Kalophrynus anya, Zug, 2015

Zug, George R., 2015, Morphology and Systematics of Kalophrynus interlineatus-pleurostigma Populations (Anura: Microhylidae: Kalophryninae) and a Taxonomy of the Genus Kalophrynus Tschudi, Asian Sticky Frogs, Proceedings of the California Academy of Sciences 62 (5), pp. 135-190 : 150-154

Kalophrynus anya Zug, sp. nov.

Northern Burmese Sticky Frog

Figure 5 , 11A–B.

TYPE MATERIAL.— HOLOTYPE: USNM 537420 , adult male from Chatthin Wildlife Sanctuary , San Maung Camp (~ 23°35´57˝N, 95°31´13˝E) elevation ~ 200 m, approximately 2 km WNW of Chatthin (town), Sagaing Division, Myanmar, collected 5 September 1998 by Htun Win.

PARATYPES: CAS 232488 , Moenyini , Indawgyi Wildlife Sanctuary headquarters (24°44´55.7˝N 96°20´52.6˝E) elevation 799 ft [~242 m], Kachin State, Myanmar; USNM 520321 , 523964–965 locality same as holotype ; USNM 523966 Chatthin Wildlife Sanctuary , approximately 5 km WNW (~ 23°36´06˝N, 95°31´13˝E) of Chatthin (town), Sagaing Division, Myanmar ; USNM 537419 , 537421–422 locality same as holotype .

DEFINITION.— Adult K. anya can be distinguished from all congeners by the following combination of characters: no size dimorphism, adults 34–47 mm SVL; HeadL/ SVL 30–32 %; tympanum about equal eye diameter, Tymp/EyeD 72–100 %; moderate short hindlimbs, HndlL/ SVL 132–147 %; short hindfoot, HndfL/ThghL 85–91 %; dimorphism of SnEye/HeadL (35–43 %) and EyeD/HeadL (31–37 %) with females smaller; strong supratympanic ridge dorsoposteriorly overhanging tympanic annulus; paired vomerine folds, each broad, smoothly undulatory with modest medial separation; single postorbital fold with five or six undulatory lobes; single broad buccal fold comprised of ten or more abutting rectangular lobes; forefeet without webbing; hindfeet webbed at base, highest between 3 rd and 4 rd toe, usually to base of 2 nd subarticular tubercle of 4 rd toe; head pattern variably with HeadMid 50 % present and DorsNap 33 % present; chin usually with paired, broad longitudinal bars; hindlimbs (HndlBr) seldom barred.

DESCRIPTION OF HOLOTYPE.— Moderately robust-bodied male (39.7 mm SVL) with well-developed, moderate-length fore- and hindlimbs (45 % ForarmL+HandL/ SVL, 119 % HndlL/ SVL); head ovate (25 % HeadL/ SVL, 30 % HeadW/ SVL) and continuous with body (no apparent constriction or enlargement at juncture of head and trunk). Body measurements are: HeadL 10.1 mm; HeadW 11.8 mm; SnEye 4.3 mm; NarEye 2.3 mm; EyeD 3.5 mm; Tymp 3.1 mm; ForarmL 9.8 mm; HandL 8.1 mm; ThghL 14.4 mm; CrusL 13.0 mm; TarsL 7.5 mm; HndfL 12.8 mm. Eye distinctly larger than tympanum (89 % Tymp/EyeD).

Skin strongly glandular with dorsal skin thickened and cloak-like extending from nape to posterior end of trunk, smooth surface, thickest anteriorly forming distinct supratympanic fold over dorsal and posterior edge of tympanic annulus and swollen mass above axilla; posteriorly glandular cloak merges imperceptibly into lateral trunk skin; limbs without enlarged glandular folds; chest and abdominal skin glandular and pebble-like with scattered unpigmented ‘pebbles’ on chest. Tongue large, elliptical, posterior two-thirds free. No vomerine teeth. Three sets of palatal folds on roof of mouth: vomerine paired, each long with smooth undulatory free edge and distinctly separated medially; postorbital single, indistinct, and free-edge undulatory; buccal, single continuous fold of 15 to 16 rectangular, abutting lobes, medial lobes larger than lateral ones. Fingers and toes well-developed and tips bluntly rounded; hand web free; foot basally webbed (WebIII.2 = 1, WebIV.1 = 0.5). Subarticular tubercles well developed on hand and foot ( Fig. 2A–B ); large, nearly circular palmar tubercle with small tubercle at base of each finger and only third finger with tubercle of free portion of digit; no nuptial pads or asperities evident; paired moderate-sized, circular metatarsal tubercles on edges, inner slightly smaller than outer; each toe with basal subarticular tubercle; third toe with single tubercle on free portion of digit, fourth with two tubercles, and fifth with one (hence total of two tubercles on fifth toe). Digital length formulae, hand 3>2≈4=1, foot 4>3>5>2>1.

