Japalura sagittifera Smith, 1940

Mahony, Stephen, 2009, A new species of Japalura (Reptilia: Agamidae) from northeast India with a discussion of the similar species Japalura sagittifera Smith, 1940 and Japalura planidorsata Jerdon, 1870, Zootaxa 2212, pp. 41-61 : 50-53

Japalura sagittifera Smith, 1940 .

( Figures 2 & 4 )

Lectotype (designated herein): BMNH 1946.8.13.97 formerly BMNH 1940.6.1.44 (male), type locality: “Pangnamdim, Triangle, Upper Burma ”, presented and collected by Ronald Kaulback, between 1937 and 1939.

Paralectotypes (designated herein): BMNH 1946.8.13.96 formerly BMNH 1940.6.1.43 (female), BMNH 1946.8.14.92–99 formerly BMNH 1940.6.1.45–50 and BMNH 1946.8.13.1–2 formerly BMNH 1940.6.3.34– 49 (5 males, 5 females), details as above for lectotype.

Other: BMNH 1974.844–845, “3000’, Pangnamdim, 27°42' N, 97°54' E, north Burma.” presented and collected by Ronald Kaulback.

Diagnosis: This species differs from all species of the genus Japalura by the following combination of characters: small size (SVL males 50–53.3 mm, females 56.5–63.9 mm), tail length/SVL ratio 205.1–231.3%, 7–8 SL, 7–8 IL, 53–58 MD, 17–20 4 FingLam, 25–27 4 ToeLam, tympanum concealed, axillary fold present, nuchal crest, gular fold and gular pouch absent, enlarged keeled dorsal scales present, body shape subquadrangular in cross section, dorsal colouration of males is primarily blue (= green in life).

Description of lectotype: All measurements are summarized in Table 1. Habitus subquadrangular in cross section, in profile snout steeply tapering to a pointed tip, rostral rectangular, approximately 3 times longer than deep, bordered by seven scales including the first SL and the nasal (right side), but separated from the nasal on the left by one small scale. Nostril vertically oval, small, laterally positioned, nasal large, irregularly shaped, bordered by seven scales (right side) including the rostral and SL one and two, seven smooth rectangular SL, scales of the loreal region heterogenous in size and irregularly arranged, flat, becoming weakly keeled below the orbit and on the temporal region, a row of three enlarged keeled temporal scales run posteriorly from the orbit. Scales surrounding the eye, small and granular. Tympanum concealed. Canthals and supraciliaries imbricate, forming a laterally protruding supraorbital ridge. Dorsal head and snout scales heterogenous in size, smooth anteriorly, becoming strongly keeled on the supraorbital and occipital region. Those at the rear of the head are slightly domed with a varying number of keels radiating from the center of the scale. Parietal eye oval, longitudinally oriented, clearly visible through the parietal which is subequal in size to contacting scales. Three enlarged, dome shaped scales are situated posterior to the parietal. A single upper and a pair of lower conical temporal spines with numerous keels radiating outwards from the apex of the spine. The upper and anterior most spine is slightly larger than the lower pair, which are in line with but do not contact the three enlarged temporals. A smaller spine is situated at the rear of the jowls. Seven smooth IL, mental sub-triangular, approximately as wide as long, bordered posteriorly by a pair of elongated postmentals, subequal in length to the mental but do not contact with each other, bordered posteriorly by a row of three enlarged chin shields, postmental pairs are flat. Remaining gular scales imbricate to subimbricate and weakly keeled, the largest are subequal or slightly smaller than those on the chest region. No transverse gular fold or apparent gular pouch present, however Smith (1940) states that a small gular sac is distinct in life. Ventral scales subimbricate, weakly keeled and arranged in regular rows. 53 MD, subequal in size to the largest dorsals, all with a strong median keel. An additional row of moderately to strongly keeled scales form a continuous dorsolateral ridge from the posterior temporal spines to beyond the insertion of the forearms. A second almost continuous row of keeled scales runs longitudinally on either side of the middorsal scale row. Dorsal scales are heterogenous in size, shape and texture but mainly consist of small flat scales and distinctly larger keeled scales which are variously arranged into five regular, posterior pointing “V” shaped ridges of which the apex is the middorsal scale row. A distinct shallow axillary oblique fold almost reaching the dorsolateral keel. The upper and lower forearms, hands and fingers are covered in longitudinal rows of strongly keeled distally pointed scales, those on the outer surfaces more strongly keeled grading into weakly keeled scales of the inner surfaces. Scales of the hind legs are highly heterogenous and consisting of small weakly keeled scales grading to large trihedral oblique spines. Ventral surfaces of the thigh, tibia, and dorsal and ventral surface of feet with keeled scales forming regular rows. Number of lamellae on the fingers: I(8), II(12), III(19), IV(17), V(12), and 25 4ToeLam. Scales on the tail are strongly keeled ventrally, grading to weakly keeled dorsally. Hemipenal bulge distinctly enlarged.

