Pseudocalotes rhammanotus Harvey, Hamidy, Kurniawan, Shaney & Smith

Harvey, Michael B., Hamidy, Amir, Kurniawan, Nia, Shaney, Kyle & Smith, Eric N., 2014, Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia, Zootaxa 3841 (2), pp. 211-238 : 226-235

Pseudocalotes rhammanotus Harvey, Hamidy, Kurniawan, Shaney & Smith new species

( Fig. 9 )

Holotype. An adult female ( MZB 10804) from montane forest along the ridge of a mountain south of Danau Ranau (= Lake Ranau), Lampung, Sumatra, Indonesia, 4.9394° S, 103.85292° E, 1237 m, 1954 hrs, 17 June 2013, found by Elijah Wostl, accompanied by other field party members from BC, UB, and UTA.

Diagnosis. A species of Pseudocalotes reaching at least 196 mm (67.9 mm SVL) and distinguished from congeners by the following combination of characters: (1) discrete interparietal present; (2) canthals six; (3) enlarged, heavily keeled to subpyramidal posttemporal and posttympanic scales present; postrictal modified scale absent; (4) gulars relatively small, homogenous, no transition to small scales on poorly developed gular pouch; (5) short antehumeral fold with noticeably smaller scales below it ( Fig. 7 ); (6) dorsolateral row of six heavily keeled scales; dorsolateral series reduced to single scale on neck; (7) scales on lower flanks feebly keeled and homogenous (8) 51 scales around midbody (9) dorsal crest of projecting scales extending to base of tail; all projecting scales of crest separated by medial contact between scales of paravertebral series; (10) ventrals smaller than dorsals; (11) subdigital lamellae at base of Toe III bicarinate with preaxial and postaxial keels equally developed; (12) dorsum brown and green, banded in unique holotype; (13) ventral body white with small brown spots densest laterally; (14) gular pouch white medially grading to dirty brown laterally; (15) tongue cream, throat black.

Comparisons. Pseudocalotes rhammanotus can be distinguished from other agamids on Sumatra by its clearly visible tympanum, enlarged row of scales below the orbit bordering the supralabials, relatively wide gap between the dorsal and nuchal crests, heterogenous dorsal squamation, tail about twice as long as SVL, keeled subdigital lamellae, dorsals larger than ventrals, absence of large spines in the nuchal and postorbital regions, relatively narrow head, and other characters of external morphology described in the differential diagnosis of Hallermann & Böhme (2000; see also Mahony 2010).

Three other congeners occur on Sumatra. Of these, Pseudocalotes rhammanotus is most similar to P. tympanistriga . However, unlike P. tympanistriga (characters in parentheses), P. rhammanotus has an antehumeral fold with noticeably smaller scales below it (fold and transition to smaller scales in this region absent), 6/6 heavily keeled scales between the limbs (4 or 5), a wide nuchal gap of 7 scales (narrow, 2 or 3 scales), fewer vertebrals (20 vs. 24–31, 27 ± 3, n = 5), vertebrals that project from the dorsum (vertebrals enlarged with upturned tips but without entire scale projecting from dorsum), fewer subdigital lamellae (24 vs. 26–32, 28 ± 3, n = 5), and fewer infralabials (8 vs. 9–11). Unlike P. cybelidermus and P. guttalineatus (characters in parentheses), P. rhammanotus lacks an enlarged, postrictal modified scale (present, slightly ventral and posterior to border of auditory meatus); has much smaller gulars covering most of the throat without any transition to smaller scales medially on the gular pouch (large scales laterally, transitioning to small granular scales medially on the gular pouch; Fig. 4 ); 36 gulars from the preaxial margin of the arm to the mental (31 or fewer gulars); only 20 projecting scales making up the dorsal crest (24–34), because medial contact between scales of the paravertebral series separates each vertebral from the one behind it (similar separation limited to posterior body in P. guttalineatus , absent in P. cybelidermus except as apparent anomalies); only a single heavily keeled scale in the dorsolateral series in front of the arm (four or five); a poorly developed longitudinal skin fold of smaller scales spanning the area between the posterior margin of the jaw and anterior margin of the scapula (fold and transition to smaller scales in this region absent); subequal keels on the bicarinate lamellae at the base of Toe III (preaxial keel much more developed, postaxial keel absent on some lamellae, very low on others); and cream tongue (yellow-orange).

