Microhyla berdmorei

Microhyla berdmorei (Blyth, 1856) Figures 6 A, B, 7 A, 8, 9, 10 A; Tables 3, 6, S 2 Common name. Berdmore’s narrow-mouthed frog. Synonymy and chresonymy. Engystoma berdmorei Blyth, 1856 “ 1855 ”: 720. Type (s): not stated. Type locality: “ Schwe Gyen ” (now Shwegyin) on the Sitang River, “ Pegu ” (now Bago Region), Myanmar; Sclater (1892: 23). Callula natatrix Cope, 1867: 192. Syntypes: MCZ 630 (= MCZ 1587), and MCZ A-153454 -57 (apparently renumbered), according to Barbour and Loveridge (1929) and Harvard University, USA data (http://mczbase.mcz.harvard.edu). Type locality: “ Near Rangoon, Burmah ” (now Yangon Region, Myanmar); Li et al. (2019: 568). Diplopelma (Engystoma) berdmorei — Günther (1868: 146). Diplopelma pulchrum Theobald (1868: 83, in part) (non Engystoma pulchrum Hallowell, 1861, now Microhyla pulchra; misidentification). Diplopelma berdmorei — Theobald (1873: 112). Microhyla berdmorii — Boulenger (1882: 166). Incorrect subsequent spelling. Microhyla (Engystoma) berdmorei — Mason (1882: 292). Microhyla (Engystoma) berdmorei — Boulenger in Mason (1882: 500). Microhyla berdmorei — Bourret (1942: 509, in part); Frank and Ramus (1995: 90, in part); Dutta (1997: 58–59, in part); Das and Dutta (1998: 64); Chanda (2002: 36-39, in part); Choudhury (2002: 277, in part); Devi and Shamungou (2006: 317–324); Sarkar and Ray (2006: 291, in part); Das and Dutta (2007: 154–181); Wogan et al. (2008: 88, in part); Dinesh et al. (2009: 302: in part); Mahony et al. (2009: 80–94); Mathew and Sen (2010: 65–66); Sengupta et al. (2010: 30); Bjiu et al. (2019: 103); Garg et al. (2019: 15, in part); Li et al. (2019: 566, in part); Nguyen et al. (2019: 549-580, in part); Ahmad and Mim (2020: 36–71); Gorin et al. (2020: 1–47, in part); Hakim et al. (2020: 1239-1268); Poyarkov et al. (2020 c: 1525 –1558, in part); Gorin et al. (2021: 97, in part); Kundu et al. (2021: 1586–1591, in part); Zug (2022: 30, in part); Raj et al. (2023: 57–61); Dinesh et al. (2023: 7, in part); Frost (2024, page “ Microhyla berdmorei ” in part). Microhyla (Microhyla) berdmorei — Dubois (1987: 3). Microhyla butleri (non Microhyla butleri Boulenger, 1900) — Lalremsanga et al. (2007: 348; misidentification). Microhyla sp. — Hasan et al. (2012: 162–172). Types. Not stated in the original publication; ZSIC 9718–20 (not examined by us) were listed as syntypes by Sclater (1892), although Blyth (1856) provided measurements for a single specimen. Garg et al. (2019) discussed the confusing status of the type specimens; see discussion below. Revised diagnosis. Microhyla berdmorei sensu stricto is characterized by a combination of the following morphological features: (1) large body size (SVL 33.0 mm in a male, 36.3-38.1 mm in females), with moderately slender and triangular body habitus; (2) head wider than long; (3) dorsal skin shagreened with occasional small tubercles; (4) snout rounded in dorsal and ventral views, obtusely pointed in lateral view; (5) first finger shorter than half of the second finger length; (6) finger tips with weak disks bearing wide and shallow dorsomedial grooves; (7) toes with distinct disks, each bearing a narrow and deep dorsomedial groove; (8) tibiotarsal articulation of the adpressed limb extending far beyond snout; (9) toe webbing reaching disks on all toes except toe IV; webbing formula: i 1-1 ii 1 - 2 iii 1 - 2 iv 2 - 1 v; (10) throat and chin dusty gray to almost black in breeding males; belly yellowish-white; (11) dorsal surfaces of forelimbs and hindlimbs with narrow prominent crossbars, up to 5–6 crossbars on thighs; (12) dark-brown patch above cloacal opening inverted-U-shaped or crescent-shaped; dark blotches on both sides of cloaca; (13) dorsum generally with distinct olive-brown “ teddy-bear ” - pattern edged with light brown; dark dorsal markings may be indistinct or completely absent; (14) body flanks with black irregular spots and blotches; (15) broad olive-gray lateral band extending from armpit to middle of trunk; (16) light postocular stripe yellowish-beige with no dark edging; (17) reddish spots on dorsum and dorsal surfaces of hindlimbs absent; (18) iris with black stripe below the pupil. Material examined. In this study, we used morphological data from five specimens of M. berdmorei sensu stricto from India (see Table S 2 for details). Description of the voucher specimen MZMU A-8003 (Fig. 9). An adult female in a good state of preservation. Body size large (SVL 36.4 mm; other measurements are presented in Table S 2). Body habitus slender, triangular, and dorsoventrally flattened (Fig. 9). Head triangular in dorsal view; wider than long (HW / HL ratio 1.19). Snout comparatively short and protruding (SL / HL ratio 0.46), rounded in dorsal view (Fig. 9 A), obtusely pointed in profile, noticeably extending beyond the edge of the lower jaw (Fig. 9 E). Eyes large, protruding in dorsal and lateral views, significantly shorter than the snout (EL / SL ratio 0.71), slightly longer than interorbital