Dryophytes leopardus Shimada et Matsui 2025, sp. nov.

Dryophytes leopardus Shimada et Matsui sp. nov. (English name: East Japan tree frog) (Japanese name: Higashi-Nihon-Ama-gaeru) (Fig. 8) urn:lsid:zoobank.org:act: 107E307B-3F40-4C3D-8F5A-4E4E890C9AC0 Hyla viridis?: Hallowell 1861, p. 500. “Simoda, Japan ” (=Shimoda, Shizuoka, Japan) Hyla arborea (part): Hilgendorf 1880, p. 120. “Yeso” (= Hokkaido, Japan) Hyla arborea var. japonica: Bedriaga 1889, p. 475, 487. “ Tokyo ” and “Yezo” (= Hokkaido, Japan) Hyla japonica: Camerano 1879, p. 895. “ Tokio (Giappone)” (= Tokyo, Japan) Hyla arborea japonica: Sclater 1892, p. 32. “Yezo, Japan ” (= Hokkaido, Japan); “Kiga, Japan ” (=Hamamatsu, Shizuoka, Japan) Hyla stepheni (part): Nikolski 1905, p. 401. Jesso (= Hokkaido, Japan) Hyla new taxon A: Dufresnes et al. 2016 Dryophytes japonicus (part): Duellman et al. 2016, p. 23 Holotype. KUHE 66109 (former AUEZ 3999), an adult male from Yoshiwara, Toyota City, Aichi Prefecture, Japan (35°00’56’’ N, 137°05’38’’ E, 10 m asl), collected on 25 October 2023 by Tomohiko Shimada. Paratypes. KUHE 66111 (former AUEZ 4001) one adult male and KUHE 66110, 66112–66113 (former AUEZ 4000, 4002–4003), three adult females, data same as the holotype. Referred specimens. KUHE 8812, 29190–29191, 37629, Aomori Prefecture; KUHE 29240, 29244, 29246, 32922, 32925, Iwate Prefecture; KUHE 39830, Miyagi Prefecture; KUHE 40056, T3124, Akita Prefecture; KUHE 13131–13132, 13383, 18480, 18485, Yamagata Prefecture; KUHE 43347, Fukushima Prefecture; KUHE 36528, Tochigi Prefecture; KUHE 28075–28076, Ibaraki Prefecture; KUHE 36146, 38829–38830, 38832, 39551–39552, 39554–30555, AUEZ 4026, 4036, 4039–4040, 4048, 4068, 4076, 4089–4091, 4099, Niigata Prefecture; KUHE 7950, 33017, Ishikawa Prefecture; KUHE 16099, 30510–30511, 30513, 42971–42973, 43010, AUEZ 3760–3761, Gifu Prefecture; KUHE 37382, 39976, Shizuoka Prefecture; AUEZ 2616, Aichi Prefecture; AUEZ 13455–13456, 25559, 42928–42929, 42934, 42936, 42938, 42941, Mie Prefecture, KUHE 24152, 25433, 33109, 34913, 39940, 41350–41351, Shiga Prefecture; KUHE 43235–43236, Kyoto Prefecture. Etymology. The specific epithet is a Greek (and Latin) noun denoting a leopard, alluding to the color pattern of the rear of thigh observed in this species. Diagnosis. A moderate-sized species of the genus Dryophytes, with adult SVL 25–35 mm in males and 27–42 mm in females. It shares most morphological characters with its sister species, D. japonicus, but usually differs in having the black and/or white patterns on the rear of thigh, and they definitely differ in mitochondrial and nuclear genome characters. It differs from other congeneric species in East Asia, D. suweonensis and D. immaculatus, in the presence of finger webbing and more developed toe webbing, as well as in the pattern of the rear of thigh. Description of holotype (in millimeters). Snout-vent length (SVL) 30.9; body robust; head short, wider (HW 12.2, 39.4%SVL) than long (HL 10.3, 33.4%SVL); snout truncate, tip rounded in dorsal outline; projecting beyond lower jaw, slightly rounded in lateral profile; canthus sharp; lore vertical, concave; nostril below canthus, midway between tip of snout (S-NL 1.5, 4.9%SVL) and anterior margin of upper eyelid; internarial distance (IND 2.5, 8.0%SVL) subequal to nostril to eye (N-EL 2.3, 7.3%SVL); eye large, length (EL 4.2, 13.5%SVL) 1.8 times eyenostril distance, equaling to snout length (SL 4.2, 13.5%SVL); interorbital (IOD 3.0, 9.7%SVL) wider than width of upper eyelid (UEW 2.7, 8.7%SVL) and internarial distance; pineal spot invisible; tympanum large and distinct, nearly circular (TDv=TDh 1.9, 6.0%SVL), about half eye diameter; vomerine teeth in indistinctly oval, small, and slightly oblique raised series (each of 3 teeth), the center posterior to line connecting posterior margins of choanae, connected with each other, but widely separated from choanae; tongue narrow anteriorly, moderately notched, without papilla; a pair of internal vocal sacs and vocal openings on corners of mouth. Forelimb stout (forelimb length 18.6, 60.1%SVL; LAL 15.0, 48.6%SVL); webbing poorly but distinctly developed; finger length formula: I<II<IV<III (Fig. 8C), second finger as long as third; finger tips with round adhesive discs having circummarginal grooves; disk of third finger largest (3FDW 1.23, 4.0%), two thirds of tympanum diameter; indistinct palmar tubercles and supernumerary tubercles present; subarticular tubercles prominent, circular; indistinct nuptial pads on dorsal, medial, and ventral surfaces of first finger extending from its base to subarticular tubercle, covered with minute yellow asperities; inner palmer tubercle distinct, oval (IPTL 1.7, 5.5%SVL), subequal to tympanum diameter; indistinct skin fold present in outer side of forelimb, starting from disk of fourth finger to elbow. Hindlimb long (HLL 45.8, 148.6%SVL), about 2.5 times the length of forelimb; tibia (TL 13.8, 44.7%SVL) shorter than foot (FL 13.8, 44.8%SVL); heels do not touch with each other