 4.55–5.35 kHz in A. surinamensis [n=20]). Anomaloglossus dewynteri can be distinguished from A. blanci (Figure 2) by (1) its larger size in males (X= 19.9 mm, range 19.4–20.4 mm in A. dewynteri [n=3] vs. X¯ =16.9, range 15.9–18.8 mm in A. blanci [n=8]); (2) belly from solid black to dark grey with abundant small conspicuous white spots (pale grey with small white illdefined spots in A. blanci); (3) ventral surface of thighs dark grey (yellowish in life and cream in preservative in A. blanci); (4) strong metacarpal ridge (inconspicuous in A. blanci, Figures 2F, 7F); (5) call with longer notes (X=0.228, range 0.221– 0.237 s [n=3] in A. dewynteri vs. X¯ =0.094, range 0.090– 0.103 s in A. blanci [n=6]) and higher dominant frequency (X¯ =3.52, range 3.45–3.59 kHz in A. dewynteri [n=4] vs. X¯ =4.75, range 4.48–5.41 kHz in A. blanci [n=6]). Anomaloglossus dewynteri can only be distinguished from the most similar A. degranvillei (Figure 5) by its call consisting of longer notes (X¯ =0.228, range 0.221– 0.237 s [n=3] in A. dewynteri [n=4] vs. X¯ =0.158, range 0.157– 0.160 s in A. degranvillei [n=2]) (Figure 3; Table 2). Molecular data provided by Vacher et al. (2017) demonstrated that these two species harbour well-differentiated mtDNA lineages (2.3% on 16S) and our phylogenetic analysis indicates that A. dewynteri is not the sister species of A. degranvillei (see below). Anomaloglossus dewynteri could possibly be distinguished from A. degranvillei by its more tuberculate skin and by its darker dorsal and lateral colouration. However, these two features should be taken with caution given the small number of specimens available and the paucity of available data from living specimens. Description of the holotype (Figure 7). An adult male 19.9 mm SVL; body robust; head wider than long, HL 85% of HW; HL 32% of SVL; dorsal skin tuberculate, large tubercle on eyelids, snout large, rounded to nearly truncate in dorsal view, acute in lateral view, extending past lower jaw, SL 56% of HL. Nares located anterolaterally; canthus rostralis rounded but well defined, loreal region concave; IN 40% of HW; EN 31% of HL, 77% of ED. Tympanum indistinct; supratympanic fold present, extending from ventrodorsal corner of the eye onto the upper arm; choanae round, small, located anterolaterally. Forelimb slender, skin tuberculate; metacarpal ridge present; HAND 25% of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened without webbing, lateral fringes present on preaxial edges of Fingers II and III; Finger III not distinctly swollen; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers appressed; finger discs expanded, wider than long, about 1.5X width of digit; width of disc on Finger III 0.8 mm. Relative lengths of adpressed fingers III> IV> I> II; palmar tubercle large, quadrangular. 1.0 mm in diameter (larger than Finger III disc), thenar tubercle small (equal to Finger III disc), elliptic, half the size of palmar tubercle, well separated from palmar tubercle. Only basal subarticular tubercles of Fingers III and IV are conspicuous; Finger I subarticular tubercle largest followed by Finger II subarticular tubercle, basal subarticular tubercle on Finger III and IV smaller, subequal. The holotype has a missing left hand (all the individuals collected in 2016 had some atrophied or missing fingers/toes). Hind limb robust, skin tuberculate; TL 47% of SVL; heels not in contact when hind limbs are flexed at right angle to sagittal plane of body; FL 42% of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; discs on toes larger than width of toes; disc on Toe I only slightly larger than width of digit. Size of disc on Toe IV 1.0 mm. Feet moderately webbed, webbing present between Toes I–IV, webbing without melanophores; lateral fringes present on all toes. Toe webbing formula I 0- -1- II 1 ½-2- III 1 - -3 IV 3 - -2+ V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.7 mm in length, outer metatarsal tubercle round, protuberant, 0.5 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end, extending distally to the fringe on preaxial edge of Toe I. Metatarsal fold strong. Colour of holotype in life (Figure 7). Dorsal colour dark brown, faint dorsolateral stripe, upper flanks black becoming slightly lighter ventrally with a few white-blue freckles. Supratympanic fold black bordered by a discontinuous orange postocular stripe. Upper lip brown, loreal and internarial region black. Throat solid black becoming slightly paler posteriorly, belly and ventral parts of legs dark grey with light blue freckles. Iris with reddish metallic pigmentation and pupil ring interrupted ventrally by transversal pigmentation. Upper and posterior surfaces of arm pale orange, black anteriorly (same colour as throat). Dorsal surfaces of thigh, shank and tarsus with diffuse combination of orange, pale brown and two wide poorly defined dark brown cross bands; more cross bands on tarsus. Paracloacal marks inconspicuous. Toes and digits with small light blue dots. Palms and soles black. Colour of holotype in preservative. After one year in preservative (70% ethanol), the specimen faded and he dorsal colouration now varies from brown to grey. The bluish freckles turned cream as well as the orange and reddish marks. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. There is only slight variation among specimens of the type series. However, specimens photographed in 2010 displayed a dorsal colouration going from brown with black blotches to uniform brick; flank colouration black with various pattern of brown ventroposterior marks as well as white and bluish dots (Figure 6). Advertisement call (Figure 3). Three specimens (one uncollected) calling from the leaf litter near a stream were recorded from a distance of about 1 m and at temperatures ranging from 21 to 24°C. They emitted single pulsed notes (note length X¯ =0.228, range 0.221– 0.237 s [n=3]) at a regular pace (inter-note interval X¯ =0.761, range 0.600– 0.883 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 3.52 kHz on average (range 3.45–3.59 kHz) with a slight upward modulation (ca. 0.2 kHz). Distribution and ecology. Anomaloglossus dewynteri is only known from two rocky streams close to each other at 600 m elevation, on the slope of the Itoupé massif, an isolated mountain reaching 800 m elevation in the south-eastern part of French Guiana. Males call from the banks of these streams during the day and their activity seems fostered by rainfall. Many individuals were observed during the first exploration of the massif in 2010. However, only four individuals were detected in 2016 despite eight days of searching by four herpetologists, and despite any obvious perturbation of the habitat. This species co-occurs with A. surinamensis on the massif, but the two species were not found together along the same streams. Anomaloglossus dewynteri has the same reproductive mode as A. blanci and A. degranvillei, with endotrophic tadpoles carried by the male until metamorphosis. In 2010, one individual carrying six metamorphs at similar stages (Gosner stages 42–43) was observed. We cannot exclude that these metamorphs are not from two different clutches of smaller size. Comparison with the type series of Anomaloglossus degranvillei. Our examination of the original type series used by Lescure (1975) to describe Anomaloglossus degranvillei shows that it included three distinct species of the A. degranvillei clade occurring in French Guiana. Based on the original description and according to the characters observed in the holotype, we restrict the type series to the holotype and three of the original paratypes (MNHNP 1973- 1656–58) from Atachi Bakka (3.5455 N 53.9068 W). Paratypes MNHNP 1659–63 and MNHNP 1667–70 from Eaux Clément (4.6556 N 52.2480 W) and MNHNP 1665-66 from Cacao (4.5610 N 52.4629 W) correspond to A. blanci based on the following combination of characters: (1) body size comprised between 15.9–18.8 mm in males; (2) belly pale grey with small white freckles; and (3) protruding snout in lateral view. Paratype MNHNP 1973-1672 from Galbao (3.600 N 53.2833 W), MNHNP 1664 from Saül (3.62556 N 53.2072 W) and MNHNP 1676 from Crique Grégoire (5.0975 N 53.0506 W) correspond to A. surinamensis as inferred from the following combination of characters: (1) small body size (between 14.0– 15.3 mm in males 16.0– 17.1 mm in females); (2) snout short and rounded in lateral view; (3) toes moderately webbed, with fringes (4) abdomen and ventral side of legs cream with brown freckles. Unfortunately, we could not examine LACM 44211-12 from Crique Ipoussing (4.1307 N 52.5760 W), LG870–873 from Eaux Clément (4.6556 N 52.2480 W) and LACM 44213–23 from Montagne Tortue (4.300 N 52.3667 W), but these localities lie within the range of Anomaloglossus blanci and, to our knowledge, no other species of the group occur there. These paratypes are thus excluded from the type series of A. degranvillei.

Anomaloglossus blanci sp. nov. Colostethus degranvillei ��� Lescure 1975 (in part): 414; Lescure & Marty 2001 (in part): 86 Anomaloglossus degranvillei ��� Fouquet et al. 2007 (in part): Fig. S2; Fouquet et al. 2012 (in part): 469 Anomaloglossus sp. ���north FG��� ��� Vacher et al. (2017) Holotype. MNHN2017.0103 (field no. AF0953), an adult male, collected by Michel Blanc and Ma��l Dewynter, 10 August 2014, corridor 7 on Route Nationale 2, French Guiana, 4.0639 N 52.0416 W, ~ 100 m elevation (Figure 2). Paratypes. Ten specimens: MNHN2017.0110 (field no. AF0952), an adult male, collected with the holotype (carrying two froglets on its back). MNHN2017.0105 (field no. AF0293), an adult male collected by Antoine Fouquet, 25 May 2007, crique Diamant near Patawa Camp, French Guiana, 4.5161 N 52.1005 W, ~ 200 m elevation. MNHN2017.0106���8 (field no. AF0878, AF0932, AF0933), three adult males collected by Antoine Fouquet, 0 1 February 2013, crique Diamant near Patawa Camp, French Guiana, 4.5161 N 52.1005 W, ~ 200 m elevation. MNHN2017.0104 (field no. AF0292), an adult female collected by Antoine Fouquet, 25 May 2007, crique Diamant near Patawa Camp, French Guiana, 4.5161 N 52.1005 W, ~ 200 m elevation. MNHN2017.0109 (field no. AF1332), an adult male collected by Ma��l Dewynter, 16 December 2013, Saut Grand Machicou, French Guiana, 3.8974 N 52.5836 W, ~ 150 m elevation. IRSNB4198 (field no. PK3287), an adult male collected by Philippe J. R. Kok, 18 April 2010, near Patawa Camp, French Guiana, ca. 4.5161 N 52.1005 W, ~ 200 m elevation. IRSNB4199, 4200 (field no. PK3284, 3286), two juvenile females collected by Philippe J. R. Kok, 18 April 2010, near Patawa Camp, French Guiana, ca. 4.5161 N 52.1005 W, ~ 200 m elevation. Etymology. This species is dedicated to our friend Michel Blanc, in honour of his invaluable contribution to field herpetology in French Guiana, notably the discovery of many previously undocumented species and crucial natural history observations. Definition and diagnosis. We assigned the new species to the genus Anomaloglossus based on previous studies (Fouquet et al. 2012; Vacher et al. 2017) and the presence of a median lingual process. The new species belongs to the A. degranvillei clade (Vacher et al. 2017). (1) Small-sized Anomaloglossus (average male SVL= 16.9 mm [15.9���18.8, n=8], female SVL= 17.9 mm [n=1]) (Table 1); (2) body robust; (3) skin tuberculate on dorsum (particularly the posterior half) and legs, with a larger tubercle on each eyelid, ventral skin smooth; (4) conspicuous dark brown to black glandulous supratympanic fold anteroventrally bordered by a light stripe (orange in life, cream in preservative) extending from the ventroposterior edge of the eye onto the upper arm to the anterior cubital region; (5) tympanum indistinct; (6) snout short and protruding in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II shorter than Finger I when fingers adpressed; (9) tip of Finger IV almost reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle on Finger III and IV indistinct; (11) Finger III not distinctly swollen in males; (12) Fingers II and III with preaxial fringes particularly developed in males, inconspicuous in females; (13) toes moderately webbed, with well-developed fringes on Toes II, III and IV (sensu Grant et al. 2006; keel-like lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved; (15) no black arm gland in males (sensu Grant & Castro 1998, see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark ill-defined but present (reddish in life, cream speckled with melanophores in preservative); (18) dorsolateral stripe present, faint, upper part of flanks much darker than dorsum, a thin middorsal raphe going from the tip of snout to behind the head and more rarely to the vent sometimes present; (19) ventrolateral stripe absent but ventral part of flanks lighter with white or bluish flecks (in life); (20) sexual dichromatism in throat colour pattern present, throat black in reproductive males, evenly grey with small ill-defined white dots in females; (21) no sexual dichromatism in ventral colour pattern, abdomen mostly pale grey with small ill-defined white dots in both sexes, lower abdomen and ventral surface of legs yellowish in life and cream in preservative; (22) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (23) median lingual process longer than wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (24) single note call of 0.090��� 0.103 s length and dominant frequency at 4.48���5.41 kHz (n=6) (Figure 3; Table 2). Morphological comparisons with other lowland Anomaloglossus. The only other species group cooccurring with the Anomaloglossus degranvillei group is the A. stepheni group, represented in French Guiana by A. baeobatrachus, which is readily distinguishable by conspicuous dorsolateral stripes, swollen third finger in males and absence of webbing and fringes on toes. Within the Anomaloglossus degranvillei group, A. blanci can be distinguished from A. surinamensis (Figure 4) by (1) a shorter snout, protruding in lateral view (vs. rounded in A. surinamensis); (2) a larger body size (X�� =16.9; range 15.9���18.8 mm in males [n=8] and one adult female measuring 17.9 mm in A. blanci vs. X�� =14.75, range 14.0��� 15.3 mm in males [n=8] and X�� =16.5, range 16.0��� 17.1 mm in females [n=4] in A. surinamensis); (3) large fringes on fingers and toes in males (narrow in A. surinamensis); (4) presence of a large tubercle on the top of the eyelid (inconspicuous in A. surinamensis); (5) ventral colouration entirely grey with small ill-defined white dots (vs. grey throat and cream belly, the latter with irregular grey blotches less abundant posteriorly in A. surinamensis); (6) yellowish ventral surface of thigh (vs. light grey and more heavily spotted with black in A. surinamensis); (7) large MLP (> 0.5 mm vs. A. surinamensis); (8) call characterized by longer pulsed notes (X�� =0.094, range 0.090��� 0.103 s in A. blanci [n=6] vs. tonal note X�� =0.032, range 0.028��� 0.037 s in A. surinamensis [n=20]) emitted between longer intervals (X�� =1.414, range 1.200��� 1.906 s in A. blanci [n=6] vs. X�� =0.573, range 0.372��� 0.825 s in A. surinamensis [n=20]), but both species have a similar dominant frequency (X�� =4.75, range 4.48���5.41 kHz in A. blanci [n=6] vs. X�� =4.89 kHZ, range 4.55���5.35 kHz in A. surinamensis [n=20]). Anomaloglossus blanci can be distinguished from A. degranvillei (Figure 5) by (1) a smaller size (X�� =16.9; range 15.9���18.8 mm in males [n=8] and one female measuring 17.9 mm in A. blanci vs. X�� =20.17; range 19.60���20.50 mm in males [n=3] and X�� =23.40, range 22.4���23.9 mm in females [n=5] of A. degranvillei); (2) belly pale grey with small ill-defined white dots (from solid black to dark grey with abundant small conspicuous white spots in A. degranvillei); (3) ventral surface of thighs yellowish in life and cream in preservative (dark grey in A. degranvillei); (4) call with shorter notes (0.090��� 0.103 s in A. blanci [n=6] vs. 0.157��� 0.160 s in A. degranvillei [n=2]) and higher dominant frequency (4.48���5.41 kHz in A. blanci [n=6] vs. 3.60��� 3.62 s in A. degranvillei [n=2]). Description of the holotype (Figure 2). An adult male, 17.1 mm SVL; body robust; head wider than long, HL 89% of HW; HL 33% of SVL; dorsal skin tuberculate, one enlarged tubercle on each eyelid, snout long (SL 53% of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 42% of HW; EN 28% of HL, 70% of ED. Tympanum indistinct; supratympanic fold present extending from posteroventral corner of the eye onto the upper arm; choanae small, circular, located anterolaterally. Forelimb slender, skin tuberculate; metacarpal ridge absent; HAND 24% of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened without webbing, lateral fringes present on preaxial edges of Fingers II and III; Finger III not distinctly swollen; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5X width of digit; width of disc on Finger III 0.6 mm; discs with distinct dorsal scutes. Relative lengths of adpressed fingers III> IV> I> II; palmar tubercle large, heart-shaped, 0.8 mm in diameter (larger than Finger III disc), thenar tubercle small (equal to Finger III disc), elliptic, half the size of palmar tubercle, well separated from palmar tubercle. Only basal subarticular tubercles on Fingers III and IV are conspicuous; Finger I subarticular tubercle largest followed by Finger II subarticular tubercle, basal subarticular tubercle on Finger III and IV smaller, subequal. Hind limb robust, skin tuberculate; TL 46 % of SVL; heels not in contact when hind limbs are flexed at right angle to sagittal plane of body; FL 42% of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes; disc on Toe I only slightly larger than width of digit. Width of disc on Toe IV 0.8 mm. Feet moderately webbed, webbing present between Toes I���IV, webbing without melanophores; lateral fringes present on all toes. Toe webbing formula I 1 ��-1- II 1 ��-2- III 1 - -3 IV 3 - -2+ V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end, extending distally to the fringe on preaxial edge of Toe I. Metatarsal fold strong. Colour of holotype in life (Figure 2). Dorsal colour chestnut brown, with a diffuse dark brown hourglass pattern extending from the interorbital region to middorsum immediately followed by a diffuse dark brown patch over the sacrum. Faint dorsolateral stripes. Black lateral mask extending from tip of snout to the upper part of flanks, tapering to groin and containing most of the indistinct tympanum and supratympanic fold, only interrupted by a large postocular orange stripe extending from eye onto the upper arm down to the anterior cubital region. Upper lip with 2���3 small light blue blotches. Posteroventral part of flanks pale brown with a few small white-blue and dark brown blotches. Throat solid black becoming paler posteriorly; belly grey with ill-defined light blue and white spots, lower part of belly and ventral surface of thighs and arms orange with a few ill-defined grey spots. Iris with reddish metallic pigmentation and pupil ring interrupted dorsally and ventrally by transversal pigmentation. Upper arm pale orange dorsally, black on its anterior part (same colour as throat), and dark brown with illdefined orange marbling on its posterior part. Lower arm light brown with two ill-defined dark brown cross bands. Dorsal surfaces of thigh, shank and tarsus with diffuse combination of orange, pale brown and a wide ill-defined dark brown cross band; more cross bands on tarsus. Reddish paracloacal marks. Toes and digits with small light blue dots. Palms and soles dark grey. Colour of holotype in preservative. After three years in preservative (70% ethanol), the specimen faded and the dorsal colouration now varies from brown to grey. The bluish freckles turned cream as did the orange and reddish marks. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. Intraspecific variation is high and adult dorsal colouration varies from brown to reddish brown. Most specimens have dark brown blotches on dorsum. One male (MNHN 2017.0108/AF0933) has a middorsal raphe running from the tip of the snout to the vent; in four males (MNHN 2017.0105���7/AF0293, 0 878, 0 932 and MNHN 2017.0109/AF1332; Figure 6) that raphe is inconspicuous and extends only from the tip of the snout to the interorbital region. Flanks can be dark brown to light grey rendering the lateral black band more or less conspicuous. Additionally, overall colouration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as males carrying tadpole apparently display overall lighter colours, smoother skin and sharper contrasts while calling males are very dark and highly tuberculate. Advertisement call (Figure 3). Six specimens (three uncollected) calling from the leaf litter 1���3 meters away from a stream were recorded from a distance of about 1 m and at temperatures ranging from 23 to 26��C. They emitted single short pulsed notes (note length X�� = 0.094 s; range 0.090��� 0.103 s) at a regular pace (inter-note interval X�� = 1.414 s; range 1.200��� 1.906 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 4.75 kHz on average (range 4.48���5.41 kHz) with a slight upward modulation (ca. 0.1 kHz). Distribution and ecology. Anomaloglossus blanci inhabits small sandy or rocky streams at low elevations (from 50 to 200 m elevation) often close to larger tributaries (e.g. Crique Diamant, Approuague). Males call from stream banks during the day with peak in intensity at dawn (6���7 am) and late afternoon (4���5 pm) during the rainy season, but also during humid days of the dry season. Eighteen populations have been documented in north-eastern French Guiana (Figure 1), but despite intensive surveys of these populations in 2016, no individual could be found in three of them. No noticeable degradation of the habitat has been observed. Interestingly, this species does not co-occur with A. surinamensis, whose populations seem stable. No tadpole before Gosner stage 41 has been observed, either in the water or carried by males. However, two observations have been made of adults carrying tadpoles at a late development stage (> stage 41) (Figure 6). The tadpoles are endotrophic and lack functional mouth. If they are detached from the back of their father, they die within a few hours despite humid and ideal thermal conditions (pers. obs.). These observations suggest that the tadpoles may undergo a phase of nidicolous development after hatching and only complete their development on the back of the adult from an advanced stage. Only 1���4 tadpoles of the same stage were observed at a time carried by adults. This reduced clutch size is likely linked to the fact that large eggs are needed to complete endotrophic development. However, adults were often observed carrying tadpoles at two different stages, demonstrating that they can keep mating while already attending a clutch. This is likely linked to the extended time needed for the tadpoles to develop until complete metamorphosis., Published as part of Antoine, Fouquet, Vacher, Jean-Pierre, Courtois, Elodie A., Villette, Benoit, Reizine, Hugo, Gaucher, Philippe, Jairam, Rawien, Ouboter, Paul & Kok, Philippe J. R., 2018, On the brink of extinction: two new species of Anomaloglossus from French Guiana and amended definitions of Anomaloglossus degranvillei and A. surinamensis (Anura: Aromobatidae), pp. 1-23 in Zootaxa 4379 (1) on pages 5-10, DOI: 10.11646/zootaxa.4379.1.1, http://zenodo.org/record/1172286, {"references":["Lescure, J. (1975) Contribution a l'etude des amphibiens de Guyane francaise. III. Une nouvelle espece de Colostethus (Dendrobatidae): Colostethus degranvillei nov. sp. Bulletin du Museum National d'Histoire Naturelle, Zoologie, 203, 413 - 420.","Vacher, J. - P., Kok, P. J. R., Rodrigues, M. T., Dias Lima, J., Lorenzini, A., Martinez, Q., Fallet, M., Courtois, E. A., Blanc, M., Gaucher, P., Dewynter, M., Jairam, R., Ouboter, P., Thebaud, C. & Fouquet, A. (2017) Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield. Molecular Phylogenetics and Evolution, 112, 158 - 173. https: // doi. org / 10.1016 / j. ympev. 2017.04.017","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262. https: // doi. org / 10.1206 / 0003 - 0090 (2006) 299 [1: PSODFA] 2.0. CO; 2","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147. https: // doi. org / 10.1206 / 308.1","Grant, T. & Castro, F. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology, 32, 378 - 392. https: // doi. org / 10.2307 / 1565452"]}

Anomaloglossus dewynteri sp. nov. Anomaloglossus sp. ��� Itoup����� Vacher et al. 2017 Holotype. MNHN2017.0111 (field no. AF3644/APA-973-23-2), an adult male, collected by Antoine Fouquet, 11 January 2016, near base camp on Mont Itoup��, French Guiana, 3.0230 N 53.0955 W, elevation ca. ~ 600 m (Figure 7). Paratopotypes. Three specimens: MNHN2017.0112 (field no. AF0574; not measured because badly preserved), an adult male, collected by Ma��l Dewynter, 0 5 April 2010, near base camp on Mont Itoup��, French Guiana, 3.0230 N 53.0955 W, ~ 600 m elevation. MNHN2017.0113���4 (field no. AF3686/APA-973-23-1, AF3645/ APA-973-23-3), two adult males, collected by Antoine Fouquet, with the holotype. Etymology. This species is dedicated to our friend Ma��l Dewynter for his invaluable contribution to the herpetology of French Guiana. Adult definition and diagnosis. We assigned the new species to the genus Anomaloglossus based on previous studies (Fouquet et al. 2012; Vacher et al. 2017) and the presence of a median lingual process. The new species belongs to the A. degranvillei clade according to Vacher et al. (2017). (1) Medium-sized Anomaloglossus (average male SVL= 19.9 mm [19.4���20.4, n=3], female unknown) (Table 1); (2) body robust; (3) skin tuberculate on dorsum (particularly the posterior half) and legs, with a larger tubercle on each eyelid, ventral skin smooth; (4) conspicuous dark brown to black glandulous supratympanic fold flanked anteroventrally by a discontinuous stripe (orange in life, cream in preservative) running from the ventroposterior edge of the eye and extending onto the upper arm; (5) tympanum indistinct; (6) snout short and acute in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II shorter than Finger I when fingers adpressed; (9) tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle on Finger III and IV indistinct; (11) Finger III not distinctly swollen in males; (12) fingers with fringes particularly developed on preaxial edges of Fingers II and III; (13) toes moderately webbed, with well-developed fringes (sensu Grant et al. 2006; keel-like lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved, slightly tuberclelike; (15) no black arm gland in males (sensu Grant & Castro 1998. see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark inconspicuous; (18) dorsolateral stripe absent or inconspicuous, dorsal part of flanks darker than dorsum; (19) ventrolateral stripe sometimes present as a broken, poorly defined pale brown blotch, ventral part of flanks with flecks white or bluish (in life); (20) sexual dichromatism in throat colour pattern unknown (although likely because present in other species of the clade), throat solid black in reproductive males; (21) abdomen and ventral side of legs dark grey with small ill-defined white dots; (22) iris with metallic pigmentation and pupil ring interrupted ventrally by transversal black pigmentation; (23) median lingual process longer than wide, tapered, bluntly pointed, smooth (nonpapillate), reclined in pit; (24) single note call of 0.221��� 0.237 s length and dominant frequency at 3.45���3.59 kHz (n=3) (Table 2; Figure 3). Morphological comparisons with other lowland Anomaloglossus. The only other species group cooccurring with the Anomaloglossus degranvillei species group is the A. stepheni species group, represented by A. baeobatrachus in French Guiana, which is readily distinguishable by conspicuous dorsolateral stripes, swollen third fingers in males and absence of webbing and fringes on toes. Within the Anomaloglossus degranvillei group, A. dewynteri can be distinguished from A. surinamensis (Figure 4) by (1) a shorter snout, acute in lateral view (vs. rounded in A. surinamensis); (2) a larger body size in males (X�� = 19.9 mm, range 19.4���20.4 mm in A. dewynteri [n=3] vs. X�� =14.75, range 14.0��� 15.3 mm in A. surinamensis [n=8]); (3) more developed fringes on fingers and toes; (4) presence of a large tubercle on the top of the eyelid (inconspicuous in A. surinamensis); (5) belly dark grey with small white spots (vs. cream with irregular grey blotches less abundant posteriorly in A. surinamensis); (6) ventral surface of thigh dark grey (vs. cream with dark spots on the edges in A. surinamensis); (7) larger MLP (> 0.5 mm vs A. surinamensis; (8) call characterized by much longer pulsed notes (pulsed note X�� =0.228, range 0.221��� 0.237 s [n=3] in A. dewynteri vs. tonal note X�� =0.032, range 0.028��� 0.037 s in A. surinamensis [n=20]) emitted between longer intervals (X�� =0.76, range 0.60��� 0.88 s in A. dewynteri [n=3] vs. X�� =0.573, range 0.372��� 0.825 s in A. surinamensis [n=20]) and with lower dominant frequency (X�� =3.52, range 3.45���3.59 kHz in A. dewynteri [n=4] vs. X�� =4.89 kHZ, range 4.55���5.35 kHz in A. surinamensis [n=20]). Anomaloglossus dewynteri can be distinguished from A. blanci (Figure 2) by (1) its larger size in males (X= 19.9 mm, range 19.4���20.4 mm in A. dewynteri [n=3] vs. X�� =16.9, range 15.9���18.8 mm in A. blanci [n=8]); (2) belly from solid black to dark grey with abundant small conspicuous white spots (pale grey with small white illdefined spots in A. blanci); (3) ventral surface of thighs dark grey (yellowish in life and cream in preservative in A. blanci); (4) strong metacarpal ridge (inconspicuous in A. blanci, Figures 2F, 7F); (5) call with longer notes (X=0.228, range 0.221��� 0.237 s [n=3] in A. dewynteri vs. X�� =0.094, range 0.090��� 0.103 s in A. blanci [n=6]) and higher dominant frequency (X�� =3.52, range 3.45���3.59 kHz in A. dewynteri [n=4] vs. X�� =4.75, range 4.48���5.41 kHz in A. blanci [n=6]). Anomaloglossus dewynteri can only be distinguished from the most similar A. degranvillei (Figure 5) by its call consisting of longer notes (X�� =0.228, range 0.221��� 0.237 s [n=3] in A. dewynteri [n=4] vs. X�� =0.158, range 0.157��� 0.160 s in A. degranvillei [n=2]) (Figure 3; Table 2). Molecular data provided by Vacher et al. (2017) demonstrated that these two species harbour well-differentiated mtDNA lineages (2.3% on 16S) and our phylogenetic analysis indicates that A. dewynteri is not the sister species of A. degranvillei (see below). Anomaloglossus dewynteri could possibly be distinguished from A. degranvillei by its more tuberculate skin and by its darker dorsal and lateral colouration. However, these two features should be taken with caution given the small number of specimens available and the paucity of available data from living specimens. Description of the holotype (Figure 7). An adult male 19.9 mm SVL; body robust; head wider than long, HL 85% of HW; HL 32% of SVL; dorsal skin tuberculate, large tubercle on eyelids, snout large, rounded to nearly truncate in dorsal view, acute in lateral view, extending past lower jaw, SL 56% of HL. Nares located anterolaterally; canthus rostralis rounded but well defined, loreal region concave; IN 40% of HW; EN 31% of HL, 77% of ED. Tympanum indistinct; supratympanic fold present, extending from ventrodorsal corner of the eye onto the upper arm; choanae round, small, located anterolaterally. Forelimb slender, skin tuberculate; metacarpal ridge present; HAND 25% of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened without webbing, lateral fringes present on preaxial edges of Fingers II and III; Finger III not distinctly swollen; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers appressed; finger discs expanded, wider than long, about 1.5X width of digit; width of disc on Finger III 0.8 mm. Relative lengths of adpressed fingers III> IV> I> II; palmar tubercle large, quadrangular. 1.0 mm in diameter (larger than Finger III disc), thenar tubercle small (equal to Finger III disc), elliptic, half the size of palmar tubercle, well separated from palmar tubercle. Only basal subarticular tubercles of Fingers III and IV are conspicuous; Finger I subarticular tubercle largest followed by Finger II subarticular tubercle, basal subarticular tubercle on Finger III and IV smaller, subequal. The holotype has a missing left hand (all the individuals collected in 2016 had some atrophied or missing fingers/toes). Hind limb robust, skin tuberculate; TL 47% of SVL; heels not in contact when hind limbs are flexed at right angle to sagittal plane of body; FL 42% of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; discs on toes larger than width of toes; disc on Toe I only slightly larger than width of digit. Size of disc on Toe IV 1.0 mm. Feet moderately webbed, webbing present between Toes I���IV, webbing without melanophores; lateral fringes present on all toes. Toe webbing formula I 0- -1- II 1 ��-2- III 1 - -3 IV 3 - -2+ V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.7 mm in length, outer metatarsal tubercle round, protuberant, 0.5 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end, extending distally to the fringe on preaxial edge of Toe I. Metatarsal fold strong. Colour of holotype in life (Figure 7). Dorsal colour dark brown, faint dorsolateral stripe, upper flanks black becoming slightly lighter ventrally with a few white-blue freckles. Supratympanic fold black bordered by a discontinuous orange postocular stripe. Upper lip brown, loreal and internarial region black. Throat solid black becoming slightly paler posteriorly, belly and ventral parts of legs dark grey with light blue freckles. Iris with reddish metallic pigmentation and pupil ring interrupted ventrally by transversal pigmentation. Upper and posterior surfaces of arm pale orange, black anteriorly (same colour as throat). Dorsal surfaces of thigh, shank and tarsus with diffuse combination of orange, pale brown and two wide poorly defined dark brown cross bands; more cross bands on tarsus. Paracloacal marks inconspicuous. Toes and digits with small light blue dots. Palms and soles black. Colour of holotype in preservative. After one year in preservative (70% ethanol), the specimen faded and he dorsal colouration now varies from brown to grey. The bluish freckles turned cream as well as the orange and reddish marks. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. There is only slight variation among specimens of the type series. However, specimens photographed in 2010 displayed a dorsal colouration going from brown with black blotches to uniform brick; flank colouration black with various pattern of brown ventroposterior marks as well as white and bluish dots (Figure 6). Advertisement call (Figure 3). Three specimens (one uncollected) calling from the leaf litter near a stream were recorded from a distance of about 1 m and at temperatures ranging from 21 to 24��C. They emitted single pulsed notes (note length X�� =0.228, range 0.221��� 0.237 s [n=3]) at a regular pace (inter-note interval X�� =0.761, range 0.600��� 0.883 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 3.52 kHz on average (range 3.45���3.59 kHz) with a slight upward modulation (ca. 0.2 kHz). Distribution and ecology. Anomaloglossus dewynteri is only known from two rocky streams close to each other at 600 m elevation, on the slope of the Itoup�� massif, an isolated mountain reaching 800 m elevation in the south-eastern part of French Guiana. Males call from the banks of these streams during the day and their activity seems fostered by rainfall. Many individuals were observed during the first exploration of the massif in 2010. However, only four individuals were detected in 2016 despite eight days of searching by four herpetologists, and despite any obvious perturbation of the habitat. This species co-occurs with A. surinamensis on the massif, but the two species were not found together along the same streams. Anomaloglossus dewynteri has the same reproductive mode as A. blanci and A. degranvillei, with endotrophic tadpoles carried by the male until metamorphosis. In 2010, one individual carrying six metamorphs at similar stages (Gosner stages 42���43) was observed. We cannot exclude that these metamorphs are not from two different clutches of smaller size. Comparison with the type series of Anomaloglossus degranvillei. Our examination of the original type series used by Lescure (1975) to describe Anomaloglossus degranvillei shows that it included three distinct species of the A. degranvillei clade occurring in French Guiana. Based on the original description and according to the characters observed in the holotype, we restrict the type series to the holotype and three of the original paratypes (MNHNP 1973- 1656���58) from Atachi Bakka (3.5455 N 53.9068 W). Paratypes MNHNP 1659���63 and MNHNP 1667���70 from Eaux Cl��ment (4.6556 N 52.2480 W) and MNHNP 1665-66 from Cacao (4.5610 N 52.4629 W) correspond to A. blanci based on the following combination of characters: (1) body size comprised between 15.9���18.8 mm in males; (2) belly pale grey with small white freckles; and (3) protruding snout in lateral view. Paratype MNHNP 1973-1672 from Galbao (3.600 N 53.2833 W), MNHNP 1664 from Sa��l (3.62556 N 53.2072 W) and MNHNP 1676 from Crique Gr��goire (5.0975 N 53.0506 W) correspond to A. surinamensis as inferred from the following combination of characters: (1) small body size (between 14.0��� 15.3 mm in males 16.0��� 17.1 mm in females); (2) snout short and rounded in lateral view; (3) toes moderately webbed, with fringes (4) abdomen and ventral side of legs cream with brown freckles. Unfortunately, we could not examine LACM 44211-12 from Crique Ipoussing (4.1307 N 52.5760 W), LG870���873 from Eaux Cl��ment (4.6556 N 52.2480 W) and LACM 44213���23 from Montagne Tortue (4.300 N 52.3667 W), but these localities lie within the range of Anomaloglossus blanci and, to our knowledge, no other species of the group occur there. These paratypes are thus excluded from the type series of A. degranvillei., Published as part of Antoine, Fouquet, Vacher, Jean-Pierre, Courtois, Elodie A., Villette, Benoit, Reizine, Hugo, Gaucher, Philippe, Jairam, Rawien, Ouboter, Paul & Kok, Philippe J. R., 2018, On the brink of extinction: two new species of Anomaloglossus from French Guiana and amended definitions of Anomaloglossus degranvillei and A. surinamensis (Anura: Aromobatidae), pp. 1-23 in Zootaxa 4379 (1) on pages 10-13, DOI: 10.11646/zootaxa.4379.1.1, http://zenodo.org/record/1172286, {"references":["Vacher, J. - P., Kok, P. J. R., Rodrigues, M. T., Dias Lima, J., Lorenzini, A., Martinez, Q., Fallet, M., Courtois, E. A., Blanc, M., Gaucher, P., Dewynter, M., Jairam, R., Ouboter, P., Thebaud, C. & Fouquet, A. (2017) Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield. Molecular Phylogenetics and Evolution, 112, 158 - 173. https: // doi. org / 10.1016 / j. ympev. 2017.04.017","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262. https: // doi. org / 10.1206 / 0003 - 0090 (2006) 299 [1: PSODFA] 2.0. CO; 2","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147. https: // doi. org / 10.1206 / 308.1","Grant, T. & Castro, F. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology, 32, 378 - 392. https: // doi. org / 10.2307 / 1565452","Lescure, J. (1975) Contribution a l'etude des amphibiens de Guyane francaise. III. Une nouvelle espece de Colostethus (Dendrobatidae): Colostethus degranvillei nov. sp. Bulletin du Museum National d'Histoire Naturelle, Zoologie, 203, 413 - 420."]}

Anomaloglossus blanci sp. nov. Colostethus degranvillei — Lescure 1975 (in part): 414; Lescure & Marty 2001 (in part): 86 Anomaloglossus degranvillei — Fouquet et al. 2007 (in part): Fig. S2; Fouquet et al. 2012 (in part): 469 Anomaloglossus sp. “north FG” — Vacher et al. (2017) Holotype. MNHN2017.0103 (field no. AF0953), an adult male, collected by Michel Blanc and Maël Dewynter, 10 August 2014, corridor 7 on Route Nationale 2, French Guiana, 4.0639 N 52.0416 W, ~ 100 m elevation (Figure 2). Paratypes. Ten specimens: MNHN2017.0110 (field no. AF0952), an adult male, collected with the holotype (carrying two froglets on its back). MNHN2017.0105 (field no. AF0293), an adult male collected by Antoine Fouquet, 25 May 2007, crique Diamant near Patawa Camp, French Guiana, 4.5161 N 52.1005 W, ~ 200 m elevation. MNHN2017.0106–8 (field no. AF0878, AF0932, AF0933), three adult males collected by Antoine Fouquet, 0 1 February 2013, crique Diamant near Patawa Camp, French Guiana, 4.5161 N 52.1005 W, ~ 200 m elevation. MNHN2017.0104 (field no. AF0292), an adult female collected by Antoine Fouquet, 25 May 2007, crique Diamant near Patawa Camp, French Guiana, 4.5161 N 52.1005 W, ~ 200 m elevation. MNHN2017.0109 (field no. AF1332), an adult male collected by Maël Dewynter, 16 December 2013, Saut Grand Machicou, French Guiana, 3.8974 N 52.5836 W, ~ 150 m elevation. IRSNB4198 (field no. PK3287), an adult male collected by Philippe J. R. Kok, 18 April 2010, near Patawa Camp, French Guiana, ca. 4.5161 N 52.1005 W, ~ 200 m elevation. IRSNB4199, 4200 (field no. PK3284, 3286), two juvenile females collected by Philippe J. R. Kok, 18 April 2010, near Patawa Camp, French Guiana, ca. 4.5161 N 52.1005 W, ~ 200 m elevation. Etymology. This species is dedicated to our friend Michel Blanc, in honour of his invaluable contribution to field herpetology in French Guiana, notably the discovery of many previously undocumented species and crucial natural history observations. Definition and diagnosis. We assigned the new species to the genus Anomaloglossus based on previous studies (Fouquet et al. 2012; Vacher et al. 2017) and the presence of a median lingual process. The new species belongs to the A. degranvillei clade (Vacher et al. 2017). (1) Small-sized Anomaloglossus (average male SVL= 16.9 mm [15.9–18.8, n=8], female SVL= 17.9 mm [n=1]) (Table 1); (2) body robust; (3) skin tuberculate on dorsum (particularly the posterior half) and legs, with a larger tubercle on each eyelid, ventral skin smooth; (4) conspicuous dark brown to black glandulous supratympanic fold anteroventrally bordered by a light stripe (orange in life, cream in preservative) extending from the ventroposterior edge of the eye onto the upper arm to the anterior cubital region; (5) tympanum indistinct; (6) snout short and protruding in lateral view; (7) nares oriented ventrolaterally, situated near tip of snout; (8) Finger II shorter than Finger I when fingers adpressed; (9) tip of Finger IV almost reaching distal subarticular tubercle on Finger III when fingers adpressed; (10) distal subarticular tubercle on Finger III and IV indistinct; (11) Finger III not distinctly swollen in males; (12) Fingers II and III with preaxial fringes particularly developed in males, inconspicuous in females; (13) toes moderately webbed, with well-developed fringes on Toes II, III and IV (sensu Grant et al. 2006; keel-like lateral folds sensu Myers & Donnelly 2008); (14) tarsal keel well-defined, curved; (15) no black arm gland in males (sensu Grant & Castro 1998, see also Grant et al. 2006); (16) cloacal tubercles present; (17) paracloacal mark ill-defined but present (reddish in life, cream speckled with melanophores in preservative); (18) dorsolateral stripe present, faint, upper part of flanks much darker than dorsum, a thin middorsal raphe going from the tip of snout to behind the head and more rarely to the vent sometimes present; (19) ventrolateral stripe absent but ventral part of flanks lighter with white or bluish flecks (in life); (20) sexual dichromatism in throat colour pattern present, throat black in reproductive males, evenly grey with small ill-defined white dots in females; (21) no sexual dichromatism in ventral colour pattern, abdomen mostly pale grey with small ill-defined white dots in both sexes, lower abdomen and ventral surface of legs yellowish in life and cream in preservative; (22) iris with metallic pigmentation and pupil ring interrupted ventrally and dorsally by transversal black pigmentation; (23) median lingual process longer than wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit; (24) single note call of 0.090– 0.103 s length and dominant frequency at 4.48–5.41 kHz (n=6) (Figure 3; Table 2). Morphological comparisons with other lowland Anomaloglossus. The only other species group cooccurring with the Anomaloglossus degranvillei group is the A. stepheni group, represented in French Guiana by A. baeobatrachus, which is readily distinguishable by conspicuous dorsolateral stripes, swollen third finger in males and absence of webbing and fringes on toes. Within the Anomaloglossus degranvillei group, A. blanci can be distinguished from A. surinamensis (Figure 4) by (1) a shorter snout, protruding in lateral view (vs. rounded in A. surinamensis); (2) a larger body size (X¯ =16.9; range 15.9–18.8 mm in males [n=8] and one adult female measuring 17.9 mm in A. blanci vs. X¯ =14.75, range 14.0– 15.3 mm in males [n=8] and X¯ =16.5, range 16.0– 17.1 mm in females [n=4] in A. surinamensis); (3) large fringes on fingers and toes in males (narrow in A. surinamensis); (4) presence of a large tubercle on the top of the eyelid (inconspicuous in A. surinamensis); (5) ventral colouration entirely grey with small ill-defined white dots (vs. grey throat and cream belly, the latter with irregular grey blotches less abundant posteriorly in A. surinamensis); (6) yellowish ventral surface of thigh (vs. light grey and more heavily spotted with black in A. surinamensis); (7) large MLP (> 0.5 mm vs. A. surinamensis); (8) call characterized by longer pulsed notes (X¯ =0.094, range 0.090– 0.103 s in A. blanci [n=6] vs. tonal note X¯ =0.032, range 0.028– 0.037 s in A. surinamensis [n=20]) emitted between longer intervals (X¯ =1.414, range 1.200– 1.906 s in A. blanci [n=6] vs. X¯ =0.573, range 0.372– 0.825 s in A. surinamensis [n=20]), but both species have a similar dominant frequency (X¯ =4.75, range 4.48–5.41 kHz in A. blanci [n=6] vs. X¯ =4.89 kHZ, range 4.55–5.35 kHz in A. surinamensis [n=20]). Anomaloglossus blanci can be distinguished from A. degranvillei (Figure 5) by (1) a smaller size (X¯ =16.9; range 15.9–18.8 mm in males [n=8] and one female measuring 17.9 mm in A. blanci vs. X¯ =20.17; range 19.60–20.50 mm in males [n=3] and X¯ =23.40, range 22.4–23.9 mm in females [n=5] of A. degranvillei); (2) belly pale grey with small ill-defined white dots (from solid black to dark grey with abundant small conspicuous white spots in A. degranvillei); (3) ventral surface of thighs yellowish in life and cream in preservative (dark grey in A. degranvillei); (4) call with shorter notes (0.090– 0.103 s in A. blanci [n=6] vs. 0.157– 0.160 s in A. degranvillei [n=2]) and higher dominant frequency (4.48–5.41 kHz in A. blanci [n=6] vs. 3.60– 3.62 s in A. degranvillei [n=2]). Description of the holotype (Figure 2). An adult male, 17.1 mm SVL; body robust; head wider than long, HL 89% of HW; HL 33% of SVL; dorsal skin tuberculate, one enlarged tubercle on each eyelid, snout long (SL 53% of HL), rounded to nearly truncate in dorsal view, protruding in lateral view, extending past lower jaw. Nares located anterolaterally; canthus rostralis rounded, loreal region concave; IN 42% of HW; EN 28% of HL, 70% of ED. Tympanum indistinct; supratympanic fold present extending from posteroventral corner of the eye onto the upper arm; choanae small, circular, located anterolaterally. Forelimb slender, skin tuberculate; metacarpal ridge absent; HAND 24% of SVL; Finger I longer than Finger II when fingers adpressed; fingers large and flattened without webbing, lateral fringes present on preaxial edges of Fingers II and III; Finger III not distinctly swollen; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.5X width of digit; width of disc on Finger III 0.6 mm; discs with distinct dorsal scutes. Relative lengths of adpressed fingers III> IV> I> II; palmar tubercle large, heart-shaped, 0.8 mm in diameter (larger than Finger III disc), thenar tubercle small (equal to Finger III disc), elliptic, half the size of palmar tubercle, well separated from palmar tubercle. Only basal subarticular tubercles on Fingers III and IV are conspicuous; Finger I subarticular tubercle largest followed by Finger II subarticular tubercle, basal subarticular tubercle on Finger III and IV smaller, subequal. Hind limb robust, skin tuberculate; TL 46 % of SVL; heels not in contact when hind limbs are flexed at right angle to sagittal plane of body; FL 42% of SVL; relative length of adpressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes adpressed; toe discs larger than width of toes; disc on Toe I only slightly larger than width of digit. Width of disc on Toe IV 0.8 mm. Feet moderately webbed, webbing present between Toes I–IV, webbing without melanophores; lateral fringes present on all toes. Toe webbing formula I 1 ½-1- II 1 ½-2- III 1 - -3 IV 3 - -2+ V. One to three subarticular tubercles on toes as follows: one on Toes I and II, two on Toes III and V, three on Toe IV. Inner metatarsal tubercle protuberant elliptical, 0.5 mm in length, outer metatarsal tubercle round, protuberant, 0.3 mm in diameter. Tarsal keel well defined, tubercle-like and strongly curved at proximal end, extending distally to the fringe on preaxial edge of Toe I. Metatarsal fold strong. Colour of holotype in life (Figure 2). Dorsal colour chestnut brown, with a diffuse dark brown hourglass pattern extending from the interorbital region to middorsum immediately followed by a diffuse dark brown patch over the sacrum. Faint dorsolateral stripes. Black lateral mask extending from tip of snout to the upper part of flanks, tapering to groin and containing most of the indistinct tympanum and supratympanic fold, only interrupted by a large postocular orange stripe extending from eye onto the upper arm down to the anterior cubital region. Upper lip with 2–3 small light blue blotches. Posteroventral part of flanks pale brown with a few small white-blue and dark brown blotches. Throat solid black becoming paler posteriorly; belly grey with ill-defined light blue and white spots, lower part of belly and ventral surface of thighs and arms orange with a few ill-defined grey spots. Iris with reddish metallic pigmentation and pupil ring interrupted dorsally and ventrally by transversal pigmentation. Upper arm pale orange dorsally, black on its anterior part (same colour as throat), and dark brown with illdefined orange marbling on its posterior part. Lower arm light brown with two ill-defined dark brown cross bands. Dorsal surfaces of thigh, shank and tarsus with diffuse combination of orange, pale brown and a wide ill-defined dark brown cross band; more cross bands on tarsus. Reddish paracloacal marks. Toes and digits with small light blue dots. Palms and soles dark grey. Colour of holotype in preservative. After three years in preservative (70% ethanol), the specimen faded and the dorsal colouration now varies from brown to grey. The bluish freckles turned cream as did the orange and reddish marks. Variation among type specimens. Measurements (range, mean, and standard deviation) of the type series are provided in Table 1. Intraspecific variation is high and adult dorsal colouration varies from brown to reddish brown. Most specimens have dark brown blotches on dorsum. One male (MNHN 2017.0108/AF0933) has a middorsal raphe running from the tip of the snout to the vent; in four males (MNHN 2017.0105–7/AF0293, 0 878, 0 932 and MNHN 2017.0109/AF1332; Figure 6) that raphe is inconspicuous and extends only from the tip of the snout to the interorbital region. Flanks can be dark brown to light grey rendering the lateral black band more or less conspicuous. Additionally, overall colouration and tuberculation may vary with light intensity, time of the day and probably reproductive activity as males carrying tadpole apparently display overall lighter colours, smoother skin and sharper contrasts while calling males are very dark and highly tuberculate. Advertisement call (Figure 3). Six specimens (three uncollected) calling from the leaf litter 1–3 meters away from a stream were recorded from a distance of about 1 m and at temperatures ranging from 23 to 26°C. They emitted single short pulsed notes (note length X¯ = 0.094 s; range 0.090– 0.103 s) at a regular pace (inter-note interval X¯ = 1.414 s; range 1.200– 1.906 s). The spectral structure of the note has a developed harmonic structure and the dominant frequency is 4.75 kHz on average (range 4.48–5.41 kHz) with a slight upward modulation (ca. 0.1 kHz). Distribution and ecology. Anomaloglossus blanci inhabits small sandy or rocky streams at low elevations (from 50 to 200 m elevation) often close to larger tributaries (e.g. Crique Diamant, Approuague). Males call from stream banks during the day with peak in intensity at dawn (6–7 am) and late afternoon (4–5 pm) during the rainy season, but also during humid days of the dry season. Eighteen populations have been documented in north-eastern French Guiana (Figure 1), but despite intensive surveys of these populations in 2016, no individual could be found in three of them. No noticeable degradation of the habitat has been observed. Interestingly, this species does not co-occur with A. surinamensis, whose populations seem stable. No tadpole before Gosner stage 41 has been observed, either in the water or carried by males. However, two observations have been made of adults carrying tadpoles at a late development stage (> stage 41) (Figure 6). The tadpoles are endotrophic and lack functional mouth. If they are detached from the back of their father, they die within a few hours despite humid and ideal thermal conditions (pers. obs.). These observations suggest that the tadpoles may undergo a phase of nidicolous development after hatching and only complete their development on the back of the adult from an advanced stage. Only 1–4 tadpoles of the same stage were observed at a time carried by adults. This reduced clutch size is likely linked to the fact that large eggs are needed to complete endotrophic development. However, adults were often observed carrying tadpoles at two different stages, demonstrating that they can keep mating while already attending a clutch. This is likely linked to the extended time needed for the tadpoles to develop until complete metamorphosis.