Colouration in ethanol: In accordance with Smith (1940), the original drawings of Ronald Kaulback show the blue colouration observable in preserved specimens as green in life. Dorsal surface of the head, body to the lower flanks, fore limbs, hind limbs and tail light blue. An oblique stripe extends from the center of the lower eyelid to the rear axis of the upper and lower mandible, white centrally bordered by dark brown. A short dark brown line on the lower half of the dorsolateral keel of the neck. Two parallel thin brown transverse stripes on the dorsal surface of the left forearm and both tibia, an additional thin brown line extends from the proximal spines of the posterior thigh to the base of the tail, not extending as far as the vent. Dorsal “V” shaped keels bordered posteriorly by a thin brown line, the posterior most backed by a lighter, whitish patch on either side of the middorsal keel. Entire ventral surface of the body, throat, tail and limbs is creamy white with dark brown speckling on the chin and torso and a thin brown midventral longitudinal stripe on the chest. Smith (1940) states that Ronald Kaulback’s original drawings show the male as having a scarlet patch on the throat in the position of the gular pouch, now completely absent in the preserved specimen.

Condition of type specimens: All specimens examined are in good condition and all fixed with their mouths open. The male lectotype is the only specimen that has no incisions made to the body. All other specimens examined have a single longitudinal midventral incision and appear to be stuffed with cotton wool. This wool was not removed to determine whether the specimens were entirely gutted or merely stuffed for the purpose of filling the body cavity to provide a more natural appearance to the specimens. All specimens appear to have varying amounts of unsloughed scales, primarily on the head of some but also on the bodies and extremities of others, giving a patchy appearance to the colouration.

Variation: Variation of meristic and mensural characters in type series of J. sagittifera are summarized in Table 1. Due to the small sample size examined of the type series, conclusions on sexual dimorphism can not be reliably made, however the difference of male proportions relative to females in this series are consistent with known sexual dimorphism in other species of Japalura . Differences include a longer hind leg length, smaller SVL and general colouration of males. The original description correctly states that the axillary fold is not present on the female (= paralectotype BMNH 1946.8.13.96), however one female paratype (BMNH 1946.8.14.99) does possess shallow oblique axillary folds which extend to the dorsolateral keels.

The male paratype (BMNH 1946.8.15.1) differs in colouration from that of the holotype in details of the oblique eye to jaw stripe whereby the white and anterior dark border of the stripe are less defined and the brown speckles and stripe is absent from the ventral surface of the trunk. Dark speckles on the chin virtually absent, however the central portion of the throat is lightly speckled. Brown transverse lines of the forearms and tibia are absent. Dorsal surface of the tail appears faded to a bluish-beige. Like the holotype, the head, body, limbs and tail appear unevenly marbled with brown which is the result of unsloughed dead scales. “V” oriented series of keeled dorsal scales indistinct and thin brown posterior borders absent. On the right side, nasal in contact with the first SL only and separated from the rostral by a single small scale. The shallow oblique axillary folds become apparent when its head is held in line with the body. The three females examined show a large variation in colouration. The female paralectotype (BMNH 1946.8.13.96) is light blue on the head, lores and upper temporal region, all enlarged keeled scales of the dorsum (including “V” ridges) and flanks are light blue. Dorsal surface of the upper and lower forelimbs with two light blue bands, the remaining surface is brown densely transfused with blue distally to the base of the fingers, which are plain brown. Dorsal surface of the hind limbs also brown and densely transfused with blue. Crus and foot each with a single light transverse band with dark edges. Remaining dorsal surface of the body and tail dark brown. Ventral colouration as on the holotype. The third to fifth dorsal “V” ridges do not meet on the middorsal line. Both nasals in contact with first SL only and separated from the rostral by a single small scale.

