Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia Author Harvey, Michael B. Author Hamidy, Amir Author Kurniawan, Nia Author Shaney, Kyle Author Smith, Eric N. text Zootaxa 2014 3841 2 211 238 journal article 10.11646/zootaxa.3841.2.3 08395a3b-782f-4aef-b473-221544edcd21 1175-5326 252328 0788BB44-9FC5-49CD-9519-22D8FAA09458 Pseudocalotes guttalineatus Harvey, Hamidy, Kurniawan, Shaney & Smith new species ( Fig. 6 ) Holotype . An adult female ( MZB 9796) from montane forest above Ngarip, Lampung, Sumatra, Indonesia , 5.28°S , 104.56° E , 1391 m , collected 10 June 2013 by field parties from BC , MZB , UB, and UTA . Paratypes . Thirteen specimens ( MZB 9652, 9765, 9792–9795, 9801, UTA 60536, 60540–60543, 60554) from the same mountain as the holotype , 1341–1521 m , collected 10–12 June 2013 by field parties from BC , MZB , UB, and UTA . Two specimens ( UTA 60500–60501) from Maura Dua, Remanan Jaya, Gunung Pesagi (localy known as Masagi), Sumatera Selatan, Indonesia , 4.91°S , 104.13°E , 1574–1643 m , collected 17 June 2013 by field parties from BC , MZB , UB, and UTA . Diagnosis. A species of Pseudocalotes reaching at least 316 mm ( 95 mm SVL) and distinguished from congeners by the following combination of characters: (1) discrete interparietal absent (2) canthals 5–7, usually six; (3) enlarged, heavily keeled to subpyramidal nuchal, posttemporal, posttympanic, and postrictal scales present; (4) gulars medium-sized with sharp transition to smaller scales on well-developed gular pouch; (5) antehumeral skin fold absent; (6) 6–7 enlarged, heavily keeled scales in dorsolateral series between limbs, extending onto neck as short row of closely spaced scales; (7) scales on lower flanks feebly keeled and homogenous (8) 45–55 scales around midbody (9) dorsal crest of projecting scales, extending to tail of male and midbody of females; (10) ventrals smaller than dorsals; (11) subdigital lamellae at base of Toe III modified: prominent preaxial keels pointed; postaxial keels absent or greatly reduced; (12) dorsum green with dorsolateral series of pale grayish blue, heavily keeled scales; (13) venter light green, without pale ventrolateral stripe between limbs; (14) gular region with grayish blue lines evident when gular pouch extended; (15) tongue and floor of mouth yellow-orange, throat black. Comparisons. Pseudocalotes guttalineatus can be distinguished from other agamids on Sumatra by its clearly visible tympanum, enlarged row of scales below orbit bordering supralabials, absence of skin folds, broad separation of 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, yellow-orange buccal epithelium, black throat, and other characters of external morphology described in the differential diagnosis of Pseudocalotes ( Hallermann & Böhme 2000 , as expanded by Mahony 2010 ). Pseudocalotes guttalineatus is most similar to P. cybelidermus with which it has been found in microsympatry. Unlike P. cybelidermus (characters in parentheses), P. guttalineatus lacks an interparietal (present) and has relatively small gulars numbering 23–31 ( Fig. 4 ; relatively large gulars, 17–25) from the preaxial margin of the arm to the mental, homogenous and weakly keeled dorsals below the dorsolateral row of heavily keeled scales (dorsals on lower flanks heterogenous, with scattered heavily keeled scales among feebly keeled scales, Fig. 5 ), and ventrals smaller than dorsals (larger). FIGURE 6. Pseudocalotes cybelidermus (A, MZB 9767, SVL 52.5 mm. B, MZB 9760, SVL 40.4 mm) from montane rainforest on Gunung Pesagi and P. guttalineatus from Gunung Pesagi (C, UTA 60501, SVL 73.6 mm) and forests above Ngarip (D, MZB 9796, SVL 75.2 mm). Of the 15 species of Pseudocalotes recognized before this study, only P. tympanistriga has been reported from Sumatra. Unlike P. tympanistriga (characters in parentheses), P. guttalineatus has a serrate dorsal crest in males extending onto the base of the tail (nuchal crest projecting, but dorsal crest of low, heavily keeled scales), a gap of 4–8 scales between the nuchal and dorsal crests (2–3 scales), flanks with six or seven enlarged keeled scales in a single dorsolateral row between limbs (four or five), much more developed preaxial than postaxial keels on subdigital lamellae at the base of the third toe (preaxial and postaxial keels equally developed), and yellow-orange tongue (cream). Most other congeners appear to be restricted to high elevations and are unlikely to be discovered on Sumatra. Unlike the new species, Pseudocalotes austeniana , P. kakhienensis , and P. kingdonwardi have well-developed antehumeral folds and dorsals that are strongly heterogenous in size and shape. With 45–55 scales around midbody, P. guttalineatus has more scales around midbody than P. flavigula ( Smith 1924; 37–40 ) and fewer scales around midbody than P. andamanensis (57–62), P. brevipes ( Werner 1904; 65–80 ), P. khaonanensis Chan-ard et al. 2008 (72–75) P. microlepis (Boulenger 1888; 65–72), P. poilani (56–60), P. saravacensis Inger & Stuebing 1994 (68), and P. ziegleri Hallermann et al. 2010 (57–64). Unlike P. laurutensis Hallermann & McGuire 2001 (characters in parentheses), the new species has 5–7 canthals (four), a projecting dorsal crest (scales keeled but not projecting), a bright green dorsum with a dorsolateral row of contrasting pale spots (dorsum tan-yellow with slight greenish cast), and