Anomaloglossus apiau sp. nov. (Figs. 5���7; Table 1) Holotype. MZUSP 155833 (SMS 962), an adult male collected by Antoine Fouquet, Sergio Marques de Sousa and Pedro Murilo Sales Nunes in October 2011 at ~ 580 m at Serra do Apia��, Roraima State, Brazil (2.430168 N, 61.411715 W). Paratopotypes. (n = 35): MZUSP 155814 ���15, 155818, 155826, 155828��� 29 155831 ��� 32, eight males, and MZUSP 155812 ���13, 155816���17, 155819���25, 11 females, collected in October 2011 by AF, SMS and PMSN, and MZUSP 155834, 155836, 155838, 155840, 155842 ���43, 155851��� 52, eight males, and MZUSP 155837, 155839, 155841, 155844���46, 155850, 155853, eight females, collected in June 2012 by AF and PMSN. Etymology. The specific epithet is a noun in apposition and refers to the name of the region where the type locality lies (Serra do Apia��, Roraima State, Brazil). Adult definition and diagnosis. (1) a small Anomaloglossus (average male SVL = 17.1 mm [15.5 ���20.0, n = 14], average female SVL = 20.2 mm [18.4���21.8, n = 15]) (Table 1); (2) body robust; (3) skin on dorsum granular, especially posteriorly, venter smooth; (4) Finger II shorter than I when fingers are appressed, but Finger I shorter than II (3.3 vs. 3.5 mm in holotype) using the method of Kaplan (1997); (5) tip of Finger IV surpassing the base of the distal subarticular tubercle on Finger III when fingers appressed; (6) distal subarticular tubercle on Finger III indistinct; (7) Finger III distinctly swollen in males; (8) fingers with keel-like lateral folds particularly developed preaxially on Finger III; (9) toe webbing basal, with well developed fringes (sensu Grant et al. 2006; keel-like lateral folds sensu Myers & Donnelly 2008); (10) tarsal keel well defined, short, strongly curved, strongly elevated and tubercle-like proximally; (11) black arm gland in males (sensu Grant & Castro-Herrera 1998, see also Grant et al. 2006) and cloacal tubercles absent; (12) pale paracloacal mark present; (13) dorsolateral stripe absent; (15) ventrolateral stripe present in males as a series of more or less conspicuous white dots; (16) oblique lateral stripe present as conspicuous white dots extending from groin midway along flank and inconspicuous to the upper posterior edge of the orbit; (17) sexual dichromatism in throat and venter colour pattern present, with only a few scattered melanophores in females while males possess marbled ventral skin; (18) iris with metallic pigmentation and pupil ring; (20) median lingual process short, slightly longer than wide, tapered, bluntly pointed, smooth (non-papillate), reclined in pit (Fig. 5); (21) maxillary teeth present, small; (22) testes immaculate white, mature ova very large (3 mm in diameter), yellowish brown. Morphological comparisons with congeneric species. Anomaloglossus apiau can easily be distinguished from A. ayarzaguenai, A. breweri, A. guanayensis, A. megacephalus, A. moffetti, A. murisipanensis, A. parimae, A. parkerae, A. shrevei, A. tamacuarensis, A. tepequem, A. tepuyensis, A. triunfo, A. verbeeksnyderorum, and A. wothuja in having basal toe webbing (moderate to extensive in the aforementioned species). Anomaloglossus apiau can be readily distinguished from A. beebei, A. kaiei, A. rufulus and A. roraima in having folded fringes on fingers, best developed preaxially on Finger III (fringes barely distinct, not distinctly folded in the aforementioned species). Anomaloglossus apiau can be distinguished from A. baeobatrachus, A. leopardus, and A. stepheni by its faint, discontinuous oblique ventrolateral stripe (conspicuous and continuous in A. baeobatrachus, A. leopardus, and A. stepheni). Anomaloglossus apiau can be distinguished from A. praderioi by lacking a conspicuous light dorsolateral stripe (present in A. praderioi). Anomaloglossus apiau can be distinguished from Anomaloglossus degranvillei, A. surinamensis and A. apiau by its Finger II longer than Finger I (equal in size in A. degranvillei and A. surinamensis), and by a third finger swollen in male (not swollen in A. degranvillei and A. surinamensis). From all the other nominal species of Anomaloglossus it can be distinguished by its long advertisement call composed of pairs of short pulses (see below). Description of the holotype. An adult male of 17.7 mm SVL. Dorsal skin granular on posterior dorsum, thighs, and shanks; ventral skin smooth (Fig. 6). Cloacal tubercles absent. Head wider than long. Snout bluntly pointed in lateral view, extending past the lower jaw, bluntly pointed in ventral and dorsal views. Nares located close to tip of snout, directed posterolaterally; nares visible in frontal view, barely visible in dorsal or ventral view; IN 39 % of HW. Canthus rostralis well defined; loreal region straight, sloping outward to lip. IO 69 % of ED. SL 96 % of ED, 45 % of HL; EN 62 % of ED. Tympanic membrane inconspicuous, round, concealed posterodorsally by a diffuse supratympanic swelling; tympanic annulus not visible posterodorsally and partly unpigmented; TYM 35 % of eye length. Maxillary teeth present, small. Median lingual process short, about as wide as long, tapered, bluntly pointed, smooth (non-papillate), reclined in lingual pit. Vocal sac and vocal slits visible. Hand moderate in size, 27 % of SVL, 79 % of HW. Relative length of fingers III> IV> I> II. Fingers unwebbed. Discs of fingers expanded, disc on Finger III and IV widest (state 2 sensu Grant et al. 2006), discs on other fingers sub-equal (state 1 sensu Grant et al. 2006). Fingers with fringes, best developed preaxially on Finger II and III (Fig. 6). Palmar tubercle large, rounded, thenar tubercle smaller, elliptical; one or two round to ovoid subarticular tubercles (one each on Fingers I and II, two each on Fingers III and IV). Tip of Finger IV distinctly surpassing the base of distal subarticular tubercle on Finger III when fingers appressed. No fleshy supracarpal fold atop wrist (Fig. 6 b). Finger III strongly swollen dorsally and preaxially, swelling extending from the wrist to the third subarticular tubercle [state 3 sensu Grant et al. (2006)]. Hind limbs robust, moderately long, TIL 49 % of SVL. Relative lengths of appressed toes IV> III> V> II> I; Toe I short, not reaching the base of subarticular tubercle of Toe II. Toe discs expanded, distinctly larger than finger discs, largest on Toes II, III and IV. Feet weakly webbed; all toes with well-developed fringes. Webbing formula I 1 ����� 2 - II 2 +��� 3 - III 2 ����� 4 IV 4 -��� 2 -V (Fig. 6 b). Inner metatarsal tubercle small, elliptical; outer metatarsal tubercle small, round, about half the size of the inner metatarsal tubercle. One to three round to ovoid subarticular tubercles (one each on Toes I and II, two each on Toes III and V, and three on Toe IV with proximal tubercle on Toe IV the smallest and least conspicuous). A strong outer metatarsal fold is present, coextensive with the fringe on Toe V and almost reaching the outer metatarsal tubercle. Tarsal keel weakly curved proximally, discontinuous with the fringe along the outer edge of the first toe, strongly elevated and tubercle-like proximally (Fig. 6 b). Colour of holotype in life. Dorsal ground colour brown with a light brown interorbital triangle (Fig. 6 c). Upper surface of arm with a bright yellow mark with ill defined dark brown and white marks on forearm and wrist; upper surface of thigh yellow with well defined dark brown transverse bands, shank and foot light brown also with well defined dark brown transverse bands. Flanks dark brown, with some blue/whitish irregular iridescent blotches on the lower part, and a lateral stripe of yellowish spots from the inguinal region to the middle of the flank becoming conspicuous again with a short orange stripe on the upper posterior corner of the eye. Dorsolateral stripe absent. Upper lip brown with iridescent white marks below the eye; loreal region and side of head dark; a small whitish stripe is present from arm insertion to tympanum partly depigmented. A dark brown stripe is present on the anterior edge of upper arm, tapering from arm insertion to forearm. Throat, belly and undersurface of arm, thigh and shank marbled with patches of dark brown melanophores. Rear of thigh and cloacal region brown. Paracloacal marks orange. Palms and soles brown (Fig. 6 c). Colour of holotype in preservative. Dorsal ground colour dark brown with a thin light gray to light brown interorbital stripe and a similarly coloured small spot behind the eyes (Fig. 6 a). Dorsal surface of arm brown with dark brown transverse bands on forearm and wrist, upper arm cream; dorsal surface of leg brown with well-defined dark brown transverse bands on thigh, shank, and foot. Flanks dark brown with some irregular spots and a few whitish irregular blotches ventrolaterally. A few white spots (covering small tubercles) form a dotted oblique lateral stripe that only extends from the inguinal region to the middle of the flanks. Throat, belly, and under-surface of arms and legs marbled with patches of melanophores; rear of thigh and cloacal region brown. Paracloacal marks whitish. Palms and soles dark brown (Fig. 6 b). Variation. Males are smaller than females (average male SVL = 17.1 mm 15.5 ���20.0 mm, n = 14; average female SVL = 20.2 mm, 17.7���21.8 mm, n = 15), usually with third Finger III distinctly swollen. There is marked sexual dichromatism (Fig. 7). Dorsal colouration of females is light to dark brown with small diffuse black marks and no lateral dotted stripe. Males, however, are generally darker with a highly variable and conspicuous pattern as well as a lateral stripe consisting of white, bluish and even sometimes orange spots. Whitish-blue marks also occur ventrally on flank, shoulder and lip in males, but found in one female only. Hind limbs are also generally markedly sexually dichromatic, being of similar colouration to the dorsum in females, but with orange markings of various extents in males (from inexistent to small patches, or even sometimes becoming the main background colour). Ventral colouration of females is immaculate white, sometimes with diffuse melanophores and a yellowish throat. Males, however, generally have extensively pigmented ventral surfaces (with one exception; Fig. 7, MZUSP 155836, SVL 16.8 mm, despite conspicuous dorsal colouration and swollen third finger) giving a marbled aspect, especially on throat. In addition to sexual dimorphism in size and colour pattern, variation is extensive within sexes. Among females, dorsal colouration varies from light to dark brown and from immaculate to vermiculate with dark patches. The dark bands on thigh and shank vary from conspicuous to very faint. Lip pattern varies from almost immaculate cream to covered with dark brown marks. Shoulder is orange to light brown. In males, variation in colouration is even more striking and might depend partly on reproductive activity given that more conspicuously coloured males also have swollen third fingers. Dorsal colouration in males varies from immaculate dark brown to light brown with various different patterns of dark brown markings. The distribution of bluish white iridescent dots ranges from a few large ones on the ventral part of the flanks and lips to completely peppering flanks and lips. Limbs and ventral surface are also highly variable in colouration (see above). Advertisement calls. The call is a long trill of 19.6 s on average (sd = 11.0, range = 7.0��� 39.4, n = 10) consisting of pairs of very short pulses (average duration of pulses = 0.03 s, sd = 0.005, 0.02���0.04, n = 50; interpulse interval within pair 0.03 s and between pairs 0.073 s on average) (Fig. 8). Pulse rate per second 11.3 on average (sd = 0.55). Four harmonics are visible, with the dominant frequency located in the lowest harmonic (= fundamental frequency). The dominant frequency increases during the first seconds of the call (~ 4 s) but reaches a plateau of 4.3 kHz on average (sd = 0.129, 4.147���4.526, n = 5; Fig. 8; Table 2). FIGURE 8. Advertisement call of Anomalogossus apiau (IND 5) showing call length (> 17 s) and spectral structure. Note the paired pulses (inset). The specimen was recorded in October 2011 around 11 am at the type locality (21 ��C). The temporal structure of the call differs markedly from that of congeneric species in the duration of the call. Anomaloglossus degranvillei (Marty & Gaucher 1999), A. surinamensis (Ouboter & Jairam 2012) and A. praderioi (Kok 2010) emit a single note at rates between 50 and 100 calls/min; A. beebei (Kok et al. 2006 a), A. tamacuarensis (Myers & Donnelly 1997) and A. kaiei (Kok et al. 2006 b) emit short series of pulses (4 ���5, 2, and 1���2 pulses/call, respectively); A. baeobatrachus (Marty & Gaucher 1999) and A. stepheni (Lima et al. 2006; Martins 1989) emit short trills of 7���19 pulses and less than 1 s of duration. Anomaloglossus rufulus emits shorter (Anomaloglossus call that has a similar temporal structure to the call of A. apiau is the one of A. verbeeksnyderorum (Barrio-Amor��s et al. 2010). However pulse rate of A. apiau is higher (11.27 pulses/s vs. 8.87 in A. verbeeksnyderorum) and the pulses are paired (vs. regularly spaced in A. verbeeksnyderorum). MEAN SD IND 1 REC 1 IND 2 REC 1 IND 3 REC 1 Distribution and ecology. Anomaloglossus apiau sp. nov. was observed exclusively along rocky streams (only a few meters away from water maximal level), on the slopes of Serra de Apia�� above 500 m elevation and up to the summit at ~ 1400 m. These streams are usually shallow, but they can be fast flowing and up to 15 m wide (but unlikely torrential) or reduced to rivulets with only a film of water running on the rocks (Fig. 6 d). Males call from the stream banks or on rocks; they only call from the banks alongside larger streams. The species is locally abundant with one calling male found every ~ 5 m along the stream when calling activity is at its peak, i.e. during late afternoon or after rainfall., Published as part of Fouquet, Antoine, Souza, Sergio Marques, Sales Nunes, Pedro M., Kok, Philippe J. R., Curcio, Felipe Franco, Carvalho, Celso Morato De, Grant, Taran & Rodrigues, Miguel Trefaut, 2015, Two new endangered species of Anomaloglossus (Anura: Aromobatidae) from Roraima State, northern Brazil, pp. 191-210 in Zootaxa 3926 (2) on pages 200-206, DOI: 10.11646/zootaxa.3926.2.2, http://zenodo.org/record/243597, {"references":["Kaplan, M. (1997) A new species of Colostethus from the Sierra Nevada de Santa Marta (Colombia) with comments on intergeneric relationships within the Dendrobatidae. Journal of Herpetology, 31, 369 - 375. http: // dx. doi. org / 10.2307 / 1565665","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F., Kok, P. J., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262. http: // dx. doi. org / 10.1206 / 0003 - 0090 (2006) 299 [1: PSODFA] 2.0. CO; 2","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147. http: // dx. doi. org / 10.1206 / 308.1","Grant, T. & Castro, F. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology, 32, 378 - 392. http: // dx. doi. org / 10.2307 / 1565452","Marty, C. & Gaucher, P. (1999) Guide sonore des amphibiens anoures de Guyane. Sittelle Ceba, Le Verdier, CD. [Centre bioacoustique alpin]","Ouboter, P. E. & Jairam, R. (2012) Amphibians of Suriname. Brill, Leiden, 388 pp.","Kok, P. J. R., Sambhu, H., Roopsind, I., Lenglet, G. & Bourne, G. (2006 a) A new species of Colostethus (Anura: Dendrobatidae) with maternal care from Kaieteur National Park, Guyana. Zootaxa, 1238, 35 - 61.","Myers, C. W. & Donnelly, M. A. (1997) A tepui herpetofauna on a granitic mountain (Tamacuari) in the borderland between Venezuela and Brazil: report from the Phipps Tapirapeco Expedition. American Museum novitates, 3213, 1 - 71.","Kok, P. J. R., MacCulloch, R. D., Gaucher, P., Poelman, E. H., Bourne, G. R., Lathrop, A. & Lenglet, G. L. (2006 b) A new species of Colostethus (Anura, Dendrobatidae) from French Guiana with a redescription of Colostethus beebei (Noble, 1923) from its type locality. Phyllomedusa, 5, 43 - 66. http: // dx. doi. org / 10.11606 / issn. 2316 - 9079. v 5 i 1 p 43 - 66","Lima, A. P., Magnusson, W., Menin, M., Erdtmann, L., Rodrigues, D., Keller, C. & Hodl, W. (2006) Guia de Sapos da Reserva Adolpho Ducke: Amazonia Central. Attema Design Editorial, Manaus, 168 pp.","Martins, M. (1989) Nova especie de Colostethus da Amazonia central (Amphibia: Dendrobatidae). Revista Brasileira de Biologia, 49, 1009 - 1012.","Barrio-Amoros, C. L. & Santos, J. C. (2011) Redescription and generic assignation of Dendrobates rufulus Gorzula (Anura, Dendrobatidae) from Chimanta tepui, Venezuela. Salamandra, 47 (3), 155 - 160.","Barrio-Amoros, C. L., Santos, J. C. & Jovanovic, O. (2010) A new dendrobatid frog (Anura: Dendrobatidae: Anomaloglossus) from the Orinoquian rainforest, southern Venezuela. Zootaxa, 2413, 37 - 50."]}

Anomaloglossus tepequem sp. nov. (Figs. 2���4; Table 1) Holotype. MZUSP 63310, an adult female collected by M��rcio Martins, 20 July 1986 at ~ 700 m elevation at Tepequ��m, Roraima State, Brazil (3.750866 N, 61.705084 W). Paratopotypes (n = 14). MZUSP 63308 ���09, a female and a male, respectively, collected with the holotype by M��rcio Martins. MZUSP 70692 ���95, 70701, 70703, six males, and MZUSP 70588, 70691, 70696 ��� 99, six females, collected by Miguel T. Rodrigues, Celso Morato de Carvalho, Dante Pavan, and Gabriel Skuk, 12 December 1992 (Table 1). Etymology. The specific epithet is a noun in apposition and refers to the type locality (Tepequ��m, Roraima State, Brazil). Adult definition and diagnosis. (1) medium-sized Anomaloglossus (average male SVL = 19.2 mm [18.2���20.1, n = 4], average female SVL = 22.9 mm [21.7���24.5, n = 5]) (Table 1); (2) body robust; (3) skin on dorsum and venter smooth; (4) Finger II appears shorter than I when fingers are appressed, but actually Finger I is slightly shorter than II (4.2 vs. 4.4 mm in holotype) using the method of Kaplan (1997); (5) tip of Finger IV surpassing the base of the distal subarticular tubercle on Finger III when fingers appressed; (6) distal subarticular tubercle on Finger III and IV indistinct; (7) Finger III not swollen in males; (8) fingers with fringes particularly developed on preaxial edges of Fingers II and III; (9) toes moderately webbed, with well developed folded, flapped fringing (sensu Myers & Donnelly 2008); (10) tarsal keel well defined, straight or very weakly curved, at most weakly elevated proximally, not forming tubercle; (11) black arm gland in males (sensu Grant & Castro 1998, see also Grant et al. 2006) and cloacal tubercles absent; (12) pale paracloacal mark present; (13) dorsolateral stripe absent; (15) ventrolateral stripe absent; (16) oblique lateral stripe present, short, extending from inguinal region to approximately mid-flank; (17) sexual dichromatism in throat colour pattern present, throat evenly covered with darker and larger patches in all males and in some females but almost no pigmentation in most females; (18) no sexual dichromatism in abdomen colour pattern, which is immaculate cream; (19) iris with metallic pigmentation and pupil ring; (20) median lingual process (Fig. 2) longer than wide, tapered, bluntly pointed, smooth (nonpapillate), reclined in pit; (21) maxillary teeth present, small; (22) testes immaculate white, mature ova small (ca. 1 mm) yellowish brown. Morphological comparisons with congeneric species. Anomaloglossus tepequem is immediately distinguished from A. baeobatrachus (Boistel & de Massary 1999), A. beebei (Noble 1923), A. degranvillei (Lescure 1975), A. kaiei (Kok et al. 2006 b), A. leopardus (Ouboter & Jairam 2012), A. praderioi (La Marca 1996), A. rufulus (Gorzula 1988), A. roraima (La Marca 1996, Kok et al. 2013), A. stepheni (Martins 1989), and A. surinamensis (Ouboter & Jairam 2012) in having moderate toe webbing (toe webbing absent or basal in the aforementioned species). Anomaloglossus tepequem can be distinguished from A. breweri (Barrio-Amor��s 2006), A. murisipanensis (La Marca 1996), A. parimae (La Marca 1996), A. parkerae (Meinhardt & Parmelee 1996), A. tamacuarensis (Myers & Donnelly 1997), A. tepuyensis (La Marca 1996), A. triunfo (Barrio-Amor��s et al. 2004), A. verbeeksnyderorum (Barrio-Amor��s et al. 2010), and A. wothuja (Barrio-Amor��s et al. 2004) in having distinctly more developed toe webbing, particularly between Toes IV���V, with the webbing surpassing the distal subarticular tubercle on Toe V (toe webbing never surpasses the distal subarticular tubercle on Toe V in the aforementioned species). Anomaloglossus tepequem can be distinguished from A. ayarzaguenai (La Marca 1996), A. guanayensis (La Marca 1996), and A. moffetti (Barrio-Amor��s & Brewer-Carias 2008) by having less extensive toe webbing, especially between Toes I���III, and in always having an immaculate cream abdomen colouration in both sexes (at least covered by small melanophores, which sometimes form spots or even extensive marbling in A. ayarzaguenai, A. guanayensis, and A. moffetti). Anomaloglossus tepequem mainly differs from A. megacephalus (Kok et al. 2010) by its smaller size (maximum SVL = 24.5 mm in A. tepequem vs. 28.3 mm in A. megacephalus), a relative shorter head (HW 108 % of HL in A. tepequem vs. 90 % in A. megacephalus), and in having an unpigmented ventral colouration in both sexes (at least covered by small melanophores, sometimes forming spots or even extensive marbling in female A. megacephalus [male unknown]). Comparison with Anomaloglossus shrevei (Rivero 1961) is hindered by the brevity of the original description and the lack of comparative material from the type locality. However, Myers & Donnelly (1997) provided extensive additional information based on examination of the holotype and two additional specimens. Anomaloglossus tepequem can be distinguished from A. shrevei by a much smaller SVL in adult females (maximum female SVL = 24.5 mm in A. tepequem, 36.0 mm in A. shrevei) and an elongate, tapered median lingual process (Fig. 2; low, rounded bump or vertically aligned, blunt cone in A. shrevei; see Grant et al., 1997: 15���16; Myers & Donnelly 1997: 26, fig. 17 A���B). Description of holotype. An adult female of 21.7 mm SVL. Dorsal skin entirely smooth; ventral skin smooth (Fig. 3). Cloacal tubercles absent. Head wider than long. Snout bluntly pointed in lateral view, extending past the lower jaw, bluntly pointed in ventral and dorsal views. Nares located close to snout tip (EN 60 % of SL), directed posterolaterally; nares visible in frontal view, barely visible in dorsal and ventral view; IN 41 % of HW. Canthus rostralis well defined; loreal region straight, sloping outward to lip. IO 64 % of ED. SL 103 % of ED, 49 % of HL; EN 61 % of ED. Tympanic membrane inconspicuous, round, concealed posterodorsally by a diffuse supratympanic swelling; tympanic annulus visible; TYM 50 % of ED. Maxillary teeth present, small. Median lingual process elongate, longer than wide, tapered, bluntly pointed, reclined in pit. Hand moderate in size, 27 % of SVL, 81 % of HW. Relative length of fingers III> IV> I> II. Fingers unwebbed. Discs of fingers expanded, disc on Finger II, III, IV widest (state 2 sensu Grant et al. 2006), discs on Finger I correspond to state 1 (sensu Grant et al. 2006). Fingers with fringes [sensu Grant et al. (2006: 66���67); keel-like lateral folds sensu Myers & Donnelly (2008)], best developed on pre- and postaxial edges of Finger II and preaxial edge of Finger III (Fig. 3). Palmar tubercle large, rounded, slightly heart-shaped; thenar tubercle smaller, elliptical; one or two round to ovoid subarticular tubercles (one each on Fingers I and II, two each on Fingers III and IV, with distal tubercles on Finger III and IV inconspicuous). Tip of Finger IV distinctly surpassing the base of distal subarticular tubercle on Finger III when fingers appressed. No fleshy supracarpal fold atop wrist (Fig. 3). Hind limbs robust, moderately long, TIL 51 % of SVL. Relative lengths of appressed toes IV> III> V> II> I; Toe I short, slightly surpassing the base of subarticular tubercle of Toe II. Toe discs expanded, slightly larger than finger discs, largest on Toes II and IV. Feet moderately webbed; all toes with well developed folded, flapped fringing. Webbing formula I 0+��� 2 - II 1 -��� 2 �� III 2���3 - IV 3 -��� 1 ��V (Fig. 3). Inner metatarsal tubercle small, elliptical; outer metatarsal tubercle small, round, about half the size of inner tubercle. One to three round to ovoid subarticular tubercles (one each on Toes I and II, two each on Toes III and V, and three on Toe IV, with distal tubercle on Toe IV the smallest and less conspicuous). A strong outer metatarsal fold is present, coextensive with the fringe (sensu Grant et al. 2006) on the postaxial edge of Toe V and almost reaching the outer metatarsal tubercle. Tarsal keel strong, continuous with fringe along the outer edge of the first toe, straight or very weakly curved at proximal end, lacking pronounced elevation or tubercle proximally (Fig. 3). Colour of holotype in preservative. Dorsal ground colour brown with a dark brown interorbital bar followed by another dark brown ���M-shaped��� mark covering the scapular region. Dorsal surface of arm light brown with illdefined dark brown transverse bands on forearm and wrist and a cream upper arm; upper surface of leg light brown with well defined dark brown transverse bands on thigh, shank, and foot. Flanks dark brown, with some irregular spots and a few whitish irregular blotches on the lower part. A few white spots (covering small tubercles) form a broken oblique lateral stripe that extends from the groin midway along flank. No dorsolateral stripe. Upper lip brown, delimited by a clear line and with some whitish marks below eye; loreal region and side of head dark; small whitish stripe from arm insertion to tympanum. A dark brown stripe is present on the anterior edge of upper arm, tapering from arm insertion to forearm. Abdomen immaculate cream, throat with patches of melanophores. Ventral surface of upper arm whitish; ventral surface of forearm brown on postaxial side; ventral surface of thigh and shank whitish; rear of thigh and cloacal region brown. Paracloacal marks whitish. Palms and soles brown (Fig. 3). Measurements of holotype (in mm). SVL = 21.7; FA = 4.7; Ha = 5.9; TL = 10.5; FL = 9.9; HW = 7.3; HL = 7.1; ED = 3.4; EN = 2.1; IN = 3.0; SL = 3.5; IO = 2.2; TD = 1.7; WFD = 0.8; WTD = 0.8; ThL = 10.8; 1 FiL = 2.5. Variation. The type series comprises five adult females, three juvenile females, four adult males, and three juvenile males (Table 1). Except for the sexual dimorphism in size and throat colour pattern (see above), dorsal colour varies from light brown to chestnut-brown; dorsal pattern varies slightly in shape and conspicuousness; brown transverse bands on shanks vary from four to three in number, and also vary in conspicuousness with sometimes only the central band being visible; ventral colour variation remains subtle (Fig. 4). Distribution and ecology. Anomaloglossus tepequem was observed and collected only twice, once in 1986 and again in 1992. This is a diurnal species associated with fast flowing streams (Fig. 3 d) and probably restricted to the slopes of Tepequ��m at> 500 m elevation. In 1992, it was one of the most common frog species in the area. Dozens of males were seen actively calling from the banks and upon stones along streams, and were frequently spotted at night sleeping on rocks or leaves above or near water. Additional diurnal and nocturnal investigations performed during five days in May 2011 by seven herpetologists at the same sites where the species was previously found failed to detect it, despite the fact that the time of the year and the weather conditions were favorable. SVL FA Ha TL FL HW HL ED A. tepequem Females A (n= 5) 22.90 4.86 6.06 11.10 10.02 7.54 7.06 3.26 sd 1.05 0.18 0.21 0.42 0.39 0.39 0.30 0.17 FemalesJ(n= 3) 18.87 4.40 5.33 10.30 9.20 6.70 6.37 2.83 sd 1.15 0.10 0.29 0.44 0.36 0.46 0.38 0.29 Males A (n= 4) 19.20 4.63 5.53 10.30 9.48 6.88 6.30 3.15 sd 0.86 0.36 0.36 0.22 0.45 0.22 0.16 0.10 Males J (n= 3) 16.43 3.80 4.37 8.87 7.93 6.10 5.57 2.57 sd 0.15 0.17 0.25 0.25 0.45 0.10 0.32 0.06 A. apiau Females A (n= 15) 20.16 4.29 5.05 9.55 8.72 7.27 6.31 2.89 sd 1.17 0.27 0.20 0.42 0.43 0.41 0.38 0.17 FemalesJ(n= 4) 15.48 3.63 4.10 7.85 7.15 5.65 5.13 2.23 sd 0.59 0.26 0.29 0.56 0.81 0.39 0.38 0.15 Males A (n= 14) 17.09 3.73 4.46 8.48 7.69 6.07 5.55 2.54 sd 0.99 0.16 0.22 0.52 0.52 0.27 0.35 0.18 Males J (n= 3) 14.53 3.43 3.87 7.70 6.50 5.37 4.80 2.23 sd 0.06 0.06 0.21 0.26 0.44 0.21 0.20 0.15 continued EN IN SL IO TD WFD WTD ThL 1 FiL A. tepequem Females A (n= 5) 2.16 3.02 3.58 2.30 1.40 0.74 0.80 11.55 2.49 sd 0.09 0.19 0.16 0.19 0.19 0.04 0.04 0.63 0.06 Females J (n= 3) 1.93 2.40 3.10 2.17 1.30 0.64 0.71 10.52 2.12 sd 0.06 0.17 0.46 0.06 0.17 0.09 0.02 0.59 0.20 Males A (n= 4) 1.80 2.83 3.20 2.25 1.33 0.72 0.76 10.84 2.34 sd 0.08 0.10 0.12 0.06 0.05 0.07 0.04 0.36 0.18 Males J (n= 3) 1.67 2.40 2.73 1.70 1.40 0.49 0.59 9.18 1.78 sd 0.06 0.17 0.15 0.10 0.26 0.08 0.03 0.40 0.13 A. apiau Females A (n= 15) 1.73 2.80 3.09 2.21 1.07 0.82 0.91 10.31 2.16 sd 0.08 0.17 0.27 0.13 0.10 0.08 0.10 0.39 0.18 Females J (n= 4) 1.35 2.20 2.40 1.78 0.85 0.59 0.66 8.70 1.75 sd 0.06 0.12 0.14 0.10 0.10 0.10 0.07 0.75 0.20 Males A (n= 14) 1.54 2.44 2.67 1.90 0.97 0.67 0.79 9.06 1.91 sd 0.13 0.15 0.17 0.15 0.08 0.08 0.07 0.47 0.18 Males J (n= 3) 1.30 2.20 2.27 1.70 0.77 0.56 0.67 7.49 1.55 sd 0.10 0.10 0.12 0.10 0.06 0.05 0.02 0.43 0.04, Published as part of Fouquet, Antoine, Souza, Sergio Marques, Sales Nunes, Pedro M., Kok, Philippe J. R., Curcio, Felipe Franco, Carvalho, Celso Morato De, Grant, Taran & Rodrigues, Miguel Trefaut, 2015, Two new endangered species of Anomaloglossus (Anura: Aromobatidae) from Roraima State, northern Brazil, pp. 191-210 in Zootaxa 3926 (2) on pages 194-197, DOI: 10.11646/zootaxa.3926.2.2, http://zenodo.org/record/243597, {"references":["Kaplan, M. (1997) A new species of Colostethus from the Sierra Nevada de Santa Marta (Colombia) with comments on intergeneric relationships within the Dendrobatidae. Journal of Herpetology, 31, 369 - 375. http: // dx. doi. org / 10.2307 / 1565665","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147. http: // dx. doi. org / 10.1206 / 308.1","Grant, T. & Castro, F. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology, 32, 378 - 392. http: // dx. doi. org / 10.2307 / 1565452","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F., Kok, P. J., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262. http: // dx. doi. org / 10.1206 / 0003 - 0090 (2006) 299 [1: PSODFA] 2.0. CO; 2","Boistel, R. & de Massary, J. - C. (1999) Les amphibiens veneneux de la famille des dendrobatides. Le Courrier de la Nature, 176, 34 - 37.","Noble, G. K. (1923) New batrachians from the tropical research station British Guiana. Zoologica 3, 288 - 305.","Lescure, J. (1975) Contribution a l'etude des amphibiens de Guyane francaise. III. Une nouvelle espece de Colostethus (Dendrobatidae): Colostethus degranvillei nov. sp. Bulletin du Museum National d'Histoire Naturelle, Zoologie, 203, 413 - 420.","Kok, P. J. R., MacCulloch, R. D., Gaucher, P., Poelman, E. H., Bourne, G. R., Lathrop, A. & Lenglet, G. L. (2006 b) A new species of Colostethus (Anura, Dendrobatidae) from French Guiana with a redescription of Colostethus beebei (Noble, 1923) from its type locality. Phyllomedusa, 5, 43 - 66. http: // dx. doi. org / 10.11606 / issn. 2316 - 9079. v 5 i 1 p 43 - 66","Ouboter, P. E. & Jairam, R. (2012) Amphibians of Suriname. Brill, Leiden, 388 pp.","La Marca, E. (1996) Ranas del genero Colostethus (Amphibia: Anura; Dendrobatidae) de la Guayana venezolana, con la descripcion de siete especies nuevas. Publicaciones de la Asociacion de los Amigos de Donana, 9, 1 - 64.","Gorzula, S. (1988) Una nueva especie de Dendrobates (Amphibia: Dendrobatidae) del Macizo del Chimanta, Estado Bolivar, Venezuela. Memoria de la Sociedad de Ciencias Naturales La Salle, 48, 143 - 149.","Martins, M. (1989) Nova especie de Colostethus da Amazonia central (Amphibia: Dendrobatidae). Revista Brasileira de Biologia, 49, 1009 - 1012.","Meinhardt, D. J. & Parmelee, J. R. (1996) A new species of Colostethus (Anura: Dendrobatidae) from Venezuela. Herpetologica, 52 (1), 70 - 77.","Myers, C. W. & Donnelly, M. A. (1997) A tepui herpetofauna on a granitic mountain (Tamacuari) in the borderland between Venezuela and Brazil: report from the Phipps Tapirapeco Expedition. American Museum novitates, 3213, 1 - 71.","Barrio-Amoros, C. L., Fuentes, O. & Rivas, G. (2004) Two new species of Colostethus (Anura: Dendrobatidae) from the Venezuelan Guayana. Salamandra, 40, 183 - 200.","Barrio-Amoros, C. L., Santos, J. C. & Jovanovic, O. (2010) A new dendrobatid frog (Anura: Dendrobatidae: Anomaloglossus) from the Orinoquian rainforest, southern Venezuela. Zootaxa, 2413, 37 - 50.","Barrio-Amoros, C. L. & Brewer-Carias, C. (2008) Herpetological results of the 2002 Expedition to Sarisarinama, a tepui in Venezuelan Guayana, with the description of five new species. Zootaxa, 1942, 1 - 68.","Rivero, J. A. (1961) Salientia of Venezuela. Bulletin of the Museum of Comparative Zoology, 126, 1 - 207."]}

Anomaloglossus tepequem sp. nov. (Figs. 2–4; Table 1) Holotype. MZUSP 63310, an adult female collected by Márcio Martins, 20 July 1986 at ~ 700 m elevation at Tepequém, Roraima State, Brazil (3.750866 N, 61.705084 W). Paratopotypes (n = 14). MZUSP 63308 –09, a female and a male, respectively, collected with the holotype by Márcio Martins. MZUSP 70692 –95, 70701, 70703, six males, and MZUSP 70588, 70691, 70696 – 99, six females, collected by Miguel T. Rodrigues, Celso Morato de Carvalho, Dante Pavan, and Gabriel Skuk, 12 December 1992 (Table 1). Etymology. The specific epithet is a noun in apposition and refers to the type locality (Tepequém, Roraima State, Brazil). Adult definition and diagnosis. (1) medium-sized Anomaloglossus (average male SVL = 19.2 mm [18.2–20.1, n = 4], average female SVL = 22.9 mm [21.7–24.5, n = 5]) (Table 1); (2) body robust; (3) skin on dorsum and venter smooth; (4) Finger II appears shorter than I when fingers are appressed, but actually Finger I is slightly shorter than II (4.2 vs. 4.4 mm in holotype) using the method of Kaplan (1997); (5) tip of Finger IV surpassing the base of the distal subarticular tubercle on Finger III when fingers appressed; (6) distal subarticular tubercle on Finger III and IV indistinct; (7) Finger III not swollen in males; (8) fingers with fringes particularly developed on preaxial edges of Fingers II and III; (9) toes moderately webbed, with well developed folded, flapped fringing (sensu Myers & Donnelly 2008); (10) tarsal keel well defined, straight or very weakly curved, at most weakly elevated proximally, not forming tubercle; (11) black arm gland in males (sensu Grant & Castro 1998, see also Grant et al. 2006) and cloacal tubercles absent; (12) pale paracloacal mark present; (13) dorsolateral stripe absent; (15) ventrolateral stripe absent; (16) oblique lateral stripe present, short, extending from inguinal region to approximately mid-flank; (17) sexual dichromatism in throat colour pattern present, throat evenly covered with darker and larger patches in all males and in some females but almost no pigmentation in most females; (18) no sexual dichromatism in abdomen colour pattern, which is immaculate cream; (19) iris with metallic pigmentation and pupil ring; (20) median lingual process (Fig. 2) longer than wide, tapered, bluntly pointed, smooth (nonpapillate), reclined in pit; (21) maxillary teeth present, small; (22) testes immaculate white, mature ova small (ca. 1 mm) yellowish brown. Morphological comparisons with congeneric species. Anomaloglossus tepequem is immediately distinguished from A. baeobatrachus (Boistel & de Massary 1999), A. beebei (Noble 1923), A. degranvillei (Lescure 1975), A. kaiei (Kok et al. 2006 b), A. leopardus (Ouboter & Jairam 2012), A. praderioi (La Marca 1996), A. rufulus (Gorzula 1988), A. roraima (La Marca 1996, Kok et al. 2013), A. stepheni (Martins 1989), and A. surinamensis (Ouboter & Jairam 2012) in having moderate toe webbing (toe webbing absent or basal in the aforementioned species). Anomaloglossus tepequem can be distinguished from A. breweri (Barrio-Amorós 2006), A. murisipanensis (La Marca 1996), A. parimae (La Marca 1996), A. parkerae (Meinhardt & Parmelee 1996), A. tamacuarensis (Myers & Donnelly 1997), A. tepuyensis (La Marca 1996), A. triunfo (Barrio-Amorós et al. 2004), A. verbeeksnyderorum (Barrio-Amorós et al. 2010), and A. wothuja (Barrio-Amorós et al. 2004) in having distinctly more developed toe webbing, particularly between Toes IV–V, with the webbing surpassing the distal subarticular tubercle on Toe V (toe webbing never surpasses the distal subarticular tubercle on Toe V in the aforementioned species). Anomaloglossus tepequem can be distinguished from A. ayarzaguenai (La Marca 1996), A. guanayensis (La Marca 1996), and A. moffetti (Barrio-Amorós & Brewer-Carias 2008) by having less extensive toe webbing, especially between Toes I–III, and in always having an immaculate cream abdomen colouration in both sexes (at least covered by small melanophores, which sometimes form spots or even extensive marbling in A. ayarzaguenai, A. guanayensis, and A. moffetti). Anomaloglossus tepequem mainly differs from A. megacephalus (Kok et al. 2010) by its smaller size (maximum SVL = 24.5 mm in A. tepequem vs. 28.3 mm in A. megacephalus), a relative shorter head (HW 108 % of HL in A. tepequem vs. 90 % in A. megacephalus), and in having an unpigmented ventral colouration in both sexes (at least covered by small melanophores, sometimes forming spots or even extensive marbling in female A. megacephalus [male unknown]). Comparison with Anomaloglossus shrevei (Rivero 1961) is hindered by the brevity of the original description and the lack of comparative material from the type locality. However, Myers & Donnelly (1997) provided extensive additional information based on examination of the holotype and two additional specimens. Anomaloglossus tepequem can be distinguished from A. shrevei by a much smaller SVL in adult females (maximum female SVL = 24.5 mm in A. tepequem, 36.0 mm in A. shrevei) and an elongate, tapered median lingual process (Fig. 2; low, rounded bump or vertically aligned, blunt cone in A. shrevei; see Grant et al., 1997: 15–16; Myers & Donnelly 1997: 26, fig. 17 A–B). Description of holotype. An adult female of 21.7 mm SVL. Dorsal skin entirely smooth; ventral skin smooth (Fig. 3). Cloacal tubercles absent. Head wider than long. Snout bluntly pointed in lateral view, extending past the lower jaw, bluntly pointed in ventral and dorsal views. Nares located close to snout tip (EN 60 % of SL), directed posterolaterally; nares visible in frontal view, barely visible in dorsal and ventral view; IN 41 % of HW. Canthus rostralis well defined; loreal region straight, sloping outward to lip. IO 64 % of ED. SL 103 % of ED, 49 % of HL; EN 61 % of ED. Tympanic membrane inconspicuous, round, concealed posterodorsally by a diffuse supratympanic swelling; tympanic annulus visible; TYM 50 % of ED. Maxillary teeth present, small. Median lingual process elongate, longer than wide, tapered, bluntly pointed, reclined in pit. Hand moderate in size, 27 % of SVL, 81 % of HW. Relative length of fingers III> IV> I> II. Fingers unwebbed. Discs of fingers expanded, disc on Finger II, III, IV widest (state 2 sensu Grant et al. 2006), discs on Finger I correspond to state 1 (sensu Grant et al. 2006). Fingers with fringes [sensu Grant et al. (2006: 66–67); keel-like lateral folds sensu Myers & Donnelly (2008)], best developed on pre- and postaxial edges of Finger II and preaxial edge of Finger III (Fig. 3). Palmar tubercle large, rounded, slightly heart-shaped; thenar tubercle smaller, elliptical; one or two round to ovoid subarticular tubercles (one each on Fingers I and II, two each on Fingers III and IV, with distal tubercles on Finger III and IV inconspicuous). Tip of Finger IV distinctly surpassing the base of distal subarticular tubercle on Finger III when fingers appressed. No fleshy supracarpal fold atop wrist (Fig. 3). Hind limbs robust, moderately long, TIL 51 % of SVL. Relative lengths of appressed toes IV> III> V> II> I; Toe I short, slightly surpassing the base of subarticular tubercle of Toe II. Toe discs expanded, slightly larger than finger discs, largest on Toes II and IV. Feet moderately webbed; all toes with well developed folded, flapped fringing. Webbing formula I 0+– 2 - II 1 -– 2 ½ III 2–3 - IV 3 -– 1 ½V (Fig. 3). Inner metatarsal tubercle small, elliptical; outer metatarsal tubercle small, round, about half the size of inner tubercle. One to three round to ovoid subarticular tubercles (one each on Toes I and II, two each on Toes III and V, and three on Toe IV, with distal tubercle on Toe IV the smallest and less conspicuous). A strong outer metatarsal fold is present, coextensive with the fringe (sensu Grant et al. 2006) on the postaxial edge of Toe V and almost reaching the outer metatarsal tubercle. Tarsal keel strong, continuous with fringe along the outer edge of the first toe, straight or very weakly curved at proximal end, lacking pronounced elevation or tubercle proximally (Fig. 3). Colour of holotype in preservative. Dorsal ground colour brown with a dark brown interorbital bar followed by another dark brown “M-shaped” mark covering the scapular region. Dorsal surface of arm light brown with illdefined dark brown transverse bands on forearm and wrist and a cream upper arm; upper surface of leg light brown with well defined dark brown transverse bands on thigh, shank, and foot. Flanks dark brown, with some irregular spots and a few whitish irregular blotches on the lower part. A few white spots (covering small tubercles) form a broken oblique lateral stripe that extends from the groin midway along flank. No dorsolateral stripe. Upper lip brown, delimited by a clear line and with some whitish marks below eye; loreal region and side of head dark; small whitish stripe from arm insertion to tympanum. A dark brown stripe is present on the anterior edge of upper arm, tapering from arm insertion to forearm. Abdomen immaculate cream, throat with patches of melanophores. Ventral surface of upper arm whitish; ventral surface of forearm brown on postaxial side; ventral surface of thigh and shank whitish; rear of thigh and cloacal region brown. Paracloacal marks whitish. Palms and soles brown (Fig. 3). Measurements of holotype (in mm). SVL = 21.7; FA = 4.7; Ha = 5.9; TL = 10.5; FL = 9.9; HW = 7.3; HL = 7.1; ED = 3.4; EN = 2.1; IN = 3.0; SL = 3.5; IO = 2.2; TD = 1.7; WFD = 0.8; WTD = 0.8; ThL = 10.8; 1 FiL = 2.5. Variation. The type series comprises five adult females, three juvenile females, four adult males, and three juvenile males (Table 1). Except for the sexual dimorphism in size and throat colour pattern (see above), dorsal colour varies from light brown to chestnut-brown; dorsal pattern varies slightly in shape and conspicuousness; brown transverse bands on shanks vary from four to three in number, and also vary in conspicuousness with sometimes only the central band being visible; ventral colour variation remains subtle (Fig. 4). Distribution and ecology. Anomaloglossus tepequem was observed and collected only twice, once in 1986 and again in 1992. This is a diurnal species associated with fast flowing streams (Fig. 3 d) and probably restricted to the slopes of Tepequém at> 500 m elevation. In 1992, it was one of the most common frog species in the area. Dozens of males were seen actively calling from the banks and upon stones along streams, and were frequently spotted at night sleeping on rocks or leaves above or near water. Additional diurnal and nocturnal investigations performed during five days in May 2011 by seven herpetologists at the same sites where the species was previously found failed to detect it, despite the fact that the time of the year and the weather conditions were favorable. SVL FA Ha TL FL HW HL ED A. tepequem Females A (n= 5) 22.90 4.86 6.06 11.10 10.02 7.54 7.06 3.26 sd 1.05 0.18 0.21 0.42 0.39 0.39 0.30 0.17 FemalesJ(n= 3) 18.87 4.40 5.33 10.30 9.20 6.70 6.37 2.83 sd 1.15 0.10 0.29 0.44 0.36 0.46 0.38 0.29 Males A (n= 4) 19.20 4.63 5.53 10.30 9.48 6.88 6.30 3.15 sd 0.86 0.36 0.36 0.22 0.45 0.22 0.16 0.10 Males J (n= 3) 16.43 3.80 4.37 8.87 7.93 6.10 5.57 2.57 sd 0.15 0.17 0.25 0.25 0.45 0.10 0.32 0.06 A. apiau Females A (n= 15) 20.16 4.29 5.05 9.55 8.72 7.27 6.31 2.89 sd 1.17 0.27 0.20 0.42 0.43 0.41 0.38 0.17 FemalesJ(n= 4) 15.48 3.63 4.10 7.85 7.15 5.65 5.13 2.23 sd 0.59 0.26 0.29 0.56 0.81 0.39 0.38 0.15 Males A (n= 14) 17.09 3.73 4.46 8.48 7.69 6.07 5.55 2.54 sd 0.99 0.16 0.22 0.52 0.52 0.27 0.35 0.18 Males J (n= 3) 14.53 3.43 3.87 7.70 6.50 5.37 4.80 2.23 sd 0.06 0.06 0.21 0.26 0.44 0.21 0.20 0.15 continued EN IN SL IO TD WFD WTD ThL 1 FiL A. tepequem Females A (n= 5) 2.16 3.02 3.58 2.30 1.40 0.74 0.80 11.55 2.49 sd 0.09 0.19 0.16 0.19 0.19 0.04 0.04 0.63 0.06 Females J (n= 3) 1.93 2.40 3.10 2.17 1.30 0.64 0.71 10.52 2.12 sd 0.06 0.17 0.46 0.06 0.17 0.09 0.02 0.59 0.20 Males A (n= 4) 1.80 2.83 3.20 2.25 1.33 0.72 0.76 10.84 2.34 sd 0.08 0.10 0.12 0.06 0.05 0.07 0.04 0.36 0.18 Males J (n= 3) 1.67 2.40 2.73 1.70 1.40 0.49 0.59 9.18 1.78 sd 0.06 0.17 0.15 0.10 0.26 0.08 0.03 0.40 0.13 A. apiau Females A (n= 15) 1.73 2.80 3.09 2.21 1.07 0.82 0.91 10.31 2.16 sd 0.08 0.17 0.27 0.13 0.10 0.08 0.10 0.39 0.18 Females J (n= 4) 1.35 2.20 2.40 1.78 0.85 0.59 0.66 8.70 1.75 sd 0.06 0.12 0.14 0.10 0.10 0.10 0.07 0.75 0.20 Males A (n= 14) 1.54 2.44 2.67 1.90 0.97 0.67 0.79 9.06 1.91 sd 0.13 0.15 0.17 0.15 0.08 0.08 0.07 0.47 0.18 Males J (n= 3) 1.30 2.20 2.27 1.70 0.77 0.56 0.67 7.49 1.55 sd 0.10 0.10 0.12 0.10 0.06 0.05 0.02 0.43 0.04

We describe two new species of Anomaloglossus from Roraima State, Brazil, that are likely endemic to single mountains currently isolated among lowland forest and savanna ecosystems. The first species, Anomaloglossus tepequem sp. nov. was collected in 1986 and 1992 along a single stream at >500 m elevation on a tepui-like mountain named Tepequém, but was not detected during recent investigations. It is mainly diagnosed from other Anomaloglossus species by its well developed foot webbing, immaculate cream abdomen colouration and small body size (males: 18.2–20.1 mm, females: 21.7–24.5). The second species, Anomaloglossus apiau sp. nov. was found along several streams between 500 and 1400 m elevation on Serra do Apiaú, and is mainly diagnosed from congeners by its weakly webbed feet, males with swollen third finger and ventrolateral stripe formed by white dots, and its advertisement call; a long trill (up to almost 40 s) consisting of pairs of very short pulses. The discovery of these two apparently microendemicspecies suggests that additional Anomaloglossus species remain to be described in the Guiana Shield. Both species should be considered critically endangered given their seemingly reduced range size, association with highland habitat, and the anthropogenic pressure they currently face.

Microcaecilia iwokramae (Wake & Donnelly, 2010) new combination Holotype. Center for Biological Diversity, University of Guyana (CSBD) reported as number HA 1500, bearing tag MAD 2018. Diagnosis. A Microcaecilia with fewer secondary annuli than all other Microcaecilia except M. taylori and M. dermatophaga Wilkinson et al., 2013; differs from M. dermatophaga in having a shorter tooth row on the paired nasopremaxillary bones with fewer (13) teeth in total; differs from most M. taylori specimens in lacking a dorsal transverse groove on the first nuchal collar and in usually having fewer primary annuli (Remarks. All the new specimens were collected by digging soil with hoes, particularly under rotting wood or leaf litter or between the buttress roots of trees. We collected 13 specimens (11 subsequently preserved) of Microcaecilia iwokramae in an estimated total of approximately 56 person hours of digging across altitudes of c. 100��� 850 m. All specimens were swabbed and found to be negative in a PCR-assay for the presence of chytrid (Gower et al., 2013). These findings suggests that there is a healthy population of M. iwokramae and that the species is reasonably common in the vicinity of the type locality. Wilkinson et al. (2013) recently provided a key to the species of Microcaecilia and a generic diagnosis that included reference to the presence of lungs (to distinguish Microcaecilia from Caecilita) that is no longer necessary., Published as part of Wilkinson, Mark, Kok, Philippe J. R., Ahmed, Farah & Gower, David J., 2014, Caecilita Wake & Donnelly, 2010 (Amphibia: Gymnophiona) is not lungless: implications for taxonomy and for understanding the evolution of lunglessness, pp. 383-388 in Zootaxa 3779 (3) on page 386, DOI: 10.11646/zootaxa.3779.3.6, http://zenodo.org/record/4910439, {"references":["Wake, M. H. & Donnelly, M. A. (2010) A new lungless caecilian (Amphibia: Gymnophiona) from Guyana. Proceedings of the Royal Society B, 277, 915 - 922. http: // dx. doi. org / 10.1098 / rspb. 2009.1662","Wilkinson, M., O'Connor, A. & Nussbaum, R. A. (2013) Taxonomic status of the neotropical caecilian genera Brasilotyphlus Taylor, 1968, Microcaecilia Taylor, 1968 and Parvicaecilia Taylor, 1968 (Amphibia: Gymnophiona: Siphonopidae). Occasional Papers of the Museum of Zoology, University of Michigan, 744, 1 - 12.","Gower, D. J., Doherty-Bone, T. M., Loader, S. P., Wilkinson, M., Kouete, T. M., Tapley, B., Orton, F., Daniel, O. Z., Wynne, F., Flach, E., Muller, H., Menegon, M., Stephen, I., Browne, R. K., Fisher, M. C., Cunningham, A. A. & Garner, T. W. J. (2013) Batrachochytrium dendrobatidis infection and lethal chytridiomycosis in caecilian amphibians (Gymnophiona), EcoHealth, 10, 173 - 178. http: // dx. doi. org / 10.1007 / s 10393 - 013 - 0831 - 9"]}

According to current understanding, five lineages of amphibians, but no other tetrapods, are secondarily lungless and are believed to rely exclusively on cutaneous gas exchange. One explanation of the evolutionary loss of lungs interprets lunglessness as an adaptation to reduce buoyancy in fast-flowing aquatic environments, reasoning that excessive buoyancy in such an environment would cause organisms being swept away. While not uncontroversial, this hypothesis provides a plausible potential explanation of the evolution of lunglessness in four of the five lungless amphibian lineages. The exception is the most recently reported lungless lineage, the newly described Guyanan caecilian genus and species Caecilita iwokramae Wake & Donnelly, 2010, which is inconsistent with the reduced disadvantageous buoyancy hypothesis by virtue of it seemingly being terrestrial and having a terrestrial ancestry. Re-examination of the previously only known spec- imen of C. iwokramae and of recently collected additional material reveal that this species possesses a reasonably well-developed right lung and is a species of the pre-existing caecilian genus Microcaecilia Taylor, 1968. We therefore place Caecilita in the synonymy of Microcaecilia, and re-evaluate the plausibility of the reduced disadvantageous buoyancy hypothesis as a general explanation of the evolution of lunglessness.