In contrast the paralectotype (BMNH 1946.8.14.99) possess no blue pigmentation and is instead patterned with various shades of light and dark brown. Additional patterns not discernable on BMNH 1946.8.13.96 include, a dark brown crescentic bar which transverses the supraorbital region, two dark bars extend from the upper eyelids to the supraciliaries, “V” shaped dorsal ridges are bordered posteriorly by a wide dark brown bar and tail mottled with dark and light brown. Right side, nasal in contact with first SL only and directly in contact with the rostral. The third female examined, paralectotype (BMNH 1946.8.14.97) shares with the males the primarily solid light blue colouration on the limbs, flanks and head, it differs however by its dorsal colouration which from the back of the head to the base of the tail between the dorsolateral keels, is a sharply contrasting plain pale brown. Large patches where the outer scale layer has been shed are a lighter creamywhite on the dorsum and tail, and blue on the head. Tail with two wide blue blotches anteriorly, the remainder of the tail speckled blue to grayish blue. Keeled “V” shaped ridges are indistinct. Dorsolateral keels greatly reduced and barely distinguishable. On the right side, nasal in contact with the first SL only and separated from the rostral by a single small scale.

Distribution: Currently known from the type locality “Pangnamdim, Dadung” [Nam Tamai Valley (27°42'N, 97°54'E), The Triangle, Upper Burma] ( Smith, 1940). Pangnamdim is located in Kachin State, northern Myanmar at ca. 1,166 m asl. Dadung could not be located on gazetteers but may refer to Dazung Dam (ca. 1,230 m asl) nearby Pangnamdim. A population of this species has recently been found in eastern Arunachal Pradesh, northeast India ( Kunte and Manthey, 2009).

Etymology: The specific epithet sagittifera is an adjective in feminine form referring to sagitta (“arrow”) and ferō (“carry or bear”). Smith’s choice of this name is not obvious from the description but may have reference to the prominent dorsal “V” arrangements of enlarged scales apparent on most individuals.

Comparison: J. sagittifera is morphologically most similar to J. otai sp. nov. and J. planidorsata . It can be distinguished from J. otai sp. nov. as described above, from J. planidorsata (in brackets) based on the following combination of characteristics: gular pouch present (vs. absent) in life, mature females SVL 56.5– 63.9 mm, males 46.4–56.3 mm (vs. female SVL 49.1–52.7 mm, males 33–38 mm), HL/SVL 28–29.2% (vs. 30.4–32.3%), longer tail in females TailL/SVL 208.2–231.3% (vs. female 148.2–148.6%), longer hind foot, FootL/SVL 37.7–42.1% (vs. 32.7–33.2%), longer 4ToeL/SVL, 22.1–25.8% (vs. 17.9–18%), less SL and IL 7– 8 (vs. 10–11 and 9–11 respectively), more 4ToeLam 25–27 (vs. 17–20) and presence (vs. absence) of green colouration on most individuals in life.