COLORATION IN PRESERVATIVE.— Dorsum from tip of snout to thigh medium rufous brown with faded brown middorsal stripe on head, bifurcating at nape into diagonally diverging stripes; parasagittal stripes from nape to trunk also diagonal; laterally neck and trunk dark brown, fading ventrally; narrow white dorsolateral stripe evident only on neck and shoulder; face (loreal) dark brown at canthus rostralis, lighter brown to mouth; vague banding on dorsal of thighs. Venter dark brown to nearly black on chin to mid throat, fading gradually thereafter to creamy white in prepubic area; vague lighter longitudinal stripe from chin to anterior chest.

DESCRIPTION AND INTRAPOPULATIONAL VARIATION.— Moderate-sized adults, not sexually dimorphic although females average larger (mean 41.6, 36.4–47. 1 mm SVL) than males (40.7, 34.0–41. 3 mm). These differences are reflected in the other measurements: HeadL 9.9–13. 8 mm ♀♀, 9.6–10. 7 mm ♂♂; HeadW 10.5–14. 8mm ♀♀, 10.3–12.0 mm ♂♂; SnEye 3.7–5.0 mm ♀♀, 3.8– 4.4 mm ♂♂; NarEye 2.2–3. 5 mm ♀♀, 2.2– 2.7 mm ♂♂; EyeD 3.3–4. 6 mm ♀♀, 3.2–4.0 mm ♂♂; Tymp 2.0–3. 9 mm ♀♀, 2.5– 3.3 mm ♂♂; ForarmL 9.3–12. 1 mm ♀♀, 9.0–10. 6 mm ♂♂; HandL 7.0–10. 7 mm ♀♀, 8.1–9. 1 mm ♂♂; ThghL 13.1–17. 8 mm ♀♀, 11.7–15. 5 mm ♂♂; CrusL 11.7–15. 8 mm ♀♀, 12.1–13. 3 mm ♂♂; TarsL 7.0–10. 5 mm ♀♀, 6.9– 8.4 mm ♂♂; HndfL 11.3–14. 9 mm ♀♀, 10.3–13. 4 mm ♂♂. Body proportions differ between females and males, although not greatly so (all values are percent): HeadL/ SVL 25– 32 ♀♀, 25– 28 ♂♂; HeadW/HeadL 97– 118 ♀♀, 105– 117 ♂♂; SnEye/HeadL 35– 41 ♀♀, 40– 43 ♂♂; NarEye/SnEye 56– 65 ♀♀, 50– 61 ♂♂; EyeD/HeadL 31– 33 ♀♀, 34– 37 ♂♂;Tymp/EyeD 61– 93 ♀♀, 78– 89 ♂♂; Forarm/ SVL 24– 26 ♀♀, 22– 26 ♂♂; Forarm/CrusL 73– 85 ♀♀, 69– 80 ♂♂; HndlL/ SVL 110– 125 ♀♀, 119– 122 ♂♂; CrusL/ SVL 30– 32 ♀♀, 32– 33 ♂♂; CrusL/ThghL 81– 96 ♀♀, 86– 103 ♂♂; TarsL/ThghL 52– 62 ♀♀, 52– 59 ♂♂; HndfL/ SVL 27– 32 ♀♀, 30– 32 ♂♂; HndfL/ThghL 79– 96 ♀♀, 82– 88 ♂♂.

Tongue large, obovate, posterior half to two-thirds free. Vomerine teeth absent. Palatal fold morphology is moderately variable although these data are not quantified. The vomerine folds range from smooth-edged, undulatory folds to folds with broad lobular appearance ( USNM 523964, 537419, 537422); usually folds moderately separated on midline, rarely narrowly separated. Postorbital folds are usually well developed and higher than buccal folds; also most variable of three palatal folds and usually continuous although uncommonly narrow separated at midline; fold free edge ranges from smooth, undulatory to broad, round-edged lobes. Buccal fold is low, continuous with ten to 14 smooth-edged rectangular lobes.