Pristimantis espedeus sp. nov. (Figs 2, 4��� 5, Table 1) Eleutherodactylus sp. 1 Marty & Gaucher (1999) Eleutherodactylus sp. 2 Lescure & Marty (2001) Pristimantis sp. 2 Dewynter et al. (2008); Fouquet et al. (2012) Holotype. R 121 (AF 1520), an adult male collected by EC and AF in Montagne Grande Tortue, (4.30901, - 52.357163; 370 m elevation), municipality of R��gina, French Guiana, on 13 June 2013 at 18 h 40. The specimen was recorded and photographed prior to capture (Fig. 2). Paratypes (n= 17). R 116���117 (AF0277���0278), two males collected on 10 May 2007 near the Nouragues station in the R��serve Naturelle des Nouragues (4.091667, - 52.7; 390 m elevation), municipality of R��gina, French Guiana; R 118���120 (AF1156, 1190, 1203), three males collected between 24 April and 3 May 2013 near the Aya camp in the R��serve Naturelle de la Trinit�� (4.619673, - 53.408797; 320 m elevation), municipality of St Elie, French Guiana; R 122���123 (AF 1521���1522), two males collected with the holotype; R 124 (AG 508) a male collected on 15 December 2012 in Monts Belv��d��re (3.719347, - 53.412809; 350 m elevation), municipality of Sa��l, French Guiana; R 125���131 (PG 452���455, 475 ��� 477), seven males collected on 1 February 2005 in Montagne Kotika (3.933333, - 54.1972222; 370 m elevation), municipality of Papa��chton, French Guiana; R 132 (PG 524), a male collected on 27 March 2006 on Pic Coudreau (3.306667, - 53.1472222; 400 m elevation), municipality of Sa��l, French Guiana; and R 133 (AG 270), a female collected on 12 November 2005 in Mont Lucifer (4.766667, - 53.9166667; 370 m elevation), municipality of St Laurent du Maroni, French Guiana (Fig. 1). Etymology. The specific name is considered a noun in apposition and derives from the old Norman language espedeus meaning ���special��� referring to the peculiar ecology and call of the new species (see Advertisement call, and Distribution and ecology sections below). Moreover, espedeus phonetically resembles ���sp. 2 ��� in French, which is how the species has been named for almost two decades. Definition and Diagnosis. The new species is characterized by the following unique combination of characters: (1) SVL small, adult males 23.0 �� 1.1 mm (range 20.7���24.8 mm, n= 17), adult female 29.4 mm (n= 1) (Table 1); (2) dorsal skin granular, slightly more granular posteriorly, with a few enlarged tubercles sometimes forming a W-shaped scapular fold and/or dorsal dermal folds, dorsal skin less tuberculate in females, ventral skin granular; (3) tympanic membrane differentiated, tympanic annulus visible externally, supratympanic fold obscuring upper and posterodorsal edges of tympanum, tympanum 40 % of eye length; (4) tibia length 1.9 X hand length (5) snout broadly rounded in profile and dorsal view; (6) each upper eyelid with two prominent tubercles; (7) choanae round and large (0.7 mm for the holotype), dentigerous processes of vomers oblique, narrowly separated, each bearing four teeth; (8) vocal slits present, vocal sac median, subgular, nuptial pads rugous translucid, barely visible, located on the dorsal surface of the proximal segment of Finger I; (9) Finger I shorter than the second, 72 % of Finger II; (10) fingers and toes with lateral fringes; (11) axillary tubercle absent; (12) ulnar tubercles barely visible or absent; (13) calcars absent, heel tubercles absent, inner metatarsal tubercle oval, much larger than the round outer tubercle; (14) discs broadly expanded, elliptical with broad lateral fringes, outer tubercle round, palmar tubercle heart-shaped; (15) in preservative, dorsum tan, gray and brown, three colour patterns exist: dorsally banded, dorsolaterally banded, and streaked, ventral colouration creamish white with small dark spots becoming denser on throat; (16) iris golden or silver with a dark copper horizontal band; (17) anterior surface of thighs and groin reddish in life and brown in preservative, fingers and toes with red spots in life, lack of yellow circumscribed spot on groin; (18) advertisement call characterized by clusters (0.2��� 0.5 s long) of 3���5 very short notes (0.02��� 0.03 s long) with a dominant frequency ranging between 2.7���2.8 kHz and emitted every 1.6��� 5.3 s; (19) calling activity exclusively crepuscular; (20) occurring at elevations between 200���700 m elevation on massifs reaching> 400 m. SVL TiL FeL TaL FL HeL HW Ind IOD P. espedeus sp. nov. M (17) 22.95 12.46 11.69 6.32 9.72 8.56 8.23 2.11 2.51 sd 1.07 0.68 0.53 0.46 0.49 0.41 0.33 0.21 0.23 F (1) 29.40 16.30 16. 0 0 8.30 13.40 11.20 11.00 2.50 3.50 sd NA NA NA NA NA NA NA NA NA P. inguinalis M (11) 20.23 10.34 9.90 5.48 8.12 7.60 7.07 1.74 2.04 sd 1.09 0.66 0.67 0.45 0.49 0.36 0.51 0.17 0.17 P. sp. 1 M (13) 17.66 9.51 9.01 5.09 7.35 6.69 5.95 1.62 2.06 sd 1.08 0.46 0.58 0.18 0.52 0.59 0.50 0.20 0.17 F (2) 22.20 11.50 11. 0 0 6.40 8.95 8.4 7.45 1.80 2.3 sd 0.42 0.42 0.28 0. 0 0 0.07 0.14 0.49 0.14 0. 0 0 P. sp. 4 M (5) 16.36 9.36 8.68 5.32 7.38 6.28 5.30 1.74 1.96 sd 0.85 0.57 0.54 0.43 0.37 0.33 0.46 0.17 0.22 F (6) 22.69 12.20 11.47 7.10 9.57 8.43 7.85 2.05 2.52 sd 1.57 0.51 0.67 1.24 1.43 0.58 0.47 0.21 0.17 EN ED TD FD 4 TD ETS 1 FiL 2 FiL HL P. espedeus sp. nov. M (17) 2.74 3.06 1.19 1.39 1.29 4.05 3.01 4.19 6.56 sd 0.26 0.22 0.25 0.09 0.11 0.24 0.49 0.49 0.41 F (1) 4.30 4.01 2.30 1.70 1.60 5.20 4.50 6.10 8.60 sd NA NA NA NA NA NA NA NA NA P. inguinalis M (11) 2.44 2.83 0.97 1.10 1.02 3.49 2.47 3.31 5.62 sd 0.27 0.12 0.09 0.12 0.10 0.25 0.15 0.15 0.34 P. sp. 1 M (13) 2.18 2.52 0.85 0.78 0.75 3.24 2.21 2.94 4.63 sd 0.32 0.21 0.14 0.12 0.13 0.19 0.24 0.34 0.24 F (2) 3.00 2.80 0.95 1.00 1.00 4.15 2.60 3.40 5.25 sd 0.00 0.00 0.21 0.14 0.14 0.35 0.00 0.28 0.49 P. sp. 4 M (5) 2.10 2.32 0.62 0.84 0.80 3.18 2.00 2.58 4.48 sd 0.32 0.25 0.04 0.11 0.12 0.23 0.23 0.28 0.33 F (6) 3.05 3.33 0.73 1.15 1.20 4.34 2.85 3.65 6.02 sd 0.22 0.14 0.10 0.18 0.14 0.21 0.33 0.58 0.34 Morphological comparisons with congeneric species. Using the raw data, MDFA clearly segregates well the four species (Fig. 3 a). Three variables notably contribute to the discrimination in opposite ways: HL and HW on the one hand, and TiL on the other. When the MDFA is based on the residuals, discrimination is less clear, especially between Pristimantis espedeus sp. nov. and P. sp. 1 (Fig. 3 b), suggesting that most of the morphological variation is size-based between these species (Fig. 3 c). Nevertheless, the variables HL and TiL still contribute appreciably and in opposite ways. Therefore, we plotted the ratio between these two variables. In P. espedeus sp. nov. and P. inguinalis, TiL is less than 50 % of HL, while it accounts for> 50 % in most specimens of P. sp. 1 and P. sp. 4 (Fig. 3 d, Table 2). Compared to the three other species of the unistrigatus group occurring in French Guiana (i.e. Pristimantis inguinalis, P. sp. 1 and P. sp. 4, the last two were previously partly misidentified with P. marmoratus in French Guiana e.g. in Lescure & Marty 2001), Pristimantis espedeus sp. nov. differs mainly by its larger SVL in males: mean of 23.0 mm (n= 17) in P. espedeus sp. nov. vs. mean of 20.2 mm (n= 11) in P. inguinalis, mean of 17.7 mm (n= 13) in P. sp. 1, and mean of 16.4 mm (n= 5) in P. sp. 4 (Table 1; Fig. 3). Pristimantis espedeus sp. nov. further differs from P. sp. 1 and P. sp. 4 by its red-orange inner thighs and groin in life (dark grey in P. sp. 1 and P. sp. 4), a more granular skin, a less clear W-shaped scapular fold, a proportionally larger head (Fig. 4), a less acuminate snout, more prominent tubercles on arms, larger finger discs, and a heart-shaped palmar tubercle (elliptical in P. sp. 1 and P. sp. 4). From P. inguinalis the new species mainly differs by its red-orange inner thighs and groin in life (a bright, well circumscribed yellow spot in P. inguinalis), and a cream ventral colouration (black in P. inguinalis). From the species of the unistrigatus group reported to occur below ca. 700 m elevation in the Guiana Shield [i.e. P. ockendeni (Boulenger, 1912) and P. marmoratus (Boulenger, 1900)], P. espedeus sp. nov. is readily diagnosable by its body size: P. ockendeni is larger (adult female holotype = 34.0 mm vs. 29.4 mm in P. espedeus sp. nov. female), and P. marmoratus is obviously smaller (adult male holotype [with large vocal slits present] = 18.0 mm vs. min. 20.7 mm in P. espedeus sp. nov. males). Pristimantis espedeus sp. nov. further differs from P. ockendeni by its reddish groin and thighs (absent in P. ockendeni), and from P. marmoratus by its cream venter (the preserved holotype of P. marmoratus has a dark grey venter suggestive of a dark ventral colouration in life). Pristimantis ockendeni from Manaus (Lima et al. 2006) apparently corresponds to a still undescribed species given its smaller body size compared to the holotype of P. ockendeni, and can be easily distinguished from P. espedeus sp. nov. by call characteristics (see below). Three other species associated with the name P. ockendeni have been described by Elmer and Cannatella (2008) from the lowlands of Ecuador and do not correspond to P. espedeus sp. nov. given their smaller SVL and the absence of reddish colour on groin and thighs. All the other species of the unistrigatus group known from the Guiana Shield occur in the Pantepui region and are generally associated with highlands (i.e. > 700 m elevation). Among the Pristimantis species found at midelevation in Pantepui, P. espedeus sp. nov. can mainly be distinguished from P. jester (Means & Savage, 2007) by the presence of a tympanum (absent in P. j e s t e r) and presence of fringes on fingers (absent in P. jester), from P. saltissimus (Means & Savage, 2007) by a slightly larger body size (16 mm ��� 27.1 mm in P. saltissimus vs. 20.7���29.4 mm in P. espedeus sp. nov.), a large, prominent tympanum (present, but indistinct in P. saltissimus), and presence of fringes on fingers (absent in P. saltissimus), from P. guaiquinimensis (Schl��ter & R��dder, 2007) by its smaller body size (females 32.4���33.6 mm in P. guaiquinimensis [see Kok & Barrio 2013] vs. 29.4 mm in P. espedeus sp. nov. female), a distinct tympanum (weakly distinct), and presence of fringes on fingers (absent), from P. sarisarinama (Barrio-Amor��s & Brewer-Carias, 2008) by the presence of nuptial pads in males (reported to be absent in P. sarisarinama), tubercles on eyelids (absent in P. sarisarinama), and distinct vocalisation (1���2, rarely 3 notes in P. sarisarinama vs. 3���5 notes in P. espedeus sp. nov.), from P. pulvinatus (Rivero, 1968) by its slightly smaller body size (males 23.0��� 26.1 mm in P. pulvinatus vs. 20.7���24.8 mm in P. espedeus sp. nov.) and distinct vomerine teeth (indistinct or absent in P. pulvinatus), from P. memorans (Myers & Donnelly, 1997) by the presence of fringes on fingers (absent in P. memorans) and a higher internote interval (0.20��� 0.29 s in P. memorans vs. 0.066��� 0.124 s in P. espedeus sp. nov.). Pristimantis espedeus sp. nov. can also be distinguished from the upper Amazonian species of the unistrigatus group Pristimantis lirellus (Dwyer, 1995), P. croceoinguinis (Lynch, 1968), and P. imitatrix (Duellman, 1978), with which it shares phylogenetic affinities according to previous analyses (Fouquet et al. 2012) as well as from P. carvalhoi (Lutz & Kloss, 1952) and P. variabilis (Lynch, 1968) that are related to the previous species according to Dwyer (1995), by the absence of a bright yellow spot in the groin region [present in the aforementioned species according to Dwyer (1995)], and in having a reddish groin region instead. Description of Holotype (Figs. 2, 4��� 5). An adult male 23.8 mm SVL in excellent condition. Head longer than wide (HW 8.4 mm, HeL 9.3 mm; HL 90.3 % of HeL), cranial crests not detectable. In dorsal view, snout longer than eye length (ETS 130 % of ED), broadly rounded, canthus rostralis straight and loreal region slightly concave, eyenaris distance slightly shorter than eye length (EN 94 % of ED). Nares slightly protuberant, directed posterolaterally, visible from frontal and dorsal views. Upper eyelid tubercles slightly prominent on the granular skin. Tympanum distinct, 1.6 mm long, ca. 50 % of eye length (3.3 mm). Supratympanic fold conspicuous in life, originating at posterior corner of eye, failing to reach shoulder; post-rictal tubercles absent. Vocal sac present. Throat and under surface of thighs smooth, belly granular; posteroventral surface of thigh and cloacal region granular; cloacal sheath absent. Dentigerous processes of the vomers oblique, narrowly separated, each bearing four teeth. Vocal slits visible on the floor of the mouth. Dorsal skin granular (more granular posteriorly) with enlarged tubercles, including dermal folds on dorsal portion of the flanks. Hand length 29 % of SVL. Finger I 79 % of II. Relative length of fingers III> IV> II> I; adpressed Finger I fails to reach proximal edge of digital pad of Finger II; adpressed Finger IV reaches the intercalary cartilage of Finger III on the left side, the base of the disc of Finger III on the right side. Large, barely visible rugous translucid nuptial pad on the dorsal surface of the proximal segment of Finger I. Lateral fringes on all fingers, best developed pre-axially on Fingers II and III (Fig. 3). Finger discs broadly expanded, elliptical, broader than long, circumferential groove conspicuous, distal edge of disc rounded; disc of Finger III (1.4 mm) 2.1 times wider than t distal end of adjacent phalanx. Palmar tubercle large, well defined, not fully pigmented, deeply bifid; thenar tubercle large, protuberant, ovoid; supernumerary tubercles few, large (almost equal in size to subarticular tubercles), slightly protuberant; subarticular tubercles large, round and protuberant, one each on FI and FII, two each on FIII and FIV. Hind limbs moderate in length, heels slightly overlap when held at right angles to sagittal plane; TiL 54 % of SVL; FL 44 % of SVL. Relative length of toes IV> V> III> II> I; tip of Toe V extends to the distal edge of the distal subarticular tubercle on Toe IV; tip of Toe III extends to the proximal half of the penultimate subarticular tubercle on Toe IV. Lateral fringes on all toes, best developed pre-axially on Toes III���IV; webbing basal between Toe IV���V (Fig. 2 c). Toe discs mostly equal in size to finger discs, WTD/WFD = 1; toe discs broadly expanded, elliptical, broader than long, circumferential groove conspicuous, distal edge of disc rounded. Inner metatarsal tubercle elongate, oval, about four times the size of the projecting, round outer metatarsal tubercle; subarticular tubercles round, large and protuberant; supernumerary plantar tubercles small, low and round, increasing in size distally. Calcars absent; no outer tarsal tubercles detectable in preservative; inner tarsal fold not detectable. Colour of the Holotype in preservative (Fig. 2 a���f). Dorsal background colour light brown, with a broad dark brown vertebral stripe extending from snout to vent, and two large laterodorsal dark brown bands extending from anterior corner of eye to cloaca. Two dark brown bands on the forearms and three well-defined brown bands on shanks and tarsi, ill-defined bands on thighs. Cloacal area dark brown. Five ill-defined dark bands on the upper lip and the loreal region; black supratympanic ridge obscuring upper part of tympanum; black band extending from tip of snout, tapering to anterior corner of eye. Dark stripe on lower edge of orbit. Ventral colouration creamish white with small dark spots denser on throat and thighs. Colour of the Holotype in life (Fig. 2 g). Dorsal background colour chestnut brown, with a broad dark brown stripe extending from snout to vent flanked by two large laterodorsal dark brown bands extending from anterior corner of eye to cloaca. Dorsal half of flanks yellowish brown with ill-defined oblique dark brown reticulations. Loreal region yellowish brown with a series of ill-defined greyish brown markings on the upper lip. Arms and legs yellowish brown with transversal dark bands. Groin and anterior surface of thighs bright red. Red markings also visible on fingers and toes. Belly creamish white with yellowish granular tubercles; throat similarly coloured, but with small dark brown spots. Variation. Pristimantis espedeus is highly polychromatic dorsally. Three main patterns exist: a dorsally banded morph (cf. holotype), a dorsolaterally banded morph (two dorsolateral reddish brown bands in life; R 120, 124, 133 in Fig. 5), and a streaked morph (with more or less well-defined chevron-like markings; R 118, 122, 123 in Fig. 5). A W-shaped dark brown scapular fold is sometimes visible particularly in the streaked morph. Sexual dimorphism can be evaluated based on the only female collected. The female is much larger than the largest male, the skin is less granular, and thumbs lack nuptial pads. Female colouration is similar to the dorsolaterally banded morph observed in some males. No additional significant difference was detected. Advertisement call (Fig. 6, Table 3). The following description is based on three recordings, one of the holotype (Montagne Grande Tortue) and two from uncollected individuals from the R��serve Naturelle des Nouragues and the R��serve Naturelle de la Trinit��, respectively. The advertisement call consists of short clusters (X= 0.33 s; range= 0.24��� 0.5 s) of generally 4 (3 to 5) short notes (X= 0.02 s; range= 0.019��� 0.025 s) having an internote interval of 0.09 s (range= 0.066��� 0.124 s) emitted between silent intervals of 3.21 s (range= 1.58��� 5.29 s) (Fig. 6). The spectral structure consists of three harmonics, with the fundamental frequency dominating (X= 2.7 kHz, range= 2.68���2.84 kHz, Fig. 6). Two secondary harmonics are visible (but not shown in Fig. 6), having a frequency of ca. 5.4 and 8.2 kHz, respectively. The vocalization of Pristimantis espedeus sp. nov. is easily distinguished from that of the two sympatric species with known advertisement calls (the call of P. sp. 4 remains undocumented): P. inguinalis emits a single note of ca. 3 kHz every 3���5 s, and P. sp. 1 emits clusters of 7���10 very short notes of ca. 4 kHz. The species of the unistrigatus group occurring in Manaus referred to as P. ockendeni by Lima et al. (2006) emits shorter clusters (0.20&ndash, Published as part of Fouquet, Antoine, Martinez, Quentin, Courtois, Elodie A., Dewynter, Ma��l, Pineau, K��vin, Gaucher, Philippe, Blanc, Michel, Marty, Christian & Kok, Philippe J. R., 2013, A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately evelated massifs of French Guiana, pp. 569-586 in Zootaxa 3750 (5) on pages 572-581, DOI: 10.11646/zootaxa.3750.5.8, http://zenodo.org/record/218859

Fouquet, Antoine, Martinez, Quentin, Courtois, Elodie A., Dewynter, Maël, Pineau, Kévin, Gaucher, Philippe, Blanc, Michel, Marty, Christian, Kok, Philippe J. R. (2013): A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately evelated massifs of French Guiana. Zootaxa 3750 (5): 569-586, DOI: http://dx.doi.org/10.11646/zootaxa.3750.5.8

Pristimantis imthurni sp. nov. Figs 8-10, Table 1 urn:lsid:zoobank.org:act: 9AFC18C4-23A0-4C20-96EB-480F0F512791 Eleutherodactylus sp. H ��� McDiarmid & Donnelly 2005: 514 [table 18A.1], 521. Definition and diagnosis A small species of the genus Pristimantis currently not assigned to any species group, but morphologically most similar to species of the polyphyletic unistrigatus species group (sensu Hedges et al. 2008, see also discussion in Kok et al. 2011) mainly characterized in having Finger I shorter than II, Toe V longer than III, extending to the distal edge of the distal subarticular tubercle of Toe IV when toes are adpressed, and by the absence of cranial crests and the presence of vomerine teeth. The new species is characterized by the following unique combination of characters: (1) body small, male 22.9 mm SVL; (2) dorsal skin smooth, belly skin coarsely areolate; (3) tympanum absent (tympanic membrane not differentiated and tympanic annulus not visible externally); (4) tiny pharyngeal ostia present; (5) snout broadly rounded in dorsal view and in profile, canthus rostralis nearly straight in dorsal view, rounded in cross section, loreal region concave, flaring slightly at upper lip; (6) upper eyelid smooth with one barely distinct enlarged tubercle; (7) choanae very small, round, dentigerous processes of vomers inconspicuous, ovoid, posterior and medial to choanae, each bearing 3-4 teeth; (8) absence of vocal slits in male; (9) tongue cordiform; (10) two large, broadly connected, mostly unpigmented nuptial pads on each thumb in male; (11) Finger I shorter than II; (12) fingers with lateral fringes, best developed preaxially on Fingers II-III; (13) ulnar tubercles absent; (14) tarsal tubercles absent, one inconspicuous calcar tubercle present; (15) inner metatarsal tubercle oval, about four times the size of the round, projecting outer metatarsal tubercle; (16) Toe V longer than III, extending to the distal edge of the distal tubercle on Toe IV when toes are adpressed; (17) toes with lateral fringes, best developed preaxially on Toes III-IV, webbing basal between Toes IV-V; (18) in life dorsal background colouration is vivid yellow with a chocolate brown middorsal stripe and chocolate brown flanks, ventral colouration is translucent brown with a few white spots and flecks; (19) in preservative dorsal background colouration is pale yellow with a light brown middorsal stripe and light brown flanks, ventral colouration is light grey with a few white spots and flecks. Etymology The specific epithet is a noun in the genitive case, honouring Sir Everard F. im Thurn (1852-1932), British colonial official, author, explorer, botanist, and photographer. Everard im Thurn was the first to climb a major tepui (Mount Roraima in December 1884), along with British surveyor Harry Perkins, a Pomeroon Amerindian named Gabriel, and five other unnamed Amerindians (Dalziell 2007). Im Thurn���s expedition on Roraima and his numerous discoveries were partly eclipsed by the popular novel that they inspired: ���The Lost World��� by Sir Arthur Conan Doyle (see Introduction; Dalziell 2007). Material examined Holotype VENEZUELA: an adult male collected by Philippe J. R. Kok, 20 Jun. 2012 at 20h37, summit of Ptaritepui, Bol��var state, 5��46���09��� N, 61��49���02��� W, 2471 m elevation, IRSNB 4165 (field number PK 3671). Morphological comparisons with congeneric species For the same reasons mentioned above, comparisons with congeners only focus on the 21 Pristimantis species known to occur in the Pantepui region. Pristimantis imthurni sp. nov. is most easily distinguished from P. abakapa, P. aureoventris, P. avius, P. cantitans, P. dendrobatoides, P. guaiquinimensis, P. marmoratus, P. memorans, P. pruinatus, P. pulvinatus, P. saltissimus, P. sarisarinama, P. vilarsi, P. yuruaniensis, and P. zeuctotylus by lacking a differentiated tympanic membrane and an external tympanic annulus (at least one of these structures is detectable in the latter 15 species). It further differs from P. vilarsi and P. zeuctotylus in having Finger I II in P. vilarsi and P. zeuctotylus). Pristimantis imthurni sp. nov. is immediately distinguished from P. auricarens, P. jester, P. marahuaka, P. muchimuk, and P. yaviensis in having lateral fringes on most fingers and toes (absent or limited to a weak keel in the latter 5 species). By lacking a differentiated tympanic membrane and an external tympanic annulus, and in having lateral fringes on fingers and toes, Pristimantis imthurni sp. nov. is morphologically most similar to P. jamescameroni sp. nov., described earlier in this paper. These two species are mainly distinguished based on distinct dorsal and ventral colour patterns (compare Fig. 2 with Fig. 8), skin texture (smooth in P. imthurni sp. nov. vs. shagreen in P. jamescameroni sp. nov.), condition of the supratympanic fold (inconspicuous in P. imthurni sp. nov. vs. conspicuous in P. jamescameroni sp. nov.), and head proportion (shorter, with a more rounded snout in P. imthurni sp. nov.). Besides being morphologically reliably diagnosable, the two species are currently geographically isolated (isolated lineage segments). Description of the holotype An adult male 22.9 mm SVL (Figs 8-10, see Table 1 for measurements), in very good condition, except a large scar under the right thigh where a piece of muscle was removed prior to preservation. Head slightly longer than wide (HW 97.7% of HL), wider than body; HW 37.1% of SVL; HL 38.0% of SVL; cranial crests absent. Snout longer than eye length (SL 112.1% of EL), broadly rounded in dorsal view and in profile; canthus rostralis nearly straight in dorsal view, rounded in cross section, loreal region concave, flaring slightly at upper lip; eye-naris distance shorter than eye length (EN 72.7% of EL). Nares slightly protuberant, directed posterolaterally, visible in frontal and dorsal views. Widest upper eyelid width narrower than interorbital distance (upper eyelid width 81.5% of IO), upper eyelid smooth with one barely visible enlarged tubercle on each eyelid. Tympanum absent (tympanic membrane not differentiated and tympanic annulus not visible externally); tiny pharyngeal ostia present. Supratympanic fold inconspicuous in life, barely visible and fragmented in preservative, slightly arched, apparently originating at posterior corner of eye, failing to reach shoulder; a few low and small post-rictal tubercles. Choanae very small, round; dentigerous processes of vomers inconspicuous, ovoid, posterior and medial to choanae, each bearing 3-4 teeth. Tongue cordiform, slightly wider than long, rounded posteriorly, posterior one-third free. Vocal slits and vocal sac absent. Dorsal skin smooth, including on head; middorsal raphe barely detectable in preservative, well visible in life; no dermal folds or ridges visible on dorsal surface; flanks slightly granular. Throat and upper chest surfaces smooth; no thoracic fold detectable; belly skin coarsely areolate; no discoidal fold detectable; posteroventral thigh and cloacal region coarsely areolate; cloacal sheath absent, two distinctly enlarged tubercles below cloaca. Hand length 27.1% of SVL. Finger I 88.6% of II. Relative length of fingers III> IV> II> I; adpressed Finger I fails to reach proximal edge of digital pad of Finger II; adpressed Finger IV reaches proximal edge of digital pad of Finger III. Two large, broadly connected, non-spinous, mostly unpigmented (translucent when wet, white when dry) nuptial pads on each thumb, extending from the preaxial surface of the thenar tubercle and invading most of it, along the dorsal and the preaxial surface of the thumb. Lateral fringes on all fingers, best developed preaxially on Fingers II-III (Fig. 9). Finger discs expanded, elliptical, broader than long, circumferential groove conspicuous, distal edge of disc rounded; disc of Finger III 1.8 times wider that the distal end of the adjacent phalanx. Palmar tubercle large, well defined, fully pigmented, deeply bifid; thenar tubercle large, protuberant, ovoid; supernumerary tubercles few, of variable sizes, slightly protuberant; subarticular tubercles large, round and protuberant, one each on FI and FII, two each on FIII and FIV. Ulnar tubercles absent; presence of a small antebrachial tubercle. Hind limbs moderate in length, heels slightly overlap when held at right angles to sagittal plane; TIL 50.2% of SVL; FL 40.6% of SVL. Relative length of Toes IV> V> III> II> I; tip of Toe V extends to the distal edge of the distal subarticular tubercle on Toe IV; tip of Toe III extends to the distal edge of the penultimate subarticular tubercle on Toe IV. Lateral fringes on all toes, best developed preaxially on Toes III-IV (Fig. 9); webbing basal between Toe IV-V. Toe discs slightly larger in size than finger discs, WTD/WFD = 1.08; toe discs expanded, elliptical, broader than long, circumferential groove conspicuous, distal edge of disc rounded. Inner metatarsal tubercle elongate, oval, about four times the size of the projecting, round outer metatarsal tubercle; subarticular tubercles round, large and protuberant; supernumerary plantar tubercles small, low and round, increasing in size distally. Single, inconspicuous, very small, round calcar tubercle; no outer tarsal tubercles detectable; inner tarsal fold not detectable. Colour of holotype in life (see Fig. 8) Dorsal background colouration vivid yellow, top of head greyish brown; chocolate brown middorsal stripe and chocolate brown band on flanks; ill-defined, irregular reddish orange dorsolateral line; a few reddish orange blotches on the posterior part of the dorsum; a few reddish orange minute speckles on flanks; brown facemask; enlarged tubercle on eyelid whitish. Lower flanks yellow; groins, anterior thighs, and armpits brown. Arms and legs greyish brown with a few ill-defined white and dark yellow blotches. Ventral surfaces translucent greyish brown with a few irregular white spots and flecks; posterior surface of thighs and cloacal area brown. Upper face of hands and feet greyish brown with a few white and yellow markings; upper face of tip of fingers white; upper face of tip of Toes I-III white, brown on Toes IV-V. Palms, soles, undersurface of fingers and toes brown; discs greyish brown. Iris silver with dark brown venation and an ill-defined broad horizontal brownish stripe. Colour of Holotype in preservative (see Fig. 10) After 13 months in 70% ethanol (July 2013). Vivid yellow faded to pale yellow. Chocolate brown and brown faded to light brown; top of head is dark brown; ill-defined white and dark yellow blotches on arms and legs are slightly more conspicuous and form irregular, ill-defined transverse bands; dorsal pattern turned generally more conspicuous. Ventral colouration is light grey with a few white spots and flecks. Granules in the cloacal area, as well as subarticular and supernumerary tubercles became dark brown. Discs are grey. Distribution and ecology Pristimantis imthurni sp. nov. is only known from the summit of Ptari-tepui (Figs 1B, 6B, 11), Bol��var state, Venezuela, where it occurs at 2471 m elevation. Ptari-tepui lies in the Ptari massif, in the Los Testigos subdistrict (McDiarmid & Donnelly 2005). It is located ca. 47 km NE of the Chimant�� massif, and ca. 22 km SE of Kamarkawarai-tepui (airline). According to the GPS, Ptari-tepui reaches a maximal elevation of about 2471 m above sea level. The summit area of Ptari-tepui is ca. 1 km 2 and is dominated by flat open rock surfaces and small islands of low meadow-like tepui vegetation (Huber 1995; Fig. 11B). The male holotype ��� and single specimen collected ��� was found at night while calling partly concealed in the muddy soil in a small vegetation patch. A few other males were heard calling very sporadically from other small vegetation islands by day or night, but could not be located. Pristimantis imthurni sp. nov. is not abundant. The only other anuran reported from the summit of Ptari-tepui is Tepuihyla rimarum Ayarzag��ena, Se��aris & Gorzula, 1993, which apparently also occurs in low density (pers. obs.)., Published as part of Kok, Philippe J. R., 2013, Two new charismatic Pristimantis species (Anura: Craugastoridae) from the tepuis of " The Lost World " (Pantepui region, South America), pp. 1-24 in European Journal of Taxonomy 60 on pages 15-21, DOI: 10.5852/ejt.2013.60, http://zenodo.org/record/3824956, {"references":["McDiarmid R. W. & Donnelly M. A. 2005. The herpetofauna of the Guayana Highlands: amphibians and reptiles of the Lost World. In: Donnelly M. A., Crother B. I., Guyer C., Wake M. H. & White M. E. (eds) Ecology and Evolution in the Tropics: A Herpetological Perspective: 461 - 560. University of Chicago Press, Chicago.","Hedges S. B., Duellman W. E. & Heinicke M. P. 2008. New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa 1737: 1 - 182.","Kok P. J. R, Means D. B. & Bossuyt F. 2011. A new highland species of Pristimantis Jimenez de la Espada, 1871 (Anura: Strabomantidae) from the Pantepui region, northern South America. Zootaxa 2934: 1 - 19.","Dalziell R. 2007. Everard im Thurn in British Guiana and the western Pacific. In: Hulme P. & McDougall R. (eds.) Writing, Travel and Empire: 97 - 116. I. B. Tauris & Co. Ltd, London.","Huber O. 1995. Geographical and physical features. In: Berry P. E., Holst B. K. & Yatskievych K. (eds) Flora of the Venezuelan Guayana. Volume 1. Introduction: 1 - 61. Missouri Botanical Garden Press, St Louis."]}

Pristimantis jamescameroni sp. nov. Figs 2-5, Table 1 urn:lsid:zoobank.org:act: 67EB6C36-0F31-4587-9805-3136BEF4E010 Eleutherodactylinae series b (in part) ��� Gorzula & Se��aris 1999: 55 + plate 46. Eleutherodactylus sp. B ��� McDiarmid & Donnell, 2005: 514 [table 18A.1], 524. Pristimantis sp. ��� Hedges et al. 2008: 179. Pristimantis sp. ���Aprada��� ��� Kok et al. 2012: Suppl. Information: 13. Definition and diagnosis A small species of the genus Pristimantis currently not assigned to any species group, but morphologically most similar to species of the polyphyletic unistrigatus species group (sensu Hedges et al. 2008, see also discussion in Kok et al. 2011) mainly characterized in having Finger I shorter than II, Toe V longer than III, extending to the distal edge of the distal subarticular tubercle of Toe IV when toes are adpressed, and by the absence of cranial crests and the presence of vomerine teeth. The new species is characterized by the following unique combination of characters: (1) body small, male 22.8 mm SVL, females 26.3-27.5 mm (n=3); (2) dorsal skin shagreen, belly skin coarsely areolate; (3) tympanum absent (tympanic membrane not differentiated and tympanic annulus not visible externally); (4) tiny pharyngeal ostia present; (5) snout rounded to subovoid in dorsal view, rounded to slightly sloping in profile, canthus rostralis nearly straight in dorsal view, rounded in cross section, loreal region slightly concave, flaring slightly at upper lip; (6) upper eyelid shagreen with 1-2 distinctly enlarged tubercles on each eyelid; (7) choanae small, oval, dentigerous processes of vomers very small, barely visible, slightly oblique, ovoid to triangular, posterior and medial to choanae, each bearing 1-5 teeth; (8) absence of vocal slits in male; (9) tongue cordiform; (10) two large, mostly unpigmented nuptial pads on each thumb in male; (11) Finger I shorter than II; (12) fingers with lateral fringes, best developed preaxially on Fingers II-III; (13) ulnar tubercles absent or inconspicuous, when present low, not forming a distinct line; (14) tarsal tubercles absent, one inconspicuous calcar tubercle present; (15) inner metatarsal tubercle oval, about four times the size of the round, projecting outer metatarsal tubercle; (16) Toe V longer than III, extending to the distal edge of the distal tubercle on Toe IV when toes are adpressed; (17) toes with lateral fringes, best developed preaxially on Toes III-IV, webbing basal between Toes IV-V; (18) in life main dorsal background colouration is orange to orangish dark brown, either with irregular dark brown chevrons or a darker middorsal band, and/or a light middorsal stripe, ventral colouration is white with brown reticulation and an orange patch posteriorly; (19) in preservative dorsal background colouration is light to dark brown, either with irregular brown chevrons or a darker middorsal band, and/or a light middorsal stripe, ventral colouration is cream with brown reticulation. Etymology The specific epithet is a noun in the genitive case, honouring the Canadian film director, producer, environmentalist and explorer Mr. James F. Cameron in recognition of his efforts to alert the general public to environmental problems through pioneering high quality ���blockbuster��� movies and adventurous documentaries. James Cameron also encourages people to go vegan (a diet excluding animal products), one of the effective ways to reduce human environmental impacts such as global climate change, identified as a serious threat to tepuis ecosystems (see Nogu�� et al. 2009). Material examined Holotype VENEZUELA: an adult male collected by Philippe J. R. Kok, 15 Jun. 2012 at 08h18, summit of Apradatepui, Bol��var state, 5��24���42���N, 62��27���00��� W, 2570 m elevation, IRSNB 4160 (field number PK 3636). Paratopotypes (n=4) VENEZUELA: three adult females (IRSNB 4161-63, field numbers PK 3632, PK 3637, PK 3635) and one juvenile (IRSNB 4164, field number PK 3639), collected by Philippe J. R. Kok (except IRSNB 4161, collected by Brad Wilson) 14-15 Jun. 2012, all from the summit of Aprada-tepui between 2557- 2571 m elevation. Description of the holotype An adult male 22.8 mm SVL (Figs 2 A-B, 3, 4D, H, see Table 1 for measurements), in very good condition, except a large scar under the right thigh where a piece of muscle was removed prior to preservation. Head slightly longer than wide (HW 94.7% of HL), wider than body; HW 39.0% of SVL; HL 41.2% of SVL; cranial crests absent. Snout longer than eye length (SL 117.1% of EL), rounded to subovoid in dorsal view, rounded to slightly sloping in profile; canthus rostralis nearly straight in dorsal view, rounded in cross section, loreal region slightly concave, flaring slightly at upper lip; eye-naris distance shorter than eye length (EN 71.4% of EL). Nares slightly protuberant, directed posterolaterally, visible in frontal and dorsal views. Widest upper eyelid width narrower than interorbital distance (upper eyelid width 82.7% of IO), upper eyelid shagreen with 1-2 enlarged tubercles on each eyelid. Tympanum absent (tympanic membrane not differentiated and tympanic annulus not visible externally); tiny pharyngeal ostia present. Supratympanic fold conspicuous in life, slightly arched, originating at posterior corner of eye, failing to reach shoulder; post-rictal tubercles evident. Choanae small, oval, not concealed by palatal shelf of maxillary arch; dentigerous processes of vomers very small, barely visible, slightly oblique, ovoid to triangular, posterior and medial to choanae, each bearing 3-4 teeth. Tongue cordiform, slightly longer than wide, rounded posteriorly, posterior half free. Vocal slits and vocal sac absent. Dorsal skin shagreen, including on head; middorsal raphe detectable; no dermal folds or ridges visible on dorsal surface; flanks granular. Throat surface shagreen; upper chest shagreen, slightly ���wrinkled���; weak thoracic fold; belly skin coarsely areolate; weak discoidal fold anterior to groin; posteroventral thigh and cloacal region coarsely areolate; cloacal sheath absent. Hand length 27.2% of SVL. Finger I 80.0% of II. Relative length of fingers III> IV> II> I; adpressed Finger I fails to reach proximal edge of digital pad of Finger II; adpressed Finger IV reaches the intercalary cartilage of Finger III on the left side, the base of the disc of Finger III on the right side. Two large, not connected, non-spinous, mostly unpigmented (translucent when wet, white when dry) nuptial pads on each thumb, one extending along the preaxial surface of the thenar tubercle and invading most of it, the other one extending along the dorsal and the preaxial surface of the thumb. Lateral fringes on all fingers, best developed preaxially on Fingers II-III (Fig. 3). Finger discs broadly expanded, elliptical, broader than long, circumferential groove conspicuous, distal edge of disc rounded; disc of Finger III 2.1 times wider that the distal end of the adjacent phalanx. Palmar tubercle large, poorly defined, not fully pigmented, deeply bifid; thenar tubercle large, protuberant, ovoid; supernumerary tubercles few, large (almost equal in size to the subarticular tubercles), slightly protuberant; subarticular tubercles large, round and protuberant, one each on FI and FII, two each on FIII and FIV. Ulnar tubercles few, inconspicuous, forming an ill-defined line; presence of a small antebrachial tubercle. Hind limbs moderate in length, heels slightly overlap when held at right angles to sagittal plane; TIL 50.4% of SVL; FL 41.7% of SVL. Relative length of Toes IV> V> III> II> I; tip of Toe V extends to the distal edge of the distal subarticular tubercle on Toe IV; tip of Toe III extends to the distal edge of the penultimate subarticular tubercle on Toe IV on the left side, to the proximal half of the penultimate subarticular tubercle on Toe IV on the right side. Lateral fringes on all toes, best developed preaxially on Toes III-IV; webbing basal between Toe IV-V (Fig. 3). Toe discs mostly equal in size to finger discs, WTD/WFD = 1; toe discs broadly expanded, elliptical, broader than long, circumferential groove conspicuous, distal edge of disc rounded. Inner metatarsal tubercle elongate, oval, about four times the size of the projecting, round outer metatarsal tubercle; subarticular tubercles round, large and protuberant; supernumerary plantar tubercles small, low and round, increasing in size distally. Single, inconspicuous and very small round calcar tubercle; no outer tarsal tubercles detectable; inner tarsal fold not detectable. Colour of holotype in life (see Fig. 2 A-B) Dorsal background colour orangish brown covered with numerous tiny white flecks, top of head slightly darker; inconspicuous darker (than adjacent dorsum) middorsal band outlined by ill-defined dark brown dorsolateral lines; irregular white band on the canthus rostralis; bluish white line on outer edge of upper eyelids; upper lips white; most post-rictal tubercles white; enlarged tubercles on eyelids light brown. Flanks orangish brown, with numerous tiny white flecks and some irregular white spots of variable sizes; groins, anterior thighs, and armpits bright orange. Arms and legs coloured as dorsal surfaces, with a few small irregular white spots on upper arms and on the distal part of tarsi. Throat, chest, and lower flanks white with brown anastomosed reticulation; belly and most of the undersurface of thighs bright orange; undersurface of distal thighs, shanks, and tarsi brown with a few small irregular white spots; posterior surface of thighs and cloacal area brown. Upper face of hands and feet orangish brown, except tip of Fingers I-II, which is bright orange, and tip of toes, which is whitish. Palms and soles brown; undersurface of Fingers I-III bright orange, including discs; undersurface of tip of Toes I-III bright orange including discs. Iris silver with dark brown venation and an ill-defined, broad horizontal brownish stripe; greenish silver spot on the posterior upper part of the iris. Colour of holotype in preservative (see Fig. 4D, H) After 13 months in 70% ethanol (July 2013), orangish brown faded to light brown. Orange and white faded to cream. Dorsal pattern turned generally more conspicuous. Brown reticulation on throat, chest, and lower flanks faded to light brown. Granules in the cloacal area, as well as subarticular and supernumerary tubercles became dark brown. Discs are greyish brown. Sexual dimorphism and variation among paratypes See Table 1 for measurements of the type series, and Figs 2, 4, 5 for intraspecific variation. Sexual dimorphism evident in size, with adult females being larger than the adult male (26.3-27.5 vs. 22.8 mm SVL), and by the presence of two nuptial pads in the male. A sexual dichromatism in belly colouration seems to occur, with the presence of an orange spot restricted to the middle area of the belly in living females, whereas the orange area covers the belly, the posterior part of the ventral surface, and the undersurface of thighs in a more continuous way in the living male (see Fig. 2B, D, F). Hands and feet slightly longer in the male than in the females (HAND III 27.2% vs. 24.0-25.1% of SVL; FL 41.7% vs. 37.8-40.3% of SVL). No additional significant difference is detected in other size ratios. No significant variation in skin texture occurs among the preserved specimens, but in life IRSNB 4163 (adult female, Fig. 2C) had a conspicuous dorsolateral fold extending from behind the eye to the end of the body; that fold totally disappeared in preservative. Eyelid tubercles are more difficult to detect in preserved specimens and range from 1 to 2 on each eyelid. Number of teeth on dentigerous processes of vomers varies from 2 to 6. Weak thoracic and discoidal folds detectable in IRSNB 4163 (female) and IRSNB 4164 (juvenile), not seen in the other paratypes. Tip of Toe V fails to reach the distal edge of the distal subarticular tubercle on Toe IV on the left side in IRSNB 4161 (female) only. Colour pattern variable. In life, dorsal background colouration varies from bright orange to orangish dark brown, usually covered with tiny white flecks; dorsal pattern varies from irregular brown chevrons (e.g., in IRSNB 4161 and IRSNB 4162) to a slightly darker (than adjacent dorsum) middorsal band outlined by ill-defined dark brown dorsolateral lines (e.g., in IRSNB 4160) and/or a pale middorsal stripe (e.g., in IRSNB 4163); pattern on flanks varies from a few irregular oblique white (e.g., IRSNB 4163) or brown/white (e.g., IRSNB 4162) stripes to white spots (holotype); colour of ventral reticulation varies from brown to dark brown; inconspicuous transverse bands on arms and legs occur in two female paratypes (IRSNB 4162 and IRSNB 4163). Enlarged tubercles on eyelids are usually white (light brown in the holotype). The juvenile (IRSNB 4164, Fig. 5) has a complex and more marked dorsal pattern consisting of a greyish brown interorbital line preceded by a greyish brown circle and followed by a wide W-shaped marking outlined by greyish brown in the scapular region; the W-shaped marking is followed by a greyish brown arch, itself followed by two back to back C-shaped light grey lines. It also differs from adults in having an irregular greyish band on the canthus rostralis, the upper lip not completely white, granules in the loreal region, dark brown labial stripes, the supratympanic fold underlined by a dark brown band, brown transverse bands on the forearm, and reddish transverse bands on the legs. Iris varies from silver or greenish silver to bronze, with dark brown venation and an illdefined, broad horizontal brownish stripe. Morphological comparisons with congeneric species Available data suggest that tepui summit Pristimantis species have relatively restricted distributions in the Guinea Shield highlands; therefore, comparisons with congeners only focus on Pristimantis species known to occur in the Pantepui region (as defined by Kok 2013). Comparisons of external character states are based both on original descriptions and examination of museum specimens (see Appendix for material examined). Twenty species of Pristimantis are currently reported from the Pantepui region: P. aureoventris Kok, Means & Bossuyt, 2011, only known from the summit of Wei-Assipu-tepui and the upper slopes of Mount Roraima, Cuyuni-Mazaruni District, Guyana; P. abakapa Rojas-Runjaic, Salerno, Se��aris & Pauly, 2013, described from the summit of Abakap��-tepui, Bol��var state, Venezuela; P. auricarens (Myers & Donnelly, 2008), known only from Auy��n-tepui, Bol��var state, Venezuela; P. avius (Myers & Donnelly, 1997), reported from Pico Tamacuari, Amazonas state, Venezuela; P. cantitans (Myers & Donnelly, 1996), known from Cerro Yav�� and Cerro Yutaj��, Amazonas state, Venezuela; P. dendrobatoides Means & Savage, 2007, only known from the Wokomung massif, Potaro-Siparuni District, Guyana; P. guaiquinimensis (Schl��ter & R��dder, 2007), described from Guaiquinima-tepui, Bol��var state, Venezuela; P. jester Means & Savage, 2007, known from the Wokomung massif, Potaro-Siparuni District, Guyana, and the slopes of Maringma-tepui, Cuyuni-Mazaruni District, Guyana (Kok, pers. obs. 2007); P. marahuaka (Fuentes-Ramos & Barrio-Amor��s, 2004), endemic to Cerro Marahuaka, Amazonas state, Venezuela; P. marmoratus (Boulenger, 1900), reported as widespread in the Guiana Shield from eastern Venezuela to French Guiana and northern Brazil (Frost 2013), but several species may hide under this name; P. memorans (Myers & Donnelly, 1997), known only from the Sierra Tapirapec��, Amazonas state, Venezuela; P. muchimuk Barrio-Amor��s, Mesa, Brewer-Car��as & McDiarmid 2010, apparently endemic to the summit of Chur��-tepui, Bol��var state, Venezuela; P. pruinatus (Myers & Donnelly, 1996), known only from Cerro Yav��, Amazonas state, Venezuela; P. pulvinatus (Rivero, 1968), reported from the Gran Sabana region and Auy��n-tepui, Bol��var state, Venezuela to western Guyana (Frost 2013), but more than one species may hide under this name (see Kok et al. 2012); P. saltissimus Means & Savage, 2007, known from the Wokomung massif, Potaro-Siparuni District, Guyana, and the slopes of Maringma-tepui, Cuyuni-Mazaruni District, Guyana (Kok pers. obs. 2007); P. sarisarinama Barrio- Amor��s & Brewer-Car��as, 2008, endemic to Sarisari��ama-tepui, Bol��var state, Venezuela; P. vilarsi (Melin, 1941), known from upper Amazonian Brazil, Colombia, Venezuela and Peru (Frost 2013); P. yaviensis (Myers & Donnelly, 1996), reported from Cerro Yav�� and Cerro Yutaj��, Amazonas state, Venezuela; P. yuruaniensis R��dder & Jungfer, 2008, known with certainty only from the summit of Yuruan��-tepui, Bol��var state, Venezuela (possible occurrence on the neighbouring Kuken��n-tepui, see M��gdefrau & M��gdefrau 1994; R��dder & Jungfer 2008; Kok et al. 2011); and P. zeuctotylus (Lynch & Hoogmoed, 1977), widespread in northeastern South America (Frost 2013). Pristimantis jamescameroni sp. nov. is most easily distinguished from P. abakapa, P. aureoventris, P. avius, P. cantitans, P. dendrobatoides, P. guaiquinimensis, P. marmoratus, P. memorans, P. pruinatus, P. pulvinatus, P. saltissimus, P. sarisarinama, P. vilarsi, P. yuruaniensis and P. zeuctotylus by lacking a differentiated tympanic membrane and an external tympanic annulus (at least one of these structures is detectable in the latter 15 species). It further differs from P. vilarsi and P. zeuctotylus in having Finger I II in P. vilarsi and P. zeuctotylus). Pristimantis jamescameroni sp. nov. is immediately distinguished from P. auricarens, P. jester, P. marahuaka, P. muchimuk, and P. yaviensis (the only known Pantepui ���earless��� Pristimantis) in having conspicuous lateral fringes on fingers and toes (absent or limited to a weak keel in the latter 5 species). Distribution and ecology Pristimantis jamescameroni sp. nov. is only known from the summit of Aprada-tepui (Figs 1B, 6A, 7 A-B), Bol��var state, Venezuela, where it occurs from 2557 to 2571 m elevation. Aprada-tepui lies in the Aprada massif (Fig. 6A), in the Chimant�� subdistrict (McDiarmid & Donnelly 2005). It is located ca. 22 km NW of the Chimant�� massif and ca. 30 km S of Auy��n-tepui (airline). According to the GPS, Aprada-tepui reaches a maximal elevation of about 2575 m above sea level. The summit area of Aprada-tepui is ca. 4.3 km 2, and is mainly covered by open rock vegetation and small islands of tepui forests (Huber 1995, Fig. 7B); it is characterized by a high number of small lakes (Fig. 7B) and some deep canyons. Aside from the male holotype, which was collected by day on the ground while calling in a small shallow rock crevice, hidden by the veget, Published as part of Kok, Philippe J. R., 2013, Two new charismatic Pristimantis species (Anura: Craugastoridae) from the tepuis of " The Lost World " (Pantepui region, South America), pp. 1-24 in European Journal of Taxonomy 60 on pages 5-15, DOI: 10.5852/ejt.2013.60, http://zenodo.org/record/3824956, {"references":["Gorzula S. & Senaris J. C. 1999 \" 1998 \". Contribution to the herpetofauna of the Venezuelan Guayana. I. A data base. Scientia Guaianae 8: 1 - 269.","McDiarmid R. W. & Donnelly M. A. 2005. The herpetofauna of the Guayana Highlands: amphibians and reptiles of the Lost World. In: Donnelly M. A., Crother B. I., Guyer C., Wake M. H. & White M. E. (eds) Ecology and Evolution in the Tropics: A Herpetological Perspective: 461 - 560. University of Chicago Press, Chicago.","Hedges S. B., Duellman W. E. & Heinicke M. P. 2008. New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa 1737: 1 - 182.","Kok P. J. R., MacCulloch R. D., Means D. B., Roelants K., Van Bocxlaer I. & Bossuyt F. 2012. Low genetic diversity in tepui summit vertebrates. Current Biology 22 (15): 589 - 590 + supplementary information. http: // dx. doi. org / 10.1016 / j. cub. 2012.06.034","Kok P. J. R, Means D. B. & Bossuyt F. 2011. A new highland species of Pristimantis Jimenez de la Espada, 1871 (Anura: Strabomantidae) from the Pantepui region, northern South America. Zootaxa 2934: 1 - 19.","Nogue S., Rull V. & Vegas-Vilarrubia T. 2009. Modeling biodiversity loss by global warming on Pantepui, northern South America: projected upward migration and potential habitat loss. Climatic Change 94: 77 - 85. http: // dx. doi. org / 10.1007 / s 10584 - 009 - 9554 - x","Kok P. J. R. 2013. Islands in the Sky: Species Diversity, Evolutionary History, and Patterns of Endemism of the Pantepui Herpetofauna. PhD thesis, Leiden University, The Netherlands.","Rojas-Runjaic F. J. M., Salerno P. E., Senaris J. C. & Pauly G. B. 2013. Terraranans of the Lost World: a new species of Pristimantis (Amphibia, Craugastoridae) from Abakapa-tepui in the Chimanta massif,","Myers C. W. & Donnelly M. A. 2008. The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History 308: 1 - 147. http: // dx. doi. org / 10.1206 / 308.1","Means D. B. & Savage J. M. 2007. Three new malodorous rainfrogs of the genus Pristimantis (Anura: Brachycephalidae) from the Wokomung Massif in west-central Guyana, South America. Zootaxa 1658: 39 - 55.","guaiquinimensis (Schluter & Rodder, 2007). Zootaxa 3694 (1): 75 - 80. http: // dx. doi. org / 10.11646 / zootaxa. 3694.1.6","Fuentes-Ramos O. & Barrio-Amoros C. L. 2004. A new Eleutherodactylus (Anura, Leptodactylidae) from Marahuaka Tepui, Amazonas, Venezuela. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 28: 285 - 290.","Frost D. R. 2013. Amphibian Species of the World: an Online Reference. Version 5.5 (9 Jan. 2013). American Museum of Natural History, New York [online]. Available from http: // research. amnh. org / herpetology / amphibia / index. php [accessed 20 Jul. 2013].","Barrio-Amoros C. L., Mesa J., Brewer-Carias C. & McDiarmid R. W. 2010. A new Pristimantis (Anura, Terrarana, Strabomantidae) from Churi-tepui in the Chimanta massif, Venezuelan Guayana. Zootaxa 2483: 35 - 44.","Rodder D. & Jungfer K-H. 2008. A new Pristimantis (Anura, Strabomantidae) from Yuruani-tepui, Venezuela. Zootaxa 1814: 58 - 68.","Magdefrau H. & Magdefrau K. 1994. Biologie von Anuren auf Tepuis der Roraima-Gruppe in Venezuela. In: Herrmann H. - J. & Zimmermann H. (eds) Beitrage zur Biologie der Anuren: 84 - 102. Tetra Werke, Melle.","Lynch J. D. & Hoogmoed M. S. H. 1977. Two new species of Eleutherodactylus (Amphibia, Leptodactylidae) from northeastern South America. Proceedings of the Biological Society of Washington 90: 424 - 439.","Huber O. 1995. Geographical and physical features. In: Berry P. E., Holst B. K. & Yatskievych K. (eds) Flora of the Venezuelan Guayana. Volume 1. Introduction: 1 - 61. Missouri Botanical Garden Press, St Louis."]}

