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Additional file 2: Appendix. List of additional museum specimens examined.

Additional file 1: Table S1. µCT scan data for the holotype of Stefania maccullochi sp. nov. and for comparative species in the S. riveroi clade.

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 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 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. 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