when limbs are held at right angles to body; tibiotarsal articulation of adpressed limb reaching posterior corner of eye; toe tips with round adhesive discs having circummarginal grooves; toe length formula I<II<III<V<IV; third toe subequal to fifth; toes moderately webbed, formula I 1–2 II 0–2 1/2 III 0–2 IV 2–1 V (Fig. 8D); excision of membrane between two outer toes reaching middle subarticular tubercle of fourth when toes in contact; webs thick, not crenulate; subarticular tubercles prominent, rounded; inner metatarsal tubercle distinct, oblong (IMTL 1.2, 4.0%SVL), equaling to 3FDW, less than half length of first toe (1TL 3.5, 11.2%SVL); outer metatarsal tubercle small but distinct; indistinct tarsal fold present, connecting tibiotarsal articulation to inner metatarsal tubercle. Two distinct skin folds connecting both arms present on pectoral region, starting from anterior and posterior edges of arms, respectively. Head and dorsum smooth; no dorsolateral fold; a supratympanic fold from eye, curving to axilla; ventral and ventrolateral side of trunk and ventral side of thigh coarsely granular. Color in life. Dorsum green fringed with a narrow yellowish white line on canthus, eyelid, supratympanic fold, and lateral side of trunk; a dark brown band running below canthus and supratympanic fold, vaguely expanding to lateral side of trunk; lores green below canthus; upper lip white with dark dots; upper half of tympanum surrounded by the band below supratympanic fold; dorsal side of thighs marked with dark crossbars; rear of thighs marked with several white blotches with reticulated pattern formed by cluster of pigmentation; anterior half of ventrum white; vocal sac and posterior half of ventrum yellowish white; ventral surfaces of thigh and tibia yellowish pink. Variation. Females are significantly larger in SVL (mean ± SD = 33.0 ± 3.2 mm, n = 40) than males (30.4 ± 2.0 mm, n = 40; t-test, P <0.01). Size of N-LL, IND, and some characters of distal part of forelimbs (1FL, HAL, 3FDW, 1TL, and FL), all relative to SVL, tended to be greater in females than males (Mann-Whitney U-test, two-tailed, P <0.05). Similar sexual differences tended to be seen in 4TDW and LAL, but p-values were slightly larger than 0.05 (p=0.052 and 0.058, respectively). Males tended to have more developed toe webbing than females. In the second and the third toes, outer edge of web reach at toe disk in males, but reach at the first articulation in females. In the fourth toes, two phalanges are free from inner webs in males, while three phalanges are free in females. Although the center of vomerine teeth lays posterior to the line connecting the posterior borders of the choanae in the holotype, it lays anterior the line in a part of paratypic specimens (i.e. KUHE 66110), and the latter status seems to be common in this species (Matsui & Matsui, 1982). Eggs and larvae. In the topotypic population, Aichi Prefecture, eggs laid at a time ranged from 109–161 (mean ± SD = 124 ± 25, n = 4) (Motohiro Nakamura, personal communication). Eggs each 1.2 mm in diameter and light brown in animal hemisphere. Eggs are usually laid in small clumps. A total of eight tadpoles in stages 31–35 (total length [TOTL] = 25.8–38.3 [mean ± SD = 33.5 ± 3.8] mm, head body length [HBL] = 11.1–14.0 [mean ± SD = 12.3 ± 1.0] mm), and three in stages 36–41 (TOTL 33.8–40.9 [mean = 37.4] mm, HBL=14.5–15.3 [mean = 14.8] mm), from the type locality were closely examined. Head and body slightly flattened above, spheroidal below; head body width (HBW) maximum slightly anterior to level of spiracle 54–68% (median = 62%) of HBL; head body depth (HBD) 89–108% (median = 97%) of HBW; snout rounded; eyes dorsolateral, visible from below; nostril open, dorsal, rim raised, midway between tip of snout and eye; internarial 29–58% (median = 49%) of interorbital. Oral disk anteroventral, emarginate, width 31–43% (median = 39%) of HBW; marginal papillae on upper labium with wide gap; lower labium with a continuous row of papillae, submarginal papillae present near corners; denticles 2(2)/3 (Fig. 9D) or 2(1–2)/3; beaks with black outer margins; outer surface smooth; margin finely serrate; upper beak weekly convex medially; neither beak divided. Spiracle sinistral, tube pointing upward and backward, almost completely attached to body wall. Anal tube dextral, attached to ventral fin; loops of gut visible ventrally in young larvae. Tail moderately long and lanceolate, both margins weakly convex, tapering gradually to slightly rounded tip; tail length 178–219% (median = 197%; only specimens with non-damaged tail) of HBL, maximum depth 30–36% (median = 36%) of length; dorsal fin origin at midpoint of body, deeper than ventral fin except near tail tip and body; ventral fin origin continuous to vent; caudal muscle moderately strong, maximum tail width 36–51% (median = 44%) of HBW; muscle depth at anterior one-third of tail 34–45% (median = 41%) of tail depth, steadily narrowed posteriorly, shallower than either fin in distal ha