Pristimantis aureoventris sp. nov. (Figs. 2���7, Table 1) Holotype. IRSNB 4152 (field number PK 2152), an adult male collected by Philippe J. R. Kok, Paul Benjamin and Claudius Perry, 17 November 2009 at 20 h00, summit of Wei Assipu Tepui, Cuyuni-Mazaruni District, Guyana (05�� 13 ��� 05���N, 060�� 42 ��� 15 ���W, 2210 m elevation). Paratopotypes (n= 2). IRSNB 4153 (field number PK 2066), a female collected by Philippe J. R. Kok, Paul Benjamin and Claudius Perry, 3 November 2009, and IRSNB 4154 (field number PK 2087), a subadult female collected by Philippe J. R. Kok, Paul Benjamin and Claudius Perry, 4 November 2009. Referred specimen (n= 10). IRSNB 15821 ��� 22 (field numbers PK 2162 ��� 63), females; IRSNB 15820, IRSNB 15823 (field numbers PK 2085, PK 2086), males; IRSNB 15824 ��� 27 (field numbers PK 2107 ���08, PK 2114, PK 2117), males; IRSNB 15637 (field number PK 2158), a very small juvenile, all collected by Philippe J. R. Kok, Paul Benjamin and Claudius Perry on the summit of Wei Assipu Tepui between 3���19 November 2009. IRSNB 15643 (field number CPI 10484), a male collected by D. Bruce Means, 17 November 2006, from ca. 300 m below the base of the ultimate cliff of the northern ���Prow��� of Mount Roraima, Cuyuni-Mazaruni District, Guyana (05�� 15 ���N, 060�� 43 ���W, 2305 m elevation). Etymology. The specific name aureoventris is a noun in apposition derived from the Latin words ��� aureus ��� meaning golden, and ��� ventris ��� meaning venter, and refers to the golden ventral face of the new species. Generic allocation. To date, there is no identifiable morphological synapomorphy supporting the genus Pristimantis (Hedges et al. 2008). We assign the new taxon to the genus Pristimantis based on molecular phylogenetic relationships (the authors, unpublished data) as well as on its morphological characteristics, which fall into the range of other Pristimantis species. Definition and diagnosis. A small species of the genus Pristimantis currently not assigned to any species group, but morphologically most similar to species of the non-monophyletic unistrigatus species group (sensu Hedges et al. 2008, see Discussion) mainly characterized in having Finger I shorter than II, Toe V longer than III, extending to the distal edge of the distal subarticular tubercle of Toe IV when toes are adpressed, and by the absence of cranial crests and the presence of vomerine teeth. The new species is characterized by the following combination of characters: (1) dorsal skin shagreened to slightly granular, rarely with some conical tubercles, faint middorsal raphe in life, ventral skin areolate; (2) tympanum distinct, less than half the size of the eye; (3) snout rounded to subovoid in dorsal view, rounded to slightly sloping in profile, canthus rostralis nearly straight to concave, round; (4) upper eyelid smooth to slightly granular, usually with one or two distinctly enlarged tubercles on each eyelid; (5) choanae small, oval, dentigerous processes of vomers small, oblique, V-shaped, posterior and medial to choanae, each bearing 2���5 teeth; (6) absence of vocal slits in males, but presence of a shallow vocal sac and of two white non-spinous nuptial pads; (7) Finger I shorter than II; (8) fingers with broad lateral fringes; (9) ulnar tubercles low, inconspicuous, not forming a distinct line in preservative, but more conspicuous in life, sometimes forming a row; (10) tarsal tubercles low, inconspicuous, not forming a distinct line, calcar tubercles present, not pronounced; (11) inner metatarsal tubercle oval, two to four times the size of the round outer metatarsal tubercle; (12) toes with broad lateral fringes, webbing basal between Toes IV-V, Toe V longer than III, usually surpassing the proximal edge of the distal tubercle on Toe IV when toes are adpressed; (13) in life dorsal colouration highly variable, ranging from light golden brown, medium or dark brown to dark brown with two broad light brown longitudinal stripes mottled with dark brown and finely edged by a cream line; a W-shaped marking may be present on scapula; ill-defined light brown oblique bands are usually present on flanks with similarly coloured illdefined cross-bands on legs; broad light greyish brown bands may be present on flanks, dark brown dorsolateral bands may be present as well; a faint dark interorbital bar is usually visible; the snout is sometimes paler than body with some dark markings; a dark brown to black postocular stripe across upper and posterior parts of tympanum is present in most specimens; the groin is usually black or brown, sometimes with some small golden spots; a conspicuous reddish orange flashmark on groin may be visible; arms and legs often have a reddish or orangish tint. In preservative dorsal colouration brown to dark brown, the same patterns as described in life faded, but are still visible; (14) males 19.6���22.7 mm SVL, females 24.0���29.0 mm. Compared to the 21 Pristimantis species distributed in the Pantepui region, P. aureoventris is immediately distinguished from P. stegolepis (Schl��ter & R��dder, 2007), P. v il ar s i (Melin, 1941), and P. zeuctotylus (Lynch & Hoogmoed, 1977) in having F I II in P. stegolepis, P. vilarsi and P. zeuctotylus); from P. auricarens, P. jester Means and Savage, 2007, P. marahuaka, P. m u c h i m u k, and P. yaviensis in having a distinct tympanum (indistinct in P. auricarens, P. j e s t e r, P. marahuaka, P. m u c h i m u k, and P. yaviensis); from P. a v i u s (Myers & Donnelly, 1997), P. cantitans, P. inguinalis (Parker, 1940), P. marmoratus (Boulenger, 1900), P. memorans (Myers & Donnelly, 1997), P. pruinatus, P. pulvinatus (Rivero, 1968), and P. sarisarinama Barrio-Amor��s and Brewer-Car��as, 2008 in lacking vocal slits in male (vocal slits present in male of P. a v iu s, P. cantitans, P. inguinalis, P. marmoratus, P. memorans, P. pruinatus, P. pulvinatus, and P. sarisarinama); from P. dendrobatoides Means and Savage, 2007 in having lateral fringes on fingers and toes (absent in P. dendrobatoides), in having a shagreened to slightly granular dorsal skin (covered with large granular tubercles in P. dendrobatoides), and by the presence of nuptial pads in male (absent in P. dendrobatoides); from P. guaiquinimensis (Schl��ter & R��dder, 2007) and P. tepuiensis (Schl��ter & R��dder, 2007) by its much smaller size in adult male (male SVL max 22.7 mm [n= 8] in P. a u re o v e n t r i s vs. 33.6 mm [n= 2] in P. guaiquinimensis, and 34.7 mm [n= 1] in P. tepuiensis), by the presence of nuptial pads in male (absent in P. guaiquinimensis and P. tepuiensis), by the presence of vomerine teeth (absent in P. guaiquinimensis and P. tepuiensis), and in having lateral fringes on fingers and toes (absent in P. guaiquinimensis and P. tepuiensis); from P. saltissimus Means and Savage, 2007 in having lateral fringes on fingers and toes (absent in P. saltissimus), and by the presence of nuptial pads in male (absent in P. saltissimus); from P. yuruaniensis by its smaller size in adult female (female SVL max 29.0 mm [n= 3] in P. a u re o v e n t r i s vs. 32.2 mm [n= 5] in P. yuruaniensis), by having the ventral part golden yellow in life, usually with reddish brown to dark brown mottling, opaque with internal organs not, or barely visible through the skin (ventral part whitish, orangish or brown speckled with minute brownish dots in life, translucent, internal organs well visible through the skin in P. yuruaniensis), by having a higher degree of pattern polymorphism, and by having a less tuberculate skin. Description of the holotype. An adult male 20.4 mm SVL (Fig. 2) in very good condition of preservation, except a few small scars on back and upper arms. Head slightly longer than wide (HW 90.4 % of HL), wider than body, HW 39.9 % of SVL, cranial crests absent. Snout rounded in dorsal view, slightly sloping in profile; canthus rostralis nearly straight, round, loreal region slightly concave, sloping outward to lip; nares slightly protuberant, directed posterolaterally. Upper eyelid width narrower than IO; upper eyelid slightly granular with two barely discernible enlarged tubercles on each eyelid (better visible in life). Tympanum distinct, but not conspicuous, slightly obscured posteriorly, vertically oval, TYM 29 % of EL, separated from eye by about half its length; supratympanic fold originating at corner of eye, failing to reach shoulder, slightly arched; small postrictal tubercles evident. Choanae small, oval, slightly kidney-shaped, not concealed by palatal shelf of maxillary arch; dentigerous processes of vomers slightly smaller than choanae, oblique, V-shaped, posterior and medial to choanae, each bearing 3 to 4 teeth. Tongue much longer than wide, rounded posteriorly, posterior 2 / 3 free. Vocal slits absent, shallow subgular vocal sac present, seemingly not very distensible. Dorsal skin shagreened, almost smooth on head; middorsal raphe faint; no scapular folds or ridges visible; dorsal surfaces of limbs shagreened; flanks granular. Throat smooth; venter areolate (coarsely granular); posteroventral thigh and cloacal region areolate; weak discoidal fold anterior to groin; ulnar tubercles low, inconspicuous, not forming a distinct line in preservative, but more visible in life with three of them more prominent, forming a row. Finger I 85 % of II; relative length of adpressed fingers III> IV> II> I; adpressed Finger I failing to reach proximal edge of subdigital pad of Finger II; broad pre- and postaxial lateral fringes on fingers; presence of two whitish non-spinous nuptial pads, one adjacent to thenar tubercle and having about the same size, the other on the posterodorsal side of the thumb, having about the same size as the subarticular tubercle on Finger I. Finger discs expanded, with wide digital pads, broader than long, slightly rounded; distal median edge rounded. Palmar tubercle large, deeply bifid, V-shaped; thenar tubercle large, ovoid; subarticular tubercles large, round; supernumerary tubercles present, relatively large and protuberant (Fig. 3). Hind limbs relatively long; heels overlap when held at right angles to sagittal plane; TIL 55.7 % SVL; FL 45.8 % SVL. Relative length of adpressed toes IV> V> III> II> I; tip of Toe V extends to distal edge of ultimate subarticular tubercle of Toe IV, tip of III to the distal edge of the penultimate subarticular tubercle of IV. Toe discs as wide or narrower than finger discs, WFD/WTD = 1.11, with wide digital pads. Toes with broad pre- and postaxial lateral fringe, except on Toe I where fringes are barely visible; webbing basal between Toes IV-V. Inner metatarsal tubercle elongate, oval, about twice the size of the round outer metatarsal tubercle; supernumerary plantar tubercles low and round; subarticular tubercles large and protuberant. Calcar tubercles indistinct. Tarsal tubercles low, inconspicuous, not forming a distinct line; inner tarsal fold not visible (Fig. 3). Measurements of the Holotype (mm). SVL 20.4; HL 9.0; HW 8.2; SL 3.5; EN 2.3; IN 1.8; EL 3.2; IO 2.5, TYM 0.9; HAND I 3.6; HAND II 4.2; HAND III 5.8; HAND IV 4.7; WFD 1.1; FL 9.4; WTD 1.0; TIL 11.4. Colour of the Holotype in life. Dorsal ground colour dark brown with two broad light brown longitudinal stripes mottled with dark brown and finely edged by a cream line, extending from tip of snout to vent. Flanks dark brown with ill-defined light brown oblique bands; limbs adorned with similarly coloured ill-defined cross-bands. Arms and legs have a reddish tint. Broad dark brown postocular stripe across upper and posterior parts of tympanum, not contacting eye and failing to reach shoulder; broad dark brown canthal stripe; ill-defined dark brown labial bars from eye to upper lip. Posterior and anterior surfaces of thighs reddish brown, slightly mottled with golden. Throat, chest, and upper belly golden yellow with reddish brown mottling, lower belly and under surfaces of limbs reddish brown; internal organs not visible through the skin. Palms and soles dark brown. Iris greenish bronze, with fine black venation and a median horizontal reddish streak (Fig. 2). Colour of the Holotype in preservative. After 14 months in preservative, patterns and colours faded, but remain roughly the same. The golden ventral colour and the light brown stripes and bands became greyish. Arms and legs still have a slightly reddish tint (Fig. 2). Variation. See Table 1 for measurements, and Figs. 4���6 for intrapopulational variation. The most striking variation occurs in colour and pattern, which are very variable among living individuals (Fig. 4). The holotype is the only specimen having a striped dorsal pattern. Among the other specimens available, the dorsal colouration in life ranges from light golden brown (e.g. IRSNB 15823, adult male) to dark brown (e.g. IRSNB 15827, adult male), sometimes with a W-shaped marking on scapula (e.g. IRSNB 15820 and IRSNB 15643, adult males, and IRSNB 4154, subadult female). One male (IRSNB 15825) has a dark brown dorsum with broad light greyish brown bands on flanks; two individuals (IRSNB 4153, adult female, and IRSNB 15826, adult male) have a brownish dorsum with ill-defined dark brown dorsolateral bands. The ill-defined light brown oblique bands on flanks and the similarly coloured ill-defined cross-bands on legs may be absent (e.g. IRSNB 15820, adult male). A faint dark interorbital bar is usually visible (absent in the holotype), this bar being conspicuous in IRSNB 4154 (subadult female). In three individuals (IRSNB 4154, subadult female, IRSNB 15824 adult male, IRSNB 15825, adult male) the snout is paler than body (tan) with some dark markings (a black inverted triangle in IRSNB 4154). A dark brown to black postocular stripe across upper and posterior parts of tympanum is present in all specimens from Wei Assipu Tepui, that stripe is poorly detectable in the Roraima specimen. Groin is usually black or brown, sometimes with some small golden spots; one male (IRSNB 15827) had a conspicuous reddish orange flashmark on its groin in life (Fig. 7), which became white after preservation. Arms and legs usually have a reddish or orangish tint; IRSNB 15824 (adult male) had the posterior and anterior surfaces of thighs, the tibio-tarsal articulation, and part of the feet including Toes III���IV bright red; IRSNB 15825 (adult male) had the tibio-tarsal articulation distinctly paler (light grey) than the legs (dark brown); IRSNB 4154 (subadult female) had the anterior surface of thighs, the posterior surface of shanks, and part of the feet including Toes III���IV orange, a similar pattern is visible in IRSNB 15826 (adult male), but much less vivid. In other specimens, the posterior and anterior surfaces of thighs are reddish brown to dark brown, slightly mottled with golden. Under surfaces of limbs are reddish brown to dark brown, often heavily spotted with golden. Mottling on venter varies from reddish brown to dark brown, almost black, and is variably extensive; two individuals (IRSNB 15823, adult male from Wei Assipu Tepui, and IRSNB 15643, adult male from Mount Roraima, see Figs. 4, 6) lack mottling on venter. Due to the golden colouration of the venter, internal organs are little (IRSNB 15822, adult female) or not (holotype and other specimens) visible through the ventral skin in life (Figs. 2, 4). Iris colouration is variable, some individuals lack the median horizontal reddish streak (e.g. IRSNB 15822, adult female); IRSNB 15820 (adult male) has a blue-green iris (Fig. 4). In preservative all specimens became darker and variation in colour and pattern is less visible, but is still discernible (Fig. 5). Dorsal skin texture is slightly variable, most individuals having the dorsal skin shagreened to slightly granular. One specimen however (IRSNB 15827, adult male) has small conspicuous conical tubercles on anterior dorsum and legs, as is the case in the tiny (7.1 mm SVL) referred juvenile (IRSNB 15637). Few small conical tubercles are also visible on legs of most of the specimens. Small scapular tubercles were visible in seven individuals of both sexes in life (IRSNB 15822 ��� 24, IRSNB 15827, IRSNB 15643), but these tubercles are barely seen on the preserved animals and are absent in the holotype. Eyelid tubercles can be conspicuous (e.g. in IRSNB 15824, adult male) or barely visible (e.g. in IRSNB 15820, adult male). Number of teeth on dentigerous processes of vomers varies from two to five; they are absent in the very small juvenile (IRSNB 15637). A weak discoidal fold anterior to groin is visible only in the holotype and in two other specimens (IRSNB 15826, adult male, IRSNB 4153, adult female). The tip of Toe V fails to reach the distal edge of the ultimate subarticular tubercle of Toe IV on both sides in one female (IRSNB 4153), and on one side in four individuals of both sexes (IRSNB 15822 ��� 24, IRSNB 15643). No obvious sexual dimorphism is noted, except in size (females are larger than males). Other differences among available specimens are considered minor. Advertisement call. The following description is based on a sample of 21 advertisement calls from the male holotype (IRSNB 4152), recorded on the summit of Wei Assipu Tepui (05�� 13 ��� 05���N, 060�� 42 ��� 15 ���W, 2210 m elevation), on 17 November 2009 at 20 h00, air temperature 15.0��C. The male was calling concealed between the basal leaves of an Orectanthe sceptrum (Xyridaceae) growing ca. 150 cm above the ground on the wall of a 300 -cm deep small crevice. Temporal structure. The advertisement call of Pristimantis aureoventris consists of a soft, single, unpulsed note (perceived by the human ear as a small drop falling in the water). Calls are apparently emitted in bouts of about 20 calls (variation of the number of calls in each bout is unknown). The call rate was 18 calls/min based on a 1 -min period. The mean call duration is 0.021 �� 0.002 and varies from 0.017 to 0.023 s. The inter-call interval is rather uniform and has a mean of 3.097 �� 0.275 and a range of 2.730��� 3.610 s (Fig. 8 A). Spectral structure. Three harmonics are apparently developed, with the fundamental frequency dominating (mean: 2295, range: 2180���2430 Hz) (Fig. 8 A). The third harmonic contains slightly more sound energy than the second. The three harmonics show a feeble ���U-shaped��� frequency modulation (Fig. 8 A). Comparisons. Unfortunately, very few detailed descriptions of Pristimantis advertisement calls are available from the highlands of the Pantepui region. Myers and Donnelly (1996) provided analyses of unvouchered calls of different individuals of two highland Pristimantis species (tentatively assigned to P. cantitans and P. pruinatus) from Cerro Yav��, a massif located ca. 680 km W airline from Wei Assipu Tepui, and R��dde, Published as part of Kok, Philippe J. R., Means, Bruce & Bossuyt, Franky, 2011, A new highland species of Pristimantis Jim��nez de la Espada, 1871 (Anura: Strabomantidae) from the Pantepui region, northern South America, pp. 1-19 in Zootaxa 2934 on pages 4-17, DOI: 10.5281/zenodo.278071, {"references":["Hedges, S. B., Duellman, W. E. & Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182.","Schluter, A. & Rodder, D. (2007 \" 2006 \") Three new frogs of the genus Eleutherodactylus (Amphibia, Leptodactylidae) from Guaiquinima Table Mountain, Bolivar, Venezuela. Herpetotropicos, 3, 88 - 99.","Melin, D. (1941) Contributions to the knowledge of the Amphibia of South America. Goteborgs Kungliga Vetenskaps och Vitter-Hets Samhalles Handlingar, Ser. B, 1, 1 - 71.","Lynch, J. D. & Hoogmoed, M. S. H. (1977) Two new species of Eleutherodactylus (Amphibia, Leptodactylidae) from northeastern South America. Proceedings of the Biological Society of Washington, 90, 424 - 439.","Means, D. B. & Savage, J. M. (2007) Three new malodorous rainfrogs of the genus Pristimantis (Anura: Brachycephalidae) from the Wokomung Massif in west-central Guyana, South America. Zootaxa, 1658, 39 - 55.","Myers, C. W. & Donnelly, M. A. (1997) A tepui herpetofauna on a granitic mountain (Tamacuari) in the borderland between Venezuela and Brazil: Report from the Phipps Tapirapeco Expedition. American Museum Novitates, 3213, 1 - 71.","Parker, H. W. (1940) Undescribed anatomical structures and new species of reptiles and amphibians. Annals and Magazine of Natural History, Series 11 (5), 257 - 274.","Boulenger, G. A. (1900) Batrachians. Report on a collection made by Messrs. F. V. McConnell and J. J. Quelch at Mount Roraima in British Guiana. Transactions of the Linnean Society of London, Zoology, Series 2, 8, 55 - 56.","Rivero, J. A. (1968) A new species of Eleutherodactylus (Amphibia, Salientia) from the Guayana Region, Edo. Bolivar, Venezuela. Breviora, 306, 1 - 10.","Barrio-Amoros, C. L. & Brewer-Carias, C. (2008) Herpetological results of the 2002 expedition to Sarisarinama, a tepui in Venezuelan Guayana, with the description of five new species. Zootaxa, 1942, 1 - 68.","Myers, C. W. & Donnelly, M. A. (1996) A new herpetofauna from Cerro Yavi, Venezuela: first results of the Robert G. Goelet American Museum-Terramar Expedition to the northwestern tepuis. American Museum Novitates, 3172, 1 - 56.","Rodder, D. & Jungfer, K-H. (2008) A new Pristimantis (Anura, Strabomantidae) from Yuruani-tepui, Venezuela. Zootaxa, 1814, 58 - 68.","Duellman, W. E. & Trueb, L. (1986) Biology of amphibians. McGraw-Hill Book Company, New York, 670 pp.","Carreno, R., Nolla, J. & Astort, J. (2002) Cavidades del Wei-Assipu-Tepui, Macizo del Roraima, Brasil. Boletin de la Sociedad Venezolana de Espeleologia, 36, 36 - 45.","Huber, O. (1995 a) Geographical and physical features. In: Steyermark, J. A., Berry, P. E. & Holst, B. K. (Eds.), Flora of the Venezuelan Guayana, Vol. 1. Introduction, pp. 1 - 61. Missouri Botanical Garden, Timber Press, Portland, Oregon.","Huber, O. (1995 b) Vegetation. In: Steyermark, J. A., Berry, P. E. & Holst, B. K. (Eds.), Flora of the Venezuelan Guayana, Vol. 1. Introduction, pp. 97 - 160. Missouri Botanical Garden, Timber Press, Portland, Oregon.","MacInnes, H. (1974) Climb to the Lost World. Hodder and Stoughton, Ltd., London, 224 pp.","Villarreal, O., Senaris, C. & DoNascimento, C. (2002) Contribucion al conocimiento faunistico del Wei-Assipu-Tepui, Macizo del Roraima, con enfasis en la anurofauna y opiliofauna. Boletin de la Sociedad Venezolana de Espeleologia, 36, 46 - 50."]}

Gonatodes timidus sp. nov. (Figs. 1���4, Table 1) Holotype. IRSNB 2664 (field number PK 3555), an adult male collected by Philippe J. R. Kok, 22 March 2011 at 15 h00, base of Iwokrama Mountains, Potaro-Siparuni District, Guyana (04�� 19 ��� 52 ���N, 058�� 47 ��� 58 ���W, 209 m elevation). Paratypes (n= 9). two adult males (IRSNB 2665 ��� 66, field numbers PK 3538, PK 3557), four adult females (IRSNB 2667 ��� 70, field numbers PK 3539 ��� 41, PK 3556), two juveniles (IRSNB 2671 ��� 72, field numbers PK 3522 ��� 23) with same data as holotype, collected between 19 to 23 March 2011, and one juvenile (IRSNB 2673, field number PK 3553) collected by M. Wilkinson, 21 March 2011 in early afternoon, base of Iwokrama Mountains, opposite side of river of Tortoise camp (04�� 19 ��� 48 ���N, 058�� 47 ��� 54 ���W, 225 m elevation). Etymology. The specific name is considered a noun in apposition and is derived from the Latin adjective timidus meaning ���shy��� or ���fearful���. It refers to the tendency of individuals of the new species to avoid to be seen by hastily escaping between rocks, making them very difficult to collect. Definition and diagnosis. The new species differs from all known congeners by the following unique combination of characters: (1) moderate body size, with adult males ranging from 46.0��� 46.9 mm SVL, and adult females from 44.1���51.6 mm SVL; (2) absence of a clearly differentiated elongate supraciliary spine; (3) absence of clusters of distinctly enlarged conical scales on sides; (4) 89���97 scales around midbody; (5) 46���50 ventral scales counted in a longitudinal row; (6) escutcheon scales on posterior belly and ventral surfaces of thighs in males evident; (7) three to four lateral rows of scales on distal part of fingers and toes; (8) medial subcaudal scales distinctly differentiated from adjacent scales on non-regenerated tail, the pattern of scales consisting in repetitive series of a single midventral scale in contact laterodistally with one scale per side followed by a single midventral in contact with two scales per side (1 ��� 1 ��� sensu ��vila-Pires 1995); (9) sexual dichromatism obvious, males with upper surface of head black with bluish white to vivid yellow irregular stripes and blotches, and underside of head orange, females lack any colourful head ornamentation, but usually have two thin, black-edged white crescent-shaped collars on the neck region, and the underside of head black with bluish white reticulation. Compared to the six Gonatodes species known to occur in the Guiana Shield, Gonatodes timidus is readily distinguished by its radically different male pattern and colouration. It further differs from G. alexandermendesi Cole and Kok, 2010, G. astralis Schargel, Rivas, Makowsky, Se��aris, Natera, Barros, Molina and Barrio-Amor��s, 2010, and G. superciliaris Barrio-Amor��s and Brewer-Carias, 2008 by the absence of a distinctly elongate supraciliary spine (present in G. alexandermendesi, G. astralis and G. superciliaris), from G. annularis Boulenger, 1887 by having the medial subcaudal scales distinctly differentiated from adjacent scales on non-regenerated tail (not distinctly differentiated in G. annularis) and a dark copper coloured iris in males (always blue in G. annularis males), from G. infernalis Rivas and Schargel, 2008 by its smaller size in female (maximum 51.6 mm in G. timidus vs. minimum 55.6 mm in G. infernalis), the presence of markings on head and body in females (absence of dorsal pattern in G. infernalis), and in having the throat black with bluish white irregular stripes and blotches in females (immaculate in G. infernalis), from G. h u m e r a l i s (Guichenot, 1855) by its larger size in females (minimum 44.1 mm SVL in G. t i m - idus vs. maximum 40.5 mm SVL in G. humeralis), the presence of 3���4 lateral rows of scales at each side on distal part of fingers and toes (2 in G. humeralis), and the absence of a black antehumeral spot (always present in G. humeralis). Although absent from the Guiana Shield region, Gonatodes hasemani Griffin, 1917 (a southwestern Amazonian species) was shown to be the sister species of G. annularis and to be part of the ��� Guiana Shield clade��� sensu Schargel et al. (2010); G. hasemani is immediately distinguished from G. t i m i d u s in having a distinctly elongate supraciliary spine (absent in G. timidus), and in having the medial subcaudal scales not differentiated from adjacent scales on non-regenerated tail (distinctly differentiated in G. timidus). Description of the Holotype. An adult male in very good condition (Figs. 1���4), with SVL of 46.9 mm. TL 52.5 mm, tail complete, but with distal part regenerated. Head 1.2 times longer than wide (HL: 11.4 mm; HW: 9.6 mm). Snout short (2.2 mm), 0.19 times HL, acutely rounded in dorsal view, rounded in profile, gently sloping towards top of head. Neck slightly narrower than head and body. Body nearly cylindrical, but wider than high; AXG 18.8 mm. Limbs well developed with moderately long digits, ToIV 6.5 mm, 0.74 times ShL (8.8 mm). Tail round in section, tapering towards tip, which is bluntly rounded. Tongue relatively wide, narrowing anteriorly, covered by small scale-like papillae that become much less defined posteriorly; a bump bearing larger papillae is present in the middle of the tongue; posterior lateral margins indented; tip rounded with a short median cleft. Teeth small, subequal in length, conical. Rostral large, visible from above, roughly heptagonal, with a small indentation on top margin from which a short cleft extends forward about halfway through rostral. Three postrostrals, lateral ones (supranasals) distinctly larger than median, which is partially contained in the indentation of the rostral scale, and slightly larger than adjacent posterior scales on snout. Rostrals and postrostrals with many minute tubercles. Nostril bordered by rostral, lateral postrostral, three postnasals, and first supralabial. Dorsal postnasal slightly larger than lower two on the left side, noticeably larger than lower two on the right side; dorsal postnasals noticeably larger than scales in the loreal region; lower two postnasals about same size as scales in the loreal region. Scales on top of snout roughly round and granular, juxtaposed, gradually becoming more oval-shaped and conical towards loreal region. Loreal scales number about 14 (both sides) in a straight line between postnasals and anterior margin of orbit. Scales decrease slightly in size from the posterorostrals towards posterior part of the head. Scales on supraorbital region similar (in size and shape) to and continuous with those on top of head. Supraciliary flap without a distinctly elongate supraciliary spine, but three (both sides) larger conical supraciliary scales projecting posterolaterally. Pupil round. Supralabials 6 (both sides), first largest, last smallest, followed by 10 / 10 much smaller scales along the lip to rictus of mouth, all granular and similar in size to scales on temporal region. Fifth supralabial below centre of eye. Scales on temporal region similar (in size and shape) to those on posterior top of head, but scales contained in white stripes and blotches on head slightly larger and sometimes more spiny (especially on snout) than adjacent scales. Ear opening deep, very small (0.3 mm), much smaller than eye horizontal length (2.5 mm), obliquely oval. Mental large with a round posterior margin. Postmentals three, distinctly larger than adjacent posterior scales. Scales on chin located directly behind postmentals small and polygonal, granular and tiny posteriorly, a few larger, polygonal, juxtaposed scales adjacent to infralabials. Infralabials 5 (right side)/ 4 (left side), decreasing in size posteriorly, first two very large and projecting onto ventral plane, suture between third and fourth below centre of eye. Scales on nape and sides of neck continuous with those on head, granular, slightly conical anteriorly, becoming more conical posteriorly. Scales on throat smooth, imbricate, with round posterior margin, transition between juxtaposed granular, conical gular scales and scales on throat somewhat abrupt. Dorsals subconical to conical, round to oval-shaped at base, slightly larger than scales on top of head, somewhat projecting posteriorly; dorsolaterally and on flanks slightly larger than dorsally, distinctly larger proximal to ventrolateral region. Transition between scales on flanks and ventrals somewhat abrupt, but not clearly demarcated. Ventral region with scales distinctly larger than dorsals, slightly smaller on chest than on belly, smooth, with round to slightly acuminate posterior margin; ventrals in oblique rows also forming rather regular longitudinal rows on belly, with 46 scales along the midventral line between an imaginary line between the anterior part of the arm insertions with the body to the vent (not including small granules lining the edge of the cloaca). Scales around midbody about 94, of which about 15 are ventrals. Escutcheon scales evident: posterior belly (between thighs) composed of a cluster of about 45 scales distinctly different from other ventrals, larger, unpigmented except slightly on their lateral margins (under high magnification all ventrals are covered by tiny flecks of dark pigments, except scales on chest, which are also unpigmented). The same unpigmented scales are present under the thighs where they are arranged in three transverse rows of about 9���11 scales each (Fig. 4). Scales on dorsum of tail slightly larger than on body, flat, smooth, imbricate, with posterior margin rather acuminate; transition area between the conical body scales and the caudal scales somewhat abrupt, but not clearly demarcated. Scales on underside of tail smooth, flat, imbricate, with round posterior margin, increasing in size towards midventral line; first six small subcaudals posterior to vent on midventral line increasing in size posteriorly, but not clearly differentiated from adjacent scales, followed by a midventral line of transversely enlarged scales clearly differentiated from adjacent scales, the pattern of these scales consisting in repetitive series of a single midventral scale in contact laterodistally with one scale per side followed by a single midventral in contact with two scales per side (1 ��� 1 ��� sequence sensu ��vila-Pires 1995), although the first enlarged subcaudal is divided; scales on that midventral line entirely cover the underside of the distal portion of the tail (about 18.7 mm long, which corresponds to a regenerated part). Scales on limbs conical and juxtaposed, except ventrally and on anterior surface of forearms and thighs where they are smooth, flat, with round to acuminate posterior margin. Lamellae under first (I) through fifth (V) finger (right/left side): I: 14 /- (first finger missing on left side), II: 15 / 12 (claw, and possibly distal part of second finger missing on left side), III: 18 / 19, IV: 21 / 21, V: 17 / 17. Lamellae under first (I) through fifth (V) toe (right/left side): I: 11 / 12, II: 16 / 17, III: 20 / 20, IV: 22 / 23, V: 20 / 20. Fingers and toes with three to four lateral rows of scales distally. Claws exposed, non-retractile, between two basal scales (one dorsal and one ventral). Colour of the Holotype in life. The dorsal surface of the head is black with bluish white irregular stripes and blotches, which are distributed as follows: one broken (left side) or incomplete (right side) stripe on each canthus, one small blotch located medially between the posterior corner of the eyes, one large oblique blotch located posterodorsally to eye on each side, and one small blotch above ear opening on each side; a very small yellowish white spot is present about 1 mm posterior to rictus on each side, three similar tiny spots are found on the side of the neck on each side, and another tiny yellowish white spot is found below the last infralabials on each side. The supralabials are iridescent black. Two bluish white broken, incomplete crescent-shaped collars are present on the neck region, which is black dorsally and orange ventrolaterally. The underside of the head, including the infralabials and the chin, is orange with some scales partially or completely dark pigmented resulting in a faint reticulated pattern; the imbricate scales on the throat and the chest are orange, as well as most of the anterior surface of arms; all other ventral parts, including palms, soles, legs and tail are grey, except escutcheon scales, which are paler. A faint greyish brown, moderately broad vertebral stripe extends from posterior neck to near tail insertion. This stripe is irregularly margined by blackish blotches (five on the right side, three on the left side) and a few inconspicuous greenish brown to light brown suffusions. The flanks are chestnut brown with a few black flecks and small dark blotches. A darker patch is located on the right anterior flank, just posterior to arm insertion, and might correspond to a regenerated patch of skin. The limbs are similar in colour to the flanks, but with more dark markings. The tail is similar in colour to the dorsum, but slightly darker and without any longitudinal stripe. The iris is dark copper with a thin golden ring around the pupil. The claws are greyish white. Colour of the Holotype in preservative. After one month in preservative, patterns are essentially the same as in life, but the orange colour became white and the bluish tint in the white blotches and stripes disappeared. The tongue is whitish, with its first one-third dark grey. Variation in paratypes. The paratypes consist in two adult males (SVL 46.0 and 46.7 mm), four adult females (SVL from 44.1 to 51.6 mm), and three unsexed juveniles (SVL from 29.1 to 32.9 mm). Table 1 provides morphometric measurements of the type series, and Figs. 1���2 show variation and sexual dichromatism. Supralabial counts vary from 6 (n= 7) to 7 (n= 2, one side only), and infralabials counts vary from 4 (n= 1, one side only) to 5 (n= 8). Loreal scales counted in straight line between postnasals and anterior border of orbit range from 12 to 15, 14 being the most common state (n= 5). Postrostrals range from 3 (n= 7) to 4 (n= 2). Postmentals range from 2 (n= 7) to 3 (n= 2). Scales around midbody range from 89 to 97, of which 16 to 19 are ventrals. There are 46 to 50 scales between an imaginary line between the anterior part of the arm insertions with the body to the vent. Clusters of escutcheon scales on lower belly range from about 45 to 50 scales; under the thighs these scales are arranged in three to four transverse rows of about 2 (most anterior row) to 13 scales each, these scales are absent in females (see Fig. 4). The pattern of subcaudals (sensu ��vila-Pires 1995) is consistent, with the first distinctly enlarged subcaudal divided only in the holotype and in one juvenile. Variation in the number of lamellae under first (I) through fifth (V) finger is as follows: I: 12���16, II: 16���19, III: 17���22, IV: 19���23, V: 16���20. Variation in the number of lamellae under first (I) through fifth (V) toe is as follows: I: 11���14, II: 16���18, III: 19���23, IV: 22���25, V: 19���23. All paratypes have a few larger conical supraciliary scales projecting posterolaterally, but a distinct elongate supraciliary spine as in Gonatodes alexandermendesi, G. astralis, G. hasemani and G. superciliaris is always absent. There is an evident sexual dichromatism (see Figs 1���2). In life, the two male paratypes are very similar to the holotype in colour and pattern, except that IRSNB 2665 has vivid yellow stripes and blotches on head (instead of bluish white), a less contrasted pattern on body, and a paler body colour. Stripes on canthus extend from postnasals to near the eye in both male paratypes (broken and incomplete in the holotype), and the two crescent-shaped collars are more complete. Sides of neck are less orangish in the male paratypes than in the holotype. In preservative, the male paratypes are almost identical to the holotype, except in the shape of blotches and stripes on the head. IRSNB 2666 has a slightly pinkish throat (instead of white in the other paratype and in the holotype). In life, the female paratypes are brownish dorsally and laterally, with some whitish markings on tail, flanks, and limbs. Like in males a faint greyish brown, moderately broad vertebral stripe extends from posterior neck to near tail insertion and is irregularly margined by inconspicuous blackish blotches (more conspicuous in IRSNB 2668 and IRSNB 2669, which are the two smallest adult females). That vertebral stripe is also much more conspicuous in IRSNB 2668 and IRSNB 2669, and probably faints with age. The four female paratypes lack any colourful head ornamentation, but have two (only one in IRSNB 2670) thin, black-edged white crescent-shaped collars on the neck region, and usually a faint oblique stripe/blotch located posterodorsally to eye on each side (inconspicuous in IRSNB 2667). The dorsal surface of the head is slightly lighter than the body (distinctly lighter in IRSNB 2670). The supralabials are iridescent light grey, sometimes whitish posteriorly. The underside of the head, including the throat and the upper chest is black with bluish white reticulation (see Fig. 2). The infralabials are dark grey, usually with a variably large white area posteriorly. The ventral face is light grey with a suffusion of light orangish yellow (see Fig. 2); the ventral surface of the tail is dark grey. In preservative the female paratypes are dorsally darker than in life, but patterns remain essentially the same as in life. The bluish white colour became white, and the venter became very light grey. The three unsexed juveniles have essentially the same pattern and colouration as the adult females, including on throat, but patterns are more conspicuous and usually more contrasted. Two juveniles, IRSNB 2671 and IRSNB 2673, have a less contrasted throat pattern compared to the other juvenile. Females and juveniles usually have lighter iris. Distribution and ecology. The new species is currently known only from the type locality, between 209���225 m elevation, in lowland primary rainforest at the base of the Iwokrama Mountains in the Iwokrama Forest Reserve, central Guyana (Fig. 5). Gonatodes timidus is diurnal, locally abundant, but difficult to collect because specimens are very shy and tend to quickly escape in cracks and holes between boulders when disturbed. Most adult specimens have been collected on, or between relatively large boulders located about 10���30 m from a stream (Fig. 6). One adult female (IRSNB 2669) was collected between cavities on a tree trunk, close to large boulders. Two juveniles (IRSNB 2671 ��� 72) have been collected among the leaf litter close to boulders, an additional juvenile (IRSNB 2673) was collected in a rotten fallen tree trunk. No juveniles have been observed on rocks. As reported in other, Published as part of Kok, Philippe J. R., 2011, A new species of the genus Gonatodes Fitzinger, 1843 (Reptilia: Sphaerodactylidae) from central Guyana, northern South America, pp. 1-12 in Zootaxa 3018 on pages 2-9, DOI: 10.5281/zenodo.205720, {"references":["Avila-Pires, T. S. C. (1995) Lizards of Brazilian Amazonia (Reptilia: Squamata). Zoologische Verhandelingen, 299, 1 - 706.","Schargel, W. E., Rivas, G. A., Makowsky, R., Senaris, J. C., Natera, M. A., Barros, T. R., Molina, C. R. & Barrio-Amoros, C. L. (2010) Phylogenetic systematics of the genus Gonatodes (Squamata: Sphaerodactylidae) in the Guayana region, with description of a new species from Venezuela. Systematics and Biodiversity, 8, 321 - 339.","Barrio-Amoros, C. L. & Brewer-Carias, C. (2008) Herpetological results of the 2002 expedition to Sarisarinama, a tepui in Venezuelan Guayana, with the description of five new species. Zootaxa, 1942, 1 - 68.","Boulenger, G. A. (1887) On a new geckoid lizard from British Guiana. Proceedings of the Zoological Society of London, 1887, 153 - 154.","Rivas, G. A. & Schargel, W. E. (2008) Gecko on the rocks: an enigmatic new species of Gonatodes (Sphaerodactylidae) from Inselbergs of the Venezuelan Guayana. Zootaxa, 1925, 39 - 50.","Guichenot, A. (1855) Reptiles. In: Castelnau F. (Ed), Animaux Nouveaux ou Rares Recueillis pendant l'Expedition dans les Parties Centrales de l'Amerique du Sud, de Rio de Janeiro a Lima, et de Lima au Para; Executee par Ordre du Gouvernement Francais pendant les Annees 1843 a 1847, sous la Direction du Comte Francis de Castelnau. Reptiles. Tome second. P. Bertrand, Paris, pp. 1 - 95.","Griffin, L. E. (1917) A list of the South American lizards of the Carnegie Museum, with descriptions of four new species. Annals of the Carnegie Museum, 11, 304 - 320.","Cole, C. J. & Kok, P. J. R. (2006) A new species of gekkonid lizard (Sphaerodactylinae: Gonatodes) from Guyana, South America. American Museum Novitates, 3524, 1 - 13.","Bauer, A. M., Russell, A. P. & Shadwick, R. E. (1989) Mechanical properties and morphological correlates of fragile skin in gekkonid lizards. The Journal of Experimental Biology, 145, 79 - 102."]}

Anadia mcdiarmidi sp. nov. Figs 1-7, Table 1 Anadia breweri (nomen nudum) ��� Gorzula 1992: 276, figs 168-171 Anadia species a ��� Gorzula & Se��aris 1999: 114-115, fig. 88 Anadia sp. A ��� McDiarmid & Donnelly 2005: 514 Anadia breweri (nomen nudum) ��� Brewer-Car��as & Audy 2010: 210, unnumbered figure Holotype IRSNB 2677 (field number PK 3565), 1 adult ♂ collected by Philippe J. R. Kok, 03-05-2011 at 11h39, summit of Abakap��-tepui, Bol��var State, Venezuela, 05��11'09''N 062��17'36''W, 2201 m elevation. Paratopotypes (n=3) 1 adult ♂ (IRSNB 2674, field number PK 3578) collected by Philippe J.R. Kok, 06-05-2011 at 9h40, summit of Abakap��-tepui, Bol��var State, Venezuela, 05��10'50''N 062��17'49''W, 2242 m elevation); 1 juvenile ♂ (IRSNB 2675, field number PK 3577) collected by Philippe J.R. Kok, 06-05-2011 at 9h10, summit of Abakap��-tepui, Bol��var State, Venezuela, 05��11'00''N 062��17'34''W, 2200 m elevation; and 1 other juvenile ♂ (IRSNB 2676, field number PK 3589) collected by Philippe J.R. Kok, 09-05-2011 at 9h00, summit of Abakap��-tepui, Bol��var State, Venezuela, 05��11'07''N 062��17'19''W, 2204 m elevation. Etymology The specific epithet is a noun in the genitive case, honouring Roy McDiarmid, North American herpetologist (Smithsonian Institution), for his contribution to the knowledge of the Pantepui herpetofauna. Generic allocation The new species is placed in the genus Anadia because it fits the 11 diagnostic traits for the genus proposed by Oftedal (1974): dorsal scales smooth, essentially homogeneous; no distinct rows of much smaller scales between lateral scales; ventral scales smooth, quadrangular; head scales smooth, without striation; single frontonasal separating nasals; paired prefrontals; several large pigmented palpebrals; gular scales smooth, essentially homogeneous, with no distinctly enlarged longitudinal rows; 7-10 large preanal scales [maximum 9 in Oftedal (1974)]; femoral pores conspicuous in adult males; and limbs well developed, pentadactyl with all toes clawed. Definition and diagnosis A fairly robust species currently not assigned to any species group. The new species is characterized by the following combination of characters: (1) size small, body robust (maximum known SVL 62.2 mm); (2) tail longer than SVL; (3) dorsal scales small, quadrangular; (4) middorsal scales 53-57; (5) suboculars large, unequal in size, with sometimes one scale slightly protruding downward between 4 th and 5 th supralabial, suboculars separated from palpebrals by 2-3 rows of mostly inconspicuous tiny scales, a few of them distinctly enlarged with angular downward protrusion between suboculars; (6) nasal entire, without sub-nostril groove; (7) 3-4 large pigmented palpebrals; (8) body uniform beige or greyish to bluish brown in life, devoid of any conspicuous pattern in males; (9) absence of any dark line on side of head in adult males; (10) venter immaculate golden grey in life; (11) femoral pores 9-10 on each side in males, unknown in females; (12) preanal pores absent; (13) hemipenis globose, weakly bilobed, bordered by numerous flounces (>20, including about eight medioproximal asulcate flounces) bearing comblike rows of minute weakly mineralized spinules, sulcus spermaticus divided in the crotch by a small fleshy bumplike structure, sulcus branches barely detectable. The new taxon is immediately distinguished from Anadia ocellata, A. vittata, A. rhombifera, and A. petersi [all members of the ocellata group of Oftedal (1974)] and A. bogotensis [single member of the bogotensis group of Oftedal (1974)] in having quadrangular dorsal scales (subhexagonal in species of the ocellata group, imbricate in A. bogotensis), and in lacking a subnostril groove or divided nasal (nasal divided or subnostril groove present in species of the ocellata group and in A. bogotensis); from A. altaserrania [a member of the bitaeniata group of Oftedal (1974) according to Harris & Ayala (1987)] in having prefrontals (lacking in A. altaserrania), in having more femoral pores in males (9-10 in A. mcdiarmidi vs. 3-4 in A. altaserrania), and in having a higher number of middorsal scales (53-57 in A. mcdiarmidi vs. 45-47 in A. altaserrania); from A. bitaeniata, A. brevifrontalis, A. hobarti, A. pamplonensis, and A. pulchella [all members of the bitaeniata group of Oftedal (1974) according to La Marca & Garc��a-P��rez (1990)] in having a higher number of middorsal scales (53-57 in A. mcdiarmidi vs. maximum 50 in species of the bitaeniata group); from A. blakei and A. marmorata [members of the marmorata group of Oftedal (1974)] in having a higher number of middorsal scales (more than 50 in A. mcdiarmidi vs. less than 35 in A. blakei and A. marmorata); from A. bumanguesa [a member of the bitaeniata group of Oftedal (1974) according to Rueda-Almonacid & Caicedo (2004), but a possible synonym of A. steyeri and thus a member of the steyeri group of Oftedal (1974) according to Rivas et al. (in press)] in having a lower number of scales around midbody (35-36 in A. mcdiarmidi vs. 41 in A. steyeri), and in having pigmented palpebrals (unpigmented in A. bumanguesa); from A. pariaensis (not assigned to any species group in the original description) in having a much lower number of middorsal scales (54- 57 in A. mcdiarmidi vs. 70-72 in A. pariaensis); from A. steyeri [single member of the steyeri group of Oftedal (1974)] in having a lower number of scales around midbody (35-36 in A. mcdiarmidi vs. 40-45 in A. steyeri); and from A. escalerae (not assigned to any species group in the original description), the geographically closest relative, in having a robust body (slender in A. escalerae), in having smooth dorsal scales (weakly keeled in A. escalerae), in lacking dorsolateral stripes (present in A. escalerae), in having a lower number of middorsal scales (54-57 in A. mcdiarmidi vs. 63 in A. escalerae), and in having a lower number of scales around midbody (35-36 in A. mcdiarmidi vs. 40 in A. escalerae). Description of the Holotype An adult male in good condition (Figs 1A, C, 2, 3, 4, 6), with 62.2 mm SVL and 110.0 mm TL (see also Table 1). A rather robust Anadia species with moderately long limbs (digits overlap when limbs are adpressed along body). Snout attenuate, flat in profile. HL 25% of SVL, 1.5 times longer than wide, 1.6 times wider than high; head conspicuously wider than neck. Neck long, 68% of HL, 35% of AXG. SAL 93% of AXG, 46% of SVL. Body wider than deep. Tail dorsoventrally flattened, about 1.8 times longer than SVL. Limbs pentadactyl with all digits clawed. Forelimb 27% of SVL, 52% of AXG; hind limb 36% of SVL, 73% of AXG. Scutellation on the dorsal surface of head is typical for the genus [see fig. 1 in Oftedal (1974)], except the condition of the parietals and interparietal (see below and Figs 2, 7). The frontal is damaged, the frontoparietals are slightly damaged. Head scales are smooth with a few minute scattered pits, those pits are especially concentrated on the upper and posterior margins of supralabials, on the lower and posterior margins of infralabials, and on the posterior margin of rostral. Rostral much wider than deep, laterally in contact with nasals and first supralabial, dorsally in contact with large frontonasal. Frontonasal with nearly straight anterior margin, posteriorly pointed. Paired prefrontals in contact, medial suture short. Frontal damaged, but obviously wider anteriorly and with blunt anterior point. Paired frontoparietals slightly damaged, with long medial suture, in contact with interparietal, parietals, and two posterior supraoculars. Three large supraoculars subequal in size. Interparietal divided into about two scales unequal in size. Parietals not extending as far as interparietal posteriorly, divided into two scales on the left side, into four scales on the right side. A series of six small to large occipitals (postparietals) circling the common posterior margin of parietals and interparietal. Nasal scale entire, with a small indentation situated anterodorsally to nostril on the left side. Nasal scale in contact with rostral, posteriorly in contact with prefrontal, and in broad contact with loreal. Loreal large, pentagonal, in broad contact with first superciliary (= presuperciliary) and frenocular, in point contact with preocular. One small preocular, three postoculars. First superciliary large, followed by four shorter superciliaries. No small azygous scale between superciliaries and supraoculars. Frenocular followed posteriorly by three suboculars. Third subocular slightly extending to lip between supralabials 4-5 on the left side only. Suboculars separated from palpebrals by 2-3 poorly defined rows of mostly inconspicuous tiny scales, two of them distinctly enlarged with angular downward protrusion between suboculars. Seven supralabials. Eight ciliaries along upper eyelid. Lower eyelid scales translucent, slightly pigmented, with four higherthan-wide palpebrals. Temporal scales subimbricate, smooth, with flat surfaces, slightly larger above and smaller below. Ear opening broad, ovoid, slightly inclined posterodorsally, edged with small, slightly pebblelike scales; tympanum recessed, pigmented. Underside of head with six infralabials on each side. A large mental followed by a large postmental in contact with first two infralabials. Two pairs of large genials in broad contact medially, in lateral contact with infralabials 3-4; two pair of postgenials, the first pair large, each scale in contact with last genial and infralabials 4-5, the second pair much smaller, each scale in contact with first postgenial and in point contact with fifth infralabial. Gulars arranged in transverse rows, becoming slightly larger posteriad, culminating in a well-defined collar row of eight subequal scales; no distinctly enlarged longitudinal rows. Side of neck between ear and collar pebbled with subequal irregularly shaped juxtaposed scales. Middorsal scales 54. Dorsal scales on neck subimbricate, irregularly shaped, with rounded surfaces. Dorsal body scales smooth, juxtaposed, mainly quadrangular, a few irregularly shaped, but most are longer than wide, in transverse rows only. Lateral scales smooth, similar to dorsals, but smaller. Ventral scales much wider than dorsals, smooth, juxtaposed, quadrangular, slightly longer than wide except for ca. four median longitudinal rows of square to wider than long scales on belly; in 14 longitudinal rows at midbody and 31 transverse rows between collar and vent. Eight marginal preanal scales anterior to vent, the middle two narrower than the adjacent lateral ones, most outer ones very small. Femoral pores well developed, in swollen pore scales mostly in linear contact; 9 pores on left thigh, 10 on right. Pores do not extend onto preanal area (Fig. 3). Caudal scales smooth, subimbricate, rectangular, in transverse rows around tail. Scales on dorsal surfaces of arm large, smooth, imbricate; ventral surfaces of arm with similar, but smaller and less imbricate scales. Hind limbs with large, smooth subimbricate to imbricate scales on anterior face of thighs and on lower legs; scales similar but smaller on ventral side of thigh. Dorsal and posterior sides of thigh with much smaller, irregularly shaped juxtaposed scales. Moderate-size, imbricate scales atop hands and feet. Supradigital scales single; upper and lower ungualsheath scales covering base of claws, leaving tips well exposed. Palms and soles with small slightly raised juxtaposed scales (Fig. 4). Two enlarged smooth thenar scales at base of pollex. Subdigital lamellae mostly single. Lamellae under first (I) through fifth (V) finger (right/left side): I: 7/6, II: 9/9, III: 11/11, IV: 13/14, V: 10/9. Lamellae under first (I) through fifth (V) toe (right/left side): I: 6/6, II: 10/10, III: 14/12, IV: 17/15, V: 15/6 (large part of Toe V missing on left side). Tongue lanceolate; proximal two-third unpigmented, distal one-third heavily pigmented, including fork. Upper surface behind fork covered with imbricate scalelike papillae that extend onto ventrolateral sides of tongue. Raised medioventral side of tongue with numerous (>10) of thin, oblique, anteriorly converging and pointed (chevronlike) infralingual plicae; anterior pair of plicae swollen, bluntly pointed and larger than those following. Very shallow medioventral groove. Anterior maxillary and dentary teeth conical, unicuspid, with no or very feeble recurvature, becoming larger posteriorly. The left hemipenis (Fig. 5) extends about four subcaudal rows when adpressed to the tail, and is about 7.0 mm in length and 4.0 mm across its widest point when fully everted (organ not maximally expanded because one of the lobes was slightly damaged during the eversion process). Hemipenial body globose, tapered near the base, weakly and symmetrically bilobed. Sulcus spermaticus broader near crotch, relatively shallow, curving about halfway around the base of the organ, then running medially towards the lobes. Sulcus flanked by a narrow nude area on each side, and divided in the crotch by a small fleshy bumplike structure, from where two narrow, ill-defined and barely detectable sulcus branches seem to extend laterally into the folded thickened 'ear-like' tissues of each lobe towards the apices of the lobes. Lobes are complexly folded, not flattened terminally, and sulcus branches seem to end in small depressions (from which residual seminal fluid was removed before hemipenis preparation). Very base of hemipenis nude, pigmented in life. Lateral and asulcate faces of the organ with a series of roughly equidistant oblique plicae or flounces with weakly calcified spinules. Twenty-three rows of flounces extending along the organ. Basal flounces shorter, almost in straight transverse lines, the three first rows almost continuous across the hemipenial base, except for narrow nude areas on each lateral face and for nude areas edging each side of the sulcus spermaticus; five subsequent rows continuous across hemipenial body (except for nude areas edging each side of the sulcus spermaticus), with the seventh and eight rows distinctly pointing medially towards the apex on the asulcate face (those eight flounces proximal to the median nude space of the asulcate face correspond to the 'medioproximal asulcate flounces' sensu Myers et al. 2009); fifteen remaining rows chevron-shaped, also circling the organ, but separated by a broad median nude space on the asulcate face and by the nude areas edging each side of the sulcus spermaticus. The plicae bear comblike rows of minute weakly mineralized spinules that project slightly from the edges of the plicae and usually increase in size towards the lateral faces and the basal part of the organ (staining of the spinules by Alizarin Red is well visible under the microscope although not observable in Fig. 5). Colour of the Holotype in life Dorsum greyish to bluish brown depending on light intensity, devoid of any conspicuous pattern, but with a few scattered black scales on flanks. Arms and legs, including digits, greyish brown with a few scattered black scales; rear of thighs with a few ill-defined dark markings. Ventral surfaces of head and body golden grey, ventral surfaces of legs and tail bluish grey, palms and soles lemon yellow speckled with brown and yellowish orange scales. Tympanum dark brown, iris pale yellow (Figs 1A, C, 4). Colour of the Holotype in preservative After approximately two months in preservative, the dorsal surfaces became dark brown (looking slightly peppered when specimen is in liquid), the ventral surfaces are metallic blue, except the anterior surface of head, which is light grey (Fig. 6). Palms and soles are grey. Tympanum is dark brown. It is worthwhile to note that the ventral skin, and some parts that are light grey (like the anterior underside of head), turn blue or bluer when pressure is applied to the skin. Variation in paratypes The paratypes consist in 1 adult ♂ (SVL 60.4 mm), and 2 juvenile ♂ (SVL from 38.2 to 42.6 mm). Table 1 provides measurements of the type series, and Fig. 1B shows one of the juvenile paratypes in life (IRSNB 2675). One of the main differences among specimens of the type series is the occurrence of a presupraocular scale in the three paratypes (absent in the holotype); that presupraocular scale is divided on both sides in IRSNB 2675. IRSNB 2676 has four suboculars on both sides (vs. three in the holotype and other paratypes). Supralabial counts are consistent, but infralabial counts vary from 5 (n=1) to 7 (n=1, one side only). Pits on head scales are inconspicuous in the two juveniles (vs. well visible in the adult males). Suboculars are always separated from palpebrals by 2-3 poorly defined rows of inconspicuous tiny scales. In IRSNB 2674 three of these scales (vs. two in the holotype) are distinctly enlarged (larger than in holotype) with angular downward protrusion between suboculars. In both juvenile paratypes, only one of these scales is only slightly enlarged and slightly protrudes between suboculars. The condition of the parietals and interparietal is highly variable among specimens and none of the specimens examined has complete interparietal and parietals, which are always divided in several smaller scales (see Fig. 7 for comparison). Number of middorsal scales varies from 53 to 57. Ventrals are in 11 to 14 longitudinal rows and in 29 to 30 transverse rows. The number of femoral pores varies from 8 to 10. Preanals vary from 7 to 10. The fully everted, maximally expanded, right hemipenis of the adult ♂ paratype (IRSNB 2674) is identical to the holotype's organ, except that maximal expansion makes the sulcus spermaticus bifurcation and the sulcus branches even more difficult to appreciate. Maximal expansion does not highlight hemipenial features that would have been overlooked in the not maximally expanded hemipenis of the holotype. The hemipenis is 6.6 mm in length and 4.0 mm across its widest point when everted; there are twentyfour rows of flounces extending along the organ, ca. eight of which are medioproximal asulcate flounces. Colour in life ranges from beige to dark greyish, sometimes bluish, brown dorsally, depending on light intensity. Rear of thighs usually has a few ill-defined dark markings. Some blue markings may be visible around ear, or at the base of the tail, or on ventral parts. Ventral parts range from golden grey to pale yellow. One of the juvenile paratypes (IRSNB 2675, see Fig. 1B) has two poorly defined dark brown stripes between eye and ear. In preservative all specimens became dark brown dorsally, looking slightly peppered when in liquid. Ventral parts are always metallic blue (Fig. 6). Tympanum in preserved specimens varies from dark brown to blue or golden blue. Distribution and ecology The new species is known only from the Chimant�� Massif in Venezuela (Fig. 8) where it has been reported from Abakap��-tepui (type locality), Amur��-tepui, and Murei-tepui (sometimes named Eruod, Published as part of Kok, Philippe J. R. & Rivas, Gilson A., 2011, A new species of Anadia (Reptilia, Squamata) from the Venezuelan ' Lost World', northern South America, pp. 1-18 in European Journal of Taxonomy 3 on pages 3-15, DOI: 10.5852/ejt.2011.3, http://zenodo.org/record/3711435, {"references":["Gorzula S. 1992. La herpetofauna del macizo del Chimanta. In: Huber O (ed.), El Macizo del Chimanta: 267 - 280, 304 - 310. Oscar Todtmann Editores, Caracas.","Gorzula S. & Senaris J. C. 1999 \" 1998 \". Contribution to the herpetofauna of the Venezuelan Guayana: I. A data base. Scientia Guaianae 8: 1 - 269.","McDiarmid R. W. & Donnelly M. A. 2005. The herpetofauna of the Guayana Highlands: amphibians and reptiles of the Lost World. In: Donnelly M. A., Crother B. I., Guyer C., Wake M. H. & White M. E. (eds.) Ecology and Evolution in the Tropics: A Herpetological Perspective: 461 - 560. University of Chicago Press, Chicago, Illinois.","Brewer-Carias C. & Audy M. 2010. Entranas del Mundo Perdido. Carlos Capriles de Altolitho C. A., Caracas.","Oftedal O. T. 1974. A revision of the genus Anadia (Sauria, Teiidae). Arquivos de Zoologia, Sao Paulo 25 (4): 203 - 265.","Harris D. M. & Ayala S. C. 1987. A new Anadia (Sauria: Teiidae) from Colombia and restoration of Anadia pamplonensis Dunn to species status. Herpetologica 43 (2): 182 - 190.","La Marca E. & Garcia-Perez J. E. 1990. A new species of high Andean Anadia (Sauria: Teiidae) from Paramo El Riecito, Estado Trujillo, Venezuela. Herpetologica 46 (3): 275 - 282.","Rueda-Almonacid J. V. & Caicedo J. R. 2004. Una nueva especie de Anadia (Reptilia: Sauria: Gymnophthalmidae) para el noroccidente de la cordillera oriental de Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 28 (107): 281 - 284.","Myers C. W., Rivas Fuenmayor G. & Jadin R. C. 2009. New species of lizards from Auyantepui and La Escalera in the Venezuelan Guayana, with notes on \" microteiid \" hemipenes (Squamata: Gymnophthalmidae). American Museum Novitates 3660: 1 - 31. http: // dx. doi. org / 10.1206 / 657.1"]}