Like Pseudocalotes rhammanotus and P. tympanistriga , P. dringi , P. khaonanensis , and P. saravacensis lack modified subdigital lamellae at the base of Toe III. Although reported to have bicarinate subdigital lamellae at the base of Toe III, P. flavigula actually lacks or has greatly reduced postaxial lamellae similar to P. cybelidermus and P. guttalineatus . With 51 scales around midbody, P. rhammanotus has more midbody scales than P. flavigula (38–40) and fewer than either P. khaonanensis (72–75) or P. saravacensis (68). Unlike P. dringi , P. rhammanotus has six canthals (five), heavily keeled gulars (smooth), projecting scales forming the dorsal crest (described as “very low dorsal crest formed by the keeled but not enlarged middorsal scale row” in the original description), and an antehumeral fold (antehumeral fold absent). Like P. rhammanotus , P. austeniana , P. kakhienensis , and P. kingdonwardi have antehumeral folds, however the dorsals of these species are strongly heterogenous in size and shape. All remaining species of Pseudocalotes have modified lamellae at the bases of their third toes.

Description of Holotype. Adult female 195.6 mm (SVL 67.9 mm) in length; SVL 34.7% and tail 65.3% of total length; tail 1.88X as long as SVL; distance from axilla to groin accounting for 48.2% of SVL; head 66.4% as wide as long, accounting for 22.4% of SVL; snout subacuminate in dorsal view and in profile, sloping upward at about 20° to horizontal ( Fig. 7 ); dorsal head scales imbricate, smooth on snout, keeled posteriorly; rostral not visible from above, wider than mental, about three times as wide as tall, contacting first supralabials (though very narrowly on left side) and five small postrostrals; postrostral series separating nasal from rostral; in prefrontal region, five noticeably enlarged scales in line down center of head; scales of frontal and parietal region small, keeled; key-hole shaped depression in interparietal region poorly developed, enclosed posteriorly by high keels of U-shaped parietal series; parietal eye not visible; interparietal scale narrow, diamond-shaped; supranasal scale single, elongate; circumorbital scales 9/10, distinctly enlarged, roughly pentagonal or hexagonal, contacting canthal series laterally; canthals keeled, 6/6; supraciliaries 6/7, first three elongate, others shorter; 1/1 large subpyramidal scale behind supraciliaries and at end of circumorbital series; small angulate scales of temporal region ending posteriorly in 2/2 larger scales with noticeably higher keels; single subpyramidal posttemporal scale on neck just behind bulge produced by hypertrophy of the jaw muscles, not followed posteriorly by similar enlarged scales.

Nasal trapezoidal, contacting supralabials 1–2/1–2; nostril large, oval, directed laterally, positioned at dorsal edge in center of nasal with its upper margin contributing to canthus; distance from posterior border of nostril to anterior ocular angle 42.1% of head length; loreal region slightly sloping; scales of loreal region smooth; eye 22.4% of head length; palpebrals granular; second row of palpebrals above eye with 2/3 larger, heavily keeled scales in center; large subocular scale row bordering supralabials; row of 2/2 distinctly enlarged, roughly hexagonal, keeled scales between external auditory meatus and orbit; external auditory meatus subcircular, shallow, 17.1% as long as head and 76.5% as long as ocular aperture, its distance from orbit 24.3% of head length; tympanic membrane clearly visible, opaque throughout, though darker where it attaches to extracolumella; scales surrounding meatus small, granular; prominent post-tympanic subpyramidal scale separated from meatus by four scales; similar large postrictal modified scale absent from both sides.

Supralabials smooth, 9/9; infralabials smooth, 8/8; 4/5 enlarged postmentals contacting infralabials, thereafter, sublabial scales separating enlarged postmentals from infralabials; first pair of postmentals separated medially by single small gular; gulars keeled, 36 from posterior tip of mental to preaxial margin of arm; gular pouch poorly developed, longitudinal; scales in center of gular pouch same size as gulars lateral to it; lateral-most four longitudinal rows of gulars wider than rest, about 2– 3 X as large as small gulars covering most of throat.