Fingers lack webbing. Both finger and toe tips are bluntly rounded. Subarticular tubercles are well developed on digits; only third finger bears subarticular tubercle on free portion of digit (low on fifth); all fingers have a tubercle at their base and another row between a large, usually circular (rarely elliptical), nearly medial outer palmar tubercle. Nuptial pads not present. For hindfoot, each toe has a basal subarticular tubercle, third and fifth toes have an additional tubercle on free portion of digit, two tubercles on fourth toe (fifth toe of USNM 523965 lacks tubercles). Inner and outer metatarsal tubercles are present; inner is small to moderate-sized, nearly circular to elliptical; outer moderate-sized, circular to elliptical; inner and outer tubercles equal-sized to outer larger than inner. Toes lightly webbed WebIII2 median 1.0 (0.5–1.0), WebIV1 1.0 (0.5–1.0). Digit lengths nearly constant for fore- and hindfeet; finger formula 3>1≈2>4; toe formula 4>3>5>2>1.

Color pattern variation statistics for adults are (median and range): HeadMid 1, 0–2; HeadPsag 1, 0–2; DorsNap 1, 0–2; DorsPsag 1, 0–2; IngSpt 2, 0–2; HndlBr 0, 0–2; DlatSt 2, 1–2; Loreal 1, 0–2; LatTrnk 2, 0–2; Chin 1, 0–2; Chest 1, 0–2. Kalophrynus anya is highly variable, commonly with distinct middorsal stripe and faded parasagittal stripes on head, faded to distinct nape stripe in most individuals and bifurcating and quickly fragmenting or disappearing; inguinal spot almost always present and usually as ocelli, dorsolateral narrow white stripe rarely absent or indistinct, hindlimbs uncommonly barred, loreal area and lateral trunk usually uniformly dark, chin and throat always dusky and often with pair of darker longitudinal bars, and chest commonly dusky.

ETYMOLOGY.— All readers of Burmese cultural history are familiar with the country’s division into Upper and Lower Burma. This species occurs in the more northern portion of Upper Burma, and the Burmese a-nya tha refers to the human inhabitants of Upper Burma (auk tha, Lower Burma inhabitants). Because tha references human populations, I have taken the liberty of subverting the meaning of a-nya to the region of Upper Burma, hence the name for this Upper Burma endemic species.

DISTRIBUTION.— Myanmar. Kalophrynus anya is presently known from recent vouchers in east-central Sagaing Division (Chatthin) and southcentral Kachin State (Indawgyi) ( Fig. 6 ). Bourret (1942) reported specimens from Bhamo and Teinzo in Upper Burma. I have not located the source of these records; see discussion in the distribution section of the K. interlineatus account.

NATURAL HISTORY.— Many authors (e.g., Fei et al. 2010, Ohler and Grosjean 2005) report K. interlineatus and K. pleurostigma as a common frog and regularly calling in choruses. Our ( MHS teams) observations found K. anya an uncommon species, and we heard no choruses. Our assessment is that it is an uncommon species. We found individuals in open In ( Dipterocarpus tuberculatus ) forest with slight to moderate ground vegetation (grass), seasonally with heavy leaf litter.

Figures

FIGURE 5 (left). Images of holotype of Kalophrynus anya (USNM 537420, male, 39.7 mm SVL) in (A) dorsal, (B) lateral, and (C) ventral views [photographer, J.A. Poindexter].

FIGURE 11. Coloration of living representatives of pleurostigma-interlineatus group. (A) Dorsolateral view of a K. anya (USNM 520321) from the Chatthin Wildlife Sanctuary, Sagaing, Myanmar [photographer, C. Hansen]; (B) dorsolateral view of a K. anya (USNM 523966) from the Chatthin Wildlife Sanctuary, Sagaing, Myanmar [photographer, G.R. Zug]; (C) dorsolateral view of a K. interlineatus (CAS) from the Tanintharyi National Park, Tanintharyi, Myanmar [photograph- er, J.V. Vindum]; (D) dorsolateral view of a K. meizon (FMNH 242796) from Poring Station, Mt. Kinabulu Park, Ranua District, Sabah [photographer, R.F. Inger]; (E) dorsolateral view of a K. sinensis (UK 333148) from Agusan del Norte, Mindanao, Philippines [photographer, R.M. Brown]; (F) dorsolateral view of a K. pleurostigma (UK 333148) from Gunung Kunyit, Jambi, Sumatra [photographer, E.N. Smith].

FIGURE 2. Schematic ventral view of the right (A) fore- and (B) hindfoot of Kalophrynus anya sp. nov. (USNM 537420). Scale bar equals ~1 mm.

FIGURE 6. Burmese distribution of voucher specimens of Kalophrynus anya sp. nov. in northern Myanmar and K. inter- lineatus in southern and peninsular Myanmar. The three report- ed localities of K. orangensis are also shown; all lie within the Brahmaputra river valley. Solid circles denote specimens exam- ined in this study; open circles are literature or unexamined museum records; and stars represent type localities of available names.