Microcaecilia iyob sp. nov. (Figs. 1; Table 1) Gymnopis unicolor (Dum��ril, 1864) in part; Dunn (1942: 471���472) Microcaecilia unicolor (Dum��ril, 1864) in part; Taylor (1968: 542���546, fig. 297) Holotype. FMNH 35117, a mature male from ���Oko River at Cuyuni River��� Guyana; collected by Neal A. Weber in June 1936. Diagnosis. A Microcaecilia that differs from M. taylori Nussbaum & Hoogmoed, 1979 and M. grandis Wilkinson et al., 2009 in having fewer PAs (M. rabei (Roze & Solano, 1963) in having fewer PAs undivided by SAGs ( 8); and from M. albiceps (Boulenger, 1882), M. supernumeraria Taylor, 1970, and M. unicolor (Dum��ril, 1864) in having more numerous premaxillary-maxillary teeth (> 20) in long series that extend posteriorly beyond the posterior margin of the choanae, and bicuspid vomeropalatine teeth. Description of the holotype. Some morphometric and meristic data are given in Table 1. In poor condition in places, incisions extending from CMs across C 1 and C 2, almost broken into two parts at the 12 th PA, less substantial breaks at about 25 th and 60 th PA, small (5mm) midventral incisions just anterior and posterior to second of these breaks, a 12 mm incision beginning 45 mm in front of terminus, some open scale pockets and patches of abraded skin, rusty hole on 36 th PA another 20 mm anterior to vent. An arthropodan exoskeletal element embedded in the skin of the lower jaw on the left side. Mature male with multiple large testis lobes. Body dorsoventrally flattened throughout, relatively uniform, narrowing slightly onto collars anteriorly and more substantially behind the level of the vent; L/W = c. 32. In dorsal view, sides of head straight, converging steadily to level of nares, snout tip between nares rather blunt. In lateral view, top of head weakly convex; margins of upper jaw (lip) concave especially anterior to the level of the TAs; ridge bearing vomeropalatine teeth just visible close to CM; lower jaw half the height of upper jaw at CM. In ventral view, mouth moderately recessed, margins of lower jaw and upper lips not much more blunt than tip of snout. Eyes not visible. TAs very slightly elevated, the elevations visible dorsally and ventrally, lie just above imaginary lines between nares and CMs, distinctly closer to CMs than to nares. Nares small, dorsolateral, circular depressions with anteroventral ovate aperture, somewhat closer to level of AM than to ST, much closer to top than to bottom of snout in lateral view, not visible from below. Teeth pointed, gently recurved, lacking serrations or flanges; elements of outer series much smaller posteriorly, monocuspid, dentaries largest, premaxillary-maxillary teeth large; vomeropalatines much smaller, more uniform in size, bicuspid, about as many in front of the anterior margins of the choanae as behind, vomerine series forming gentle arch anteromedially; distance between vomeropalatine and premaxillary-maxillary series anteriorly almost equal to projection of snout; series converging posteriorly to about half that distance, upper series extending posteriorly well (5���7 elements or more on each side) past choanae, approaching CMs, palatine series extending slightly further posteriorly. Palate moderately arched. Choanal apertures subcircular, separated from each other by twice their individual widths, anterior margins approximately level with tentacles; valves not visible. Tongue simple (no surface architecture) with broad attachment anteriorly. Nuchal region barely more massive than adjacent body. C 1 shorter than C 2, well-marked, NG 1 and NG 2 completely encircling body, bending slightly anteromedially on dorsum, NG 3 incomplete ventrally; substantial TG on dorsum of C 2, just visible laterally. Behind collars, 97 PAs and a small unsegmented terminal cap. AGs slightly raised posteriorly, complete or nearly so dorsally, mostly separated mid-ventrally, 31 AGs complete ventrally anterior to vent; no AGs posterior to vent on venter. First SAG present mid-dorsally on 52 nd PA, all PAs behind 55 th divided by mid-dorsally complete SAGs, 12 of which are complete ventrally (on PAs 84���95) anterior to vent region. Vent region interrupts last four to five AGs. Scattered small scales in a single incomplete row present dorsally as far anterior as 25 th PA, near terminus scale pockets as deep as the width of a PA, with multiple (8���9) rows of either slightly ovate or rhomboidal (largest 1.0 x 0.8 mm) scales. No subdermal connective tissue scales. Body terminus acuminate with no indication of terminal keel. In lateral view, ventral surface concave/indented at level of vent. Vent within a fairly discrete, approximately circular, depressed cloacal disc, with slightly transverse opening, its lips formed by about 13 somewhat irregular, short denticulations; a pair of cloacal papillae on anterior denticulations. Gray-brown, more brown anteriorly and where superficial epidermis missing, darker, more plumbous grey posteriorly, perhaps slightly paler on lateral flanks especially posteriorly where each annulus has a paler anterior and darker posterior half. Head paler than body especially ventrally, except for narrow darker lines approximately level with medial aspect of mandibles, central pale colour extending onto C 1. Collar grooves and AGs edged in white, especially laterally. TAs and nares lie within small pale spots. Cloacal disc, including small ventral area of terminal cap, pale. Etymology. The species is named to commemorate the International Year of Biodiversity (2010) to serve as a reminder that much biodiversity is undescribed and much sampled biodiversity is poorly understood taxonomically. The epithet iyob is designated a genderless noun in apposition. Suggested English common name. Commemorative microcaecilia. Edward Harrison Taylor was by far the most important figure in the history of caecilian taxonomy. In his then comprehensive taxonomic study of caecilians, Taylor (1968) provided accounts for some 158 species, of which he was the authority for almost half (73). These and other prodigious feats of species description led to Taylor having a reputation as something of a ���splitter���, and a not insubstantial number of his species have been subsequently relegated to synonyms. Although Taylor occasionally explicitly held back from describing possible new species for variant specimens, this was the exception rather than the rule. In his account of Microcaecilia unicolor, Taylor (1968: 544) gave data for seven specimens from French Guiana, including the lectotype, and for a single specimen from Guyana that is now the holotype of M. iyob. These data reveal substantial differences in annulation and dentition between the M. unicolor from French Guiana and the Guyana specimen, that result in some very high ranges for these characters in Taylor���s diagnosis of M. unicolor. Of these differences Taylor (1968: 546) writes only that the Guyana specimen ���differs in having a primary count of about eight less than the lowest French Guiana specimen, and 18 lower than the highest count. The secondaries are reported to be more erratic, varying from 22 to 74? (fide Dr. E. R. Dunn), the number of complete secondaries��� [sic]. Thus, Taylor (1968) was following Dunn (1942) in including the Guyana specimen with the species unicolor, and was perhaps dissuaded from considering the distinctiveness of the Guyana specimen within his sample any further by Dunn���s report of a very wide range in secondary annuli. Uncharacteristically, Taylor (1968) did not comment on the clear differences in the numbers and disposition of teeth in the upper jaw that are evident in his data and that, by themselves, are strong evidence that the samples from French Guiana and from Guyana are not conspecific. Two years later, Taylor (1970: 313) highlighted that the Guyana specimen differed from the type in the number of premaxillary-maxillary teeth and cautioned ���Material from this area should be acquired and carefully examined, as I may have erred in so disposing of this specimen.��� Unfortunately, Taylor (1968) used the holotype of Microcaecilia iyob as the basis for his descriptive account of M. unicolor and, as Wilkinson et al. (2009) noted, this has contributed to taxonomic confusion regarding M. unicolor (e.g. Nussbaum & Hoogmoed 1979). There are some minor differences between the data reported by Taylor (1968) and those reported here on the holotype of Microcaecilia iyob (Table 1). Additionally, Taylor (1968) reported that the PAGs are mostly incomplete dorsally whereas we find them to be mostly complete or nearly so, and he also reported that the first and second nuchal grooves curve forward ventrally whereas we find them to be mostly straight (Fig. 21. A syndrome of dental characters seemingly divides Microcaecilia into two groups. Microcaecilia albiceps, the type species, and M. unicolor have short premaxillary-maxillary teeth series that do not extend posteriorly beyond the choanae, bicuspid vomeropalatine teeth and serrated dentary teeth. The enigmatic M. supernumeraria, known only from its holotype, appears to belong with these species. It shares the first two of these dental characters but the condition of the third is unknown. Differently, M. iyob resembles M. rabei, M. grandis and M. taylori in having long premaxillary-maxillary teeth series and all teeth are monocuspid and unserrated. The new species is readily distinguished from these other Microcaecilia by its pattern of annulation with relatively few PAs and many SAGs and consequently relatively small numbers of undivided PAs. Several museum specimens of Microcaecilia that we have seen, including the holotype of M. iyob, have pointed and recurved arthropodan exoskeletal remains embedded in the skin of the head. We suspect this is evidence of frequent and aggressive encounters with either ants or termites, and possibly the result of preying upon them. The new species is known from only a single specimen collected more than 70 years ago and, given that there have been no dedicated attempts to recollect or otherwise survey its status in the wild, it must be considered data deficient for conservation purposes. Targeted fieldwork is needed to shed light on the distribution and abundance of this very poorly known species, and to provide material for both morphological and molecular systematic study. Microcaecilia iyob is only the third new species of caecilian described from Guyana since Taylor (1968). That these descriptions have appeared within the last two years and have been based on both newly collected (Wake & Donnelly 2009, Gower et al. 2010) and older material (this work), suggests both that the caecilian fauna of Guyana remains very incompletely known, and that further species new to science probably remain to be discovered from this region., Published as part of Wilkinson, Mark & Kok, Philippe J. R., 2010, A new species of Microcaecilia (Amphibia: Gymnophiona: Caeciliidae) from Guyana, pp. 35-40 in Zootaxa 2719 on pages 36-39, DOI: 10.5281/zenodo.199833, {"references":["Dumeril, A. (1864) Catalogue methodique de la collection des batraciens du Museum d´Histoire Naturelle de Paris. Memoires de la Societe Imperiale des Sciences Naturelles de Cherbourg 1, 307 - 321.","Dunn, E. R. (1942) The American caecilians. Bulletin of the Museum of Comparative Zoology, Harvard 91, 439 - 540.","Taylor, E. H. (1968) The caecilians of the world: A taxonomic review. The University of Kansas Press, Lawrence, KS.","Nussbaum, R. A. & Hoogmoed, M. S. (1979) Surinam caecilians, with notes on Rhinatrema bivittatum and the description of a new species of Microcaecilia (Amphibia, Gymnophiona). Zoologische Mededelingen Leiden 54, 217 - 235.","Kamei, R. G., Wilkinson, M., Gower, D. J. & Biju, S. D. (2009) Three new species of striped Ichthyophis (Amphibia: Gymnophiona: Ichthyophiidae) from the northeast Indian states of Manipur and Nagaland. Zootaxa 267, 26 - 42.","Roze, J. A. & Solano, H. (1963) Resumen de la familia Caeciliidae (Amphibia: Gymnophiona) de Venezuela. Acta Biologica Venezuelica 3, 287 - 300.","Boulenger, G. A. (1882) Catalogue of the Batrachia Gradientia s. Caudata and Batrachia Apoda in the Collection of the British Museum (2 nd Ed.), Taylor & Francis, London.","Wake, M. H. & Donnelly, M. A. (2009) A new lungless caecilian (Amphibia: Gymnophiona) from Guyana. Proceedings of the Royal Society of London B 227 (1683), 915 - 922.","Gower, D. J., Wilkinson, M., Sherratt, E. & Kok, P. J. R. (2010) A new species of Rhinatrema Dumeril & Bibron (Amphibia: Gymnophiona: Rhinatrematidae) from Guyana. Zootaxa 2391, 47 - 60."]}

Eastern Pantepui Anomaloglossus species Anomaloglossus megacephalus can be distinguished from A. beebei by (characters of A. beebei in parentheses, see also Kok et al. 2006 b and Kok & Kalamandeen 2008) its larger size, female SVL max 28.3 mm [n= 3] in A. megacephalus (18.7 mm in A. beebei, n= 27), fingers with keel-like lateral folds, best developed preaxially on Fingers II and III (fringes not folded), toes with folded flaplike fringing (fringes not folded), throat in adult female blotched (immaculate), palm dark brown to black (yellowish), distinct dark bands on thigh and shank (absent), dark interorbital V-shaped band (absent). Anomaloglossus megacephalus can be distinguished from A. breweri (Barrio-Amorós, 2006) by (characters of A. breweri in parentheses) its larger size, female SVL max 28.3 mm [n= 3] in A. megacephalus (23.8 mm in A. breweri, n= 1), throat in adult female blotched (immaculate), median lingual process short, wider than long, tapered (distinctly longer than wide, tip pointed). Anomaloglossus megacephalus can be distinguished from A. kaiei by (characters of A. kaiei in parentheses, see also Kok & Kalamandeen 2008) its larger size, female SVL max 28.3 mm [n= 3] in A. megacephalus (19.8 mm in A. kaiei, n= 25), fingers with keel-like lateral folds, best developed preaxially on Fingers II and III (fringes not folded), toes moderately webbed (basally webbed), dorsolateral stripe absent (present), oblique lateral stripe present (absent), throat in adult female blotched (immaculate). Anomaloglossus megacephalus can be distinguished from A. murisipanensis (La Marca, 1998) in having (characters of A. murisipanensis in parentheses, all based on the preserved holotype, which is a juvenile and the only known specimen) fingers with keel-like lateral folds, best developed preaxially on Fingers II and III (fringes barely detectable), more webbing on toes, symmetrical cloacal tubercles present (absent), tarsal keel weakly to distinctly curved, slightly tuberclelike (straight, not tuberclelike), oblique lateral stripe always present, even if broken in spots (not distinguishable), no white stripe between naris and eye in preservative (present). Anomaloglossus megacephalus can be distinguished from A. praderioi by (characters of A. praderioi in parentheses, see also Kok 2010) its larger size, female SVL max 28.3 mm (n= 3) in A. megacephalus (22.7 mm in A. praderioi, n= 1), Finger IV longer than I (equal), tip of Finger IV always surpassing the base of the distal subarticular tubercle on Finger III when fingers adpressed (barely reaching the base), toes moderately webbed with folded flaplike fringing (basally webbed, folded flaplike fringing not present on all toes), dorsolateral stripe absent (present), oblique lateral stripe present (absent), throat in adult female blotched (immaculate). Anomaloglossus megacephalus can be distinguished from A. roraima in having (characters of A. roraima in parentheses) fingers with keel-like lateral folds, best developed preaxially on Fingers II and III (fringes barely distinct, not distinctly folded), all toes with folded flaplike fringing (absent), webbing on toes moderate (absent). Anomaloglossus megacephalus can be distinguished from A. rufulus (Gorzula, 1990) in having (characters of A. rufulus in parentheses) webbing on toes moderate (rudimentary), and most remarkably by ventral colour pattern; light with a few dark brown blotches in A. megacephalus (dark brown marbled with white blotches in A. rufulus). The new species superficially most resembles A. parkerae, A. tepuyensis (with which it has been confused), and A. triunfo (Barrio-Amorós, Fuentes-Ramos & Rivas-Fuenmayor, 2004), in sharing a similar dorsal pattern. It should be noted that A. triunfo is a possible synonym of A. tepuyensis (Kok & Barrio- Amorós unpubl. data, see also comments by Myers & Donnelly 2008). Anomaloglossus parkerae is distinguished from A. tepuyensis and A. triunfo only by a few characters such as size, definition of dorsal pattern, and condition of finger and toe fringes (Myers & Donnelly 2008). Adults of A. megacephalus can notably be distinguished from those three similar taxa in having a more slender body (Anomaloglossus parkerae, A. tepuyensis and A. triunfo are distinctly stockier frogs), a comparatively longer snout and a larger, somewhat more massive head [See Fig. 5 for close comparison between A. megacephalus and A. tepuyensis and A. parkerae; compare also with Duellman’s (1997: 10) colour plate of A. parkerae, fig. 30 (showing A. parkerae) in Lötters et al. (2007), and figs. 13, 15 and 16 illustrating A. tepuyensis in Myers & Donnelly (2008); see also below for further comments], and in having symmetrical cloacal tubercles (absent in all females of A. parkerae, A. tepuyensis and A. triunfo examined). Direct comparison of specimens shows a difference in head size in Anomaloglossus megacephalus compared to the similar A. parkerae, A. tepuyensis and A. triunfo, as confirmed by plotting of morphometric proportions. Comparison of HL 3 with BEL indicates that the new species proportionally has a longer head than A. tepuyensis (Fig. 6). In order to distinguish between those similar species, the midline distance from the tip of the snout to an imaginary line between anterior insertions of upper arms is measured, and that distance is then extended posteriorly from the same imaginary line. In two of the three available A. megacephalus adult specimens the distance reaches or surpasses the anterior insertion of thigh, whereas in all adult specimens of A. parkerae, A. tepuyensis and A. triunfo examined (n = 20), the distance fails to reach the thigh insertion (see Fig. 5 B). It is noteworthy to mention that the only specimen of A. megacephalus in which HL 3 is not longer than BEL is in poor preservation state.

Distinguishing among Anomaloglossus species from the Pantepui region We first present a focussed comparison between the new species and the ten Anomaloglossus species known to occur in the Eastern Pantepui District (i.e. east of the Rio Caron��). The Rio Caron�� likely acts as a biogeographic barrier for Anomaloglossus species since no species has been reported to occur on both sides of the river in the Guiana highlands of Venezuela and Guyana (it must be noted that very few Pantepui anurans are known to occur on both sides of the river). But, in order to provide a more comprehensive comparison, we also compared the new species with the six known Anomaloglossus species that occur in the Western Pantepui District (i.e. west of the Rio Caron��) in the Guiana highlands of Venezuela., Published as part of Kok, Philippe J. R., Macculloch, Ross D., Lathrop, Amy, Willaert, Bert & Bossuyt, Franky, 2010, A new species of Anomaloglossus (Anura: Aromobatidae) from the Pakaraima Mountains of Guyana, pp. 18-32 in Zootaxa 2660 on page 26, DOI: 10.5281/zenodo.199004

Anomaloglossus megacephalus sp. nov. (Figs. 1���4; Table 1) Anomaloglossus tepuyensis (La Marca, 1998) in part; Grant et al. (2006: 120, 158) Anomaloglossus tepuyensis (La Marca, 1997 [sic]); MacCulloch & Lathrop (2009: 11, figs. 5���6, plate B) Holotype. IRSNB 1986 (field number PK 1881), an adult female collected by Philippe J. R. Kok, Paul Benjamin and Claudius Perry, 20 November 2007 at 16 h 30, eastern base of Mount Maringma, Cuyuni- Mazaruni District, Guyana (05�� 12 ��� 37 ���N, 060�� 33 ��� 59 ���W, 1060 m elevation). Paratypes. ROM 39637 - 38, adult females collected by Amy Lathrop and Carter Cox, 28 October 2000, northeastern plateau of Mount Ayanganna, Cuyuni-Mazaruni District, Guyana (05�� 24 ��� N, 59 �� 57 ��� W, 1490 m elevation). Etymology. The specific epithet is a combination of mega (Greek meaning "large") and cephalus (Greek meaning ���head���), which refers to the large head of the species and is used as a noun in apposition. Adult definition and diagnosis. (1) large-sized Anomaloglossus (male unknown, female 27.3���28.3 mm SVL); (2) body robust, but relatively more slender than most known congeners, head large; (3) skin on dorsum smooth to granular, more tuberculate posteriorly, skin on venter smooth to slightly granular; (4) Finger I subequal or shorter than II; (5) tip of Finger IV surpassing the base of the distal subarticular tubercle on Finger III when fingers adpressed; (6) distal tubercle on Finger IV present; (7) condition of Finger III in males unknown; (8) fingers with keel-like lateral folds (sensu Myers & Donnelly 2008), best developed preaxially on Fingers II and III; (9) toes moderately webbed, with well-developed folded flaplike fringing (sensu Myers & Donnelly 2008); (10) tarsal keel well-defined, weakly curved, slightly tuberclelike; (11) presence of black arm gland in males (sensu Grant & Castro-Herrera 1998, see also Grant et al. 2006) unknown; (12) symmetrical cloacal tubercles present; (13) pale paracloacal mark present; (14) dorsolateral stripe absent in female (condition unknown in males); (15) ventrolateral stripe present, inconspicuous; (16) oblique lateral stripe present, broken in spots or relatively well-defined; (17) condition of sexual dichromatism in throat colour unknown; (18) condition of sexual dichromatism in ventral colouration unknown; (19) iris with metallic pigmentation and pupil ring; (20) large intestine anteriorly pigmented; (21) colour of testes unknown; (22) mature oocytes partly pigmented; (23) median lingual process short, wider than long, tapered (24) maxillary teeth present, small. Description of the Holotype. An adult female 27.3 mm SVL in excellent condition of preservation. Dorsal skin granular, more tuberculate posteriorly; ventral skin slightly granular. In preservative, a barely distinct epidermal ridge borders the tip of snout dorsally. Dorsal surface of hind limbs granular and tuberculate, with two distinct symmetrical enlarged tubercles located laterally between urostyle and vent (Fig. 1; see Grant et al. 1997 for a description and illustration of the cloacal tubercles). Head longer than wide (HL 2> HW), greatest width 34 % of SVL. Snout bluntly pointed in lateral view, extending past the lower jaw, bluntly pointed in ventral and dorsal views. Nares located close to tip of snout, directed posterolaterally; nares visible from front, barely visible from above and below; posterior rim of naris bordered behind by a crescent-shaped ridge; rim bearing a small tuberclelike prominence posterodorsally, this ���bump��� is distinct in frontal, dorsal and ventral views; internarial distance 40 % of greatest head width. Canthus rostralis well defined; loreal region straight, sloping outward to lip. Interorbital distance 83 % of eye length, 90 % of upper eyelid width. Snout length 123 % of eye length, 49 % of head length; distance from anterior corner of eye to posterior margin of naris 60 % of eye length. Postrictal tubercles inconspicuous. Tympanic membrane inconspicuous, round, concealed posterodorsally by a diffuse supratympanic swelling; tympanic annulus prominent anteroventrally; tympanum 43 % of eye length, separated from eye by a distance equal to 23 % of eye length. Midline distance from tip of snout to arm insertion greater than distance from arm insertion to thigh insertion (HL 3> BEL). Forearm slightly longer than upper arm, no distinct ulnar fold. Hand moderate in size, HAND III 27 % of SVL, 78 % of HW. Relative length of fingers III> IV> I ��� II. Fingers unwebbed. Discs of fingers expanded, disc on Finger IV widest, discs on other fingers subequal. Fingers with keel-like lateral folds (sensu Myers & Donnelly 2008), best developed pre- and (slightly) postaxially on Finger II and preaxially on Finger III (Fig. 2 A). Palmar tubercle large, rounded, slightly heart-shaped; thenar tubercle smaller, elliptical; one or two round to ovoid subarticular tubercles (one each on Fingers I and II, two each on Fingers III and IV, with distal tubercles on Finger III and IV slightly less conspicuous). A very low, barely distinct outer metacarpal fringe. Tip of Finger IV distinctly surpassing the base of distal subarticular tubercle on Finger III when fingers adpressed. No fleshy supracarpal fold atop wrist (Fig. 2 A). Hind limbs robust, moderately long, with heel of adpressed leg reaching anterior corner of eye; TIL 49 % of SVL. Relative lengths of adpressed toes IV> III> V> II> I; first toe short, slightly surpassing the base of subarticular tubercle of second toe. Toe discs expanded, slightly larger than finger discs, largest on Toes II and IV. Feet moderately webbed; all toes with well developed folded flaplike fringing (sensu Myers & Donnelly 2008), except on postaxial edge of Toe V on the right side. Webbing formula I 0+��� 2 - II 1 -��� 2 �� III 2 -��� 3 + IV 3 ����� 2 - V (Fig. 2 A). Inner metatarsal tubercle small, elliptical; outer metatarsal tubercle small, round, about half the size of the inner; a distinct oval medial metatarsal tubercle present on both sides. One to three round to ovoid subarticular tubercles (one each on Toes I and II, two each on Toes III and V, and three on Toe IV, with distal tubercle on Toe IV the smallest and least conspicuous). A strong outer metatarsal fold coextensive with the folded flaplike fringing on Toe V, almost reaching the outer metatarsal tubercle on the left side; outer metatarsal fold absent on the right side. A slightly tuberclelike tarsal keel weakly curved at proximal end, extending proximolaterad from preaxial edge of inner metatarsal tubercle, not continuous with the fringe along the outer edge of the first toe (Fig. 2 A). Maxillary teeth present, small. Tongue longer than wide, wider and free posteriorly, with rounded margin; median lingual process short, wider than long, tapered. Colour of the Holotype in life. Dorsal ground colour medium brown, with a dark brown V-shaped interorbital bar slightly outlined with light creamish brown, followed by two less defined dark brown bars, the first more or less V-shaped, at the level of arm insertion, the second just anterior to sacrum. Small, poorly defined, dark brown flecks on snout, between and on dorsal bars. Upper surface of arm light brown with welldefined dark brown transverse bands on forearm and wrist, a less defined one on elbow; upper surface of thigh, shank, and foot light brown with well-defined dark brown transverse bands. A yellow wash on the anterior face of the thigh. Flanks dark brown, slightly lighter ventrally with a few bluish irregular spots and a few whitish irregular blotches on the lower part forming a broken inconspicuous ventrolateral stripe. A few white spots (covering small tubercles) form a broken oblique lateral stripe that does not extend to arm insertion. No dorsolateral stripe. Upper lip creamish brown suffused with melanophores, two bluish marks below eye; loreal region and side of head dark brown; tympanum area light creamish brown concealed in a poorly defined light stripe from posterior corner of eye to arm insertion. A dark brown stripe is present on the anterior edge of upper arm, tapering from arm insertion to forearm. Throat light brown, blotched with white and dark brown; belly light grey, blotched with dark brown, blotches more numerous and larger laterally. Undersurface of upper arm light brown with a few dark markings; undersurface of forearm blackish; undersurface of thigh and shank light grey blotched with dark brown; rear of thigh and cloacal region blackish. Pale reddish orange paracloacal marks present. Tip of digits whitish. Palms and soles black. Iris mostly orange-bronze, darkened with black suffusion, with two oblique metallic white marks separated by a black triangular mark below pupil (Fig. 1 A���B). Colour of the Holotype in preservative. Dorsal ground colour brown, with a dark brown V-shaped interorbital bar slightly outlined with light brown, followed by two less defined dark brown bars, the first more or less V-shaped, at the level of arm insertion, the second just anterior to sacrum. Small, poorly defined, dark brown flecks on snout, between and on dorsal bars. Upper surface of arm light brown with well-defined dark brown transverse bands on elbow, forearm and wrist; upper surface of leg light brown with well-defined dark brown transverse bands on thigh, shank, and foot. Flanks dark brown, with some irregular spots and a few whitish irregular blotches on the lower part forming a broken inconspicuous ventrolateral stripe. A few white spots (covering small tubercles) form a broken oblique lateral stripe that does not extend to arm insertion. No dorsolateral stripe. Upper lip dirty brown suffused with melanophores, two whitish marks below eye; loreal region and side of head black; distinct whitish stripe extending from posterior corner of eye to arm insertion including most of tympanum. A dark brown stripe is present on the anterior edge of upper arm, tapering from arm insertion to forearm. Throat and belly whitish, blotched with dark brown. Undersurface of upper arm whitish with a few dark markings; undersurface of forearm blackish; undersurface of thigh whitish blotched with dark brown; undersurface of shank light grey blotched with dark brown; rear of thigh and cloacal region blackish. Light brown paracloacal marks present. Palms and soles black (Fig. 1 B���C). Variation. See Table 1 for morphometric measurements of the holotype and the paratypes and Figs. 2 B and 3 for intra- and interpopulational variation. Symmetrical enlarged cloacal tubercles are clearly visible in the holotype. Those tubercles are present, but less visible in the paratypes probably due to an artefact of preservation (cloacal tubercles are most visible in well preserved specimens). Only the holotype and one paratype have a medial metatarsal tubercle, a character that is sometimes present in other Anomaloglossus species as well. Only the holotype lacks folded flaplike fringing on the postaxial edge of Toe V and a coextensive strong outer metatarsal fold on the right side, indicating an aberrant condition. An additional difference between the holotype and the paratypes is the length of Finger I, which is slightly longer than II in the holotype on the left hand (vs. shorter in the two paratypes), and subequal on the right hand (vs. shorter in the two paratypes). Character IRSNB 1986 ROM 39637 ROM 39638 Toe webbing variation in females (male unknown) is I 0+- 2 - II (1 - - 1 ��)���(2 ��- 3 -) III (2 - - 2)���(3 +- 3 ��) IV (3 ��- 3 ��)���(2 - - 2) V. The condition of the oblique lateral stripe is variable among specimens, from a series of light spots/ blotches extending diagonally from the groin across the flanks and barely reaching the level of the insertion of the arm (e.g. IRSNB 1986) to an almost solid, clearly visible stripe extending diagonally from the groin across the flanks to the level of the insertion of the arm, then breaking in a series of light spots almost reaching the eye (e.g. ROM 39638). Slight variation also occurs between the right and the left flank. In life, ROM 39638 was darker and the dark dorsal bars less detectable (Fig. 2 B). Tadpole description. Two tadpoles were collected in a stream at 1490 m on Mt. Ayanganna. Description and illustrations of one of the two stage- 25 tadpoles are provided in MacCulloch and Lathrop (2009: 11, under Anomaloglossus tepuyensis). Comparison with other known Anomaloglossus tadpoles occurring in the Pantepui region.- Tadpoles of A. megacephalus most resemble those of A. parkerae (Meinhardt & Parmalee, 1996) and A. tepuyensis, which are also deposited in streams. The tadpole of A. parkerae was described by Duellman (1997), that of A. tepuyensis by Myers and Donnelly (2008). Both descriptions contain drawings and written descriptions. Two items are noteworthy: (1) Duellman (1997) describes the oral disc of A. parkerae as ���not emarginate���, whereas Myers and Donnelly (2008) describe the oral disc of A. tepuyensis as ���emarginate���. The oral disc margins of these two species are very similar (Duellman 1997, fig. 12; Myers & Donnelly 2008, fig. 17) and we follow Grant et al. (2006) in describing the oral discs of Anomaloglossus as emarginate; (2) Duellman (1997) describes the LTRF of A. parkerae as 2 (1)/ 3, although the oral disc illustrated shows a LTRF 2 (2)/ 3. This is obviously an error and the LTRF of A. parkerae is 2 (2)/ 3, like in A. tepuyensis and A. megacephalus. Although there are few quantifiable differences among the larvae of Anomaloglossus megacephalus, A. parkerae and A. tepuyensis, the tadpole of A. megacephalus markedly differs from that of A. parkerae and A. tepuyensis in having a longer tail [tail length of stage- 25 tadpoles is 68 % of total length in A. megacephalus (n= 2) vs. 62-63 % in A. parkerae (n= 4) and A. tepuyensis (n= 29) tadpoles in the same stage], in having the maximum height of tail similar to body height (distinctly higher in A. parkerae and A. tepuyensis), and in having the upper fin noticeably lower than tail musculature at midtail (higher or equal in A. parkerae and A. tepuyensis). When compared to other known Anomaloglossus tadpoles from the Pantepui region, A. megacephalus differs from A. beebei and A. roraima by stream habitat (phytotelmata habitat in A. beebei and A. roraima); from A. kaiei and A. praderioi in having the upper fin slightly increasing in length from tail-body junction [first quarter of upper fin straight before increasing in length to about midlength in A. kaiei (n= 30) and A. praderioi (n= 11)]; and from A. tamacuarensis (Myers & Donnelly, 1997) in having a longer tail [body length of stage- 25 tadpoles is 68 % of total length in A. megacephalus (n= 2) vs. 63-65 % in A. tamacuarensis (n= 12) tadpoles in the same stage], and in having less marginal papillae and shorter lateral processes. Distribution and ecology. Anomaloglossus megacephalus is currently only known from the eastern base of Maringma Tepui, Guyana, where it occurs at 1060 m elevation in low-canopy forest, and from the northeast plateau of Mt. Ayanganna, Guyana on the basis of two specimens collected at 1490 m elevation in dense lowcanopy, high-tepui forest (Fig. 4). Anomaloglossus megacephalus appears to be uncommon and elusive, with only three specimens collected. It is a diurnal species, the holotype was found on the sandy bank of a small forest stream. The Ayanganna specimens were collected in wet forest not closely associated with stream banks. Egg deposition site unknown. Tadpoles are deposited in streams. Advertisement call is unknown., Published as part of Kok, Philippe J. R., Macculloch, Ross D., Lathrop, Amy, Willaert, Bert & Bossuyt, Franky, 2010, A new species of Anomaloglossus (Anura: Aromobatidae) from the Pakaraima Mountains of Guyana, pp. 18-32 in Zootaxa 2660 on pages 20-26, DOI: 10.5281/zenodo.199004, {"references":["La Marca, E. (1998 \" 1996 \") Ranas del genero Colostethus (Amphibia: Anura: Dendrobatidae) de la Guyana Venezolana con la descripcion de siete especies nuevas. Publicaciones de la Asociacion de Amigos de Donana, 9, 1 - 64.","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F. B., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262.","MacCulloch, R. D. & Lathrop, A. (2009) Herpetofauna of Mount Ayanganna, Guyana. Results of the Royal Ontario Museum Ayanganna Expedition 2000. Royal Ontario Museum Science Contributions, 4, 1 - 36.","Character IRSNB 1986 ROM 39637 ROM 39638 Toe webbing variation in females (male unknown) is I 0 + - 2 - II (1 - - 1 ½) - (2 ½ - 3 -) III (2 - - 2) - (3 + - 3 ½) IV (3 ½ - 3 ½) - (2 - - 2) V.","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum - Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147.","Grant, T. & Castro-Herrera, F. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology, 32, 378 - 392.","Grant, T., Humphrey, E. C. & Myers, C. W. (1997) The median lingual process of frogs: A bizarre character of Old World ranoids discovered in South American dendrobatids. American Museum Novitates, 3212, 1 - 40.","Meinhardt, D. J. & Parmelee, J. R. (1996) A new species of Colostethus (Anura: Dendrobatidae) from Venezuela. Herpetologica, 52, 70 - 77.","Duellman, W. E. (1997) Amphibians of La Escalera Region, Southeastern Venezuela: Taxonomy, Ecology, and Biogeography. Scientific Papers of the Natural History Museum of the University of Kansas, 2, 1 - 52.","Myers, C. W. & Donnelly, M. A. (1997) A tepui herpetofauna on a granitic mountain (Tamacuari) in the borderland between Venezuela and Brazil: report from the Phipps Tapirapeco expedition. American Museum Novitates, 3213, 1 - 71."]}

Western Pantepui Anomaloglossus species Among the Western Pantepui Anomaloglossus species, A. ayarzaguenai (La Marca, 1998) and A. moffetti Barrio-Amor��s and Brewer-Car��as, 2008 are the geographically closest species to A. megacephalus (ca. 400 km airline to the west). Anomaloglossus ayarzaguenai and A. moffetti are very similar to each other, geographically close (from Cerro Jaua and Cerro Sarisari��ama, respectively) and possible synonyms (Kok & Barrio-Amor��s unpubl. data). Anomaloglossus megacephalus is distinguished from A. ayarzaguenai and A. moffetti in having a comparatively longer snout, and a larger, somewhat more massive head [compare Figs. 1 and 2 B with fig. 4 a���c illustrating A. moffetti in Barrio-Amor��s & Brewer-Car��as (2008)], in having a distinct tympanum in life (barely distinct in A. ayarzaguenai and A. moffetti), separated from eye by 42���60 % of its greatest length (25 % or less in A. ayarzaguenai and A. moffetti), and in lacking a dark area spotted with white ventrolaterally at the level of arm insertion (always present in A. ayarzaguenai and A. moffetti). Anomaloglossus megacephalus mostly differs from A. guanayensis (La Marca, 1998) by (characters of A. guanayensis in parentheses) its larger size, female SVL max 28.3 mm (n= 3) in A. megacephalus (23.5 mm in A. guanayensis, n= 3), and in having less developed keel-like lateral folds on fingers (extensive, present pre- and postaxially on all fingers). Anomaloglossus megacephalus can be distinguished from A. parimae (La Marca, 1998) by (characters of A. parimae in parentheses) its larger size, female SVL max 28.3 mm (n= 3) in A. megacephalus (23.1 mm in A. parimae, n= 2), in having a short, wider than long median lingual process (slender, noticeably longer than wide), and in having distinctly more webbing on toes. Anomaloglossus megacephalus can be distinguished from A. shrevei (Rivero, 1961) in having (characters of A. shrevei in parentheses) a weakly to distinctly curved, slightly tuberclelike tarsal keel, (short, straight, not tuberclelike), less developed keel-like lateral folds on fingers (extensive, present pre- and postaxially on all fingers), and dark blotches on throat and belly (throat and belly immaculate). Anomaloglossus megacephalus mostly differs from A. tamacuarensis by (characters of A. tamacuarensis in parentheses) its larger size, female SVL max 28.3 mm (n= 3) in A. megacephalus (25.0 mm in A. tamacuarensis, n= 2), in having a comparatively longer snout, and a larger, somewhat more massive head [compare Figs. 1 and 2 B with figs. 11 and 13 illustrating A. tamacuarensis in Myers & Donnelly (1997)], less developed keel-like lateral folds on fingers (extensive, present pre- and postaxially on all fingers), a conspicuous tympanum (inconspicuous), and in having a short, wider than long median lingual process (slender, noticeably longer than wide)., Published as part of Kok, Philippe J. R., Macculloch, Ross D., Lathrop, Amy, Willaert, Bert & Bossuyt, Franky, 2010, A new species of Anomaloglossus (Anura: Aromobatidae) from the Pakaraima Mountains of Guyana, pp. 18-32 in Zootaxa 2660 on pages 28-29, DOI: 10.5281/zenodo.199004, {"references":["La Marca, E. (1998 \" 1996 \") Ranas del genero Colostethus (Amphibia: Anura: Dendrobatidae) de la Guyana Venezolana con la descripcion de siete especies nuevas. Publicaciones de la Asociacion de Amigos de Donana, 9, 1 - 64.","Barrio-Amoros, C. L. & Brewer-Carias, C. (2008) Herpetological results of the 2002 expedition to Sarisarinama, a tepui in Venezuelan Guayana, with the description of five new species. Zootaxa, 1942, 1 - 68.","Rivero, J. A. (1961) Salientia of Venezuela. Bulletin of the Museum of Comparative Zoology, 126, 1 - 207.","Myers, C. W. & Donnelly, M. A. (1997) A tepui herpetofauna on a granitic mountain (Tamacuari) in the borderland between Venezuela and Brazil: report from the Phipps Tapirapeco expedition. American Museum Novitates, 3213, 1 - 71."]}

Key to the species of Chironius from the Guiana Shield 1. 10 scale rows at midbody.............................................................................................................................................. 2 - 12 scale rows at midbody............................................................................................................................................ 3 2. Fewer than 149 ventrals ....................................................................................................................... Chironius fuscus - More than 149 ventrals................................................................................................................................................ 4 3. More than 178 ventrals............................................................................................................... Chironius cochranae - Fewer than 178 ventrals................................................................................................................................................ 5 4. Loreal distinctly longer than high; adults red, juveniles green ....................................................... Chironius scurrulus - Loreal as high as long, or slightly longer than high; adults and juveniles brownish with lighter crossbands............... ...................................................................................................................................................... Chironius challenger 5. Yellow spots on the first scale row of tail ....................................................................................... Chironius carinatus - Not as above ...................................................................................................................................... Chironius exoletus

Key to the species of Chironius from the Guiana Shield 1. 10 scale rows at midbody.............................................................................................................................................. 2 - 12 scale rows at midbody............................................................................................................................................ 3 2. Fewer than 149 ventrals ....................................................................................................................... Chironius fuscus - More than 149 ventrals................................................................................................................................................ 4 3. More than 178 ventrals............................................................................................................... Chironius cochranae - Fewer than 178 ventrals................................................................................................................................................ 5 4. Loreal distinctly longer than high; adults red, juveniles green ....................................................... Chironius scurrulus - Loreal as high as long, or slightly longer than high; adults and juveniles brownish with lighter crossbands............... ...................................................................................................................................................... Chironius challenger 5. Yellow spots on the first scale row of tail ....................................................................................... Chironius carinatus - Not as above ...................................................................................................................................... Chironius exoletus, Published as part of Kok, Philippe J. R., 2010, A new species of Chironius Fitzinger, 1826 (Squamata: Colubridae) from the Pantepui region, northeastern South America, pp. 31-44 in Zootaxa 2611 on page 43, DOI: 10.5281/zenodo.197801

Microcaecilia iyob sp. nov. (Figs. 1; Table 1) Gymnopis unicolor (Duméril, 1864) in part; Dunn (1942: 471–472) Microcaecilia unicolor (Duméril, 1864) in part; Taylor (1968: 542–546, fig. 297) Holotype. FMNH 35117, a mature male from “Oko River at Cuyuni River” Guyana; collected by Neal A. Weber in June 1936. Diagnosis. A Microcaecilia that differs from M. taylori Nussbaum & Hoogmoed, 1979 and M. grandis Wilkinson et al., 2009 in having fewer PAs (M. rabei (Roze & Solano, 1963) in having fewer PAs undivided by SAGs ( 8); and from M. albiceps (Boulenger, 1882), M. supernumeraria Taylor, 1970, and M. unicolor (Duméril, 1864) in having more numerous premaxillary-maxillary teeth (> 20) in long series that extend posteriorly beyond the posterior margin of the choanae, and bicuspid vomeropalatine teeth. Description of the holotype. Some morphometric and meristic data are given in Table 1. In poor condition in places, incisions extending from CMs across C 1 and C 2, almost broken into two parts at the 12 th PA, less substantial breaks at about 25 th and 60 th PA, small (5mm) midventral incisions just anterior and posterior to second of these breaks, a 12 mm incision beginning 45 mm in front of terminus, some open scale pockets and patches of abraded skin, rusty hole on 36 th PA another 20 mm anterior to vent. An arthropodan exoskeletal element embedded in the skin of the lower jaw on the left side. Mature male with multiple large testis lobes. Body dorsoventrally flattened throughout, relatively uniform, narrowing slightly onto collars anteriorly and more substantially behind the level of the vent; L/W = c. 32. In dorsal view, sides of head straight, converging steadily to level of nares, snout tip between nares rather blunt. In lateral view, top of head weakly convex; margins of upper jaw (lip) concave especially anterior to the level of the TAs; ridge bearing vomeropalatine teeth just visible close to CM; lower jaw half the height of upper jaw at CM. In ventral view, mouth moderately recessed, margins of lower jaw and upper lips not much more blunt than tip of snout. Eyes not visible. TAs very slightly elevated, the elevations visible dorsally and ventrally, lie just above imaginary lines between nares and CMs, distinctly closer to CMs than to nares. Nares small, dorsolateral, circular depressions with anteroventral ovate aperture, somewhat closer to level of AM than to ST, much closer to top than to bottom of snout in lateral view, not visible from below. Teeth pointed, gently recurved, lacking serrations or flanges; elements of outer series much smaller posteriorly, monocuspid, dentaries largest, premaxillary-maxillary teeth large; vomeropalatines much smaller, more uniform in size, bicuspid, about as many in front of the anterior margins of the choanae as behind, vomerine series forming gentle arch anteromedially; distance between vomeropalatine and premaxillary-maxillary series anteriorly almost equal to projection of snout; series converging posteriorly to about half that distance, upper series extending posteriorly well (5–7 elements or more on each side) past choanae, approaching CMs, palatine series extending slightly further posteriorly. Palate moderately arched. Choanal apertures subcircular, separated from each other by twice their individual widths, anterior margins approximately level with tentacles; valves not visible. Tongue simple (no surface architecture) with broad attachment anteriorly. Nuchal region barely more massive than adjacent body. C 1 shorter than C 2, well-marked, NG 1 and NG 2 completely encircling body, bending slightly anteromedially on dorsum, NG 3 incomplete ventrally; substantial TG on dorsum of C 2, just visible laterally. Behind collars, 97 PAs and a small unsegmented terminal cap. AGs slightly raised posteriorly, complete or nearly so dorsally, mostly separated mid-ventrally, 31 AGs complete ventrally anterior to vent; no AGs posterior to vent on venter. First SAG present mid-dorsally on 52 nd PA, all PAs behind 55 th divided by mid-dorsally complete SAGs, 12 of which are complete ventrally (on PAs 84–95) anterior to vent region. Vent region interrupts last four to five AGs. Scattered small scales in a single incomplete row present dorsally as far anterior as 25 th PA, near terminus scale pockets as deep as the width of a PA, with multiple (8–9) rows of either slightly ovate or rhomboidal (largest 1.0 x 0.8 mm) scales. No subdermal connective tissue scales. Body terminus acuminate with no indication of terminal keel. In lateral view, ventral surface concave/indented at level of vent. Vent within a fairly discrete, approximately circular, depressed cloacal disc, with slightly transverse opening, its lips formed by about 13 somewhat irregular, short denticulations; a pair of cloacal papillae on anterior denticulations. Gray-brown, more brown anteriorly and where superficial epidermis missing, darker, more plumbous grey posteriorly, perhaps slightly paler on lateral flanks especially posteriorly where each annulus has a paler anterior and darker posterior half. Head paler than body especially ventrally, except for narrow darker lines approximately level with medial aspect of mandibles, central pale colour extending onto C 1. Collar grooves and AGs edged in white, especially laterally. TAs and nares lie within small pale spots. Cloacal disc, including small ventral area of terminal cap, pale. Etymology. The species is named to commemorate the International Year of Biodiversity (2010) to serve as a reminder that much biodiversity is undescribed and much sampled biodiversity is poorly understood taxonomically. The epithet iyob is designated a genderless noun in apposition. Suggested English common name. Commemorative microcaecilia. Edward Harrison Taylor was by far the most important figure in the history of caecilian taxonomy. In his then comprehensive taxonomic study of caecilians, Taylor (1968) provided accounts for some 158 species, of which he was the authority for almost half (73). These and other prodigious feats of species description led to Taylor having a reputation as something of a “splitter”, and a not insubstantial number of his species have been subsequently relegated to synonyms. Although Taylor occasionally explicitly held back from describing possible new species for variant specimens, this was the exception rather than the rule. In his account of Microcaecilia unicolor, Taylor (1968: 544) gave data for seven specimens from French Guiana, including the lectotype, and for a single specimen from Guyana that is now the holotype of M. iyob. These data reveal substantial differences in annulation and dentition between the M. unicolor from French Guiana and the Guyana specimen, that result in some very high ranges for these characters in Taylor’s diagnosis of M. unicolor. Of these differences Taylor (1968: 546) writes only that the Guyana specimen “differs in having a primary count of about eight less than the lowest French Guiana specimen, and 18 lower than the highest count. The secondaries are reported to be more erratic, varying from 22 to 74? (fide Dr. E. R. Dunn), the number of complete secondaries” [sic]. Thus, Taylor (1968) was following Dunn (1942) in including the Guyana specimen with the species unicolor, and was perhaps dissuaded from considering the distinctiveness of the Guyana specimen within his sample any further by Dunn’s report of a very wide range in secondary annuli. Uncharacteristically, Taylor (1968) did not comment on the clear differences in the numbers and disposition of teeth in the upper jaw that are evident in his data and that, by themselves, are strong evidence that the samples from French Guiana and from Guyana are not conspecific. Two years later, Taylor (1970: 313) highlighted that the Guyana specimen differed from the type in the number of premaxillary-maxillary teeth and cautioned “Material from this area should be acquired and carefully examined, as I may have erred in so disposing of this specimen.” Unfortunately, Taylor (1968) used the holotype of Microcaecilia iyob as the basis for his descriptive account of M. unicolor and, as Wilkinson et al. (2009) noted, this has contributed to taxonomic confusion regarding M. unicolor (e.g. Nussbaum & Hoogmoed 1979). There are some minor differences between the data reported by Taylor (1968) and those reported here on the holotype of Microcaecilia iyob (Table 1). Additionally, Taylor (1968) reported that the PAGs are mostly incomplete dorsally whereas we find them to be mostly complete or nearly so, and he also reported that the first and second nuchal grooves curve forward ventrally whereas we find them to be mostly straight (Fig. 21. A syndrome of dental characters seemingly divides Microcaecilia into two groups. Microcaecilia albiceps, the type species, and M. unicolor have short premaxillary-maxillary teeth series that do not extend posteriorly beyond the choanae, bicuspid vomeropalatine teeth and serrated dentary teeth. The enigmatic M. supernumeraria, known only from its holotype, appears to belong with these species. It shares the first two of these dental characters but the condition of the third is unknown. Differently, M. iyob resembles M. rabei, M. grandis and M. taylori in having long premaxillary-maxillary teeth series and all teeth are monocuspid and unserrated. The new species is readily distinguished from these other Microcaecilia by its pattern of annulation with relatively few PAs and many SAGs and consequently relatively small numbers of undivided PAs. Several museum specimens of Microcaecilia that we have seen, including the holotype of M. iyob, have pointed and recurved arthropodan exoskeletal remains embedded in the skin of the head. We suspect this is evidence of frequent and aggressive encounters with either ants or termites, and possibly the result of preying upon them. The new species is known from only a single specimen collected more than 70 years ago and, given that there have been no dedicated attempts to recollect or otherwise survey its status in the wild, it must be considered data deficient for conservation purposes. Targeted fieldwork is needed to shed light on the distribution and abundance of this very poorly known species, and to provide material for both morphological and molecular systematic study. Microcaecilia iyob is only the third new species of caecilian described from Guyana since Taylor (1968). That these descriptions have appeared within the last two years and have been based on both newly collected (Wake & Donnelly 2009, Gower et al. 2010) and older material (this work), suggests both that the caecilian fauna of Guyana remains very incompletely known, and that further species new to science probably remain to be discovered from this region.

The first new species of rhinatrematid caecilian in more than 50 years, Rhinatrema shiv sp. nov., is described from Guyana. The type series of 10 specimens differs from the only other species in the genus, R. bivittatum, in having a longer tail, different colour pattern, more acuminate and relatively shorter head, and in several features of the skull and mandible including dentition. Sequence data for 1572 aligned base pairs of mitochondrial DNA are > 15% different between the two species. Rhinatrema shiv is known only from Guyana, and R. bivittatum from French Guiana and far northeastern Brazil. The only reported specimen of the genus from Suriname is not clearly referable to either of the named species.