Nuchal crest consisting of 8 elongate scales, all except first two separated from one another by low scales similar to paravertebral series; dorsal crest prominent, extending onto base of tail, consisting of transversely narrow scales (though still will the lateral portions of the scales flat against the back) with tall, pointed mucrons and keels, 20 scales to posterior margin of thigh; all scales of dorsal crest separated from one another by medial contact between small scales of paravertebral series or individual flat keeled scales similar to paravertebrals; scales of dorsal crest separated from nuchal crest by gap of 7 low, keeled dorsals; dorsals larger than ventrals, paravertebral two or three longitudinal rows pointing posteriorly, remaining dorsals directed downward and posteriorly; on neck, dorsals much smaller than those between limbs; upper four rows of dorsals on neck pointed dorsally and posteriorly, five rows below them pointed posteriorly, and remaining rows pointed downward and posteriorly; scales of flanks feebly keeled, homogenous except for single dorsolateral row of widely spaced, heavily keeled scales located in sixth row of dorsals below dorsal crest; dorsolateral keeled scales 6/6 between limbs, separated from one another by 3–7 scales; in front of arm, dorsolateral series continued by only 1/1 dorsolateral heavily keeled scale; heavily keeled scales absent from lower flanks; single heavily keeled scale on lower neck in front of arm absent; scales around midbody 51; ventrals heavily keeled, without sharp transition from scales on flanks, noticeably smaller in front of axilla, 60 from preaxial edge of arm to preaxial edge of leg; preanal scales keeled, similar to, though smaller than ventrals, seven from preaxial edge of leg to vent; scales around vent unspecialized; scales around base of tail (counted five subcaudals behind vent) 16; rictal fold very short; short longitudinal antehumeral skin fold spanning distance between posterior tip of jaw and anterior border of scapula, marking transition to smaller scales below fold.

Scales of brachium, antebrachium, thigh, and shank imbricate, keeled; some scales on middorsal and postaxial border of antebrachium modified: their keels pointed outward and blade-like toward distal tip of scales; single, similar scale with pointed blade-like keel positioned slightly postaxial to middorsal in center of thigh; this modified scale of thigh also much larger than all other scales on dorsal surface of thigh and with its large keel pointing slightly more posterior than the scales proximal and distal to it; palmar scales tiny, keeled with spinose mucrons; palmar subdigital lamellae multicarinate with spinose mucrons at base of fingers, bicarinate distally, 21/21 under Finger IV; plantar scales heavily keeled, mucronate, imbricate, those proximal to Toes III–IV noticeably larger than other plantar scales; pedal subdigital lamellae bicarinate except under Toe IV where 2/2 basal lamellae bicarinate and 2/2 lamellae distal to those (i.e., under proximal phalanx of Toe IV) with single keel; pre- and postaxial keels of bicarinate lamellae similar under all digits; preaxial and postaxial keels of subdigital lamellae at base of Toe III subequal; subdigital lamellae 23/24 under Toe IV, divided longitudinally under proximal phalangeal articulation of Toes III and IV, entire under all other phalangeal articulations; single subdigital and single supradigital ungual lamellae contacting one another on all fingers and toes; ungual lamellae much longer than other subdigital lamellae; supradigital scales proximal to supradigital ungual lamellae 4 on all fingers and toes; in relative length, Finger III = IV> II> V> I and Toe IV> III> V> II> I; when adpressed, tip of claw on Toe V slightly passing proximal phalangeal articulation of Toe IV; leg relatively long, 48.2% as long as SVL; length of foot accounting for 36.6% of length of leg.

Scale surfaces covered in macrohoneycomb; scale organs lenticular, single and positioned subterminally atop keel on body, more numerous and also concentrated on keels on head; bristled sense organs single, positioned below distal tip of keel, on digital and at least some scales of ventral surface (distribution difficult to determine since stratum corneum missing from most scale of ventral surface); callous glands, pigmented generation glands (sensu Maderson & Chiu 1970; Harvey et al. 2012), and femoral and precloacal pores absent.

Coloration: In life, dorsal surface of head reticulated green and brown, becoming mostly brown on sides of head; labials and skin of tympanum noticeably brighter green than other scales on side of head, though each labial with diffuse brown pigment as well.

Scales of nuchal and dorsal crests green and brown with distinctive, almost black apexes; dorsolateral series of heavily keeled scales with distinct pale green tips contrasting with rest of scale; about four diffuse <-shaped bands on flanks; wide, prominent band overlapping scapula much lighter green than other markings; tail with 14 dark brown bands alternating with dark green bands; both green and brown bands of tail lengthening distally; limbs (including digits) similarly banded green and brown; high keels of large postaxial scale on center of thigh white; postfemoral stripe of some congeners absent; gular pouch white medially, becoming increasingly “dirty” with brown reticulation approaching labials; ventral body white with small spots (=one or two brown scales surrounded by white scales) laterally; banding of tail continuing onto ventral surface as alternating light and dark brown areas.