distance (IOD / EL ratio 0.90). Canthus rostralis distinct, rounded; loreal area slightly concave. Nostrils oval-shaped, with lateral orientation, situated below the canthus rostralis, slightly closer to tip of snout than to eye. Interorbital distance greater than internarial distance (IND / IOD ratio 0.86). Upper eyelid length less than the interorbital distance (UEW / IOD ratio 0.79). Tympanum concealed, supratympanic fold indistinct, marked by a series of low tubercles (Fig. 9 E). Tongue slender, rounded, free for the posterior four-fifths of its length; vomerine teeth absent. Eggs seen through the incision on the left lateral side of belly; eggs small, rounded, ca. 1.3–1.5 mm in diameter, pigmented (Fig. 9 B). Forelimbs short and slender (Fig. 9 A, B); lower arm elongated and thin (LAL / FLL ratio 0.85), hand less than half of forelimb length (HAL / FLL ratio 0.48). Fingers slender and elongated; finger webbing or skin fringes absent. First finger well-developed, notably less than half of the second finger length (Fig. 9 D); the relative finger length formula: I <II ≤ IV <III. Fingertips rounded, slightly expanded into disks notably narrower than the basal phalanges of the respective fingers. Finger tips lacking peripheral grooves bearing a wide and shallow median notch on the dorsal surface of each finger (Fig. 6 B). Finger tips almost equal in width, with the first finger tip slightly narrower. Subarticular tubercles on fingers distinct, large, rounded, and protruding; the distal subarticular tubercle on the fourth finger less distinct. The subarticular tubercle formula: 1, 1, 2, 2 (Fig. 9 D). Two metacarpal tubercles: inner metacarpal tubercle distinct, protruding, rounded in shape; outer metacarpal tubercle rounded and flattened, its diameter subequal to the diameter of inner metacarpal tubercle (OPTL / IPTL ratio 0.89). Hindlimbs long, slender, almost four times longer than the forelimbs (HLL / FLL ratio 3.85). Thighs muscular, massive (Fig. 9 B), shanks elongated and slender, comprising approximately one-third of the hindlimb length (TL / HLL ratio 0.35). When the limbs are positioned at the right angle to the body, the heels significantly overlap. Tibiotarsal articulation of the adpressed limb extends well beyond the tip of the snout. Foot length comprises more than one-third of the hindlimb length, being significantly shorter than the tibia (FL / HLL ratio 0.31, TL / FL ratio 1.16). The relative toe lengths: I <II <V ≤ III <IV. Shanks smooth, inner tarsal fold absent. Tips of all toes distinctly widened into round disks, notably wider than the finger tips (4 FDD / 3 FDD ratio 1.75). Toe webbing fully developed, reaching the disks on all toes except the toe IV; webbing formula: i 1-1 ii 1 - 2 iii 1 - 2 iv 2 - 1 v (Fig. 6 A). Subarticular tubercles on toes distinct, rounded, slightly protruding, subarticular tubercle formula: 1, 1, 2, 3, 2 (Fig. 9 D). Internal metatarsal tubercle of moderate size, slightly elongated, with indistinct margins, comprising less than half the length of the first toe (IMTL / 1 TOEL ratio 0.36, Fig. 9 C). Outer metatarsal tubercle small but distinct, rounded, prominent with well-defined margins, subequal to the length of inner metatarsal tubercle (OMTL / IMTL ratio 0.94, Fig. 9 C). Skin on dorsum shagreened, with rare small tubercles getting denser laterally (Fig. 7 A). Dorsal surfaces of forelimbs almost smooth, with the exception of tiny tubercles present on forearms. Dorsal surface of hindlimbs with small tubercles sparsely scattered on thighs and shanks (Fig. 9 A). Upper eyelid smooth. Body flanks smooth, numerous small warts around the tympanal region. Ventral sides of body and limbs completely smooth (Fig. 9 B). Cloacal opening unmodified, with posterior orientation. Coloration in life. Dorsum in life olive-brown with a bronze tint laterally with a dark-brown “ teddy-bear ” pattern (see Rakotoarison et al. 2017) with irregular borders (Fig. 7 A). Dark-brown triangular interorbital bar located between the posterior parts of upper eyelids continues posteriorly, sharply narrowing at the level of head base and broadening at the scapular area. A pair of dark-brown blotches edged with beige form a trapezoidal dark marking around the cloacal opening. A yellowish-beige light postocular stripe running from the posterior corner of the eye to the axilla (Fig. 7 A). Body and head flanks slightly darker than the lateral sides of the dorsum. Snout olive-brown lacking dark markings. Broad dark olive-gray band running posteriorly from the eye. Upper jaw dark-brown with a few irregular light-brown or cream spots below the eye. Narrow cream stripe running from the posterior corner of the eye to the axilla. Except for the upper arm, limbs dorsally with indistinct brownish crossbars and spots: two crossbars visible on lower arms, three crossbars on thighs and shanks. Fingers and toes with brown transverse dorsal bars. Belly and chest yellowish-white, with a few irregular grayish spots in chest area