Western Pantepui Anomaloglossus species Among the Western Pantepui Anomaloglossus species, A. ayarzaguenai (La Marca, 1998) and A. moffetti Barrio-Amorós and Brewer-Carías, 2008 are the geographically closest species to A. megacephalus (ca. 400 km airline to the west). Anomaloglossus ayarzaguenai and A. moffetti are very similar to each other, geographically close (from Cerro Jaua and Cerro Sarisariñama, respectively) and possible synonyms (Kok & Barrio-Amorós unpubl. data). Anomaloglossus megacephalus is distinguished from A. ayarzaguenai and A. moffetti in having a comparatively longer snout, and a larger, somewhat more massive head [compare Figs. 1 and 2 B with fig. 4 a–c illustrating A. moffetti in Barrio-Amorós & Brewer-Carías (2008)], in having a distinct tympanum in life (barely distinct in A. ayarzaguenai and A. moffetti), separated from eye by 42–60 % of its greatest length (25 % or less in A. ayarzaguenai and A. moffetti), and in lacking a dark area spotted with white ventrolaterally at the level of arm insertion (always present in A. ayarzaguenai and A. moffetti). Anomaloglossus megacephalus mostly differs from A. guanayensis (La Marca, 1998) by (characters of A. guanayensis in parentheses) its larger size, female SVL max 28.3 mm (n= 3) in A. megacephalus (23.5 mm in A. guanayensis, n= 3), and in having less developed keel-like lateral folds on fingers (extensive, present pre- and postaxially on all fingers). Anomaloglossus megacephalus can be distinguished from A. parimae (La Marca, 1998) by (characters of A. parimae in parentheses) its larger size, female SVL max 28.3 mm (n= 3) in A. megacephalus (23.1 mm in A. parimae, n= 2), in having a short, wider than long median lingual process (slender, noticeably longer than wide), and in having distinctly more webbing on toes. Anomaloglossus megacephalus can be distinguished from A. shrevei (Rivero, 1961) in having (characters of A. shrevei in parentheses) a weakly to distinctly curved, slightly tuberclelike tarsal keel, (short, straight, not tuberclelike), less developed keel-like lateral folds on fingers (extensive, present pre- and postaxially on all fingers), and dark blotches on throat and belly (throat and belly immaculate). Anomaloglossus megacephalus mostly differs from A. tamacuarensis by (characters of A. tamacuarensis in parentheses) its larger size, female SVL max 28.3 mm (n= 3) in A. megacephalus (25.0 mm in A. tamacuarensis, n= 2), in having a comparatively longer snout, and a larger, somewhat more massive head [compare Figs. 1 and 2 B with figs. 11 and 13 illustrating A. tamacuarensis in Myers & Donnelly (1997)], less developed keel-like lateral folds on fingers (extensive, present pre- and postaxially on all fingers), a conspicuous tympanum (inconspicuous), and in having a short, wider than long median lingual process (slender, noticeably longer than wide).

Eastern Pantepui Anomaloglossus species Anomaloglossus megacephalus can be distinguished from A. beebei by (characters of A. beebei in parentheses, see also Kok et al. 2006 b and Kok & Kalamandeen 2008) its larger size, female SVL max 28.3 mm [n= 3] in A. megacephalus (18.7 mm in A. beebei, n= 27), fingers with keel-like lateral folds, best developed preaxially on Fingers II and III (fringes not folded), toes with folded flaplike fringing (fringes not folded), throat in adult female blotched (immaculate), palm dark brown to black (yellowish), distinct dark bands on thigh and shank (absent), dark interorbital V-shaped band (absent). Anomaloglossus megacephalus can be distinguished from A. breweri (Barrio-Amor��s, 2006) by (characters of A. breweri in parentheses) its larger size, female SVL max 28.3 mm [n= 3] in A. megacephalus (23.8 mm in A. breweri, n= 1), throat in adult female blotched (immaculate), median lingual process short, wider than long, tapered (distinctly longer than wide, tip pointed). Anomaloglossus megacephalus can be distinguished from A. kaiei by (characters of A. kaiei in parentheses, see also Kok & Kalamandeen 2008) its larger size, female SVL max 28.3 mm [n= 3] in A. megacephalus (19.8 mm in A. kaiei, n= 25), fingers with keel-like lateral folds, best developed preaxially on Fingers II and III (fringes not folded), toes moderately webbed (basally webbed), dorsolateral stripe absent (present), oblique lateral stripe present (absent), throat in adult female blotched (immaculate). Anomaloglossus megacephalus can be distinguished from A. murisipanensis (La Marca, 1998) in having (characters of A. murisipanensis in parentheses, all based on the preserved holotype, which is a juvenile and the only known specimen) fingers with keel-like lateral folds, best developed preaxially on Fingers II and III (fringes barely detectable), more webbing on toes, symmetrical cloacal tubercles present (absent), tarsal keel weakly to distinctly curved, slightly tuberclelike (straight, not tuberclelike), oblique lateral stripe always present, even if broken in spots (not distinguishable), no white stripe between naris and eye in preservative (present). Anomaloglossus megacephalus can be distinguished from A. praderioi by (characters of A. praderioi in parentheses, see also Kok 2010) its larger size, female SVL max 28.3 mm (n= 3) in A. megacephalus (22.7 mm in A. praderioi, n= 1), Finger IV longer than I (equal), tip of Finger IV always surpassing the base of the distal subarticular tubercle on Finger III when fingers adpressed (barely reaching the base), toes moderately webbed with folded flaplike fringing (basally webbed, folded flaplike fringing not present on all toes), dorsolateral stripe absent (present), oblique lateral stripe present (absent), throat in adult female blotched (immaculate). Anomaloglossus megacephalus can be distinguished from A. roraima in having (characters of A. roraima in parentheses) fingers with keel-like lateral folds, best developed preaxially on Fingers II and III (fringes barely distinct, not distinctly folded), all toes with folded flaplike fringing (absent), webbing on toes moderate (absent). Anomaloglossus megacephalus can be distinguished from A. rufulus (Gorzula, 1990) in having (characters of A. rufulus in parentheses) webbing on toes moderate (rudimentary), and most remarkably by ventral colour pattern; light with a few dark brown blotches in A. megacephalus (dark brown marbled with white blotches in A. rufulus). The new species superficially most resembles A. parkerae, A. tepuyensis (with which it has been confused), and A. triunfo (Barrio-Amor��s, Fuentes-Ramos & Rivas-Fuenmayor, 2004), in sharing a similar dorsal pattern. It should be noted that A. triunfo is a possible synonym of A. tepuyensis (Kok & Barrio- Amor��s unpubl. data, see also comments by Myers & Donnelly 2008). Anomaloglossus parkerae is distinguished from A. tepuyensis and A. triunfo only by a few characters such as size, definition of dorsal pattern, and condition of finger and toe fringes (Myers & Donnelly 2008). Adults of A. megacephalus can notably be distinguished from those three similar taxa in having a more slender body (Anomaloglossus parkerae, A. tepuyensis and A. triunfo are distinctly stockier frogs), a comparatively longer snout and a larger, somewhat more massive head [See Fig. 5 for close comparison between A. megacephalus and A. tepuyensis and A. parkerae; compare also with Duellman���s (1997: 10) colour plate of A. parkerae, fig. 30 (showing A. parkerae) in L��tters et al. (2007), and figs. 13, 15 and 16 illustrating A. tepuyensis in Myers & Donnelly (2008); see also below for further comments], and in having symmetrical cloacal tubercles (absent in all females of A. parkerae, A. tepuyensis and A. triunfo examined). Direct comparison of specimens shows a difference in head size in Anomaloglossus megacephalus compared to the similar A. parkerae, A. tepuyensis and A. triunfo, as confirmed by plotting of morphometric proportions. Comparison of HL 3 with BEL indicates that the new species proportionally has a longer head than A. tepuyensis (Fig. 6). In order to distinguish between those similar species, the midline distance from the tip of the snout to an imaginary line between anterior insertions of upper arms is measured, and that distance is then extended posteriorly from the same imaginary line. In two of the three available A. megacephalus adult specimens the distance reaches or surpasses the anterior insertion of thigh, whereas in all adult specimens of A. parkerae, A. tepuyensis and A. triunfo examined (n = 20), the distance fails to reach the thigh insertion (see Fig. 5 B). It is noteworthy to mention that the only specimen of A. megacephalus in which HL 3 is not longer than BEL is in poor preservation state., Published as part of Kok, Philippe J. R., Macculloch, Ross D., Lathrop, Amy, Willaert, Bert & Bossuyt, Franky, 2010, A new species of Anomaloglossus (Anura: Aromobatidae) from the Pakaraima Mountains of Guyana, pp. 18-32 in Zootaxa 2660 on pages 26-28, DOI: 10.5281/zenodo.199004, {"references":["Kok, P. J. R., MacCulloch, R. D., Gaucher, P., Poelman, E. H., Bourne, G. R., Lathrop, A. & Lenglet, G. L. (2006 b) A new species of Colostethus (Anura, Dendrobatidae) from French Guiana with a redescription of Colostethus beebei (Noble, 1923) from its type locality. Phyllomedusa, 5, 43 - 66.","Kok, P. J. R. & Kalamandeen, M. (2008) Introduction to the taxonomy of the amphibians of Kaieteur National Park, Guyana. Abc Taxa, 5, 1 - 278.","Barrio-Amoros, C. L. (2006) A new dendrobatid frog (Anura: Dendrobatidae: Colostethus) from Aprada tepui, southern Venezuela. Zootaxa, 1110, 59 - 68.","La Marca, E. (1998 \" 1996 \") Ranas del genero Colostethus (Amphibia: Anura: Dendrobatidae) de la Guyana Venezolana con la descripcion de siete especies nuevas. Publicaciones de la Asociacion de Amigos de Donana, 9, 1 - 64.","Kok, P. J. R. (2010) A redescription of Anomaloglossus praderioi (La Marca, 1998) (Anura: Aromobatidae: Anomaloglossinae), with description of its tadpole and call. Papeis Avulsos de Zoologia, 50 (4), 51 - 68.","Gorzula, S. (1990 \" 1988 \") Una nueva especie de Dendrobates (Amphibia: Dendrobatidae) del Macizo de Chimanta, Estado Bolivar, Venezuela. Memoria de la Sociedad de Ciencias Naturales La Salle, 48, 143 - 149.","Barrio-Amoros, C. L., Fuentes-Ramos, O. & Rivas, G. (2004) Two new species of Colostethus (Anura: Dendrobatidae) from the Venezuelan Guayana. Salamandra, 40, 183 - 200.","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum - Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147.","Lotters, S., Jungfer, K. H., Henkel, W. F. & Schmidt, W. (2007) Poison Frogs: Biology, Species & Captive Husbandry. Chimaira, Frankfurt aim Main, Germany, 668 pp."]}

Chironius challenger sp. nov. (Figs. 1, 2 A���C, 3 A���D, 4 D, 6, 7; Tables 1, 2) Chironius fuscus (Linnaeus, 1758); M��gdefrau et al. (1991: 22 ���24, figs. 22 a, b) Chironius fuscus fuscus (Linnaeus, 1758) in part; Dixon et al. (1993: 114) Chironius fuscus (Linnaeus, 1758); Myers & Donnelly (2008: 121, fig. 73) Holotype. IRSNB 2659 (field number PK 2017), a subadult female collected by Philippe J. R. Kok, 25 November 2007 at 10 h 45, southeastern slope of Maringma Tepui, Cuyuni-Mazaruni District, Guyana (05�� 12 ��� N, 060�� 35 ��� W, 1500 m elevation). Referred specimens. BMNH 1976.235, adult female collected by Adrian Warren, 17 August 1974, southern side of Auyantepui, 2088 m elevation, Venezuela; ROM 42603, juvenile collected by Samir Khan, 2 November 2004, Mount Wokomung, 1400 m elevation, Guyana; MHNLS 16088, subadult female collected by Gilson Rivas, 8 February 2002, Auyantepui, 1700 m elevation, Venezuela. Etymology. The specific epithet is considered to be a noun in apposition and refers to Professor George Edward Challenger, the fictional main character of Sir Arthur Conan Doyle's famous novel "The Lost World" published in 1912. "The Lost World" depicts an epic expedition to an isolated tepui in the middle of the tropical jungle of northern South America in search of dinosaur-like creatures and a forgotten civilisation. The reading of Sir Conan Doyle���s novel when I was a child fuelled my thirst for exploration and adventure and positively influenced my research and interest in the Pantepui region. Definition and diagnosis. A species of the genus Chironius characterized by the following combination of characters: 10 dorsal scale rows at midbody, absence of apical pits and paravertebral keels in female (male unknown), 152���159 ventrals, 99���103 subcaudals, anal plate single, loreal as long as���or slightly longer than���high, 39���41 maxillary teeth. Only five of the 20 currently known Chironius species occur in the Guiana Shield (area sensu ��vila-Pires 2005, number of species f i de Hollis 2006): C. carinatus (Linnaeus, 1758), C. cochranae Hoge and Romano, 1969, C. exoletus (Linnaeus, 1758), C. fuscus (Linnaeus, 1758), and C. scurrulus (Wagler, 1824). Among these species, only C. fuscus and C. scurrulus have 10 DSR at midbody, a condition shared by C. challenger. All other species have 12 DSR at midbody. Chironius challenger is readily distinguished from C. scurrulus by the following characters (characters of C. scurrulus in parentheses): absence of apical pits (present, at least on neck), loreal as high as long or only slightly longer than high (distinctly longer than high), juveniles brownish with lighter crossbands (uniform green in life, blue-black in preservative), adults brownish with lighter crossbands (red or reddish brown in life, colouration variable in preservative, but never with lighter crossbands). Chironius challenger is mostly distinguished from C. fuscus by the following characters (characters of C. fuscus in parentheses): absence of apical pits (present, at least on neck), absence of paravertebral keels (usually present, even if faint in females), loreal as high as long or only slightly longer than high (distinctly longer than high), higher number of ventrals - minimum 152 in C. challenger (maximum 149 in C. fuscus from the Guiana Shield), shorter tail - 30���31 % TTL in C. challenger (34���37 % TTL in C. fuscus from the Guiana Shield), lower number of subcaudals - maximum 103 in C. challenger (minimum 118 in C. fuscus from the Guiana Shield), infralabials 9 (usually 10), ventrals in adults conspicuously mottled with darker edges (not heavily mottled, no conspicuous darker edge), skin between scales blue in life (white). Further comparisons are given in the Discussion. Description of the holotype. A subadult female (according to oviduct development), in good condition, TTL 720 mm, SVL 500 mm, TL 220 mm (30.6 % TTL); head well distinct from neck, narrow anteriorly, widest at temporal region, HW 54.8 % of HL; snout rounded in dorsal and lateral view, SL 6.2 mm; eye large (EL 4.8 mm), pupil round, EN 19.3 % HL; rostral large, broader than long (width about 1.8 times height), visible from above; internasals squarish, as broad as long, slightly smaller than prefrontals, laterally in contact with nasal, their medial suture asymmetrical, noticeably sinistral to the prefrontal medial suture; prefrontals slightly larger than internasals, width 1.5 times length, irregularly hexagonal, separated from eye by preocular and supraocular; frontal pentagonal, bell-shaped, longer than wide, 3.0 times length of medial prefrontal suture; parietals large, length 1.5 times maximal width, medial suture of parietals and length of frontal equal; 5 postcephalic scales. Nasal scale partially divided dorsally and ventrally, posterior part as long as, but slightly higher than, anterior part; preocular 1 / 1, separated from frontal by prefrontal-subocular contact; loreal slightly longer than high (height 93 % length), in contact with nasal, prefrontal, preocular, second and third supralabials, separated from orbit by preocular; suboculars absent; postocular 1 / 1; temporals 2 + 2 / 1 + 1, supralabials 9 / 9, sixth largest, first contacting nasal, second contacting nasal and loreal, third contacting loreal and in point contact with preocular, fourth, fifth, and sixth entering orbit, sixth in point contact with postocular and contacting anterior temporal, seventh and eighth contacting anterior temporal, ninth contacting anterior and posterior temporals; infralabials 9 / 9, first seven in contact with genials, first pair meeting at ventral midline, separating mental from genials; 2 pairs of genials, the first shorter. Maxillary teeth 39. Dorsal scales in 10 - 10 - 10 rows, smooth without apical pits, a few weak anal ridges laterally above cloacal region; ventrals 159; caudal scale row reduction from 6 to 4 rows occurs at the 5 th subcaudal; anal single; divided subcaudals 103 (Figs. 1, 2 A���C, 3 A, 4 D). Colour of the holotype in life. Dorsolateral body ground colour light brown, darkening posteriorly, with a series of ca. 31 ill-defined greyish brown transverse bands (1, rarely 2 scales wide) outlined in black. Tail darker brown without conspicuous transverse bands. Dorsal scales with black edges, sometimes slightly peppered with dark brown or black; some dorsal scales yellow anteriorly, especially on anterior body and on lower flank. Skin between scales sky blue. Head light brown, supralabials yellow, with labials 2���3 having dark posterior edges; black postocular stripe from eye across lower part of primary temporal and upper part of ultimate supralabial reaching postcephalic scales then breaking into blotches on the lateral side of ventrals; iris dark copper; tongue black. Underside of head immaculate yellowish white; ventrals immaculate yellow until ca. the 36 th, then posteriorly bordered by black, mottled with brown and black spots increasing in number posteriorly, becoming almost completely black near the vent; subcaudals similar in colour and pattern, but less heavily mottled by black than the posteriormost ventral scales (Figs. 1 A���B, 4 D). Colour of the holotype in preservative. Pattern essentially as in life, but dorsal colours faded to grey and light grey, and yellow and sky blue faded to white (Figs. 1 C���D, 2 A���C, 3 A, 6). Variation. Three referred specimens are available, consisting of two females and a juvenile. The females vary in TTL from 720���937 mm (315 mm in the juvenile), in SVL from 450���658 mm (222 mm in the juvenile), and in TL from 201���279 mm (93 mm in the juvenile). Tail length is 42���45 % SVL. The loreal is always high, its height being 93 (in the holotype)��� 100 % (in all other specimens) its length. There is little variation in scale counts. Weak anal ridges laterally above the cloacal region are absent in one female (MHNLS 16088) and the juvenile (ROM 42603). Ventrals range from 152���159, divided subcaudals 99���103 (65 % number of ventrals), supralabials 8���9, number of infralabials in contact with genials 5���7, postcephalics 4���6. Although the holotype has only 1 postocular on both sides, all other individuals have 2. The temporal formula is quite variable and only one specimen (BMNH 1976.235) has 1 + 1 temporal on both sides. Other specimens have 1 + 1 / 1 + 2 (MHNLS 16088), 1 + 2 / 1 + 2 (ROM 42603) or 2 + 2 / 1 + 1 (holotype) temporals. Paravertebral scales are smooth and apical pits are absent in all specimens. Caudal scale row reduction from 6 to 4 rows occurs at the 5���7 th subcaudals in females, at the 10 th subcaudal in the juvenile. Maxillary teeth are 39���41 in females, 41 in the juvenile. See Table 1 for measurements and scalation for the holotype and the referred specimens. Colour and pattern are similar between the referred specimens and the holotype. Intensity of ventral mottling seems related to age and size, the two largest individuals having the most mottling on ventrals. Transverse bands are much more conspicuous and more numerous (ca. 55) in the juvenile specimen. Larger specimens mostly lack transverse bands on the tail whereas these bands are conspicuous in smaller individuals. Distribution and ecology. Based on museum specimens, Chironius challenger is known between 1400��� 2088 m elevation from three localities in the eastern Pantepui region: Auyantepui in Venezuela, Maringma Tepui and Mount Wokomung in Guyana (Fig. 5). An additional specimen collected at 1520 m elevation on the summit of Guaiquinima Tepui, Venezuela (Fig. 5) is illustrated in M��gdefrau et al. (1991: 22, 23) under C. fuscus (see Discussion and additional comparisons). The holotype was collected in the morning while it was moving in a small tree about 200 cm above the ground in low evergreen upper-montane forest. ROM 42603 was collected while moving on the ground on the northeastern slope of Mount Wokomung, also in low evergreen upper-montane forest. No data on microhabitat is available for the two other referred specimens, which are presumably from low evergreen upper-montane or high-tepui forest according to the collection localities. The specimen reported by M��gdefrau et al. (1991) from Guaiquinima Tepui was collected in the afternoon, on a rock on the bank of a stream dissecting a rocky area covered by Stegolepis (Rapateaceae) and small patches of forest (presumably high-tepui forest given the elevation). The holotype and MHNLS 16088 have few enlarged ovarian follicles (largest 3.3 and 1.9 mm in length, respectively), and BMNH 1976.235 has three eggs (11.6, 13.6, and 17.1 mm in length). Sex Female Female Female Juvenile Locality Maringma Tepui, Auyantepui, Auyantepui, Mt. Wokomung, Guyana Venezuela Venezuela Guyana, Published as part of Kok, Philippe J. R., 2010, A new species of Chironius Fitzinger, 1826 (Squamata: Colubridae) from the Pantepui region, northeastern South America, pp. 31-44 in Zootaxa 2611 on pages 32-40, DOI: 10.5281/zenodo.197801, {"references":["Magdefrau, H., Magdefrau, K. & Schluter, A. (1991) Herpetologische Daten vom Guaiquinima-Tepui, Venezuela. Herpetofauna, 13 (70), 13 - 26.","Dixon, J. R., Wiest, Jr, J. A. & Cei, J. M. (1993) Revision of the Neotropical Snake Genus Chironius Fitzinger (Serpentes, Colubridae). Monografie XIII. Museo Regionale di Scienze Naturali, Torino, Italy, 279 pp.","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum - Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147.","Avila-Pires, T. C. S. (2005) Reptiles. In: Hollowell, T., and Reynolds, R. P. (Eds.), Checklist of the terrestrial vertebrates of the Guiana Shield. Bulletin of the Biological Society of Washington, 13, pp. 24 - 40.","Hollis, J. L. (2006) Phylogenetics of the genus Chironius Fitzinger, 1826 (Serpentes, Colubridae) based on morphology. Herpetologica, 62 (4), 435 - 453."]}

Pantepuisaurus genus novum Type species. Pantepuisaurus rodriguesi species novum Etymology. A noun in apposition, derived from “Pantepui” referring to the phytogeographic province where the type species was discovered, and the Greek sauros meaning “lizard”. Gender masculine. Content. Pantepuisaurus rodriguesi species novum; monotypic. Diagnosis. An elongate gymnophthalmid (known SVL in male 58.3 mm) with a long tail (1.6 x SVL) and the unique following combination of characters: (1) distinctive ear opening and moveable eyelids, (2) limbs pentadactyl with all digits clawed, (3) nasal scales separated by a single frontonasal, (4) prefrontals present, (5) frontoparietals, parietals and interparietal present, (6) parietals and interparietal longer than wide, (7) interparietal and parietals forming a jagged, irregular posterior margin, (8) occipitals present, (9) three pairs of genials, second pair in contact with only one infralabial, (10) enlarged median pairs of gulars, (11) dorsal scales hexagonal, keeled, in transverse rows only, (12) ventral scales imbricate, hexagonal, mucronate, broadly keeled, in transverse rows only, (13) tongue anterodorsally covered by oblique, anteriorly converging plicae, posterodorsally covered by large scalelike papillae (14) hemipenis weakly bilobed with series of curved transverse plicae bearing mineralized spicules. Pantepuisaurus differs from members of Alopoglossinae in having the tongue anterodorsally covered by oblique plicae and posterodorsally covered by large scalelike papillae (entirely covered by oblique plicae in Alopoglossinae), the interparietal and the parietals forming a jagged, irregular line (straight posterior margin), and in having ventral scales in transverse rows only (in transverse and longitudinal rows in Alopoglossinae); from members of Gymnophthalminae in having hands pentadactyl with the first finger clawed (first finger absent, reduced and/or clawless in Gymnophthalminae); from Rhachisaurinae in having external ear openings (absent in Rhachisaurinae). Within Cercosaurinae (sensu Pellegrino et al. 2001 and Rodrigues et al. 2005), Pantepuisaurus differs from all genera except Ecpleopus and Leposoma of the scincoides group in having hexagonal, mucronate ventral scales arranged in transverse rows only. Pantepuisaurus is readily distinguished from Ecpleopus (characters of Ecpleopus in parentheses) notably by the presence of a broad keel on ventrals (absent), quadrangular gular scales (mucronate), complete superciliary series (incomplete), and the presence of femoral pores in males (absent). Pantepuisaurus is quickly separated from Leposoma of the scincoides group (characters of Leposoma in parentheses) in having quadrangular gular scales (mucronate), by the absence of distinct striations on head scales (present), the interparietal and the parietals forming a jagged, irregular line (round posterior margin), and the presence of occipitals (absent). Two additional gymnophthalmid genera, Adercosaurus and Kaieteurosaurus, have not been assigned to any subfamily yet, but might be related each other and with Pantepuisaurus. Pantepuisaurus is immediately distinguished from Adercosaurus (characters of Adercosaurus in parentheses) in having imbricate, hexagonal, mucronate, broadly keeled ventral scales in transverse rows only (smooth, in transverse and longitudinal rows), in having a highly reduced fourth infralabial (not reduced) and the second pair of genials partly separated from the infralabials and in contact with one infralabial only (not separated from the infralabials, in contact with two infralabials), in lacking oblique plicae at the rear of the tongue, which is posteriorly covered by large scalelike papillae (rear of tongue covered by oblique plicae), and in having mineralized spicules on hemipenes’ plicae (absent). Pantepuisaurus is most similar to Kaieteurosaurus in having hexagonal, mucronate ventral scales arranged in transverse rows only, but it is readily distinguished from it (characters of Kaieteurosaurus in parentheses) by the presence of prefrontal scales (absent), three pairs of genials (two), in having a highly reduced fourth infralabial (not reduced) and the second pair of genials partly separated from the infralabials and in contact with one infralabial only (not separated from the infralabials, in contact with two infralabials), the common suture of the interparietal and parietals forming a jagged, irregular line (more or less straight posterior margin), an entire nasal (divided), and the aspect of the ventral scales that are much less lanceolate and mucronate than in Kaieteurosaurus. It should be emphasized here that Kaieteurosaurus also has broad “keels” on ventral scales (i.e. the central part of the scale is broadly elevated), a character that was only noticed while closely comparing the two genera. This character is not conspicuous in all Kaieteurosaurus specimens and is mostly noticeable on anteriormost and posteriormost ventral scales. In Kaieteurosaurus the significant lateral extension of the flat “keel” gives a lanceolate, smooth, “unkeeled” aspect to the ventral scales as described by Kok (2005), while the keel is more prominent and conspicuous in Pantepuisaurus. Figure 4 schematically illustrates the difference in the aspect of the broad keel on ventral scales between Pantepuisaurus and Kaieteurosaurus. A summary of main diagnostic characters for Pantepuisaurus, Adercosaurus and Kaieteurosaurus is in Table 1. Distribution. The genus is currently known only from the type locality, the summit of Maringma tepui in Guyana (Fig. 1). Taxonomic comments. No less than eight new gymnophthalmid genera have been described (Myers & Donnelly 2001, Doan & Castoe 2005, Kok 2005, Rodrigues et al. 2005, 2007, Rodrigues & Maranhão dos Santos 2008) since the molecular-based phylogenetic hypothesis of Pellegrino et al. (2001), who demonstrated the monophyly of Gymnophthalmidae. Some higher-level taxonomic changes are still in debate (see Castoe et al. 2004, Rodrigues et al. 2007) and refinement of higher-level taxonomy of gymnophthalmids is still necessary. Morphologically, the new genus shows some similarities with Adercosaurus and Kaieteurosaurus. Giving the low number of specimens available, any speculation on the relationships between Adercosaurus (known only from the holotype), Kaieteurosaurus (known only from three specimens), and Pantepuisaurus (known only from the holotype) seems premature. More specimens of these three genera are clearly needed to elucidate relationships between them. However, preliminary molecular analysis and comparisons do indicate affinities between Pantepuisaurus and Kaieteurosaurus (unfortunately, molecular data on Adercosaurus are not available), but reveal significant genetic divergences between these two genera that give further support to the erection of a new genus. Molecular analysis also suggests assignment of Pantepuisaurus and Kaieteurosaurus to the tribe Ecpleopini (Cercosaurinae) (M. T. Rodrigues, pers. comm.). A more “conservative” attitude would have been to accommodate the new species in an already known monotypic genus sharing some morphological similarities (i.e. Adercosaurus or Kaieteurosaurus). However this would have given rise to two important problems: (1) assignment of the new species to either of these two genera would have been arbitrary and poorly supported because the new species differs from both of them in a combination of important characteristics currently used to diagnose gymnophthalmid genera (e.g. condition of the posterior margin of the interparietal and parietals, condition of ventral scales, number of genials, presence of prefrontals, hemipenial morphology, tongue morphology) (2) because the new species differs from these two genera in morphological characteristics used to separate other genera (especially in the tribe Ecpleopini), placing the new species in one of them would have rendered the taxonomic status of these and other genera dubious by suggesting that these characters are variable intragenerically (thus making taxonomy of the group more complicated instead of simplifying it). One could suggest that Kaieteurosaurus is a synonym of Adercosaurus, however this is not supported by any evidence, these two genera being differentiated on the basis of many important diagnostic characters (see Kok 2005).

Pantepuisaurus rodriguesi species novum Figs. 5���9 Holotype. IRSNB 2650 (field number PK 2044), an adult male collected by Paul Benjamin, Philippe J. R. Kok and Claudius Perry, 28 November 2007 at 14 h 55, summit plateau of Mount Maringma, Cuyuni-Mazaruni District, Guyana (05�� 12 ��� 57 ���N, 060�� 35 ��� 07���W, 2080 m). Etymology. The specific epithet is a noun in the genitive case, honoring Brazilian herpetologist Miguel Trefaut Rodrigues (Universidade de S��o Paulo, S��o Paulo, Brazil) for his huge contribution to the knowledge of the family Gymnophthalmidae using both morphological and molecular approaches. Diagnosis. In addition to the generic characteristics, the new species is also characterized by the following features: four supraoculars; 15 smooth temporal scales; scales around midbody 32; dorsal scales sharply keeled, in 33 transverse rows; ventral scales in 20 transverse rows; trunk length 2.6 x length of forelimb; dorsum and flanks completely black, iris red. Description of the Holotype. An adult male 58.3 mm SVL. Head length 19 % SVL, 1.38 x head width, head width 1.45 x head height. Head slightly wider than neck, temporal region not distinctly swollen, neck approximately as wide as anterior body. Snout blunt. Body oval in transverse section, slightly depressed, slightly wider than high. Tail complete, 93.3 mm, 1.60 x SVL, circular in cross-section, tapering toward the tip. Limbs well-developed, forelimbs 19 % SVL, hindlimbs 29 % SVL. Limbs pentadactyl, all fingers and toes with terminal claw. Limbs short, not overlapping when adpressed along the body. Distance between forelimbs and hindlimbs (trunk length) = 29.2 mm, 50 % SVL, 2.6 x forelimbs (Fig. 5). Tongue lanceolate, anterodorsally covered by oblique (chevron-shaped), anteriorly converging plicae, posterodorsally covered by large scalelike papillae; tip bifid, smooth; one pair of infralingual plicae after fork. Anterior teeth conical, most of posterior ones bicuspid, a few of them tricuspid. Rostral roughly hexagonal, twice as wide as high, visible from above, in broad contact with frontonasal. Frontonasal single, heptagonal, laterally in contact with nasal and loreal, not in contact with first supraocular. Prefrontals roughly pentagonal, as broad as long, with a moderately long medial suture, laterally in broad contact with first supraocular and the loreal, touching the anterior point of the second supraocular on the right, separated from it by first supraocular-frontal contact on the left. Frontal hexagonal, 1.4 x as long as wide, anteriorly as wide as posteriorly, laterally markedly concave; frontal laterally in contact over the entire length of second supraocular, touching the first supraocular on the left, separated from it by prefrontal-second supraocular contact on the right, in contact with third supraocular. Frontoparietals pentagonal, 1.3 x as long as wide with moderately long medial suture; each frontoparietal in contact with frontal, distinctly separated from second supraocular by frontal-third supraocular contact, in broad contact with third supraocular, parietal and interparietal, and narrow contact with the fourth supraocular. Interparietal heptagonal, posteriorly projecting, 1.7 x as long as wide, with parallel sides, distinctly longer than parietals. Interparietal and parietals form a jagged, irregular, posterior margin. A row of five smooth occipitals of various shapes, shorter and wider than dorsal neck scales, the central smallest. Four supraoculars, first smallest, about 3 / 4 of the fourth; second and third larger, subequal. Five superciliaries, first much larger than the others, with a short incomplete suture running dorsad from ciliaries on the right side and a similar very short suture on the left side; second and third smallest, subequal, fourth distinctly larger, about twice the size of the fifth (Fig. 6). Nasal entire, with two short divisions running ventrad from naris to supralabial on the left side, these two short divisions are incomplete, running from the first supralabial, on the right side; naris just below centre, directed laterally. Loreal quadrangular, slightly anteriorly inclined, in contact with nasal, frenocular, one preocular, first superciliary, first supraocular, prefrontal and frontonasal, separated from supralabials by frenocular-nasal contact. Two small preoculars between first superciliary and first subocular. Frenocular roughly quadrangular, longer than wide, about half-size the loreal, in contact with first supralabial. Four suboculars. Two postoculars, the lower smallest. Seven supralabials on both sides, fifth highest, fourth below middle of the eye; a very short, inconspicuous division on the first supralabial on the right side (running ventrad from nasal); suture between second and third supralabials in contact with first subocular, suture between first and second supralabials in contact with frenocular (Fig. 6). Upper eyelid with three ciliaries on both sides, second very large, apparently resulting from the fusion of several smaller scales. Ocular recess with median row of three scales separating median ciliary from superciliaries. Lower eyelid with a semi-transparent, slightly pigmented, palpebral disc of two long vertical palpebral scales (Fig. 6). Temporal region with 15 irregularly shaped, juxtaposed, smooth scales on both sides. Temporals arranged in four oblique rows. Ear opening moderately large, vertically semi-circular with a straight posterior margin, bordered by a few small, juxtaposed, smooth scales. Auditory meatus moderately deep, tympanum mostly transparent, slightly pigmented (Fig. 6). Mental trapezoidal, its posterior edge almost straight, slightly concave. Postmental large, heptagonal, laterally in contact with first and second infralabials. Three pairs of large genials, first two pairs in contact with each other medially. First pair in lateral contact with second and third infralabials; second pair in lateral contact with third infralabial only, separated from fourth and fifth infralabials by a large sublabial scale, medial suture of second pair slightly longer than that of first pair; third pair in lateral contact with fifth infralabial, separated medially by two small scales. A pair of enlarged postgenials separated medially by scales of varying sizes and widely separated from infralabials. Seven infralabials on the right side, six on the left side, fourth scale highly reduced on both sides. One row of imbricate, smooth pregulars with rounded posterior margins. Five rows of gulars, four of them with two transversely enlarged scales. Collar row with seven scales, central one distinctly larger, others decreasing in size laterally, forming a slightly scalloped margin. Gular fold distinct, concealing one row of very small scales (Figs. 6, 7 B). Head scales with numerous small pits, especially concentrated on rostral, frontonasal, supraoculars, and lateral head scales, also present on ventral head scales, and body and limbs scales. Scales on nape posterior to occipitals longer than wide, with a broad, flat keel, mucronate, in transverse rows, grading posteriorly into dorsal body scales. Sides of neck with medium-sized, roughly subequal, smooth, juxtaposed scales in about eight oblique rows, not distinctly decreasing in size posteriorly but interrupted by a row of smaller scales 2 / 3 the way before arm insertion, which weakly form annuli with nape and gular scales. Dorsals imbricate, pentagonal, some appearing hexagonal because of imbrication, elongate, sharply keeled, mucronate, except very few isolated scales. Dorsal scales in transverse rows only, 33 rows between the interparietal and the posterior edge of hindlimbs (including row of occipitals). Middorsal scales not distinctly narrower than adjacent dorsals. A zone of small juxtaposed scales before arm insertion. Flank scales similar to, but slightly narrower than, dorsal scales, most of flank scales are not mucronate but have a straight or slightly rounded posterior margin; no abrupt demarcation between dorsals and laterals. Lateral scales becoming slightly smaller just before leg insertion. No row of ventrolateral scales (see Myers & Donnelly 2008) can be detected; some very small, longitudinally oval or rounded scales intercalated in the interstitial skin between the transverse rows of lateral; these small scales are more conspicuous along the first anterior third of flank and before leg insertion. All transverse rows of dorsals correspond to one row of laterals, except at midbody on the right side where one transverse row of dorsals corresponds to two rows of laterals, and before leg insertion and after arm insertion. Ventrals imbricate, broadly keeled, longer than wide, mucronate except very few isolated scales, in transverse rows only (Fig. 7 A). Most transverse rows of ventrals correspond to a transverse row of laterals and of dorsals (exceptions occur at midbody). Twenty transverse rows between collar and preanal plate, 32 scales around midbody. Preanal plate consisting of six scales, one anterior, five posterior scales, most lateral posterior scales narrow, distinctly smaller than adjacent scales (Fig. 8). Four preanal pores. Four femoral pores on the right side, three on the left side, each femoral pore between three scales. Scales on tail keeled, not mucronate (except proximally), slightly imbricate, roughly rectangular, as long as dorsal scales, in transverse rows only, in uninterrupted annuli. Pigmented subcaudal scales keeled, slightly imbricate, slightly mucronate, roughly rectangular, in transverse rows only. Dorsal surfaces of upper and lower arms with smooth, variably polygonal scales, ventral surface of upper arms with small, juxtaposed scales, ventral surface of lower arms with scales similar to those on dorsal surface, except at the base of hands where scales are juxtaposed and distinctly smaller. Thighs with a row of enlarged, smooth, roughly hexagonal scales on anterior surface, bordered ventrally by two rows of smaller polygonal scales diminishing in size ventrad (scales of first row smooth, scales of second row broadly keeled); bordered dorsally by 2���3 rows of keeled scales diminishing in size dorsad, grading into small, smooth (but with many very small asperities and pits that give a rugose aspect), slightly granular, juxtaposed scales, which cover posterodorsal and posterior surfaces of thighs; these scales increase in size ventrally. Shanks ventrally and posteriorly with large, smooth, polygonal, imbricate scales. Scales on upper and anterior surfaces of shanks slightly smaller, imbricate, rugose (see above), and strongly keeled. Dorsal surface of hands and digits with imbricate, smooth scales, dorsal surface of feet with imbricate, broadly keeled scales, except a few anterodorsally; palm and soles covered with small, smooth, slightly protuberant, juxtaposed, polygonal scales. Two enlarged, subequal, thenar scales on inner margin of palm below pollex, each with produced inner edge, similar scales occur at the base of Toes I and III���IV; two slightly enlarged hypothenar scales on outer margin of palm. Subdigital lamellae divided basally and single on distal halves on most digits. Subdigital lamellae as follows (Roman numbers indicate digits, Arabic numbers indicate paired or single lamellae on left/right; lower ungual sheath scale is omitted from the counts): hands I 4 / 4, II 7 / 7, III 9 / 9, IV 10 / 11, V 6 / 7; feet I 5 / 5, II 8 / 8, III 11 / 10, IV 16 / 16, V 9 / 8. Color of Holotype in life. Dorsal and lateral surfaces completely black; closer examination shows that black scales on body and neck are peppered with brown or reddish brown. Upper surface of head black. Tympanum light pink. Arms, legs and tail black. Iris red (Fig. 5). Tongue dark gray in its anterior two-thirds, whitish with gray flecks on its posterior third. Palms and soles black. Underside of head and throat black with irregular light gray markings; venter orangish brown, ventral scales peppered with black; underside of upper legs black; underside of lower legs black with irregular orangish brown markings; underside of tail black, except proximally where some scales are white (Figs. 7���8). Color of Holotype in preservative. After four months in 70 % ethanol, the holotype became very dark brown, the light gray markings on the underside of head and chest are white; venter is whitish peppered with dark gray; tympanum is transparent, no other noticeable change. Hemipenis. The hemipenes of the holotype were everted in the field. The left organ was subsequently removed, sent to and prepared by Pedro Nunes (Universidade de S��o Paulo, S��o Paulo, Brazil), currently working on a PhD thesis on the hemipenial morphology in the family Gymnophthalmidae. The hemipenis was inflated with blue petroleum jelly and the spicules were stained with Alizarin Red (Fig. 9). The left hemipenis is 6.9 mm long, with a greatest width of 4.5 mm. The organ is weakly bilobed, with each of the short divisions terminating in 2���3 protuberances. Sulcus spermaticus medial, broad, poorly defined. Asulcate surface with a conspicuous series of 15���18 curved transverse plicae (tissue ridges) encircling the side of the organ and terminating laterally on the sulcate surface, most plicae being laterally shortly interrupted by a narrow nude space. A wide nude space between the ca. 8���9 distal plicae on the asulcate surface. All plicae bear mineralized spicules; plicae located on the proximal part of the hemipenis bear a single spicule, the number of spicules is increasing from the proximal part towards the tip resulting in plicae bearing a series of several uniform spicules interrupted by a larger central one (Fig. 9). Distribution and ecology. The species is currently known only from the type locality, the summit of Maringma tepui in Guyana (Fig. 1). The only specimen available was collected in the afternoon (14 h 55), during a sunny spell, in a sunny spot, on the ground, among dense vegetation (Fig. 3). No other specimen was observed. The only other lizard species recorded from the summit of Mount Maringma is Arthrosaura hoogmoedi Kok, 2008., Published as part of Kok, Philippe J. R., 2009, Lizard in the clouds: a new highland genus and species of Gymnophthalmidae (Reptilia: Squamata) from Maringma tepui, western Guyana, pp. 53-67 in Zootaxa 1992 on pages 57-66, DOI: 10.5281/zenodo.185553, {"references":["Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum - Terramar expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147.","Kok, P. J. R. (2008) A new highland species of Arthrosaura Boulenger, 1885 (Squamata: Gymnophthalmidae) from Maringma tepui on the border of Guyana and Brazil. Zootaxa, 1909, 1 - 15."]}

Oreophrynella seegobini sp. nov. Figs. 1���3 Holotype. IRSNB 1979 (field number PK 2052), an adult male collected by Philippe J. R. Kok, Paul Benjamin and Claudius Perry, 29 November 2007 at 16 h 45, summit plateau of Mount Maringma, Cuyuni- Mazaruni District, Guyana (05�� 12 ��� 59 ���N, 060�� 35 ��� 05���W, 2088 m elevation). Paratype. IRSNB 1980 (field number PK 2053), an adult male with same data as holotype. Etymology. It is my pleasure to name this species in honour of my friend Giuliano ���Kinky��� Seegobin to acknowledge him for his friendship, hospitality, and always enthusiastic help during field work in Kaieteur National Park, Guyana. Kinky (as nicknamed by his friends and diamond miner colleagues) developed a keen interest in amphibians and reptiles and is continuously ready for a collecting trip, even in the middle of the night. He unfortunately cancelled the Maringma expedition due to a health issue. Diagnosis. The new species is assigned to the genus Oreophrynella because of the following suite of characters: habitus bufoniform, parotoid glands absent, adult size less than 38 mm, Toes I���II opposed to III���V, thick skin between digits, skin tuberculate, tympanum absent. No other anuran in northern South America shares this combination of characters. In addition to the aforementioned diagnostic characters, the following features characterize the new species: (1) small size (up to 20.6 mm SVL in male, female unknown); (2) frontoparietal crests indistinct; (3) postorbital crests prominent; (4) prominent short suborbital crests; (5) dorsal skin minutely spiculate with scattered medium to large, elevated, oval and round tubercles; (6) ventral skin anteriorly rugose with few large, flat, round granules, posteriorly tuberculate; (7) webbing on hand and foot well-developed; (8) adult dorsal colour in life blackish brown; (9) adult ventral colour in life dark brownish orange; (10) one vocal slit present on the left or on the right side of the floor of the buccal cavity. Oreophrynella seegobini is immediately distinguished from all its known congeners, except O. weiassipuensis Se��aris, DoNascimiento and Villarreal, 2005 by indistinct frontoparietal crests and prominent postorbital crests. In addition to the cephalic crests condition, Oreophrynella seegobini further differs from O. dendronastes Lathrop and MacCulloch, 2007 and O. macconnelli by well-developed webbing (basal in O. dendronastes and O. macconnelli), blackish brown dorsal colour in life (light brownish orange to olive brown in O. dendronastes and O. macconnelli) and dark ventral colour in preservative (creamy yellow in O. dendronastes and O. macconnelli); from O. nigra Se��aris, Ayarzag��ena and Gorzula, 1994 by well-developed webbing (moderate in O. nigra), low density of large tubercles (high density of small closely-set tubercles in O. nigra) and dark brownish orange belly in life (blackish brown in O. nigra); from O. quelchii by well-developed webbing (moderate in O. quelchii), low density of large tubercles (high density of tubercles in O. quelchii) and dark brownish orange belly in life (bright orange with black mottling in O. quelchii); from O. vasquezi Se��aris, Ayarzag��ena and Gorzula, 1994 by well-developed webbing (moderate in O. vasquezi), low density of large tubercles (high density of closely-set tubercles in O. vasquezi) and dark flanks in life and in preservative (distinctly lighter than dorsum in O. vasquezi); from O. cryptica Se��aris, 1995 by well-developed webbing (basal to moderate in O. cryptica), blackish brown dorsal colour in life (dark reddish brown in O. cryptica), and dark ventral colour in preservative (creamy white, often with a dark patch in the centre of the belly in O. cryptica); and from O. huberi Diego-Aransay and Gorzula, 1990 by well-developed webbing (basal to moderate in O. huberi), blackish brown dorsal colour in life (rufous-orange in O. huberi), and dark ventral colour in preservative (creamy white in O. huberi). The new species is distinguished from Oreophrynella weiassipuensis by [characters of O. weiassipuensis according to Se��aris et al. (2005) are in parentheses] having more prominent postorbital crests that are straight or slightly anteriorly concave (less developed, posteriorly concave), short prominent suborbital crest (indistinct), prominent canthal ridge extending roughly from tip of snout to halfway between nostril and upper eyelid (low), eye-nostril distance 79���86 % of eye diameter (59 %), larger and more prominent dorsal tubercles, ventral skin anteriorly rugose with large, flat, round granules, posteriorly tuberculate (granular with few small tubercles), a blackish brown dorsal colour without mid-dorsal line (reddish brown with darker marks and a thin dark brown mid-dorsal line), a dark brownish orange ventral colour with a black mid-ventral line (light reddish brown without black mid-ventral line), and orange palm and sole (light brown). Description of the Holotype. Adult male (Figs. 1 A���D, 2 A, C), head slightly wider than long, HL 34 % SVL, EN 86 % EYE. Snout with a small fleshy conical projection on the tip, acuminate in profile, projecting beyond the level of the lower jaw, SL 1.4 x EYE; loreal region smooth, vertical, slightly concave; canthus rostralis distinct, angular, with a prominent canthal ridge extending roughly from tip of snout to halfway between nostril and upper eyelid; nostrils protuberant directed laterally; internarial area highly concave; IND 117 % EN; IOD 0.8 x SL, 31 % HW, with a few medium-sized tubercles; frontoparietal crests indistinct. Temporal region vertical, postorbital crest well-developed, prominent, very sharp, anteriorly inclined, almost straight, slightly concave anteriorly; tympanum absent; sharp subocular crest well visible, shorter than eye length, not connecting with postorbital crest; tongue lanceolate, wider posteriorly, 35 % longer than wide, attached anteriorly, posterior quarter free. One vocal slit on the left side (absent on the right side), short, lateral. Premaxillary and maxillary teeth absent; choanae small, round; odontophores and vomerine teeth absent. Forelimbs long, slender; axillary membrane absent; hands moderately large, relative finger lengths III>II=IV>I, fingers flattened, tips not expanded; palm, fingers and webbing covered by numerous small, round, supernumerary tubercles; subarticular tubercles larger than surrounding supernumerary tubercles; thenar and palmar tubercles conspicuous, prominent, thenar tubercle at base of first finger, ovoid, subequal to round palmar tubercle; webbing well-developed (Fig. 3 A). Hindlimbs long, slender, TIB 39 % SVL, femur slightly longer than tibia (FEM 104 % TIB), tibiotarsal articulation extending to jaw articulation when hindlimbs adpressed along body; feet moderately large; FOOT 84 % TIB; Toes I���II opposed to III���V, relative toe length IV>I>V>II=III, toes flattened, tips not expanded; sole, toes and webbing covered by numerous small, round, supernumerary tubercles; subarticular tubercles larger than surrounding supernumerary tubercles; inner and outer metatarsal tubercles small, of size similar to subarticular tubercles at the base of Toes II���V, inner metatarsal tubercle slightly larger than outer one; webbing well-developed (Fig. 3 B). Cloacal opening directed ventrally at mid-level of thighs, covered by a fleshy sheath. Dorsal skin minutely spiculate with scattered medium to large oval and round, conspicuously elevated tubercles; top of head with a few medium-sized to large tubercles, upper eyelid covered with smaller tubercles, those close to the external margin aligned in a more or less distinct row; limbs densely covered by medium-sized to large round tubercles (Fig. 1 A���C). Ventral skin anteriorly rugose with few scattered large, flat, round granules, posteriorly tuberculate (Fig. 1 D). Measurements of the Holotype (mm). SVL 20.0; HL 6.8; HW 7.8; SL 3.0; EN 1.8; EYE 2.1; IND 2.1; IOD 2.4; HAND 4.7; FEM 8.1; TIB 7.8; FOOT 6.6. Colour in life. Dorsum, top of head and flanks blackish brown, lower arm, hindlimbs and top of hands and feet slightly lighter than dorsum. A bright yellowish orange spot at arm insertion on the left side, lacking on the right side where the area is dark brownish orange. Upper eyelids and lateral surfaces of snout dark brownish orange (Fig. 1 A���C). Throat, chest and anterior part of belly dark brownish orange, posterior part of belly and area below vent blackish brown. A black mid-ventral line running from tip of lower jaw to posterior part of belly, slightly curved sinistrally. Ventral surface of limbs brownish orange. Palms and soles orange (Fig. 1 D). Iris dark greenish brown with inconspicuous dark grey streaks. Colour in preservative. Dorsal surfaces very dark brown, dorsal surfaces of hands and foot slightly lighter (Fig. 2 A). Ventral surfaces dark greyish brown, with a black patch below vent. Black mid-ventral line running from tip of lower jaw to posterior part of belly more conspicuous than in life. Palms and soles light grey (Fig. 2 C). Variation. The paratype is very similar to the holotype, except as follows: EN 79 % EYE; SL 0.8 x EYE; IND 105 % EN; IOD 1.3 x SL, 29 % HW; TIB 38 % SVL; FEM 108 % TIB; FOOT 86 % TIB; inner metatarsal tubercle proportionally larger and more conspicuous, outer indistinct; subocular crest longer and more conspicuous; tongue proportionally wider posteriorly; one vocal slit on the right side (vs. on the left side in the holotype); dorsal colour slightly darker, absence of yellowish orange spot at arm insertion (Figs. 1 E, 2 B); ventral surface of hindlimbs dark brown (instead of brownish orange); mid-ventral line slightly curved dextrally (Figs. 1 F, 2 D). Measurements of the Paratype (mm). SVL 20.6; HL 7.0; HW 7.9; SL 3.0; EN 1.9; EYE 2.4; IND 2.0; IOD 2.3; HAND 4.6; FEM 8.4; TIB 7.8; FOOT 6.7. Distribution and ecology. Oreophrynella seegobini is currently known only from the type locality, the summit of Maringma tepui in Guyana (Figs. 4 A ���B). Both specimens were collected on the same day at the same time (16 h 45), ca. 1.0 m from each other. The holotype was walking slowly on the muddy ground, while the paratype was found hidden under a rotting bromeliad leaf. Habitat consists of tepui scrub (Fig. 5, see Kok 2008 for details on the vegetation of the summit of Maringma tepui). Both specimens emitted a soft ���peep��� when handled, a similar sound that was heard around the camp in early morning and before sunset. It is thus probable that the call of the species is a soft ���peep-peep-peep��� similar to the call of other high elevation Oreophrynella species (McDiarmid & Gorzula 1989, D. B. Means, pers. comm.). The species might be more abundant than expected, but difficult to find due to minute size and cryptic colouration. More colour photographs of the habitat of the species are in Kok (2008). The only other amphibian species collected during our stay on the summit of Maringma tepui (25���30 November 2007) are Anomaloglossus cf. roraima, A. sp, and Hypsiboas sibleszi., Published as part of Kok, Philippe J. R., 2009, A new species of Oreophrynella (Anura: Bufonidae) from the Pantepui region of Guyana, with notes on O. macconnelli Boulenger, 1900, pp. 35-49 in Zootaxa 2071 on pages 36-40, DOI: 10.5281/zenodo.274789, {"references":["Senaris, J. C., DoNascimiento, C. & Villarreal, O. (2005) A new species of the genus Oreophrynella (Anura; Bufonidae) from the Guiana Highlands. Papeis Avulsos de Zoologia, 45, 61 - 67.","Senaris, J. C., Ayarzaguena, J. & Gorzula, S. (1994) Los sapos de la familia Bufonidae (Amphibia: Anura) de las tierras altas de la Guayana Venezolana: Descripcion de un nuevo genero y tres especies. Publicaciones de la Asociacion Amigos de Donana, 3, 1 - 37.","Senaris, J. C. (1995) [\" 1993 \"] Una nueva especie de Oreophrynella (Anura; Bufonidae) de la cima del Auyan-tepui, Estado Bolivar, Venezuela. Memoria de la Sociedad de Ciencias Naturales La Salle, 140, 177 - 182.","Diego-Aransay, A. & Gorzula, S. (1990) [\" 1987 \"] Una nueva especie de Oreophrynella (Anura: Bufonidae) de la Guayana Venezolana. Memoria de la Sociedad de Ciencias Naturales La Salle, 127 / 128, 233 - 238.","Kok, P. J. R. (2008) A new highland species of Arthrosaura Boulenger, 1885 (Squamata: Gymnophthalmidae) from Maringma tepui on the border of Guyana and Brazil. Zootaxa, 1909, 1 - 15.","McDiarmid, R. W. & Gorzula, S. (1989) Aspects of the reproductive ecology and behavior of the tepui toads, genus Oreophrynella (Anura, Bufonidae). Copeia, 1989, 445 - 451."]}