Iris bronze; ciliary ring pale yellow; buccal epithelium lateral to teeth sky blue at rictus becoming paler toward tip of snout; roof of mouth and throat entirely black; tongue cream.

In preservative green pigment faded, flanks uniformly darkened including heavily keeled scales of dorsolateral series; blue pigment forming wide transverse band overlapping rictus and flanked anteriorly and posteriorly by heavy deposition of brown pigment; blue rictal band interrupted by brown pigment between tympanum and eye on specimen’s right side.

Etymology. The new name rhammanotus is an adjective derived from the Greek nouns rhamma meaning suture and notos meaning back. In Pseudocalotes rhammanotus , scales of the paravertebral series separate all scales of the dorsal crest and remind one of sutures along the back of this species.

Standard English Common Name. Stitched-back False Garden Lizard.

Distribution and Natural History. When collected, the holotype of Pseudocalotes rhammanotus was sleeping on a leaf about 1.2 m above the ground in primary, upper montane rainforest just below a ridge line. For approximately three hours, E. Wostl, G. Barraza, C. Franklin, Kadafi, and M. B. Harvey searched forests at the type locality without encountering additional specimens of this species. We spent most of this time on the drier side of the mountain, and apparent scarcety of this species may be due to its preference for more humid environments just over the ridgeline from where we found the holotype.

When threatened, P. rhammanotus darkens in color and gapes its mouth exposing the sharply contrasting creamcolored tongue and black throat.

We dissected the left side of the holotype to determine reproductive condition. Each oviduct contains a single large, oval egg with a thick fibrous shell that has not yet calcified (dimensions of egg from left oviduct 17.3 X 6.8 mm; clutch size two). The left ovary contains one large and subcircular yellow egg (largest diameter 4.8 mm) and four small cream-colored eggs (0.7–1.5 mm).

......continued on the next page Nuchal crest 4–11 (8 ± 3, n = 5) 8 9–13 (10 ± 1, 10–12 (11 ± 1 8–10 (9 ± 1, 7–13 (12 ± 2,

n = 15) n = 16) n = 11) n = 6) ......continued on the next page P. cybelidermus P. flavigula P. poilani P. f l o w e r i P. microlepis P. brevipes (n = 1) (n = 2) (n = 2) (n = 2) (n = 10) ......continued on the next page While describing these new species, we discovered several new characters and unreported variation in other characters. Dring (1979) noticed that some species of Pseudocalotes have highly modified subdigital lamellae on Toe III. He illustrated the third toes of P. f l ow er i and P. dringi and first recognized Sundaland and Indochinese groups based on this character. In the Indochinese group represented by Pseudocalotes brevipes , P. f l o w e r i, P. microlepis , and P. poilani , pronounced and pointed preaxial keels extend from the base of the toe beyond the proximal phalangeal articulation, whereas the postaxial keels are absent or very reduced on the same scales. In contrast, Toe III of P. tympanistriga and P. rhammanotus has bicarinate subdigital lamellae, with well-developed preaxial and postaxial keels on all subdigital scales. One specimen in our sample (UTA 60544) of P. tympanistriga exhibits a slight modification at the proximal phalangeal articulation where the scales have divided unequally, postaxial keels are absent, and a small keeled scale on the postaxial side overlaps the position where the postaxial keel should be.