From congeners possessing a concealed tympanum, J. sagittifera differs from J. andersoniana by absence (vs. presence) of a nuchal crest and in males, body shape of males subquadrangular (vs. slightly compressed laterally), smaller adult SVL of males, 50–53.3 mm (vs. up to 75.6 mm), less MD, 53–59 (vs. 81). From J. variegata it differs by its smaller adult size, males SVL 50–53.3 mm, females 56.5–63.9 mm (vs. male 94.4 mm, females 59.6–95.4 Ota (1989c), up to 110 mm (Smith, 1935)), absence (vs. presence) of a raised nuchal crest on males and body shape of males subquadrangular (vs. slightly compressed laterally). From J. hamptoni by its smaller male adult size, SVL 50–53.3 mm (vs. to 75 mm), hind limb of males reaches beyond the snout (vs. the posterior edge of the eye), axillary fold does not extend onto the throat (vs. extends almost across the throat) and body shape of males subquadrangular (vs. laterally compressed). From J. yunnanensis , it differs by its smaller male adult size, SVL 50–53.3 mm (vs. 58.8–73.6 mm), higher number of MD, 53–58 (vs. 32–45) and shorter tail length, TailL/SVL 201.5–235.3% (vs. 265.6–299.3%) and from J. chapaensis by absence (vs. presence) of a longitudinal gular fold on males, fewer 4ToeLam, 25–27 (vs. 28–30), greater number of MD, 53–58 (vs. 35–37) and longer tail, TailL/SVL 201.5–235.3% (vs. 170.7%). From J. fasciata by absence (vs. presence) of a nuchal crest and wide light transverse band on the trunk and MD, 53–58 (vs. 35–38). From J. flaviceps by having longer hind limbs when extended forward reaching from the orbit to beyond the snout (vs. before the eye), absence (vs. presence) of a transverse gular fold, a longer tail length, TailL/SVL 201.5– 235.3% (vs. 167.3–200.8%) and greater number of MD, 53–58 (vs. 34–48). From J. splendida by absence (vs. presence) of a transverse gular fold and nuchal crest, and smaller adult size, males SVL 50–53.3 mm, females 56.5–63.9 mm (vs. male, 62.9–89.8 mm, females 71.7–76.1 mm). From J. micangshanensis and J. zhaoermii by absence (vs. presence) of a wide yellow longitudinal stripe, raised nuchal and dorsal crest in males, and male body subquadrangular (vs. moderately compressed), further from J. micangshanensis by having a longer hind limb length, reaching between the eye and snout (vs. to the neck) and from J. zhaoermii by absence (vs. presence) of a prominent green gular pouch in males. From J. batangensis by absence (vs. presence) of a wide yellow longitudinal stripe and male body subquadrangular (vs. moderately compressed).

From the east Asian insular Japalura species possessing a concealed tympanum, J. sagittifera differs from J. brevipes by having longer hind limbs when extended forward reaching the orbit to beyond the snout (vs. to the tympanic region or eye), greater number of MD, 53–58 (vs. 34–50) and shorter tail length, TailL/SVL 201.5–235.3% (vs. 241.7–268.1%), from J. luei by its general body shape, subquadrangular (vs. moderately compressed), smaller male adult size, SVL 50–53.3 mm (vs. 65.2–74.5 mm) and a greater number of MD, 53– 58 (vs. 38–44). From J. makii by absence (vs. presence) of a distinct banded dorsal pattern and subquadrangular body shape (vs. moderately compressed), longer hind limbs when extended forward reaching the orbit to beyond the snout (vs. to the tympanic region), smaller male adult size, SVL 50–53.3 mm (vs. 63– 78.8 mm) and a greater number of MD, 53–58 (vs. 38–41) and from J. polygonata and J. swinhonis by absence (vs. presence) of a wide, light lateral stripe and nuchal crest, males subquadrangular (vs. moderately compressed). The remaining species of Japalura : J. dasi , J. dymondi , J. grahami , J. kaulbacki , J., kumaoensis , J. major , J. tricarinata , and J. varcoae can be distinguished by possessing an exposed tympanum.

Figures

FIGURE 4. Dorsal view of J. sagittifera; (top) male (lectotype BMNH 1946.8.13.97), (bottom) female (paralectotype BMNH 1946.8.13.96).

FIGURE 2. J. otai sp. nov., male (holotype ZSI 25217); (A) profile view, (B) dorsal view of head, J. sagittifera, male (lectotype BMNH 1946.8.13.97); (C) profile view, (D) dorsal view of head. Images not to scale.