Pantepuisaurus genus novum Type species. Pantepuisaurus rodriguesi species novum Etymology. A noun in apposition, derived from ���Pantepui��� referring to the phytogeographic province where the type species was discovered, and the Greek sauros meaning ���lizard���. Gender masculine. Content. Pantepuisaurus rodriguesi species novum; monotypic. Diagnosis. An elongate gymnophthalmid (known SVL in male 58.3 mm) with a long tail (1.6 x SVL) and the unique following combination of characters: (1) distinctive ear opening and moveable eyelids, (2) limbs pentadactyl with all digits clawed, (3) nasal scales separated by a single frontonasal, (4) prefrontals present, (5) frontoparietals, parietals and interparietal present, (6) parietals and interparietal longer than wide, (7) interparietal and parietals forming a jagged, irregular posterior margin, (8) occipitals present, (9) three pairs of genials, second pair in contact with only one infralabial, (10) enlarged median pairs of gulars, (11) dorsal scales hexagonal, keeled, in transverse rows only, (12) ventral scales imbricate, hexagonal, mucronate, broadly keeled, in transverse rows only, (13) tongue anterodorsally covered by oblique, anteriorly converging plicae, posterodorsally covered by large scalelike papillae (14) hemipenis weakly bilobed with series of curved transverse plicae bearing mineralized spicules. Pantepuisaurus differs from members of Alopoglossinae in having the tongue anterodorsally covered by oblique plicae and posterodorsally covered by large scalelike papillae (entirely covered by oblique plicae in Alopoglossinae), the interparietal and the parietals forming a jagged, irregular line (straight posterior margin), and in having ventral scales in transverse rows only (in transverse and longitudinal rows in Alopoglossinae); from members of Gymnophthalminae in having hands pentadactyl with the first finger clawed (first finger absent, reduced and/or clawless in Gymnophthalminae); from Rhachisaurinae in having external ear openings (absent in Rhachisaurinae). Within Cercosaurinae (sensu Pellegrino et al. 2001 and Rodrigues et al. 2005), Pantepuisaurus differs from all genera except Ecpleopus and Leposoma of the scincoides group in having hexagonal, mucronate ventral scales arranged in transverse rows only. Pantepuisaurus is readily distinguished from Ecpleopus (characters of Ecpleopus in parentheses) notably by the presence of a broad keel on ventrals (absent), quadrangular gular scales (mucronate), complete superciliary series (incomplete), and the presence of femoral pores in males (absent). Pantepuisaurus is quickly separated from Leposoma of the scincoides group (characters of Leposoma in parentheses) in having quadrangular gular scales (mucronate), by the absence of distinct striations on head scales (present), the interparietal and the parietals forming a jagged, irregular line (round posterior margin), and the presence of occipitals (absent). Two additional gymnophthalmid genera, Adercosaurus and Kaieteurosaurus, have not been assigned to any subfamily yet, but might be related each other and with Pantepuisaurus. Pantepuisaurus is immediately distinguished from Adercosaurus (characters of Adercosaurus in parentheses) in having imbricate, hexagonal, mucronate, broadly keeled ventral scales in transverse rows only (smooth, in transverse and longitudinal rows), in having a highly reduced fourth infralabial (not reduced) and the second pair of genials partly separated from the infralabials and in contact with one infralabial only (not separated from the infralabials, in contact with two infralabials), in lacking oblique plicae at the rear of the tongue, which is posteriorly covered by large scalelike papillae (rear of tongue covered by oblique plicae), and in having mineralized spicules on hemipenes��� plicae (absent). Pantepuisaurus is most similar to Kaieteurosaurus in having hexagonal, mucronate ventral scales arranged in transverse rows only, but it is readily distinguished from it (characters of Kaieteurosaurus in parentheses) by the presence of prefrontal scales (absent), three pairs of genials (two), in having a highly reduced fourth infralabial (not reduced) and the second pair of genials partly separated from the infralabials and in contact with one infralabial only (not separated from the infralabials, in contact with two infralabials), the common suture of the interparietal and parietals forming a jagged, irregular line (more or less straight posterior margin), an entire nasal (divided), and the aspect of the ventral scales that are much less lanceolate and mucronate than in Kaieteurosaurus. It should be emphasized here that Kaieteurosaurus also has broad ���keels��� on ventral scales (i.e. the central part of the scale is broadly elevated), a character that was only noticed while closely comparing the two genera. This character is not conspicuous in all Kaieteurosaurus specimens and is mostly noticeable on anteriormost and posteriormost ventral scales. In Kaieteurosaurus the significant lateral extension of the flat ���keel��� gives a lanceolate, smooth, ���unkeeled��� aspect to the ventral scales as described by Kok (2005), while the keel is more prominent and conspicuous in Pantepuisaurus. Figure 4 schematically illustrates the difference in the aspect of the broad keel on ventral scales between Pantepuisaurus and Kaieteurosaurus. A summary of main diagnostic characters for Pantepuisaurus, Adercosaurus and Kaieteurosaurus is in Table 1. Distribution. The genus is currently known only from the type locality, the summit of Maringma tepui in Guyana (Fig. 1). Taxonomic comments. No less than eight new gymnophthalmid genera have been described (Myers & Donnelly 2001, Doan & Castoe 2005, Kok 2005, Rodrigues et al. 2005, 2007, Rodrigues & Maranh��o dos Santos 2008) since the molecular-based phylogenetic hypothesis of Pellegrino et al. (2001), who demonstrated the monophyly of Gymnophthalmidae. Some higher-level taxonomic changes are still in debate (see Castoe et al. 2004, Rodrigues et al. 2007) and refinement of higher-level taxonomy of gymnophthalmids is still necessary. Morphologically, the new genus shows some similarities with Adercosaurus and Kaieteurosaurus. Giving the low number of specimens available, any speculation on the relationships between Adercosaurus (known only from the holotype), Kaieteurosaurus (known only from three specimens), and Pantepuisaurus (known only from the holotype) seems premature. More specimens of these three genera are clearly needed to elucidate relationships between them. However, preliminary molecular analysis and comparisons do indicate affinities between Pantepuisaurus and Kaieteurosaurus (unfortunately, molecular data on Adercosaurus are not available), but reveal significant genetic divergences between these two genera that give further support to the erection of a new genus. Molecular analysis also suggests assignment of Pantepuisaurus and Kaieteurosaurus to the tribe Ecpleopini (Cercosaurinae) (M. T. Rodrigues, pers. comm.). A more ���conservative��� attitude would have been to accommodate the new species in an already known monotypic genus sharing some morphological similarities (i.e. Adercosaurus or Kaieteurosaurus). However this would have given rise to two important problems: (1) assignment of the new species to either of these two genera would have been arbitrary and poorly supported because the new species differs from both of them in a combination of important characteristics currently used to diagnose gymnophthalmid genera (e.g. condition of the posterior margin of the interparietal and parietals, condition of ventral scales, number of genials, presence of prefrontals, hemipenial morphology, tongue morphology) (2) because the new species differs from these two genera in morphological characteristics used to separate other genera (especially in the tribe Ecpleopini), placing the new species in one of them would have rendered the taxonomic status of these and other genera dubious by suggesting that these characters are variable intragenerically (thus making taxonomy of the group more complicated instead of simplifying it). One could suggest that Kaieteurosaurus is a synonym of Adercosaurus, however this is not supported by any evidence, these two genera being differentiated on the basis of many important diagnostic characters (see Kok 2005)., Published as part of Kok, Philippe J. R., 2009, Lizard in the clouds: a new highland genus and species of Gymnophthalmidae (Reptilia: Squamata) from Maringma tepui, western Guyana, pp. 53-67 in Zootaxa 1992 on pages 54-56, DOI: 10.5281/zenodo.185553, {"references":["Pellegrino, K. C. M., Rodrigues, M. T., Yonenaga-Yassuda, Y. & Sites Jr., J. W. (2001) A molecular perspective on the evolution of microteiids lizards (Squamata, Gymnophthalmidae), and a new classification for the family. Biological Journal of the Linnean Society, 74, 315 - 338.","Rodrigues, M. T., Freire, E. M. X., Pellegrino, K. C. M & Sites Jr., J. W. (2005) Phylogenetic relationships of a new genus and species of microteiid lizard from the Atlantic forest of north-eastern Brazil (Squamata, Gymnophthalmidae). Zoological Journal of the Linnean Society, 144, 543 - 557.","Kok, P. J. R. (2005) A new genus and species of gymnophthalmid lizard (Squamata: Gymnophthalmidae) from Kaieteur National Park, Guyana. Bulletin de L'Institut Royal des Sciences Naturelles de Belgique, Biologie, 75, 35 - 45.","Myers, C. W. & Donnelly, M. A. (2001) Herpetofauna of the Yutaje-Corocoro Massif, Venezuela: Second report from the Robert G. Goelet American Museum-Terramar Expedition to the northwestern tepuis. Bulletin of the American Museum of Natural History, 261, 1 - 85.","Doan, T. M. & Castoe, T. A. (2005) Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus. Zoological Journal of the Linnean Society, 143, 405 - 416.","Rodrigues, M. T., Pellegrino, K. C. M, Dixo, M., Kruth Verdade, V., Pavan, D., Argolo, A. J. S. & Sites Jr., J. W. (2007) A new genus of microteiid lizard from the Atlantic forests of State of Bahia, Brazil, with a new generic name for Colobosaura mentalis, and a discussion of relationships among the Heterodactylini (Squamata, Gymnophthalmidae). American Museum Novitates, 3565, 1 - 27.","Castoe, T. A., Doan, T. M. & Parkinson, C. L. (2004) Data partitions and complex models in Bayesian analysis: The phylogeny of gymnophthalmid lizards. Systematic Biology, 53, 448 - 469."]}

Centrolene gorzulae (Ayarzag��ena, 1992) Centrolenella gorzulae Ayarzag��ena 1992: 19. Centrolene gorzulai Duellman 1993: 35. Centrolenella auyantepuiana Se��aris and Ayarzag��ena 1995 ��� 1993 ���: 122. Centrolene gorzulae Ruiz-Carranza and Lynch 1995: 2. Hyalinobatrachium auyantepuiana Ayarzag��ena and Se��aris ��� 1996 ��� 1997: 13. Cochranella auyantepuiana Myers and Donnelly 1997: 16. Cochranella auyantepuiana Duellman 1999: 300. Centrolene papillahallicum Noonan and Harvey 2000: 295. Centrolene lema Duellman and Se��aris 2003: 247, new synonym. Centrolene gorzulai Myers & Donnelly 2008: 29. Holotype. MHNLS 11221 (adult male), Venezuela, Estado Bol��var: Central sector of Auyan-tepui (05�� 56 ��� N, 62 �� 34 ��� W; 1850 m). Diagnosis. This species is placed in the genus Centrolene because adult males have humeral spines (Ruiz- Carranza & Lynch 1991; Duellman & Se��aris 2003). It differs from all other species of Centrolene by having a trilobate liver covered by a white peritoneum and a mostly transparent parietal peritoneum, with a small white anterior portion (the ���bib-like white patch��� of Duellman & Se��aris 2003), which is only detectable in dissected specimens. Characterization. (1) dentigerous process on vomer and vomerine teeth absent; (2) snout subtruncate to truncate in dorsal view, truncate to slightly sloping in lateral profile; (3) tympanum distinct, small, oriented dorsolaterally with a slight posterior inclination; (4) dorsal surfaces shagreened; males with small spicules visible under magnification; (5) ventral skin strongly granular, cloacal ornamentation consisting of small enameled tubercles lateral to vent and enlarged paired round tubercles below vent; (6) parietal peritoneum mostly transparent, a small anterior portion white (the ���bib-like white patch��� of Duellman & Se��aris 2003), only detectable in dissected specimens; pericardial peritoneum white; hepatic and visceral peritonea white, urinary bladder transparent; (7) liver trilobate; (8) humeral spine in adult males present; (9) finger webbing II 2 -��� 3 III 2 +���(2 -��� 2) IV; (10) toe webbing I (1 ��� 1 ��)���(2 ��� 2 +) II 1���2 + III 1 ���(2 ��� 2 +) IV (2 ��� 2 +)��� 1 V; (11) fringe on postaxial edge of Finger IV present, enameled, metacarpal fold present, weakly enameled, ulnar fold absent; fringe on postaxial edge of Toe V present, enameled, metatarsal fold present, weakly enameled, tarsal fold usually absent or very weak; (12) nuptial excrescences finely granular, white, consisting of a dense group of glands and situated in the medial, dorsolateral internal side of Finger I (Type-V); prepollex slightly enlarged; prepollical spine projecting (spine not exposed); (13) when appressed, Finger II = Finger I; (14) diameter of eye about 2 X width of disc on Finger III; (15) in life, dorsum dark green with scattered paler flecks; (16) in preservative, dorsum lavender entirely covered with minute melanophores; (17) in life, iris metallic copper with dark brown reticulations; an incomplete pale yellow ring around pupil; (18) distribution of melanophores on digits variable, but always more numerous on Fingers III���IV than I and II; Toes IV���V with many melanophores, rarely present on Toes I���III, if present, few and restricted to the two distal phalanges; in life, hands and feet bluish green to green, tips of fingers and toes yellowish green; (19) males call from the upper side of leaves; note single and pulsed (3���7 pulses per call), advertisement call of 0.02��� 0.05 s duration, dominant frequency of 4416.97���5157.48 Hz; (20) combat behavior unknown; (21) clutches deposited on vegetation overhanging streams, in moss or between two leaves, clutch size 15��� 22 eggs (n = 4); none of the clutches observed were guarded; (22) tadpole unknown; (23) SVL in males 19.2���22.5 mm (n = 11); in females 20.9 ���22.0 mm (n = 4). Natural history and distribution. The ecology of Centrolene gorzulae was recently discussed by Kok and Castroviejo-Fisher (2008), on the basis of specimens from Kaieteur National Park, Guyana. The species was recently discovered at a new locality, the southeast slope of Mount Maringma on the border of Guyana and Brazil (ca. 1400 m elevation), and additional observations were made on clutch deposition. Four clutches were collected between 23���24 November 2007; three of them were found among moss on overhanging branches 1.5 ���2.0 m above water (Fig. 5), the fourth clutch was found deposited between two leaves (attached to both), about 1.5 m above water. Many males were calling at night in the vicinity of the clutch deposition sites, but no male-male interaction was observed. Clutches contained 15��� 22 eggs at various stages of embryogenesis, and no guarding males were detected. One female (IRSNB 14327, 20.9 mm SVL), collected on 22 November 2007, contained 13 enlarged, partly pigmented, ovarian eggs, the largest with a diameter of 2.0 mm. Centrolene gorzulae was thought to be endemic to Auyan-tepui (Ayarzag��ena 1992; Se��aris & Ayarzag��ena 2005; Myers & Donnelly 2008). Kok and Castroviejo-Fisher (2008) extended its distribution to Guyana. Here we report C. gorzulae from three localities in Venezuela (Auyan-tepui, Atapare, and Sierra de Lema) and Guyana (Peters Mountain, Kaieteur National Park, and Mount Maringma) (Fig. 6). This species probably occurs in adjacent areas of Brazil. Centrolene gorzulae has a broad altitudinal distribution (~ 450���1850 m) and has been found only on vegetation overhanging small streams., Published as part of Castroviejo-Fisher, Santiago, Guayasamin, Juan M. & Kok, Philippe J. R., 2009, Species status of Centrolene lema Duellman and Se��aris, 2003 (Amphibia: Centrolenidae) revealed by Integrative Taxonomy, pp. 16-28 in Zootaxa 1980 on pages 19-24, DOI: 10.5281/zenodo.185278, {"references":["Ayarzaguena, J. (1992) Los Centrolenidos de la Guayana Venezolana. Publicaciones de la Asociacion de Amigos de Donana, 1, 1 - 48.","Duellman, W. E. (1993) Amphibian species of the world: additions and corrections. University of Kansas Special Publications, Museum of Natural History, 21, 1 - 372.","Ruiz-Carranza, P. M. & Lynch, J. D. (1995) Ranas Centrolenidae de Colombia VIII. Cuatro nuevas especies de Centrolene de la Cordillera Central. Lozania, 65, 1 - 16.","Ayarzaguena, J. & Senaris, J. C. (\" 1996 \" [1997]) Dos nuevas especies de Cochranella (Anura; Centrolenidae) para Venezuela. Publicaciones de la Asociacion de los Amigos de Donana, 8, 1 - 16.","Myers, C. W. & Donnelly, M. A. (1997) A Tepui Herpetofauna on a Granitic Mountain (Tamacuari) in the Borderland Between Venezuela and Brazil: Report from the Phipps Tapirapeco Expedition. American Museum Novitates, 3213, 1 - 71.","Duellman, W. E. (1999) Distribution patterns of amphibians in South America. In: Duellman, W. E. (Ed.), Patterns of Distribution of Amphibians. A Global Perspective. The Johns Hopkins University Press, Baltimore and London, pp. 255 - 328.","Noonan, B. P. & Harvey, M. B. (2000) A new species of glass frog (Anura: Centrolenidae) from the Highlands of Guyana. Herpetologica, 56, 294 - 302.","Duellman, W. E. & Senaris, J. C. (2003) A new species of glass frog (Anura: Centrolenidae) from the Venezuelan Guayana. Herpetologica, 59, 247 - 252.","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: Report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147.","Ruiz-Carranza, P. M. & Lynch, J. D. (1991) Ranas Centrolenidae de Colombia I. Propuesta de una nueva clasificacion generica. Lozania, 57, 1 - 30.","Kok, P. J. R. & Castroviejo-Fisher, S. (2008) Glassfrogs (Anura: Centrolenidae) of Kaieteur National Park, Guyana, with notes on the distribution and taxonomy of some species of the family in the Guiana Shield. Zootaxa, 1680, 25 - 53.","Senaris, J. C. & Ayarzaguena, J. (2005) Revision taxonomica de la Familia Centrolenidae (Amphibia; Anura) de Venezuela. Publicaciones del Comite Espanol del Programa Hombre y Biosfera - Red IberoMaB de la UNESCO, Sevilla, 337 pp."]}

Oreophrynella macconnelli Boulenger, 1900 Oreophrynella Macconnelli [sic] Boulenger, 1900: 55 Oreophrynella quelchii macconnelli Rivero, 1961: 175 Oreophrynella macconnelli McDiarmid, 1971: 41 Oreophrynella macconnelli was described by Boulenger (1900) on the basis of a single specimen (a male, although not stated in the description) collected at ca. 1067 m elevation on Mount Roraima, Guyana. Four specimens (three males, one juvenile) of this poorly known species with characteristic basal webbing on hand and foot (Fig. 6 A���B) were collected on the southeast slope of Maringma tepui and conform to the holotype and to two topotypic specimens examined. Variation of measurements of the Maringma specimens is given in Table 1. I refrain from redescribing Oreophrynella macconnelli since a formal redescription of the species is in progress by Means (D. B. Means, pers. comm.). The following features characterize the Maringma specimens: (1) small size (up to 22.7 mm SVL in male, female unknown); (2) frontoparietal crests indistinct; (3) postorbital crests barely distinct; (4) suborbital crest indistinct; (5) dorsal skin minutely spiculate with scattered small to medium, low, round tubercles; (6) ventral skin coarsely granular; (7) webbing on hand and foot basal; (8) adult dorsal colour in life variable, from light brownish orange to olive brown, with a series of yellow spots forming a dorsolateral line, and series of yellow spots on upper arm and thigh forming short longitudinal lines; (9) adult ventral colour in life creamy yellow with brown marbling; (10) one vocal slit present on the left side of the floor of the buccal cavity. Colour in life. Adult dorsal colour is variable (Fig. 7 A���B), ranging from light brownish orange (IRSNB 14333, adult male) to olive brown with darker spotting (IRSNB 14334 ��� 5). Olive brown specimens have distinct white spots on the lower flank, while in the orange specimen these spots are yellowish and less conspicuous. All adult specimens with a series of yellow (IRSNB 14333) to brownish orange (IRSNB 14334 ��� 5) spots forming a dorsolateral line, and series of similar spots on axilla, upper arm, groin and thigh forming short longitudinal stripes on limbs. All adult specimens have a white to yellow spot below the eye, although this spot is inconspicuous in IRSNB 14333. Adult ventral colour is creamy yellow with brown marbling; palm and sole are orange (Fig. 7 D). IRSNB 14336, a juvenile, is much darker than adults in having a kaki dorsum with a few yellow spots, and dark brown flanks with a few whitish spots. A few white spots form a less conspicuous dorsolateral line than in adults, and a large longitudinal whitish spot is present on the upper arm and thigh. An anteriorly inclined whitish stripe extends from before the eye to upper lip; white spot below eye smaller than in adults (Fig. 7 C). Ventral colour similar to adults, but much darker. Colour in preservative. Dorsal colour ranges from light brown (IRSNB 14333) to dark grey, almost black (IRSNB 14334 ��� 6). All spots and stripes are creamy white (Fig. 8 A). Ventral colour is creamy yellow in adults, dark grey with white spots in the juvenile. Palm and sole are light grey (Fig. 8 B). Distribution and ecology. According to Se��aris and MacCulloch (2005) and MacCulloch et al. (2007), Oreophrynella macconnelli is reported only from the slopes of Mount Roraima in Guyana and Venezuela, between 1067���1800 m elevation. MacCulloch et al. (2007) considered O. macconnelli a highland species (i.e. typically occurring over 1500 m elevation) with highly restricted distribution. Specimens from Maringma tepui were collected at 1376 m elevation and expand the known geographic range of the species to the west (roughly 17 km airline, Fig. 4 A ���B) suggesting a wider distribution than expected. Instead of being a highland species, O. macconnelli appears rather to be an upland species occasionally invading higher elevation areas. Habitat consists of montane medium-canopy forest with abundant epiphytes and mosses, and rich undergrowth (Fig. 9 A). All the Maringma specimens were found active by night, walking on the apex of epiphytic ferns (Campyloneurum sp., Polypodiaceae, Fig 9 B) about 1.0���3.0 m above the ground, except IRSNB 14333 (adult male), which was collected on the ground at 17 h00. When handled specimens exhibited a defensive posture consisting of arching the back and flexing the head downward towards the handler (Fig. 7 B)., Published as part of Kok, Philippe J. R., 2009, A new species of Oreophrynella (Anura: Bufonidae) from the Pantepui region of Guyana, with notes on O. macconnelli Boulenger, 1900, pp. 35-49 in Zootaxa 2071 on pages 41-45, DOI: 10.5281/zenodo.274789, {"references":["Boulenger, G. A. (1900) Batrachians. Report on a collection made by Messrs. F. V. McConnell and J. J. Quelch at Mount Roraima in British Guiana. Transactions of the Linnean Society of London, Zoology, Series 2, 8, 55 - 56.","Rivero, J. A. (1961) Salientia of Venezuela. Museum of Comparative Zoology, Harvard University Bulletin, 126, 1 - 207.","McDiarmid, R. W. (1971) Comparative morphology and evolution of frogs of the Neotropical genera Atelopus, Dendrophryniscus, Melanophryniscus, and Oreophrynella. Bulletin of the Los Angeles County Museum of Natural History, 12, 1 - 66.","Senaris, J. C. & MacCulloch, R. D. (2005) Amphibians. In: Hollowell, T. & Reynolds R. P. (Eds.). Checklist of the terrestrial vertebrates of the Guiana Shield. Bulletin of the Biological Society of Washington 13, pp. 8 - 23."]}

Adelophryne gutturosa Hoogmoed and Lescure, 1984 Adelophryne gutturosa Hoogmoed and Lescure, 1984: 101; Ayarzag��ena and Diego-Aransay, 1985: 159; Hoogmoed, Borges and Cascon 1994: 272; Barrio-Amor��s 2004: 19; Se��aris and MacCulloch 2005: 21. Adelophryne gutturosa was first described by Hoogmoed and Lescure (1984) from specimens from Mount Roraima, Guyana, and Amap��, Brazil. Ayarzag��ena and Diego-Aransay (1985) reported a single specimen, a gravid female of 16 mm SVL, from La Escalera, Venezuela. Hoogmoed et al. (1994) provided an additional description based on two new specimens from Amap��. Fifteen specimens of A. gutturosa were recently collected from Guyana; two on Mount Wokomung at the same location as the holotype of A. patamona, one at Mount Ayanganna (05�� 24 ' N, 0 59 �� 57 ' W, 870 m) 10 from Kaieteur National Park (05�� 10 ' N, 059�� 29 ' W, 400���550 m), two from the vicinity of the Meamu River in the northern Pakaraima region (06�� 16 ' N, 0 60 �� 30 ' W, 664 m) and two from La Laja, Sierra de Lema, Bol��var, Venezuela (06�� 03' N, 0 61 �� 29 ' W, 450 m). The specimens from Wokomung are a female of 13 mm SVL and a male of 13.5 mm SVL. The specimen from Ayanganna is a male of 14 mm SVL. The specimens from Kaieteur National Park are six males of 13.3���14.7 mm SVL and four females of 14.6 ���15.0 mm SVL and the specimens from Meamu are two females of 15.3 and 16.0 mm SVL. The specimens from La Laja are a male of 15 mm and a juvenile of 13 mm SVL. Advertisement calls were recorded at Kaieteur National Park between 2���7 December 2005 and at two additional sites on the headwaters of the Meamu River: 0 6 �� 10 ' N, 0 60 �� 26 ' W, 703 m, on 28 December 2004 and 0 6 �� 12 ' N, 0 60 �� 29 ' W, 781 m on 2 September 2004. These 17 specimens expand the existing descriptions of A. gutturosa as follows: head width 90 % head length; shanks 45���52 % SVL; palpebrum not reticulated, a black band along its upper rim. The specimens from Ayanganna, Wokomung, Kaieteur, Meamu and La Laja differ from the previous descriptions of A. gutturosa as follows (previously described specimens in parentheses): distance from eye to tip of snout variable, 75���85 % eye diameter in specimens from Ayanganna, Wokomung, Meamu and La Laja, 100���120 % eye diameter in specimens from Kaieteur (equal); upper eyelid width variable, 70���100 % interorbital distance (50 %); teeth 7���8 in females, 8 in males at Ayanganna, Wokomung and Meamu, 4 in females, 3��� 4 in males at Kaieteur (2, 4��� 6). In some specimens from Kaieteur and La Laja the skin on the dorsum is shagreened (smooth), and the temporal region and flanks are granular. These differences are within the range of interpopulation variation. Colour in life. Considerable variation in colouration is evident, both within and among populations. Dorsal ground colour medium brown or grey. A middorsal black ���)(���, black canthal and supratympanic bars and other scattered black marks may be present; numerous irregular sky blue or white dots on dorsum and flanks; dorsal surfaces of hindlimbs and forearms medium brown or grey, with or without black marks or crossbands and small irregular sky blue or white dots; upper arm orange (Fig. 9 A���F). In the two specimens from Meamu the flanks and upper surfaces of thighs are whitish with dark stippling and no crossbands. Venter brown or grey with small irregular sky blue or white dots; iris copper, with a red ring around pupil (Fig. 9 A���F). In preservative the orange and sky blue colours become white, otherwise there is little change. Vocalisation (Fig. 10; Tables 3���4). The following description is based on a sample of 29 advertisement calls from 12 males at the three localities. Temporal structure. The advertisement call consists of a group of short notes produced in quick succession, with the interval between notes increasing progressively from the beginning to the end of the call (Fig. 10). Consequently, calls containing only a few notes have a higher note repetition rate than those containing many notes, since in the latter the notes are on average more widely spaced. Thus a 2 -note call from Kaieteur National Park (specimen IRSNB 13784) has a duration of 0.058 s and a note repetition rate of 22.22 notes/s; a 7 -note call from the same locality (specimen IRSNB 13791) has a duration of 0.85 s and a note repetition rate of 7.26 notes/s, and a 15 -note call from Meamu has a duration of 2.39 s and a note repetition rate of 6.2 notes/ s. Call duration varies considerably, both between, and within individuals participating in the same chorus and may reflect social context (eg. territorial encounters or proximity to a female). At Kaieteur, a male situated close to a female produced 2 -note calls continuously. Each note contains 4���5 high amplitude pulses which are followed by up to 7 low amplitude pulses (Fig. 10 c). The pulses vary in duration and are unevenly spaced: some are clearly separated from adjacent pulses, while others merge. Pulse rate therefore varies within and between notes and is unlikely to play a role in mate recognition. Spectral structure. Three to four harmonics are developed, with the fundamental frequency dominating (range: 3896 ��� 4979 Hz; mean: 4242 Hz). The distribution of sound energy decreases progressively through the 2 nd, 3 rd and 4 th harmonic. The frequency rises slightly from the beginning to the end of the note. Vocalisation parameters of three Adelophryne species are compared in Table 4. Natural history. Specimens were collected in forest, in late afternoon or early evening. Specimens from Ayanganna, Wokomung and Meamu were collected in leaf litter. Most specimens from Kaieteur were closely associated with formations of Monotagma spicatum (Marantaceae), locally named Dalibana (Fig. 9 G). A few individuals were observed sheltering at the base of other vegetation, although M. spicatum appears to be the preferred habitat. Males were usually found calling from the base of the plant or hidden among the rootlets; some were observed calling among dead leaves near the base of the plant. Most of the females were collected close to calling males. Small size and cryptic colouration make A. gutturosa difficult to locate, even when calling. When disturbed the frogs move slowly or remain motionless among rootlets or in leaf litter. If disturbed while away from shelter, they can rapidly jump considerable distances. A few ants were found in the stomachs of two specimens. Once we became familiar with the call of A. gutturosa, we found that this species is locally abundant. Choruses were heard between 0300���0600 and 1800���1930 in several locations in Kaieteur and Meamu. In some localities such as the Elinkwa River in Kaieteur National Park, and La Laja, A. gutturosa is the most common frog heard in late afternoon, early evening or during rains. On 2 December 2005 at 1540 a male A. gutturosa (IRSNB 13784) was located by call and a female (IRSNB 13785) was observed, immobile, in close proximity. Both specimens were collected and kept overnight in a plastic container with some dead leaves. The next morning a single large egg of 4.63 mm external diameter was found among the dead leaves (Fig. 9 H). The large size of the egg, 31 % of the female���s SVL, supports the hypothesis of direct development in this species (Ayarzag��ena and Diego-Aransay 1985). The short wet season in Guyana begins in December; Ayarzag��ena & Diego-Aransay (1985) and Hoogmoed et al. (1994) theorized that oviposition takes place at the beginning of the rainy season (to minimize desiccation of terrestrial eggs) It is likely that eggs are laid among plant roots. Our observations indicate that males exhibit site fidelity, positioning themselves as close as 2 m from adjacent calling males. Males rarely call simultaneously, although the call of one male usually prompts an adjacent male to call and there is a slight overlap between the two calls. Calls are emitted in successive waves; due to the nature of the call, this creates a very powerful ���cascading chorus���. Distribution and relationships. Adelophryne gutturosa is a widespread species, whose range extends from Amap��, Brazil, to Bol��var, Venezuela, a distance of some 1500 km. Elevation ranges from 110 and 300 m in Amap�� (Hoogmoed et al. 1994), through 400���550 m at Kaieteur, 450 m at La Laja, 664 m at Meamu, 870 m at Ayanganna, 900 m at Roraima (Hoogmoed & Lescure 1984), 950 m at Auy��n (Se��aris & Ayarzag��ena 2006), approximately 1100 m at La Escalera (Ayarzag��ena & Diego-Aransay 1985), 1234 m at Wokomung, to 1400 m at Maringma. This broad geographical and elevational range invites further study into the possible presence of cryptic species within A. gutturosa. Although they may be locally abundant, the small size and secretive habits of Adelophryne make discovery difficult. There are numerous large gaps in the known distribution of Adelophryne (Hoogmoed et al. 1994). Calls could be used to verify the presence of these frogs during biodiversity surveys. Several characters such as number of phalanges, digital discs and pads, skin texture and colour pattern have been used to distinguish among species of Adelophryne and among genera of small eleutherodactylids (Hoogmoed & Lescure 1984). Nonetheless, understanding of the phylogeny of these poorly known animals will require considerable further sampling. Relationships of this group were discussed by Lynch (1986). As pointed out by Heyer (1977) and Da Silva et al. (2007), many of the characters present in these taxa may be due to convergence, and may not reflect relationships., Published as part of Macculloch, Ross D., Lathrop, Amy, Kok, Philippe J. R., Minter, Leslie R., Khan, Samir Z. & Barrio-Amor��s, C��sar L., 2008, A new species of Adelophryne (Anura: Eleutherodactylidae) from Guyana, with additional data on A. gutturosa, pp. 36-50 in Zootaxa 1884 on pages 45-49, DOI: 10.5281/zenodo.274492, {"references":["Hoogmoed, M. S. & Lescure, J. (1984) A new genus and two new species of minute leptodactylid frogs from northern South America, with comments upon Phyzelaphryne (Amphibia: Anura: Leptodactylidae). Zoologische Mededelingen, 58, 85 - 115.","Ayarzaguena, J. & Diego-Aransay, A. (1985) Primer reporte para Venezuela de Adelophryne gutturosa (Leptodactylidae) y datos sobre su biologia. Memoria de la Sociedad de Ciencias Naturales La Salle, 45, 159 - 160.","Hoogmoed, M. S., Borges, D. M. & Cascon, P. (1994) Three new species of the genus Adelophryne (Amphibia: Anura: Leptodactylidae) from northeastern Brazil, with remarks on the other species of the genus. Zoologische Mededelingen, 68, 271 - 300.","Barrio-Amoros, C. L. (2004) Amphibians of Venezuela: systematic list, distribution and references, an update. Revista de Ecologia Latinoamericana, 9, 1 - 48.","Senaris, J. C. & MacCulloch, R. D. (2005) Amphibians. In: Hollowell, T. & Reynolds R. P. (Eds.). Checklist of the terrestrial vertebrates of the Guiana Shield. Bulletin of the Biological Society of Washington 13, pp. 8 - 23.","Senaris, J. C. & Ayarzaguena, J. (2006) A new species of Hypsiboas (Amphibia; Anura; Hylidae) from the Venezuelan Guayana, with notes on Hypsiboas sibleszi Rivero 1972. Herpetologica, 62, 308 - 318.","Lynch, J. D. (1986) New species of minute leptodactylid frogs from the Andes of Ecuador and Peru. Journal of Herpetology, 20, 423 - 431.","Heyer, W. R. (1977) Taxonomic notes on frogs from the Madeira and Purus rivers, Brasil. Papeis Avulsos de Zoologia, 31 (8), 141 - 162.","Da Silva, H. R., Campos, L. A. & Sebben, A. (2007) The auditory region of Brachycephalus and its bearing on the monophyly of the genus (Anura: Brachycephalidae). Zootaxa, 1422, 59 - 68."]}

Adelophryne patamona, sp. nov. Figs 1–7 Material. Holotype. ROM 43034, an adult male, from the north slope of Mount Wokomung, Guyana, 05º 06' 35 " N, 059º 49 ' 15 " W, 1234 m, collected by A. Lathrop, S. Khan and R. MacCulloch, 1 November 2004. Paratypes. ROM 43035, an adult female, from the north slope of Mount Wokomung, Guyana, 05º 05' 33 " N, 059º 50 ' 35 " W, 1411 m, collected by A. Lathrop, S. Khan and R. MacCulloch, 7 November 2004; IRSNB 1971 – 4, adult females, from the east base of Mount Maringma, Guyana, 05º 12 ' 38 " N, 060º 33 ' 59 " W, 1060 m, collected by P. Kok, P. Benjamin and C. Perry, 20 November 2007; IRSNB 1975, an adult female, IRSNB 1976 - 7, two juveniles and IRSNB 1978, an adult male from the southeast slope of Mount Maringma, Guyana, 05º 12 ' 16 " N, 060º 34 ' 39 " W, 1376 m, collected by P. Kok, P. Benjamin and C. Perry, 22–24 November 2007. Etymology. The specific epithet is a noun in apposition, and refers to the Patamona people, who live in the area where the holotype was collected. Diagnosis. A large species of Adelophryne, characterised by maximum SVL> 20 mm, second phalange of Finger IV longer than wide, tips of digits with small pointed discs, tympanum small, separated from eye by a distance equal to tympanum diameter, posterodorsal part of tympanum covered by a fold, and an inconspicuous glandular ridge from tympanum to arm insertion. The specimens can be placed in the genus Adelophryne because they possess the following combination of characters: small size, flattened digits with indistinct subdigital pads rather than subarticular tubercles, shortened Finger IV with single subdigital pad, discs with pointed tips and lateral grooves, vomerine teeth present, tongue narrow, tarsal tubercle absent. Of the other genera of small terrestrial frogs in the region, Noblella has a tarsal tubercle and no vomerine teeth, Phyzelaphryne has a broad tongue, round digits, and discs with circumferential grooves, and Pristimantis has two subarticular tubercles on Finger IV. The differences among these genera were elaborated upon by Hoogmoed and Lescure (1984). Adelophryne patamona can be distinguished from A. adiastola and A. pachydactyla by having 3 phalanges in Finger IV (2 phalanges in A. adiastola and A. pachydactyla) and large size, up to 23 mm SVL (largest A. adiastola 14 mm, largest A. pachydactyla 11.1 mm). Adelophryne patamona can be distinguished from A. baturitensis by the presence of subdigital pads and absence of subarticular tubercles (tubercles present, pads absent in A. baturitensis) and large size, up to 23 mm SVL (largest A. baturitensis 16.3 mm). Adelophryne patamona can be distinguished from A. maranguapensis by its posterior cloacal opening, without a transverse flap (flap present, cloaca opening ventrally in A. maranguapensis) and large size, up to 23 mm SVL (largest A. maranguapensis 17.4 mm). Adelophryne patamona can be distinguished from A. gutturosa, with which it occurs sympatrically, by the following characters: second phalange of Finger IV relatively long, 2 X its width (short, length equal to or slightly greater than width in A. gutturosa); canthus rostralis concave (straight); vomerine teeth 15–16 (2–8); large size, up to 23 mm SVL (largest A. gutturosa 16 mm) (Fig. 1). In the field these species can be easily distinguished by their distinctly different advertisement calls (see below). Characters useful for distinguishing among species of Adelophryne are in Table 1. Description of the holotype. A male, 17 mm SVL, head slightly longer than wide, as wide as adjacent part of body, head depth 50 % of width. Snout rounded in dorsal and lateral profile. Distance between snout tip and eye slightly less than eye diameter, eye-nostril distance slightly less than internarial distance, 2 X distance from nostril to snout tip, 75 % of distance from eye to snout tip. Canthus rostralis indistinct, rounded, concave; loreal region sloping steeply to upper lip. Nostrils not projecting, round, directed laterally. Interorbital distance 1.3 X internasal distance, 2 X upper eyelid width. Pupil horizontal. Temporal region vertical; tympanum small, distinct, round, 35 % of eye diameter, surrounded by a distinct annulus, partially obscured by a supratympanic fold, separated from eye by a distance equal to tympanum diameter. Low glandular ridge from tympanum to arm insertion barely visible. Choanae small, laterally placed; vomerine processes transverse, posterior to choanae, each bearing 15 teeth. Tongue narrow, sides parallel, attached anteriorly. Vocal slits small, flanking tongue, vocal sac large, subgular. Skin of dorsum, venter, and limbs smooth with scattered small tubercles; flanks and rear of thighs areolate. Cloacal opening above midpoint of thigh, directed posteriorly, dorsal flap absent. Hand with large flat oval palmar and thenar tubercles. Underside of fingers with large flat subdigital pads (1 – 1–2 – 1); no pads under ultimate phalanges. Fingers flattened, unwebbed. Tips of fingers bluntly pointed, bearing discs not wider than the adjacent phalange, discs laterally grooved. Finger lengths I Tarsus smooth, no ridge or tubercle. A large distinct oval flat inner and a smaller round conical outer metatarsal tubercle. Subdigital pads (1 – 1–2 – 3 – 2) large, flat, located just distally of digital articulations; no supernumerary pads. Toes flattened, unwebbed; toe V more slender than the others. Tips of toes dilated into small, narrow discs, those of Toes II, III, IV slightly wider than adjacent phalange, ending in asymmetrically pointed tips; those of Toes I and V equal to width of adjacent phalange, with rounded or bluntly pointed tips. Discs with lateral grooves. Toe lengths I Heel of adpressed hindlimb reaches a point between eye and tip of snout. Heels overlap slightly when hindlimbs are flexed at right angles to sagittal plane of body. Shank length 57 % of SVL. Colour of holotype in life. Ground colour of dorsum, flanks and dorsal surfaces of limbs medium brown. Central part of dorsum dark brown, bounded by a black middorsal “)(”-shaped mark and interorbital bar. Dorsum with a few other small, scattered black marks. Loreal and temporal regions dark brown. A black bar extending from tip of snout through eye to flank; another black bar in the inguinal region. Dorsal surfaces of limbs with black crossbands. Throat, chest and underside of arms dark grey with white or pale blue spots. Venter, underside of legs and rear of thighs reddish-brown with white or pale blue spots. A black anal spot. Palpebrum not reticulated, a black band along its upper rim. Iris copper with a red ring around pupil (Fig 3). In preservative the venter becomes brownish-grey with white spots. Variation. The female paratype from Wokomung is the largest specimen collected, SVL 23 mm (Fig 4). Paratypes from Maringma (Figs. 5–7) are one male 18.4 mm SVL, five females 20.7–21.5 mm SVL, and two juveniles (11.8 and 11.9 mm SVL). Dorsal skin smooth to granular; ventral tubercles variable; vomerine processes each bearing 16–18 teeth in adults, 2–3 teeth in juveniles. Colouration is variable, not sexually dimorphic (Figs. 4–7). Black dorsal markings are variable; some specimens are darker with more pale blue spots and more conspicuous tubercles on dorsum. In the specimens from Maringma the throat, chest and underside of arms are black with white or pale blue spots and a few reddish-brown dots; the venter, underside of legs and rear of thighs are reddish-brown, sometimes with very few white or pale blue spots (Figs. 6–7). Juveniles resemble adults, but the upper arm is yellowish-orange and the dorsum has more pale blue spots; ventrally they are black with numerous pale blue spots (Fig. 5 C). Vocalisation (Fig. 8; Tables 2, 4). The following description is based on a sample of six advertisement calls from a single male, IRSNB 1978, recorded on the southeast slope of Mount Maringma (05º 12 ' 16 " N, 060º 34 ' 39 " W, 1376 m), on 24 November 2007 at 14 h 15, air temperature 19.8 °C. Call parameter N Mean Range Call duration (s) 6 0.63 0.584–0.712 Inter-call interval (s) 5 3.85 3.365–4.255 Duration of first note (s) 6 0.038 0.035–0.042 Duration of second note (s) 6 0.056 0.055–0.057 Duration of third note (s) 6 0.429 0.400– 0.497 Inter-note interval between the first and the second note (s) 6 0.075 0.072–0.081 Inter-note interval between the second and the third note (s) 6 0.052 0.047–0.059 Note repetition rate (notes/s) 12 9.07 8.66–9.26 Emphasized frequency (Hz) 18 3255 3251–3269 Temporal structure. The advertisement call of Adelophryne patamona consists of a group of three soft, unpulsed, whistle-like notes (two short, followed by one long) produced in rapid succession (Fig. 8). Although a feeble isolated single whistle-like note was sometimes heard, we never observed the specimen emitting that note, and thus we cannot confidently attribute it to A. patamona. However it is possible that, like in A. gutturosa (see below), a shorter call is emitted when a female is very close to a male. Males typically call antiphonally and there was no overlapping of the calls. Spectral structure. Three harmonics are developed, with the fundamental frequency dominating (range: 3251–3269 Hz; mean: 3255 Hz). The distribution of sound energy decreases progressively through the higher harmonics. The frequency rises abruptly at the very beginning of the notes (Fig. 8). The amplitude of the third note rises sharply to a peak at the beginning of the note then drops off rapidly, rising again, slowly, to a peak near the end of the note. A smaller peak in amplitude occurs at the end of the note (Fig. 8). Vocalisation parameters of three Adelophryne species are compared in Table 4. Pulses 2 4–5 high amplitude pulses followed by up 0 to 7 low amplitude pulses Emphasized harmonic Second First First Emphasized frequency (Hz) 3200–3700 3896–4979 3251–3269 Natural history. The specimens were collected at the end of the dry season, mostly at night, in leaf litter under medium-canopy forest (Fig. 5 D). One female (IRSNB 1975) and the two juveniles (IRSNB 1976 – 7) were collected among rootlets at the base of a tree. One male, IRSNB 1978, was calling in the afternoon under leaves in a hole between roots. Many males were heard calling during afternoons and nights but were very difficult to find in their hidden calling sites. No enlarged ova were found in the females. The species was also heard on Maringma at 1414 m, and in high-canopy forest near the village of Wayalayeng (05º 14 ' 11 " N, 060º 31 ' 04" W, 678 m), but none were collected at these locations. Distribution. Known only from the Pakaraima Mountains in western Guyana. The species likely occurs in Roraima State, Brazil, only 2 km from the Maringma location. The collecting locality at 1411 m is the highest elevation reported for any Adelophryne. Relationships. Adelophryne patamona appears to be most closely related to A. gutturosa, with which it is sympatric and sometimes syntopic (see below). Colour pattern may not be useful for distinguishing among species of Adelophryne. A dark “)(” -shaped dorsal marking is present in several Adelophryne (A. baturiensis, A. gutturosa and A. maranguapensis; Hoogmoed et al. 1994). The ventral pattern of dark background with light spots is also common in the genus.