With the discovery of more species of Pseudocalotes , researchers have described various conditions intermediate between the two character states recognized by Dring (1979). Hallermann & McGuire (2001, p. 264) described and illustrated a third state for this character in P. larutensis . This species lacks postaxial keels and has “moderately enlarged and pointed preaxial keels, restricted to the base of the toe.” Later, Hallermann et al. (2010, p. 34) made a distinction between “pointed and triangular” keels in P. b rev i pe s, P. m i c ro l ep i s, and P. ziegleri and “blade-like” keels in P. f l o w e r i and P. poilani . Pseudocalotes cybelidermus and P. guttalineatus resemble P. larutensis in having slightly modified subdigital lamellae at the base of Toe III. Although thought to have bicarinate lamellae (e.g., Hallermann & McGuire 2001), P. flavigula also has an intermediate condition. Lamellae at the base of Toe III in FMNH 14903 are not bicarinate: they lack postaxial keels. The preaxial keels of this species appear subtriangular, but they are low and do not convey much of a serrated appearance to the base of the toe when viewed in profile. Manthey & Denzer (2000) and Mahony (2010) incorrectly scored this character for P. kakhienensis . In both P. kakhienensis and P. kingdonwardi , pointed preaxial lamellae and poorly developed or absent postaxial lamellae extend distal to the first phalangeal articulation. The keels are best developed in P. kakhienensis where they form a serrate preaxial edge, extending beyond the proximal phalangeal articulation. However, in both species, development of these keels is not as prominent as in species of the Indochinese group. In summary, lizards in this genus exhibit at least three states for this character: subdigital lamellae at the base of Toe III may be bicarinate, modified with relatively low preaxial keels proximal to the phalangeal articulation, or modified with triangular to blade-like preaxial keels extending beyond the phalangeal articulation. The modified lamellae differ in extent under the toe and height of the preaxial keel.

While examining subdigital lamellae, we discovered a new character in Pseudocalotes kakhienensis and P. kingdonwardi . Both species have some unicarinate lamellae under the distal phalanx of Fingers II–V and Toes II–V. The unicarinate lamellae are located in the middle of the phalanx and are noticeably longer than those at the articulations and tips of the digits. This unicarinate condition is most pronounced on Toe V of P. kakhienensis , where the only bicarinate lamellae are located at the base of the toe, proximal phalangeal articulation, and distalmost two lamellae. Among the species examined, only P. kingdonwardi and P. kakhienensis have this trait. Distal lamellae are invariably bicarinate in the other species examined.

Variation in two characters requires modification of the generic diagnosis of Pseudocalotes proposed by Hallermann & Böhme (2000). These authors (p. 201) diagnosed Pseudocalotes as having “no or weekly developed antehumeral fold (only sometimes in brevipes ), and ventral scales smaller or equal to dorsals.” However, P. brevipes , P. cybelidermus , and P. microlepis have ventrals that are distinctly larger than adjacent dorsal scales on the flanks. Antehumeral folds are present in many species we examined for purposes of comparison. We used presence of this fold to distinguish P. rhammanotus from Javan and Sumatran species, but antehumeral folds occur in several congeners on mainland SE Asia. The fold is most easily seen in a specimen of Pseudocalotes flavigula (FMNH 143903) where 3–4 oblique rows of tiny black granular scales on the fold separate large white scales around the insertion of the arm from much larger white scales above and in front of the fold. In a specimen of P. poilani (FMNH 258704), a particular heavy fold overlaps a broad patch of small granular scales equal in size to the scales covering the gular pouch. This broad patch of granular scales extends from the gular region around and above the insertion of the arm, all the way to the posterior edge of the scapula. As in other species with a similar fold, the scales above the fold are distinctly larger than those below it. Mahony (2010) called attention to the antehumeral folds of P. kingdonwardi and P. kakhienensis . The folds of these two species appear to be the same as those in P. flavigula and P. poilani .

All congeners examined by us have posttemporal tubercles, however they frequently lack the other modified scales of the neck. We only found postrictal tubercles in Pseudocalotes cybelidermus and P. guttalineatus . Posttympanic tubercles also occur in P. brevipes , P. kakhienensis , P. kingdonwardi , P. poilani , P. rhammanotus , and P. tympanistriga . In Pseudocalotes poilani a modified posttemporal scale has a similar scale positioned just above it and one or two similar scales in a horizontal row behind it, the 3–4 scales thereby forming a shallow arc of enlarged, heavily keeled scales on each side of the head. Similarly, P. kingdonwardi has two or three posttemporals and many (55%) P. cybelidermus have two, however, the modified posttemporals of these species are arranged in a horizontal row rather than a shallow arch.