Arthrosaura hoogmoedi species novum Figs. 5 ���6, 7 C, 8���9 Holotype. IRSNB 2653 (field number PK 2031), an adult female collected by Philippe J. R. Kok, Paul Benjamin and Claudius Perry, 27 November 2007 at 11 h00, summit plateau of Mount Maringma, Cuyuni-Mazaruni District, Guyana (5 �� 12 ��� 60 ��� N, 60 �� 35 ��� 06��� W, 2112 m elevation). Etymology. The specific epithet is a noun in the genitive case, honoring Dutch herpetologist Marinus Steven Hoogmoed. Marinus was curator of reptiles and amphibians at the National Museum of Natural History (Leiden, The Netherlands) for 38 years (1966���2004) before retiring as research associate at the Museu Paraense Em��lio Goeldi in Bel��m, Brazil, where he is still very active. His contribution to the knowledge of the Guiana Shield herpetofauna is extensive, and most of his papers on the lizard fauna of the area are still authoritative. Diagnosis. The new species is assigned to the genus Arthrosaura because of the following combination of characters (modified from ��vila-Pires 1995, and MacCulloch & Lathrop 2001): (1) distinctive ear opening and moveable eyelids, (2) limbs pentadactyl with all digits clawed, (3) nasal scales separated by a single frontonasal, (4) prefrontals present (although absent in Arthrosaura synaptolepis Donnelly, McDiarmid & Myers, 1992), (5) frontoparietals, parietals and interparietal present, (6) parietals and interparietal longer than wide, (7) interparietal and parietals forming a straight posterior margin, (8) occipitals present, (9) second pair of enlarged genials in contact with two infralabials, (10) enlarged median pairs of gulars, (11) dorsal scales hexagonal, keeled, in transverse rows only, (12) ventral scales quadrangular, smooth, in both transverse and longitudinal rows, (13) tongue dorsally covered by scalelike papillae (although papillae are reported as interrupting oblique plicae in Arthrosaura montigena Myers & Donnelly, 2008). In addition to the generic characteristics, the new species is characterized by the following features: four supraoculars; prefrontals in contact with loreals; palpebral disc completely opaque; 32���33 smooth temporal scales; scales around midbody 37; 3���4 middorsal scales distinctly narrower than adjacent dorsal scales; dorsal scales strongly keeled, in 29 transverse rows; trunk length 1.4 x length of forelimb; overlap of limbs equivalent to seven lateral scales when they are adpressed along the body; dark brown dorsum with a black middorsal stripe from nape to tail, and a tan ill-defined dorsolateral line from nape to midbody; flanks black; ventrally cream with dark mottling. The new species is immediately distinguished from Arthrosaura montigena, A. reticulata (O���Shaughnessy, 1881), A. synaptolepis, A. tyleri (Burt & Burt, 1931), and A. versteegii Lidth de Jeude, 1904 in having four supraoculars (vs. three) and a black middorsal stripe from nape to tail (absent in the above-mentioned species). The new species agrees with A. guianensis MacCulloch and Lathrop, 2001, A. kockii (Lidth de Jeude, 1904), and A. testigensis Gorzula and Se��aris, 1998 in having four supraoculars. Arthrosaura guianensis mainly differs (characters of A. hoogmoedi in parentheses) in having keeled parietals and interparietal (a feeble keel on interparietal only), unpigmented tympanum (pigmented), lower eyelid with semitransparent disc (disc opaque), 45 scales around midbody (37), 40���42 keeled temporals (32���33, smooth), and absence of a black middorsal stripe (present). Arthrosaura kockii mainly differs (characters of A. hoogmoedi in parentheses) in having broadly keeled temporals (smooth), unpigmented tympanum (pigmented), lower eyelid with semitransparent disc (disc opaque), a wide light vertebral stripe (black middorsal stripe), and in having an immaculate venter (dark mottled). Arthrosaura testigensis shares many characteristics with A. hoogmoedi, from which it mainly differs (characters of A. hoogmoedi in parentheses) in having less than 20 temporals (more than 30), lower eyelid with semitransparent disc (disc opaque), prefrontals separated from loreals by frontonasal-first supraocular contact (prefrontals in contact with loreals), weakly keeled dorsal scales (strongly keeled), middorsal scales not noticeably narrower than adjacent dorsal scales (3���4 middorsal scales distinctly narrower), and in having four narrow yellowish longitudinal stripes on dorsum (one middorsal black stripe and a tan ill-defined dorsolateral line). Description of the Holotype. Adult female 59.3 mm SVL. Head length 21 % SVL, 1.37 x head width, head width 1.48 x head height. Head slightly wider than neck, temporal region swollen, neck approximately as wide as anterior body. Snout blunt, rising gently posteriad. Body oval in transverse section, depressed, wider than high. Tail complete (regenerated distally), 97.0 mm, 1.63 x SVL, circular in cross-section, tapering toward the tip. Limbs well-developed, moderately long, forelimbs 31 % SVL, hindlimbs 48 % SVL. Limbs pentadactyl, all fingers and toes with terminal claw. Overlap of limbs equivalent to seven lateral scales when they are adpressed along the body. Trunk length 25.7 mm, 43 % SVL, 1.4 x forelimbs (Fig. 5). Tongue lanceolate, dorsally covered by oblique rows of small scalelike papillae; tip bifid, smooth; three well defined pairs of subequal infralingual plicae on the anterior part of the tongue, after fork. Anterior teeth conical, posterior teeth laterally compressed, tricuspid. Rostral rectangular, twice as wide as high, visible from above, in broad contact with frontonasal. Frontonasal single, heptagonal, laterally in contact with nasal and in broad contact with loreal, not in contact with first supraocular. Prefrontals roughly pentagonal, as broad as long, with a long medial suture, laterally in broad contact with first supraocular, touching the loreal and the anterior point of the second supraocular. Frontal hexagonal, 2.1 x as long as wide, slightly wider anteriorly, laterally markedly concave; laterally in contact over the entire length of second supraocular, separated from first supraocular by prefrontal-second supraocular contact, in point contact with third supraocular. Frontoparietals roughly hexagonal, 1.5 x as long as wide with relatively broad medial suture; frontoparietals in contact with frontal, distinctly separated from second supraocular by frontal-third supraocular contact, in broad contact with third and fourth supraoculars, one parietal and interparietal. Interparietal 1.6 x as long as wide, roughly pentagonal with curved sides converging anteriad, distinctly longer and narrower than parietals. Parietals and interparietal form a relatively straight posterior margin; anteriorly each is angulate. Interparietal with a short median, feeble, longitudinal keel. A row of seven occipitals of various shapes, the central largest; all of them feebly keeled, except the first left one. Four supraoculars, first smallest, about half the size of the fourth; third distinctly larger, second largest. Six superciliaries, first much larger than the others, last two smallest, others subequal (Fig. 6). Nasal entire, with a very short division running ventrad from naris to supralabial on the left side; naris just below centre; directed laterally. Loreal trapezoidal, anteriorly inclined, in contact with nasal, frenocular, first superciliary, first supraocular, and prefrontal and frontonasal, separated from supralabials by frenocular-nasal contact and an odd, very small, quadrangular, supernumerary scale located at the intersection of nasal, frenocular, and first and second supralabials, present on both sides. Three small preoculars on the right side, two on the left, between first superciliary and first subocular. Frenocular quadrangular, longer than wide, subequal to loreal, separated from the first supralabial by a small supernumerary scale. Four suboculars on the right side, five on the left. Two postoculars, the lower smallest. Seven supralabials on both sides, fifth highest, with suture between third and fourth below middle of the eye; suture between second and third supralabials in contact with first subocular, suture between first and second supralabials in contact with a small supernumerary scale on both sides (Fig. 6). Upper eyelid with 10 ciliaries. Ocular recess with median row of five small scales separating median ciliaries from superciliaries. Lower eyelid with 12 ciliaries on the right side, 11 on the left side. Lower eyelid with a palpebral disc of four large, vertical, completely opaque (pigmented), palpebral scales (Figs. 6, 7 C). Temporal region with 32 irregularly shaped, juxtaposed, smooth scales on the right side, 33 on the left. Temporals arranged in six ill-defined oblique rows, much larger in the upper part [a distinct cluster of 16 (on the right) and 18 (on the left) smaller temporals in the lower part]. Ear opening moderately large, vertically oval, posteriorly bordered by many small, smooth, mostly granular scales; anteriorly bordered by four (on the right side) or six (on the left side) smooth, enlarged, juxtaposed scales. Auditory meatus moderately deep, tympanum pigmented (Fig. 6). Mental trapezoidal, its posterior edge slightly concave. Postmental large, roughly heptagonal, laterally in contact with first and second infralabials. Three pairs of large genials, first two pairs in contact with each other medially, in lateral contact with infralabials, medial suture of second pair twice as long as that of first pair; third pair in contact with infralabials, but separated medially by small scales. A pair of enlarged postgenials separated medially by scales of varying sizes and widely separated from infralabials. Six infralabials on the right side, five on the left side. Six rows of two transversely enlarged gulars. Collar row with nine scales, decreasing in size laterally, forming a scalloped margin. Gular fold distinct, concealing three rows of very small scales (Figs. 6, 8 B). Head scales with numerous small pits, especially concentrated on rostral, frontonasal, supraoculars and lateral head scales, absent on ventral head scales and body scales. Scales on nape posterior to the row of occipitals longer than wide, keeled, mucronate, in transverse rows, grading posteriorly into dorsal body scales. Sides of neck with distinctly smaller granular scales, juxtaposed, not in rows, slightly decreasing in size posteriorly. Three (left side) or four rows (right side) of distinctly enlarged scales before arm insertion. Dorsals imbricate, appearing hexagonal because of imbrication, elongate, strongly keeled, mucronate, in transverse rows only, 29 rows between the interparietal and posterior edge of hindlimbs (including row of occipitals). Three to four middorsal scales, between anterior margin of forelimbs and anterior margin of hindlimbs, distinctly narrower than adjacent dorsals and very similar to laterals (Fig. 9 A), 37 scales around midbody. A zone of small granular scales extending from anteriorly of forelimb insertion to above arm and becoming wider in the axilla. Small granular scales at hindlimb insertion. Flank scales similar to, but narrower than, most dorsal scales (except three to four middorsal ones, see above); no abrupt demarcation between dorsals and laterals. Lateral scales becoming smaller toward groin. Below the laterals, two rows of ventrolateral scales (see Myers & Donnelly 2008); some short, smaller scales intercalated between the transverse rows of laterals; no distinct demarcation between laterals and ventrolaterals. Ventrals imbricate, smooth, longer than wide, with posterior margins slightly rounded, arranged in transverse and longitudinal rows (Fig. 8 A). Eighteen transverse rows between collar and preanal plate, six longitudinal rows of ventrals at midbody (10 rows including the ventrolaterals). Preanal plate consisting of eight scales, three anterior scales, five posterior scales (four elongate large scales and a small distal medial triangular scale); lateral posterior preanal scales narrow, almost as long as adjacent scales. No preanal or femoral pores (Fig. 9 B). Scales on tail keeled, mucronate (except distally), imbricate, appearing hexagonal because of imbrication, similar to, but slightly shorter than, dorsal scales, in transverse rows only except two middorsal caudal scales that form an almost complete longitudinal row from the seventh dorsal caudal scale (from posterior margin of hindlimbs) to tip of tail. Regenerated portion of tail (distal 31.0 mm) with much shorter scales ventrally and dorsally. Pigmented subcaudal scales smooth, imbricate, with rounded ends, in both transverse and longitudinal rows, becoming keeled laterally, grading into laterals. Dorsal surfaces of upper and lower arms with smooth, variably polygonal scales, ventral surface of upper arms with small, juxtaposed, granular scales, ventral surface of lower arms with scales similar to those on dorsal surface, except for a band on inner side with small juxtaposed granular scales. Thighs with a row of enlarged, smooth, hexagonal scales on anterior surface, bordered ventrally by two rows of slightly smaller, smooth, polygonal scales; bordered dorsally by 2���3 rows of keeled scales, diminishing in size dorsad, grading into small, granular, juxtaposed scales, which cover posterodorsal and posterior surfaces of thighs. Shanks ventrally and posteriorly with large, smooth, polygonal, imbricate scales; scales on anterior surface smaller, keeled, imbricate, grading into small, granular, juxtaposed scales, which cover distal portion of anterior side (near the ankle) and dorsal portion of shanks. Dorsal surface of hands, feet, and digits with imbricate, smooth scales, except on the posterodorsal surface of feet where 2���3 scales are keeled; palms and soles covered with small roundish or elongate scales, juxtaposed, not projecting anteriorly. Two enlarged thenar scales on inner margin of palm below pollex, two enlarged hypothenar scales on outer margin of palm. Subdigital lamellae protuberant, slightly divided basally on some digits, single on distal halves of most digits; lamellae rounded laterally. Subdigital lamellae as follows (Roman numbers indicate digits, Arabic numbers indicate paired or single lamellae on left/right; lower ungual sheath scale is omitted from the counts): hands I 4 / 4, II 8 / 8, III 14 / 13, IV 15 / 14, V 8 / 8; feet I 6 / 6, II 9 / 19, III 14 / 14, IV 20 / 20, V 11 / 11. Color of Holotype in life. Dorsum dark brown, with a black middorsal stripe (about two-three dorsal scales wide) extending from nape to tail, where it quickly becomes inconspicuous due to the dark coloration of the tail; subtle, irregular, tan dorsolateral line extending from nape to about midbody; flanks black, lower part of flanks peppered with reddish brown, and with some small pale brown spots ventrolaterally, forming an irregular ventrolateral line. Axilla region and underside of arms marbled with reddish brown and black. Head blackish brown with tan dorsolateral lines extending above eyes, from the anterior part of the loreals to the parietals. Tip of snout paler; diffuse paler blotches on lower part of lateral surface of head. Tympanum black, surrounded by some tan scales. Arms black, legs and tail blackish brown. Iris black (Fig. 5). Tongue dark gray. Palms and soles black. Chin, throat, chest, belly, and ventral surface of upper legs cream with dark mottling; lower legs ventrally reddish brown with dark mottling; tail ventrally black, except proximally where it is dirty white peppered with black (Fig. 8 A). Color of Holotype in preservative. After four months in 70 % ethanol, color pattern is similar to that in life. The dorsolateral line became white and is more conspicuous, the cream coloration of the ventral surface also became white. Distribution and ecology. The species is currently known only from the type locality, the summit of Maringma tepui in Guyana (Fig. 1). The only specimen available was collected in late morning (11 h00), during a sunny spell, in a large bromeliad (Brocchinia tatei), about 60 cm above the ground (Fig. 10). Possibly the specimen was hunting for some insects and spiders that became numerous once the sun chased away the mist generally present all over the summit. One other specimen was observed on the same day (in the afternoon) in another patch of bromeliads, but that specimen escaped. Interestingly, Gorzula and Se��aris (1998) and Fuentes and Rivas (2000) also reported collection of gymnophthalmids in Brocchinia tatei. This plant probably plays an important role as a refuge and food source for highland gymnophthalmids. Discussion. Arthrosaura hoogmoedi shares many morphological characteristics with A. testigensis from Los Testigos (Estado Bol��var, Venezuela) a group of tepuis east of Auy��n-tepui and situated about 180 km airline NWW from Maringma tepui. In having a black middorsal stripe, the new species resembles a putative new taxon (��� Arthrosaura species a���) reported by Gorzula and Se��aris (1998: 124) from the Chimant�� Massif (about 180 km airline W from Maringma tepui), unfortunately the absence of diagnostic characters and the poor quality of the photograph provided by Gorzula and Se��aris (1998, photograph # 89) does not allow further comparison. Without formally naming them, MacCulloch and Lathrop (2001) postulated the existence of two species groups of Arthrosaura, one group containing the species having four supraoculars, a relatively short body with relatively long legs (which I propose to name the kockii group), the other group containing species with three supraoculars, gracile body and relatively short legs (which I propose to name the reticulata group). Among the kockii group, the presence of keeled temporals in Arthrosaura kockii (a lowland species) and A. guianensis (an upland and highland species) might indicate a close relationship between these two taxa, while the smooth temporals could indicate a close relationship between A. hoogmoedi and A. testigensis, both being highland species. The completely opaque condition of the lower eyelid in Arthrosaura hoogmoedi is shared only with the recently described A. montigena from Auyantepui (Myers & Donnelly 2008) and is a rare trait within the family Gymnophthalmidae. It is unknown whether this is an adaptive character to high UV radiation and intense sunshine sometimes occurring on high elevation tepuis, or whether it is a plesiomorphic character state (secondarily lost in other species). Figure 7 shows a comparison of lower eyelid condition between A. hoogmoedi and two related Arthrosaura species (A. kockii and A. guianensis). Recently Myers and Donnelly (2008: 97) provided a key to the species of Arthrosaura. Unfortunately they inverted some characters of A. testigensis with those of A. guianensis and their key should be used with caution: interparietal and parietals are keeled in A. guianensis, but not in A. testigensis, and temporals are small in A. guianensis, but large in A. testigensis. The only other lizard collected on the summit of Maringma tepu, Published as part of Kok, Philippe J. R., 2008, A new highland species of Arthrosaura Boulenger, 1885 (Squamata: Gymnophthalmidae) from Maringma tepui on the border of Guyana and Brazil, pp. 1-15 in Zootaxa 1909 on pages 5-14, DOI: 10.5281/zenodo.184529, {"references":["Avila-Pires, T. C. S. (1995) Lizards of Brazilian Amazonia (Reptilia: Squamata). Zoologische Verhandelingen, Leiden, 299, 1 - 706.","MacCulloch, R. D. & Lathrop, A. (2001) A new species of Arthrosaura (Sauria: Teiidae) from the highlands of Guyana. Caribbean Journal of Science, 37, 174 - 181.","Donnelly, M. A., McDiarmid, R. W. & Myers, C. W. (1992) A new lizard of the genus Arthrosaura (Teiidae) from southern Venezuela. Proceedings of the Biological Society of Washington, 105, 821 - 833.","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: report from the Robert G. Goelet American Museum - Terramar expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147.","O'Shaughnessy, A. W. E. (1881) An account of the collection of lizards made by Mr. Buckley in Ecuador, and now in the British Museum, with descriptions of the new species. Proceedings of the Zoological Society of London, 1881, 227 - 245.","Burt, C. E. & Burt M. D. (1931) South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History, 61, 227 - 395.","Lidth de Jeude, T. W. van (1904) Reptiles and Batrachians from Surinam. Notes of the Lyden Museum, 25, 83 - 94.","Fuentes, O. & Rivas, G. (2000) Geographic distribution. Euspondylus goeleti. Herpetological Review, 31, 181 - 182."]}

Adelophryne patamona, sp. nov. Figs 1���7 Material. Holotype. ROM 43034, an adult male, from the north slope of Mount Wokomung, Guyana, 05�� 06' 35 " N, 059�� 49 ' 15 " W, 1234 m, collected by A. Lathrop, S. Khan and R. MacCulloch, 1 November 2004. Paratypes. ROM 43035, an adult female, from the north slope of Mount Wokomung, Guyana, 05�� 05' 33 " N, 059�� 50 ' 35 " W, 1411 m, collected by A. Lathrop, S. Khan and R. MacCulloch, 7 November 2004; IRSNB 1971 ��� 4, adult females, from the east base of Mount Maringma, Guyana, 05�� 12 ' 38 " N, 060�� 33 ' 59 " W, 1060 m, collected by P. Kok, P. Benjamin and C. Perry, 20 November 2007; IRSNB 1975, an adult female, IRSNB 1976 - 7, two juveniles and IRSNB 1978, an adult male from the southeast slope of Mount Maringma, Guyana, 05�� 12 ' 16 " N, 060�� 34 ' 39 " W, 1376 m, collected by P. Kok, P. Benjamin and C. Perry, 22���24 November 2007. Etymology. The specific epithet is a noun in apposition, and refers to the Patamona people, who live in the area where the holotype was collected. Diagnosis. A large species of Adelophryne, characterised by maximum SVL> 20 mm, second phalange of Finger IV longer than wide, tips of digits with small pointed discs, tympanum small, separated from eye by a distance equal to tympanum diameter, posterodorsal part of tympanum covered by a fold, and an inconspicuous glandular ridge from tympanum to arm insertion. The specimens can be placed in the genus Adelophryne because they possess the following combination of characters: small size, flattened digits with indistinct subdigital pads rather than subarticular tubercles, shortened Finger IV with single subdigital pad, discs with pointed tips and lateral grooves, vomerine teeth present, tongue narrow, tarsal tubercle absent. Of the other genera of small terrestrial frogs in the region, Noblella has a tarsal tubercle and no vomerine teeth, Phyzelaphryne has a broad tongue, round digits, and discs with circumferential grooves, and Pristimantis has two subarticular tubercles on Finger IV. The differences among these genera were elaborated upon by Hoogmoed and Lescure (1984). Adelophryne patamona can be distinguished from A. adiastola and A. pachydactyla by having 3 phalanges in Finger IV (2 phalanges in A. adiastola and A. pachydactyla) and large size, up to 23 mm SVL (largest A. adiastola 14 mm, largest A. pachydactyla 11.1 mm). Adelophryne patamona can be distinguished from A. baturitensis by the presence of subdigital pads and absence of subarticular tubercles (tubercles present, pads absent in A. baturitensis) and large size, up to 23 mm SVL (largest A. baturitensis 16.3 mm). Adelophryne patamona can be distinguished from A. maranguapensis by its posterior cloacal opening, without a transverse flap (flap present, cloaca opening ventrally in A. maranguapensis) and large size, up to 23 mm SVL (largest A. maranguapensis 17.4 mm). Adelophryne patamona can be distinguished from A. gutturosa, with which it occurs sympatrically, by the following characters: second phalange of Finger IV relatively long, 2 X its width (short, length equal to or slightly greater than width in A. gutturosa); canthus rostralis concave (straight); vomerine teeth 15���16 (2���8); large size, up to 23 mm SVL (largest A. gutturosa 16 mm) (Fig. 1). In the field these species can be easily distinguished by their distinctly different advertisement calls (see below). Characters useful for distinguishing among species of Adelophryne are in Table 1. Description of the holotype. A male, 17 mm SVL, head slightly longer than wide, as wide as adjacent part of body, head depth 50 % of width. Snout rounded in dorsal and lateral profile. Distance between snout tip and eye slightly less than eye diameter, eye-nostril distance slightly less than internarial distance, 2 X distance from nostril to snout tip, 75 % of distance from eye to snout tip. Canthus rostralis indistinct, rounded, concave; loreal region sloping steeply to upper lip. Nostrils not projecting, round, directed laterally. Interorbital distance 1.3 X internasal distance, 2 X upper eyelid width. Pupil horizontal. Temporal region vertical; tympanum small, distinct, round, 35 % of eye diameter, surrounded by a distinct annulus, partially obscured by a supratympanic fold, separated from eye by a distance equal to tympanum diameter. Low glandular ridge from tympanum to arm insertion barely visible. Choanae small, laterally placed; vomerine processes transverse, posterior to choanae, each bearing 15 teeth. Tongue narrow, sides parallel, attached anteriorly. Vocal slits small, flanking tongue, vocal sac large, subgular. Skin of dorsum, venter, and limbs smooth with scattered small tubercles; flanks and rear of thighs areolate. Cloacal opening above midpoint of thigh, directed posteriorly, dorsal flap absent. Hand with large flat oval palmar and thenar tubercles. Underside of fingers with large flat subdigital pads (1 ��� 1���2 ��� 1); no pads under ultimate phalanges. Fingers flattened, unwebbed. Tips of fingers bluntly pointed, bearing discs not wider than the adjacent phalange, discs laterally grooved. Finger lengths IColour of holotype in life. Ground colour of dorsum, flanks and dorsal surfaces of limbs medium brown. Central part of dorsum dark brown, bounded by a black middorsal ���)(���-shaped mark and interorbital bar. Dorsum with a few other small, scattered black marks. Loreal and temporal regions dark brown. A black bar extending from tip of snout through eye to flank; another black bar in the inguinal region. Dorsal surfaces of limbs with black crossbands. Throat, chest and underside of arms dark grey with white or pale blue spots. Venter, underside of legs and rear of thighs reddish-brown with white or pale blue spots. A black anal spot. Palpebrum not reticulated, a black band along its upper rim. Iris copper with a red ring around pupil (Fig 3). In preservative the venter becomes brownish-grey with white spots. Variation. The female paratype from Wokomung is the largest specimen collected, SVL 23 mm (Fig 4). Paratypes from Maringma (Figs. 5���7) are one male 18.4 mm SVL, five females 20.7���21.5 mm SVL, and two juveniles (11.8 and 11.9 mm SVL). Dorsal skin smooth to granular; ventral tubercles variable; vomerine processes each bearing 16���18 teeth in adults, 2���3 teeth in juveniles. Colouration is variable, not sexually dimorphic (Figs. 4���7). Black dorsal markings are variable; some specimens are darker with more pale blue spots and more conspicuous tubercles on dorsum. In the specimens from Maringma the throat, chest and underside of arms are black with white or pale blue spots and a few reddish-brown dots; the venter, underside of legs and rear of thighs are reddish-brown, sometimes with very few white or pale blue spots (Figs. 6���7). Juveniles resemble adults, but the upper arm is yellowish-orange and the dorsum has more pale blue spots; ventrally they are black with numerous pale blue spots (Fig. 5 C). Vocalisation (Fig. 8; Tables 2, 4). The following description is based on a sample of six advertisement calls from a single male, IRSNB 1978, recorded on the southeast slope of Mount Maringma (05�� 12 ' 16 " N, 060�� 34 ' 39 " W, 1376 m), on 24 November 2007 at 14 h 15, air temperature 19.8 ��C. Call parameter N Mean Range Call duration (s) 6 0.63 0.584���0.712 Inter-call interval (s) 5 3.85 3.365���4.255 Duration of first note (s) 6 0.038 0.035���0.042 Duration of second note (s) 6 0.056 0.055���0.057 Duration of third note (s) 6 0.429 0.400��� 0.497 Inter-note interval between the first and the second note (s) 6 0.075 0.072���0.081 Inter-note interval between the second and the third note (s) 6 0.052 0.047���0.059 Note repetition rate (notes/s) 12 9.07 8.66���9.26 Emphasized frequency (Hz) 18 3255 3251���3269 Temporal structure. The advertisement call of Adelophryne patamona consists of a group of three soft, unpulsed, whistle-like notes (two short, followed by one long) produced in rapid succession (Fig. 8). Although a feeble isolated single whistle-like note was sometimes heard, we never observed the specimen emitting that note, and thus we cannot confidently attribute it to A. patamona. However it is possible that, like in A. gutturosa (see below), a shorter call is emitted when a female is very close to a male. Males typically call antiphonally and there was no overlapping of the calls. Spectral structure. Three harmonics are developed, with the fundamental frequency dominating (range: 3251���3269 Hz; mean: 3255 Hz). The distribution of sound energy decreases progressively through the higher harmonics. The frequency rises abruptly at the very beginning of the notes (Fig. 8). The amplitude of the third note rises sharply to a peak at the beginning of the note then drops off rapidly, rising again, slowly, to a peak near the end of the note. A smaller peak in amplitude occurs at the end of the note (Fig. 8). Vocalisation parameters of three Adelophryne species are compared in Table 4. Pulses 2 4���5 high amplitude pulses followed by up 0 to 7 low amplitude pulses Emphasized harmonic Second First First Emphasized frequency (Hz) 3200���3700 3896���4979 3251���3269 Natural history. The specimens were collected at the end of the dry season, mostly at night, in leaf litter under medium-canopy forest (Fig. 5 D). One female (IRSNB 1975) and the two juveniles (IRSNB 1976 ��� 7) were collected among rootlets at the base of a tree. One male, IRSNB 1978, was calling in the afternoon under leaves in a hole between roots. Many males were heard calling during afternoons and nights but were very difficult to find in their hidden calling sites. No enlarged ova were found in the females. The species was also heard on Maringma at 1414 m, and in high-canopy forest near the village of Wayalayeng (05�� 14 ' 11 " N, 060�� 31 ' 04" W, 678 m), but none were collected at these locations. Distribution. Known only from the Pakaraima Mountains in western Guyana. The species likely occurs in Roraima State, Brazil, only 2 km from the Maringma location. The collecting locality at 1411 m is the highest elevation reported for any Adelophryne. Relationships. Adelophryne patamona appears to be most closely related to A. gutturosa, with which it is sympatric and sometimes syntopic (see below). Colour pattern may not be useful for distinguishing among species of Adelophryne. A dark ���)(��� -shaped dorsal marking is present in several Adelophryne (A. baturiensis, A. gutturosa and A. maranguapensis; Hoogmoed et al. 1994). The ventral pattern of dark background with light spots is also common in the genus., Published as part of Macculloch, Ross D., Lathrop, Amy, Kok, Philippe J. R., Minter, Leslie R., Khan, Samir Z. & Barrio-Amor��s, C��sar L., 2008, A new species of Adelophryne (Anura: Eleutherodactylidae) from Guyana, with additional data on A. gutturosa, pp. 36-50 in Zootaxa 1884 on pages 37-43, DOI: 10.5281/zenodo.274492, {"references":["Hoogmoed, M. S. & Lescure, J. (1984) A new genus and two new species of minute leptodactylid frogs from northern South America, with comments upon Phyzelaphryne (Amphibia: Anura: Leptodactylidae). Zoologische Mededelingen, 58, 85 - 115.","Hoogmoed, M. S., Borges, D. M. & Cascon, P. (1994) Three new species of the genus Adelophryne (Amphibia: Anura: Leptodactylidae) from northeastern Brazil, with remarks on the other species of the genus. Zoologische Mededelingen, 68, 271 - 300."]}

Allobates spumaponens sp. nov. Figs 1���6 Holotype. SMNS 12511, an adult male collected by Raffael Ernst, 23 /04/ 2003, Mabura Hill Forest Reserve, Upper Demerara-Berbice Region, Guyana (5 ��09'N, 58 �� 41 'W, elevation ca. 100 m). Paratypes (N= 6). SMNS 12510, an adult male collected 15 November 2002, SMNS 12512, an adult male collected 15 November 2002, SMNS 12513, an adult male collected 20 November 2002, SMNS 12514, an adult male collected 11 December 2002, SMNS 12515, an adult male collected 3 June 2003, SMNS 12516, an adult female collected 15 December 2002. All specimens with same locality data as holotype. Etymology. The specific epithet is derived from the Latin word spuma, meaning ���foam��� and the Latin verb ponere, meaning ���to place��� in reference to the atypical tadpole deposition site observed in the new species. Adult definition and diagnosis. A very small species of Allobates (males 13.2 ���17.0 mm SVL, females 17.0���19.0 mm SVL); body slender; Finger I longer than II when fingers appressed; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers appressed; no distal tubercle on Finger IV; Finger III not distinctly swollen in males; basal webbing only between Toes II���IV; lateral fringes on fingers and toes absent; throat in adult males pale with very discrete dark spotting on chin and laterally (visible under magnification only), throat in female pale, free of melanophores; belly in males hyaline-white, belly in females yellow; diffuse pale dorsolateral stripe present; diffuse, pale, partial oblique lateral stripe present; ventrolateral stripe present, with irregular dark brown blotches present ventrolaterally. Nine species of Allobates (sensu Grant et al. 2006) are reported from the Guiana Shield: A. brunneus (Cope, 1887) [although the presence of A. brunneus in the area is improbable (see Morales ��� 2000 ��� [2002], La Marca et al. 2004), we include it here to facilitate comparison], A. femoralis (Boulenger, 1884), A. granti, A. marchesianus (Melin, 1941) [several populations referred to as A. marchesianus represent a number of similar species (see Caldwell et al. 2002) and exact distribution of this species needs further investigation. Although the presence of A. marchesianus outside the type locality is uncertain, we include it here to facilitate comparison], A. myersi (Pyburn, 1981), A. rufulus (Gorzula, ��� 1988 ��� [1990]), A. sanmartini (Rivero, Langone and Prigioni, 1986), A. sumtuosus (Morales, ��� 2000 ��� [2002]), and A. undulatus (Myers and Donnelly, 2001) (Se��aris & MacCulloch 2005, Kok et al. 2006). Allobates spumaponens is easily distinguished from the more brightly coloured species of the A. femoralis group (A. femoralis, A. myersi and A. rufulus) by its cryptic colouration, smaller size (13.2 ���19.0 mm in A. spumaponens vs. 20.0��� 33.5 mm in A. femoralis, 26.4���32.8 mm in A. myersi and 20.0���23.0 mm in A. rufulus), pale throat in males (dark grey to black), and absence of mottling on chest or belly (present). Allobates brunneus (fide Cope 1887) differs by darker throat in males (pale in A. spumaponens), an ���hourglass��� pattern on the back (absent in A. spumaponens), Fingers I and II almost equal in length (Finger I much longer than II in A. spumaponens), incomplete lateral black band (complete in A. spumaponens), and call parameters. Allobates granti can be distinguished from A. spumaponens by the absence of a diffuse dorsolateral stripe (present in A. spumaponens), Finger II not reaching distal tubercle on Finger III (reaching in A. spumaponens), and call parameters. Allobates marchesianus differs in having grey to dark grey throat and chest in males (chest white and throat pale in A. spumaponens), light grey belly in males (white in A. spumaponens), wide distinct dorsolateral stripe (present but diffuse in A. spumaponens), and call parameters. Allobates sanmartini differs by larger size (maximum SVL of 25.4 mm vs. 19.0 mm in A. spumaponens), tympanum larger than half the size of eye (smaller in A. spumaponens), and Finger I shorter than second (much larger in A. spumaponens). Allobates sumtuosus differs in having Finger III swollen in males (not swollen in A. spumaponens), and lateral fringes on Toe II���IV (absent in A. spumaponens). Allobates undulatus is distinguished by larger size (maximum SVL 25.0 mm vs. 19.0 mm in A. spumaponens), swollen supracarpal pad atop wrist in males (absent in A. spumaponens), diffuse dorsolateral stripe absent (present in A. spumaponens), wavy-edged dorsal markings (absent in A. spumaponens), and different call parameters. A few extralimital species could be confused with Allobates spumaponens. These species are Allobates caeruleodactylus (Lima and Caldwell, 2001), A. crombiei (Morales, ��� 2000 ��� [2002]), A. masniger (Morales, ��� 2000 ��� [2002]), A. nidicola (Caldwell and Lima, 2003), and A. pittieri (La Marca, Manzanilla and Mijares- Urrutia, 2004). Allobates caeruleodactylus is distinguished from A. spumaponens by blue digits and blue discs on toes in males (never blue in A. spumaponens), tympanum about 50 % of eye length (less than 40 % in A. spumaponens), and different advertisement call. Allobates crombiei differs by larger size in males (17.4 ���19.0 mm vs. 13.2 ���17.0 mm in A. spumaponens), wavy-edged dorsal markings (absent in A. spumaponens), and presence of lateral fringes on toes (absent in A. spumaponens). Allobates masniger is distinguished in having a larger size in males (17.9���19.5 mm vs. 13.2 ���17.0 mm in A. spumaponens), lateral fringes on toes (absent in A. spumaponens), and dark grey belly in males (white in A. spumaponens). Allobates nidicola differs by a larger size in males (18.5���20.5 mm vs. 13.2 ���17.0 mm in A. spumaponens), a light grey to solid black throat and chest in males (chest white and throat pale in A. spumaponens), diffuse dorsolateral stripe absent (present in A. spumaponens), and different advertisement call. Allobates pittieri is distinguished from A. spumaponens by a larger size in males (16.3���18.3 mm vs. 13.2 ���17.0 mm in A. spumaponens), and black throat and grey belly in males (belly white, throat pale in A. spumaponens). Measurements of the holotype (in mm). SVL 15.7, HL 5.0, HW 5.0, SL 2.8, EN 1.4, IN 2.1, EL 2.4, IO 2.0, TYM 0.7, FAL 3.5, HAND 3.4, WFD 0.5, TL 7.5, FL 6.4, WTD 0.6. Description of the holotype. An adult male (Figs 1���2); body slender; head as wide as long; head length 31.8 % SVL; snout broadly rounded in dorsal view, acutely rounded in lateral view, extending past lower jaw, snout 56.0% head length. Nares located laterally, opening posterolaterally; canthus rostralis rounded, loreal region slightly concave, flaring slightly at upper lip; internarial distance 42.0% head width; eye-naris distance 28.0% head length, 58.3 % eye length. Tympanum subcircular, directed posterolaterally, 29.2 % of eye length; separated from eye by a distance equal to 25.7 % of tympanum diameter; supratympanic fold absent; tympanic annulus visible anteroventrally; posterodorsal aspect of tympanum barely visible; anteroventral margin of tympanum distinct. Tongue attached anteriorly, broadly rounded posteriorly, median lingual process absent. Choanae small, circular, lateral. Vocal slits long, lateral. Small teeth present on maxillary and premaxillary, dentigerous process of vomers absent. Cloacal tubercles absent; vent at level of upper thighs; a small anal flap above vent. Dorsum shagreen with some larger granules, more granular posteriorly; belly smooth. Forelimb slender, skin smooth; metacarpal ridge absent; ulnar fold absent; hand length 21.7 % SVL; Finger I longer than Finger II when fingers appressed; fingers unwebbed, lateral fringes absent; Finger III not distinctly swollen; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers appressed; tip of Finger II reaching distal subarticular tubercle on Finger III when fingers appressed; terminal discs slightly expanded, slightly wider than long, equal to or slightly larger than the width of digit; width of disc on Finger III 0.50 mm; discs with distinct dorsal scutes. Relative lengths of appressed fingers III> I> II> IV; palmar tubercle large, ovoid, 0.60 mm in diameter, 17.6 % hand length, anterior third pigmented; thenar tubercle protuberant, ovoid, anterior quarter pigmented, about half the size of the palmar tubercle and narrowly separated from it. One subarticular tubercle on Fingers II and IV; two subarticular tubercles on Finger I and III; basal subarticular tubercle on Finger I largest, subarticular tubercle on Finger II and basal subarticular tubercle on Finger III subequal; basal subarticular tubercle on Finger IV and distal subarticular tubercle on Finger III smaller, distal subarticular tubercle on Finger III smallest (Fig. 3). Hindlimb robust, skin granular; tibia length 47.8 % SVL; heels overlapping when hindlimbs are flexed at right angles to sagittal plane of body; foot length 40.8 % SVL; relative length of appressed toes IV> III> V> II> I; Toe I very short, its tip reaching the base of subarticular tubercle on Toe II when toes appressed on left foot, not reaching the base of subarticular tubercle on Toe II when toes appressed on right foot; discs on Toes II, III, IV, and V larger than width of toes; disc on Toe I equal to width of digit. Width of disc on Toe IV 0.60 mm; rudimentary webbing present only between Toes II���IV; webbing slightly pigmented; lateral fringes absent. Toe webbing formula II 2 +- 3 �� III 3 - - 4 IV. Inner metatarsal tubercle oval, 0.50 mm in length, distal portion pigmented, outer metatarsal tubercle round, 0.26 mm in diameter, entirely pigmented; medial metatarsal tubercle present on both feet, unpigmented, subequal to inner metatarsal tubercle. Two subarticular tubercles present on Toes III and V, three on Toe IV and one on Toes I and II on the right foot; distal subarticular tubercle on Toe III unpigmented and barely visible on the left foot. Subarticular tubercles on Toes I and II largest; basal subarticular tubercle on Toe IV very small on both feet. Metatarsal fold absent. Tarsal keel well defined, short, tuberclelike, directed transversely across tarsus, located 1.41 mm from proximal edge of inner metatarsal tubercle, not extending from metatarsal tubercle (Fig. 3). Colour of Holotype in life. Dorsal ground colour medium brown with several dark brown flecks on dorsum; dorsal surface of head slightly darker than dorsum. Wide lateral black band from tip of snout to vent, containing 2 / 3 of tympanum and not tapering posteriorly from axilla. Pale, narrow, diffuse dorsolateral stripe above the band, extending from tip of snout to vent. Diffuse, pale, partial oblique lateral stripe imbedded in the black band, extending from groin to about one-quarter of the distance to arm insertion. Flanks white with a few irregular brownish grey blotches; ventrolateral stripe present. Upper lip white, suffused with barely visible tiny melanophores. Throat hyaline white with barely visible tiny melanophores laterally and on the chin. Belly immaculate hyaline white. Upper surfaces of thighs and shanks brown with dark brown blotches, no crossbars discernible. Pale paracloacal mark present. Pale yellow spot on groin. Forelimbs light brown with a dark brown line posteriorly from elbow to wrist and a short dark brown line anteriorly discontinuous from arm insertion to elbow. Toes greyish with some light grey blotches. Fingers light brown with some pale blotches. Palm and sole dark brown. Iris golden bronze. Colour of Holotype in preservative. Dorsum light brown, with two diffuse whitish dorsolateral stripes suffused with melanophores (melanophores visible under magnification), extending from snout to vent. Some small, irregular, dark brown spots are visible on the dorsum (these spots correspond to darker granules). Dorsal surface of head slightly darker than body. Lateral band blackish brown extending from tip of snout to vent. Diffuse, pale, partial oblique lateral stripe visible. Belly white, flanks white with a few patches of melanophores, upper lip white suffused with melanophores (visible under magnification). Throat white with a few melanophores located on the chin and laterally (visible under magnification only). Upper surfaces of thighs and shanks same colour as dorsum with some dark brown spots. Pale paracloacal mark present. Forelimbs cream dorsally, brown with pale spots ventrally; the brown lines are still visible. Fingers cream, digits dark brown; palm and sole brown. Male secondary sexual characters. Males are smaller than females (mean SVL in males 16.16 mm; females 17.95 mm) and have a pale throat with a few melanophores located on the chin and laterally (melanophores visible under magnification only, throat immaculate in female). Under magnification melanophores are visible on lips in males, these melanophores are absent in females. Male ventral colouration is hyaline-white, that of adult females yellowish. Variation among type specimens. Descriptive statistics of the type series are given in Table 1. Excluding the sexual dimorphism, all of the paratypes conform to the description of the holotype. Two paratypes (SMNS 12515 and SMNS 12516) have a dark brown dorsum and are much darker than the other paratypes. The pale diffuse dorsolateral stripe is darker in SMNS 12515 and may sometimes be difficult to distinguish in other specimens (under magnification the stripe is always visible). In some specimens it is unclear whether the dorsolateral stripe originates from the tip of the snout or from the eye. The medial metatarsal tubercle occurs in four specimens (57 %). SMNS 12515 has a truncate snout in dorsal view, while it is broadly rounded in other type specimens. The number of small spots on the dorsum is variable (Fig. 4). Additional data taken during transect sampling. During visual transect sampling we measured the SVL of 82 individuals; 51 adult females, 21 adult males, and 10 juveniles. Average SVL in females was: 17.95 mm (SD = 0.54 mm, range = 17.0���19.0 mm, N = 51), in males: 16.16 mm (SD = 0.93 mm, range = 15.5 ���17.0 mm, N = 21) and juveniles ranged between 6.5 ���12.0 mm (mean = 10.10 mm, SD = 1.43 mm, N = 10). Tadpole description. Fourteen tadpoles were collected (SMNS 12608, subnumbered 1���14). The following description is of an individual in Stage 40, reared in the field (SMNS 12608 - 1, Fig. 5): Type 4 tadpole (Orton 1953); exotrophic; body skin with small pustules; total length 18.1 mm; body length 6.3 mm, 34.8 % of total length, 1.4 X body width, 2.3 X body height; body width 1.9 X height; body depressed, oval in dorsal view; snout round in dorsal and lateral view; eyes positioned and directed dorsolaterally; eye diameter 1.0 mm, equal to interorbital distance; interorbital distance less than internarial distance; nares positioned and directed anterodorsally; narial opening reniform in dorsal view; eye-naris distance 0.7 mm, equal to narissnout distance. Spiracular tube sinistral, short, projecting posterodorsally, its base located 54 % of body length from snout and 21 % of body height from venter. Vent tube not visible. Lateral-line system barely visible (see below). Caudal musculature highest at body-tail junction, tapering posteriorly, terminating anterior to tail tip; tail tip tapered, rounded; upper fin originating slightly posterior to junction of body and tail, gradually increasing in height to about midlength of tail, maintaining that height nearly to terminus; height of upper fin at midlength of tail 82 % of caudal musculature; comparable height of lower fin 27 % of caudal musculature; lower fin gradually increasing in height to just anterior to terminus; tail height 24 % tail length. Mouth ventral, oral disc not emarginate, laterally folded, width 1.4 mm. Teeth long, in a single row, each tooth with 15���17 cusps (Fig. 6), LTRF 2 (2)/ 3 (1). Tooth row A- 1 complete, A- 2 consisting of two short, widely separated rows at level of upper jaw sheath, A- 1 slightly longer than posterior rows; P- 1 slightly shorter than P- 2, both medially interrupted by a small gap (unnatural in P- 2); tooth row P- 3 shorter than P- 2 or P- 1. Marginal papillae moderately long, tapered, blunt-tipped, in a single row; broad median gap on upper labium approximately equal to the length of A- 1; submarginal papillae absent; jaw sheaths typical, thin, serrated, lower jaw sheath broadly V-shaped, upper jaw sheath with long slender lateral processes (Fig. 6). Variation of 14 meristic characters from tadpoles in stage 28���40 is given in Table 2. The only obvious variation is in the P- 1 gap, which is fairly variable in size and is absent in some specimens; apparently this variation is unrelated to tadpole stage. Lateral-line system evident in a few specimens, barely distinct in most, with a line originating above anterior labium, extending posteriorly above snout, passing above the eye and extending along the upper part of the body to upper tail musculature; left and right lines slightly converging between nares. A second line originates approximately above posterior labium, extending to anterior corner of eye, or stopping at mid-distance between labium and eye. In one specimen a short line is visible on the left middle part of the body. In life, tadpoles have a greyish brown ground colour with irregular dark brown mottling on body, tail musculature and fin; a broken black stripe extends from naris to eye, followed by a short black stripe posterior to eye, another black stripe extends from mouth to eye; iris silvery gold. In preservative, upper surface of body grey-brown, sides of body with grey-brown mottling; an ill-defined dark brown stripe extends from naris to eye, followed by a short dark brown stripe posterior to eye; another dark brown stripe extends from oral disc to eye. Tail musculature and fin with grey-brown mottling. Venter translucent white, suffused with brown anteriorly, intestine visible. Advertisement call. The advertisement call of Allobates spumaponens comprises a series of short-pulsed high-pitched chirps (Myers & Daly 1976); there is only one pulse per note (Fig. 7). The call consists of two harmonics, the main harmonic (dominant frequency) at about 6500 Hz and an additional harmonic (fundamental frequency) at about 3100 Hz. Duration of a regular call sequence is difficult to determine as call sequences are usually relatively long continuous periods, i.e. a call sequence may last well over four minutes in which notes are emitted with varying frequencies. The number of notes per second ranges between two and four. After the emission of 2���4 calls, a somewhat longer break of 0.49 s (mean, SD = 0.11, range = 0.277��� 0.727 based on two sequences) may occur. Table 3 summarizes measurements for the main harmoni, Published as part of Kok, Philippe J. R. & Ernst, Raffael, 2007, A new species of Allobates (Anura: Aromobatidae: Allobatinae) exhibiting a novel reproductive behaviour, pp. 21-38 in Zootaxa 1555 on pages 23-35, DOI: 10.5281/zenodo.178208, {"references":["Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F. B., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262.","Cope, E. D. (1887) Synopsis of the batrachia and reptilia obtained by H. H. Smith in the Province of Mato Grosso, Brazil. Proceedings of the American Philosophical Society, 24, 44 - 60.","Morales, V. R. (\" 2000 \") [2002] Sistematica y Biogeografia del grupo trilineatus (Amphibia, Anura, Dendrobatidae, Colostethus), con descripcion de once especies nuevas Publicaciones de la Asociacion de Amigos de Donana, 13, 1 - 59.","La Marca, E., Manzanilla, J. & Mijares - Urrutia, A. (2004) Revision taxonomica del Colostethus del norte de Venezuela confundido durante largo tiempo con C. brunneus. Herpetotropicos, 1, 40 - 50.","Caldwell, J. P., Lima, A. P. & Keller, C. (2002) Redescription of Colostethus marchesianus (Melin, 1941) from its type locality. Copeia, 2002, 157 - 165.","Myers, C. W. & Donnelly, M. A. (2001) Herpetofauna of the Yutaje-Corocoro massif, Venezuela: second report from the Robert G. Goelet American Museum - Terramar expedition to the northwestern tepuis. Bulletin of the American Museum of Natural History, 261, 1 - 85.","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2, 63 - 75.","Myers, C. W. & Daly, J. W. (1976) Preliminary evaluation of skin toxins and vocalizations in taxonomic and evolutionary studies of poison-dart frogs (Dendrobatidae). Bulletin of the American Museum of Natural History, 157, 173 - 262.","Ernst, R., Konrad, T., Linsenmair, K. E. & Rodel, M. - O. (2007) The impacts of selective logging on three sympatric species of Leptodactylus in a Central Guyana rainforest. Amphibia-Reptilia, 28, 51 - 64.","Lotters, S., Reichle, S. & Jungfer, K. - H. (2003) Advertisement calls of Neotropical poison frogs (Amphibia: Dendrobatidae) of the genera Colostethus, Dendrobates and Epipedobates, with notes on dendrobatid call classification. Journal of Natural History, 37, 1899 - 1911."]}

Atractus tamessari sp. nov. Figs 1���4 Holotype. IRSNB 2640 (field n�� PK 1365), an adult male, collected along a tributary of Elinkwa River, ESE Kaieteur National Park, ca. 500 m elevation, Potaro-Siparuni District, Guyana (5 ��08���09���N, 59 �� 25 ��� 28 ���W), on 23 March 2006 by P. J. R. Kok, Paul Benjamin and Giuliano Seegobin. Paratypes. IRSNB 2641 (field n�� PK 1372), a subadult female collected on 24 March 2006, same data as the holotype. IRSNB 2642 (field n�� PK 1550), an adult female collected at the same locality as the holotype, in May 2006 by G. Seegobin. Etymology. The specific name is a noun in the genitive case, honoring Michael Tamessar, retired Senior Scientific Officer of the Department of Biology at the University of Guyana (UG). Mike Tamessar spent most of his life encouraging Guyanese students to pursue studies in the fields of ichthyology and herpetology. Mike also collected many of the amphibians and reptiles in the collection of the Centre for Study of the Biological Diversity (CSBD) at UG. Diagnosis. An Atractus species with 15 dorsal scale rows that can be distinguished from all other congeners by the following combination of characters: (1) medium size, adults reaching about 400 mm in TTL; (2) loreal much longer than high and touching the eye; (3) two postoculars; (4) eight supralabials, fourth and fifth touching the eye; (5) seven to eight infralabials, four in contact with chinshield; (6) six maxillary teeth; (7) 153���163 ventrals; (8) 28���31 subcaudals; (9) dorsal color pattern of irregular red or pale red markings, sometimes forming an incomplete broken dorsolateral stripe, on a medium brown to brownish black ground color; (10) venter yellowish cream, heavily mottled with brownish black. The new species can be referred to the genus Atractus because it possesses the following combination of characters: medium size; head small, slightly distinct from neck; eye small, with semi-elliptical pupil; scales smooth; dorsal scales in 15 rows without reduction; loreal present, touching the eye and separating prefrontals and supralabials; nostril between two scales; single pair of chinshields; anal single; subcaudals divided; maxillary teeth decreasing in size posteriorly; occurring in northern South America (Boulenger 1894, Hoogmoed 1980). By having 15 dorsal scale rows at midbody, Atractus tamessari differs from all Atractus species having 17 scale rows in the Guiana Shield (Silva Haad 2004, Avila-Pires 2005): A. alphonsehogei Cunha & Nascimento 1983, A. badius (Boie 1827), A. charitoae Silva Haad 2004, A. duidensis Roze 1961, A. favae (Filippi 1840), A. flammigerus (Boie 1827), A. latifrons (G��nther 1868), A. lucilae Silva Haad 2004, A. major Boulenger 1894, A. riveroi Roze 1961, A. schach (Boie 1827), A. snethlageae Cunha & Nascimento 1983, A. steyermarki Roze 1958, A. torquatus Dum��ril, Bibron & Dum��ril 1854, and A. zidoki Gasc & Rodrigues 1979. Among the few Atractus species with 15 dorsal scale rows found in the region, Atractus tamessari differs from A. elaps (G��nther 1858) by having two postoculars (one in A. elaps), eight supralabials (six), and no red or yellow rings on body (present); from A. franciscopaivai Silva Haad 2004 by having eight supralabials, three contacting the loreal (five supralabials, two contacting the loreal in A. franciscopaivai), seven to eight infralabials (five), two postoculars (one), 1 + 2 temporals (1 + 1), and a yellowish cream venter heavily mottled with brownish black blotches (banded black and orange or black and yellow); from A. heliobelluomini Silva Haad 2004 by having eight supralabials, three contacting the loreal (seven supralabials, two contacting the loreal in A. heliobelluomini), two postoculars (one), 1 + 2 temporals (1 + 1), and a yellowish cream venter heavily mottled with brownish black blotches (immaculate cream); from A. insipidus Roze 1961 by having eight supralabials, three contacting the loreal (seven supralabials, two contacting the loreal in A. insipidus), and a yellowish cream venter heavily mottled with brownish black blotches (immaculate cream); from A. poeppigi (Jan 1862) by having eight supralabials (six in A. poeppigi), 1 + 2 temporals (0+ 1 or 0+ 2), the loreal much longer than high (slightly higher than long), and a yellowish cream venter heavily mottled with brownish black blotches (reddish cream or red and black); and from A. trilineatus Wagler 1828 by having 28���31 subcaudal scales (11���19 in A. trilineatus), a yellowish cream venter heavily mottled with brownish black blotches (cream with light brown spots on posteriormost ventrals and anal), and by lacking well-defined longitudinal stripes (3���4 well-defined stripes present). Description of the holotype. An adult male, TTL 397 mm, SVL 346 mm, TL 51 mm (12.8% of TTL); head slightly distinct from neck, length 1.4 times width (HW: 7.6 mm; HL: 10.8 mm); snout length (from eye to tip of snout) 4.0 mm, snout rounded in dorsal and lateral view; eye small (1.6 mm); pupil semi-elliptical; eye-nostril distance 29 % of HL; rostral small, visible in dorsal view, width about 1.8 times height; internasals small, as long as wide, laterally in contact with anterior and posterior nasals; prefrontals large, length 1.4 times width, much larger than the internasals, irregularly hexagonal, touching eye, their medial suture asymmetrical, noticeably dextral to the medial internasal suture; frontal pentagonal, slightly wider than long, 1.1 times length of medial prefrontal suture; parietals large, length 1.5 times width, medial suture of parietals 1.2 times length of frontal. Nasal divided, posterior nasal as long as anterior nasal; preocular absent; loreal long, length 2.8 times height; postoculars 2 / 2, small, upper slightly larger than lower; temporals 1 + 2, lower posterior temporal slightly longer than upper; supralabials 8 / 8, eighth largest, first contacting nasals, second contacting posterior nasal and loreal, third and fourth contacting loreal, fourth and fifth touching eye, sixth contacting postocular and anterior temporal, seventh contacting anterior and posterior temporals, eighth contacting posterior temporal; infralabials 8 / 8, first four in contact with chinshields, first pair meeting at ventral midline, separating mental from chinshields; one pair of chinshields, length 2.5 times width (Fig. 1). Maxillary teeth 6; dorsal scales in 15 - 15 - 15 rows, smooth without apical pits; ventrals 153; anal single; paired subcaudals 31. Color of holotype in life. The dorsal ground color is brownish black with numerous irregular bright red and paler red markings. The tail is brownish black with a few bright red lateral markings. The top of the head is brownish black. The snout is lighter, with a slight yellow wash. The supralabials are mostly yellow with the upper third dark brown, except for the eighth, which is almost completely dark brown with a small yellow spot anteriorly, and the first and the second, which are speckled with dark brown. The infralabials are mostly yellow with the lower third and posterior portion of each scale dark brown. The chinshields are mostly yellow with dark brown anteriorly and laterally. The mental groove area has a few dark brown speckles. The venter is yellowish cream with many irregular brownish black blotches, fewer blotches anteriorly, heavily mottled posteriorly. Underside of tail is brownish black with a few small irregular yellowish cream markings. The iris is brown (Fig. 2). Color of holotype in preservative. After one month in preservative (70 % ethanol after formalin), the red markings have faded slightly but are still well visible. The yellow coloration became white. The dark markings are unchanged. Hemipenis. Almost fully everted right hemipenis about 10.5 mm long, extending to level of seventh subcaudal when adpressed to the tail; hemipenis differentiated (sensu Savage 1960), capitate, slightly bilobed, shallowly forked; lobes distinct from the base, each lobe nearly flattened on apical portion, displaying series of spinulate calyces; capitation stronger on asulcate side; sulcus spermaticus centrifugal, bifurcate, dividing about half of organ, at the base of apex; basal portion of hemipenial body covered by large spines (Fig. 3). Variation. The type series consists of the male holotype and two female paratopotypes. The paratopotypes strongly resemble the holotype with the following exceptions (those of holotype in parentheses): TTL from 368���430 mm; SVL from 332���386 mm; TL from 36���44 mm, 9.8���10.2% of TTL (12.8%); HL from 10.6���11.5 mm; HW from 7.5���8.3 mm; 28���29 subcaudals (31); 157���163 ventrals (153); upper postocular 1.8 times the lower one in IRSNB 2641 (only slightly larger in the holotype and in IRSNB 2642); seven infralabials in IRSNB 2642 (eight in the holotype and in IRSNB 2641), apparently due to the fusion of infralabials 5 and 6. The sexual dimorphism in the number of ventrals and subcaudals is typical for this genus (Savage 1960). Prefrontal and internasal medial sutures are asymmetrical in both sexes (see Hoogmoed & Prudente 2003 and Myers 2003 for further discussion). The most striking difference between the paratopotypes and the holotype occurs in dorsal pattern and coloration, which is fairly variable. IRSNB 2641 has a medium brown dorsum with irregular pale reddish brown markings and two ventrolateral black stripes (instead of irregular bright red markings on a brownish black ground without ventrolateral stripes in the holotype and IRSNB 2642). In IRSNB 2642 the bright red markings form an incomplete broken dorsolateral stripe. The head of each paratype is medium brown and, as in the holotype, the snout is lighter with a yellow wash. The pale reddish brown markings on the dorsum of IRSNB 2641 disappeared in preservative, resulting in a uniform medium brown dorsum. Ventral patterns are similar, those of the male holotype slightly more mottled than the female paratopotypes (Fig. 4). Distribution and ecology. Atractus tamessari is known only from the type locality (Fig. 5), which is located in the southeastern part of Kaieteur National Park, at the eastern edge of the Pakaraima Mountains (Fig. 6). The holotype and IRSNB 2641 were collected at night (ca. 22: 30 h), crawling on the ground a few meters from a stream, almost exactly at the same place and the same time, at a 24 h interval. IRSNB 2642 was collected by day, apparently sleeping, on a stump in the river. The habitat can be characterized as a submontane forest of the Pakaraima uplands on white sands dominated by clump wallaba (Dicymbe spp) (ter Steege 2001). Examination of stomachs revealed no identifiable remains of prey, although X-rays of the male body showed the presence of sand in the digestive tract, which may suggest the recent ingestion of an earthworm, a very common food of Atractus (Martins & Oliveira 1993, ��� 1998 ��� [1999]). No enlarged follicles were found in IRSNB 2641 ovaries; one egg was found in the abdominal cavity of IRSNB 2642, but as the specimen was cut in two by the collector, it is highly probable that other eggs were lost. All specimens have abdominal fat deposits. Other snake species collected in the immediate vicinity are the colubrids Chironius fuscus, Erythrolamprus aesculapii, Dipsas catesbyi, Dipsas cf. pakaraima, Helicops angulatus, Pseudoboa coronata, and the viperid Bothrops brazili., Published as part of Kok, Philippe J. R., 2006, A new snake of the genus Atractus Wagler, 1828 (Reptilia: Squamata: Colubridae) from Kaieteur National Park, Guyana, northeastern South America, pp. 19-35 in Zootaxa 1378 on pages 20-25, DOI: 10.5281/zenodo.175024, {"references":["Boulenger, G. A. (1894) Catalogue of the Snakes in the British Museum (Natural History), Volume II. Taylor and Francis, London, 382 pp.","Hoogmoed, M. S. (1980) Revision of the genus Atractus in Surinam, with the resurrection of two species (Colubridae, Reptilia). Zoologische Verhandelingen, 175, 1 - 47.","Silva Haad, J. J. (2004) Las serpientes del genero Atractus Wagler, 1828 (Colubridae, Xenodontinae) en la Amazonia Colombiana. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 28, 409 - 446.","Avila-Pires, T. C. S. (2005) Reptiles. In: Hollowell, T., and Reynolds, R. P. (Eds.), Checklist of the terrestrial vertebrates of the Guiana Shield. Bulletin of the Biological Society of Washington, 13, pp. 24 - 40.","Cunha, O. R. & Nascimento, F. P. (1983) Ofidios da Amazonia XX: as especies de Atractus Wagler, 1828, na Amazonia oriental e Maranhao (Ophidia, Colubridae). Boletim do Museu Paraense Emilio Goeldi, 123, 1 - 38.","Boie, F. (1827) Bemerkungen uber Merrem's Versuch eines Systems der Amphibien. Marburg. 1820. Erste Lieferung: Ophidier. Isis von Oken, 20, 508 - 566.","Roze, J. A. (1961) El genero Atractus (Serpentes: Colubridae) en Venezuela. Acta Biologica Venezuelica, 3, 103 - 119.","Filippi, F. De (1840) Catalogo ragionato e descrittivo della raccolta de serpenti del Museo dell' I. R. Universita di Pavia. Biblioteca Italiana, 99, 65 pp.","Gunther, A. (1868) Sixth account of new species of snakes in the collection of the British Museum. Annals Magazine of Natural History, 4, 413 - 429.","Gasc, J. - P. & Rodrigues, M. T. (1979) Une nouvelle espece du genre Atractus (Colubridae, Serpentes) de la Guyane francaise. Bulletin du Museum national d'Histoire naturelle de Paris, 1, 547 - 557.","Gunther, A. (1858) Catalogue of the colubrine snakes in the collection of the British Museum. Trustees of British Museum, London, 281 pp.","Jan, G. (1862) Enumerazione sistematica delle specie d'ofidi del gruppo Calamaridae. Archivio per la Zoologia, l'Anatomia e la Fisiologia, 2, 1 - 76.","Wagler, J. G. (1828) Auzuge aus seinem Systema Aphibiorem. Isis von Oken, 21, 740 - 744.","Hoogmoed, M. S. & Prudente A. L. C. (2003). A new species of Atractus (Reptilia: Ophidia: Colubridae: Dipsadinae) from the Amazon forest region in Brazil. Zoologische Mededelingen, 77, 425 - 440.","Myers, C. W. (2003) Rare snakes - Five new species from eastern Panama: reviews of northern Atractus and southern Geophis (Colubridae: Dipsadinae). American Museum Novitates, 3391, 1 - 47.","ter Steege, H. (2001) National Vegetation map of Guyana, Guyana Forestry Commission: Georgetown.","Martins, M. & Oliveira, M. E. (1993) The snakes of the genus Atractus Wagler (Reptilia: Squamata: Colubridae) from the Manaus region, central Amazonia, Brazil. Zoologische Mededelingen, 67, 21 - 40.","Martins, M. & Oliveira, M. E. (\" 1998 \") [1999] Natural history of snakes in forests in the Manaus region, Central Amazonia, Brazil. Herpetological Natural History, 6, 78 - 150."]}