In other species, we did not find distinct dorsolateral series of enlarged, heavily keeled scales like those in the Javan and Sumatran species. Pseudocalotes brevipes , P. poilani , and P. microlepis have enlarged scales scattered among much smaller dorsals. In some specimens, the enlarged scales appear to form a dorsolateral row. We counted seven differentiated scales between the limbs of a specimen of P. poilani (FMNH 258704), 11/8 scales in a specimen of P. microlepis (FMNH 14499), and 10 scales in a specimen of P. brevipes (MVZ 224103). These counts for P. microlepis and P. brevipes are substantially higher than the 4–7 seen in the Javan and Sumatran species. Interestingly, enlarged scales on the flanks of P. ka k h i e n e ns i s and P. kingdonwardi tend to be associated with pale pigmentation. This scale distribution is most obvious in P. kakhienensis , where oblique bands of enlarged scales perfectly match oblique pale-pigmented bands. We also found this association of pale color with enlarged scales in the dorsolateral series and the modified neck scales of the three new Sumatran species. At least in preservative, enlarged scales on the flanks of P. brevipes , P. poilani , and P. microlepis are the same color as adjacent dorsals.

The new Sumatran species differ from congeners in generally having more scales in the gap between the nuchal and dorsal crests and in having more prominent dorsal crests. Like P. tympanistriga , the other species examined by us fewer than four scales in the gap and have low, crests of enlarged scales that do not project distally. The examined specimens of P. brevipes , P. microlepis and some specimens of P. kingdonwardi and P. kakhienensis lack an obvious gap, although an abrupt decrease in the size of both the last nuchal and the first vertebral sometimes marks this transition. Nuchals in these species also tend to be much narrower so that a change in shape of the scales is also evident at the transition. The vertebrals of P. kingdonwardi and, to a lesser extent, P. kakhienensis resemble those of P. cybelidermus in mostly contacting one another without or with very few interruptions by paravertebrals. As in P. guttalineatus and P. rhammanotus , middorsal contact of paravertebrals separate many of the vertebrals of the crests in the other species examined by us.

Our comparisons among congeners revealed the importance of some meristic characters in diagnosing Pseudocalotes . Regrettably, several species were not available for study, and the following discussion only applies to the species in Table 1 . Pseudocalotes tympanistriga , P. kingdonwardi , and P. kakhienensis exhibit a high frequency of two supralabials contacting the nasal, whereas only one usually contacts the nasal in the other species. Pseudocalotes kingdonwardi , P. kakhienensis , and the four species from Sumatra and Java usually have more postmentals contacting the infralabials than the remaining congeners. Pseudocalotes cybelidermus , P. guttalineatus , and P. kingdonwardi have noticeably larger and fewer gulars than the other species. Pseudocalotes kingdonwardi differs from congeners in having only 3 or 4 postrostrals (vs. usually 5 or more), fewer circumorbitals (usually fewer than 10 vs. usually 10 or more), and one very long or one elongate and one shorter enlarged scale between the eye and ear (2–4 squarish to polygonal scales). A single large white scale and a few small scales in the postero-ventral corner cover the external auditory meatus (we did not dissect any specimens to determine if a tympanum is present under these scales) of this species unlike any congener. Pseudocalotes kakhienensis and P. kingdonwardi have fewer supralabial and infralabials than congeners. Apparent fusion of the last two supralabials into an unusually long scale usually contributes to the reduced count of supralabials.

Many Pseudocalotes have narrow, elongate interparietals, although these scales may only be about twice as long as scales on either side of them and are not always obvious. Among the species we examined, P. guttalineatus and P. tympanistriga lack differentiated interparietals, and this trait is polymorphic in P. brevipes , P. microlepis , and P. poilani . This scale has not been mentioned in most recent descriptions of congeners and existing photographs of Pseudocalotes have insufficient resolution to score this character. The holotype and one new specimen of P. andamanensis illustrated by Harikrishnan & Vasudevan (2013, their figure 2A and 2B) clearly show heavily fractured scales in the parietal region and no differentiated interparietal in this species.

The four species of Javan and Sumatran Pseudocalotes have black epithelia lining the roof of the mouth and throat. In P. rhammanotus and P. tympanistriga , the tongue is cream, whereas the tongue and floor of the mouth are bright yellow-orange in P. cybelidermus and P. guttalineatus . This dichotomy in coloration of the tongue matches the distribution of other characters such as keeling at the base of the third toe and will likely prove useful in future phylogenetic analysis of small draconine agamids. Unfortunately, color of the buccal epithelium has not been reported for most congeners. In P. larutensis the tongue is red, contrasting with a black throat ( Hallermann & McGuire 2001), and it is bright yellow in P. andamanensis ( Harikrishnan & Vasudevan 2013) . A photo of P. austeniana posted on the Reptile Database ( Uetz 2014) reveals that this species has an orange tongue. Available publications and photos do not indicate whether P. andamanensis or P. austeniana have a black throat. All four of the Javan and Sumatran species opened their mouths in a threat display, and the sharply contrasting colors of the throat and tongue certainly accentuate this display. Some field observations suggest that P. austeniana does not employ a similar display ( Das & Das 2007). As field biologists encounter the rare species in this genus, we encourage them to record coloration of the mouth and throat so that the distribution of these interesting characters can be evaluated across the genus.

Many species of Pseudocalotes remain rare in collections, however variation in some of their diagnostic characters appears to be more common than previously thought ( Table 1 ). For example, Hallermann & McGuire (2001) noted that a scale positioned between the postrostral series and nasal distinguishes P. floweri from P. laurutensis . We found this character to be polymorphic in P. cybelidermus and a rare trait of P. brevipes . Evidently, P. floweri also exhibits polymorphism for this character. The nasal contacts a postrostral on both sides of a specimen (FMNH 270127) from the Cardamom Highlands of Cambodia collected after Hallermann & McGuire’s study and later discussed by Hallermann et al. (2010). Unlike typical P. floweri , this specimen also has 4/5 canthals (applying Hallermann and McGuire’s counting method, the specimen resembles P. laurutensis ), but has blade-like preaxial keels extending beyond the phalangeal articulation of Toe III. Aside from P. cybelidermus and P. f l o w e r i, other species examined by us lack a small scale positioned between the postrostral series and nasal. Nonetheless, this trait may occur in P. andamanensis if we are correctly interpreting Harikrishan & Vasudevan’s (2013, p. 5) description of the holotype and six additional specimens: “rostral in contact with 3 enlarged scales on the tip of the snout; these scales are separated from the nasal by one scale.”

Figures

FIGURE 4. Gular region of Pseudocalotes cybelidermus (A, UTA 60551, head width 7.3 mm), P. guttalineatus (B, UTA 60500, head width 8.1 mm), and P. rhammanotus (C, MZB 10804, head width 10.1 mm).

FIGURE 7. Cephalic morphology of Pseudocalotes guttalineatus (A, UTA 60554, head length 20.7 mm) and P. rhammanotus (B, MZB 10804, head length 15.2 mm). Heavily keeled scales of dorsolateral series, similar antehumeral scale, and depressed area ventral and posterior to antehumeral fold shaded gray.

FIGURE 9. Holotype of Pseudocalotes rhammanotus (MZB 10804, SVL 67.9 mm) from montane forest above Danau Ranau, Lampung, Sumatra.

Tables

TABLE 1. Comparison of selected diagnostic characters among six species of Pseudocalotes. Most common character state of polymorphic characters shown in bold. Data for P. cybelidermus repeated in second table to facilitate comparisons.

	P. tympanistriga	P. rhammanotus	P. cybelidermus	P. guttalineatus	P. kingdonwardi	P. kakhienensis
Postrostrals	4 (20%, n = 5) 5 (40%) 6 (20%) 7 (20%)	5	4 (7%, n = 15) 5 (80%) 6 (13%)	5 (81%, n = 16) 6 (13%) 7 (6%)	3 (18%, n = 11) 4 (82%)	5 (50%, n = 6) 6 (33%) 7 (17%)
Interparietal	Absent	Present	Present	Absent	Present	Present
Circumorbitals	9–13 (10 ± 1, n = 9)	9/10	9–13 (11 ± 1, n = 30)	9–13 (11 ± 1, n = 32)	7–10 (8 ± 1, n = 22)	9–12 (11 ± 1, n = 12)
Canthals	5 (10%, n = 10) 6 (90%)	6/6	5 (40%, n = 30) 6 (60%)	5 (13%, n = 32) 6 (84%) 7 (3%)	4 (9%, n == 22) 5 (77%) 6 (14%)	5 (25%, n = 12) 6 (58%) 7 (17%)
Supraciliaries	6–8 (7 ± 1, n = 10)	5/7	4–7 (5 ± 1, n = 30)	5–8 (7 ± 1, n = 32)	4–6 (5 ± 1, n = 22)	5–7 (6 ± 1, n = 12)
Scale intercalated between postrostral and nasal	Absent (n = 10)	Absent (n = 2)	Present (19%, n = 16) Absent (81%)	Absent (n = 16)	Absent (n = 22)	Absent (n = 12)