a light green gular pouch with grayish blue stripes in males (yellow central spot flanked by plum-colored patches). Unlike P. floweri ( Boulenger 1912 ) the new species has a serrate dorsal crest (low, denticulate) relatively low subdigital keels at the base of Toe III (subdigital lamellae with blade-like lateral extensions), a dorsolateral series of enlarged, heavily keeled scales between the limbs (absent), and modified posttympanic and postrictal scales (absent). Unlike P. dringi Hallermann & Böhme 2000 , P. guttalineatus has greatly reduced or absent postaxial keels at the base of Toe III (subdigital lamellae bicarinate at base of Toe III), a serrate dorsal crest of projecting scales (scales of dorsal crest not projecting, crest formed by keels of scales), and a gular pouch covered in green scales separated by pale grayish blue skin (gular sac with “an oval blue black ocellus enclosing a pinkish purple patch,” Hallermann & Böhme 2000 , p. 200). Description. Males reaching at least 315.7 mm (SVL 95.3 mm , n = 5) and females 275.8 mm (SVL 188.0 mm, n = 10) in length; smallest juvenile 128.4 mm (SVL 39.4 mm ); SVL 29.3–31.8% (30.8 ± 0.8, n = 15) and tail 68.2–70.7% (69.2 ± 0.8, n = 15) of total length; tail 2.14–2.41X (2.24 ± 0.09, n = 15) as long as SVL; distance from axilla to groin accounting for 46.4–53.7% (50.3 ± 2.3, n = 16) of SVL; head 65.5–76.8% (71.0 ± 2.8, n = 16) as wide as long, accounting for 20.3–26.6% (22.9 ± 1.8, n = 16) of SVL; snout subacuminate in dorsal view and in profile, sloping upward at about 25° to horizontal ( Fig. 7 ); dorsal head scales imbricate, smooth on snout, keeled posteriorly; rostral wider than mental, about three times as wide as tall, contacting first supralabial and five (81%, n = 16), six (13%, n = 16), or seven (6%, n = 16) small postrostrals; postrostral series separating nasal from rostral (no small scale intercalated between nasal and rostral); in prefrontal region, 2–3 larger scales at midline followed by pair of laterally positioned enlarged scales in front of orbits (arrangement of enlarged scales forming an inverted Y ); scales of frontal and parietal region small, keeled; key-hole shaped depression in interparietal region better developed in larger specimens due to hypertrophy of the jaw muscles; parietal eye not visible; discrete interparietal scale not differentiated (one specimen, UTA 60542, has a long narrow scale in the center of the parietal region that may be a differentiated interparietal); supranasal scale single, elongate; circumorbital scales 9–13 (11 ± 1, n = 32) distinctly enlarged, roughly pentagonal or hexagonal, fractured into smaller scales where they contact canthal series; canthals keeled, usually 6 (84%, n = 32), infrequently 5 (13%, n = 32) or 7 (3%, n = 32); supraciliaries 5–8 (6 ± 1, n = 32), elongate except for last one or two; 1/1 large rounded scale behind supraciliaries at dorso-posterior edge of orbit; angulate, keeled scale of temporal region interrupted by 2/2 larger scales with noticeably higher keels; single subpyramidal and heavily keeled posttemporal scale on neck just behind bulge produced by hypertrophy of the jaw muscles; second similarly enlarged and heavily keeled nuchal scale present (69%, n = 16) or absent (31%, n = 16), positioned dorsal and posterior to posttemporal scale and in fourth row of paravertebrals, separated from subpyramidal postemporal scale by one or two small dorsals. Nasal trapezoidal, contacting first and second supralabial (28%, n = 32), first supralabial only (66%, n = 32), or separated from supralabials by row of small scales (6%, n = 32); nostril large, circular, 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 36.2–48.1%(42.3 ± 2.7, n = 16) of head length; loreal region vertical; scales of loreal region weakly keeled; eye 19.3–29.5% (22.2 ± 2.5, n = 16) of head length; palpebrals granular, showing noteworthy differentiation in two places: (1) second row of palpebrals above eye with 4–6 heavily keeled scales in center, and (2) patch of thin scales, cream with melanic centers forming opaque window in lower eyelid; lower margin of orbit separated from supralabials by one small and one larger row of scales; large subocular scale row bordering supralabials; row of two (22%, n = 32) or three (78%, n = 32) distinctly enlarged, keeled, roughly hexagonal scales between external auditory meatus and orbit (when only two present, divided scale present where first scale found in specimens with three of these scales); external auditory meatus subcircular, shallow, 15.2–20.7% (17.7 ± 1.5, n = 16) as long as head and 61.3–94.4% (80.4 ± 9.7, n = 16) as long as ocular aperture, its distance from orbit 21.0–29.0% (25.5 ± 2.1, n = 16) of head length; tympanic membrane clearly visible, opaque where it attaches to extracolumella, its antero-ventral one-fourth to three-fourths semitransparent; scales surrounding meatus small, granular; prominent posttympanic subpyramidal scale separated from meatus by three scales and positioned above similar large postrictal subpyramidal scale; in most specimens, postrictal scale one scale row removed from meatus and positioned directly above it. Supralabials smooth, nine (47%, n = 32) or ten (53%, n = 32); infralabials smooth, seven (3%, n = 32), eight (56%, n = 32), nine (38%, n = 32) or ten (3%, n = 32); two (22%, n = 32), three (47%, n = 32), or four (31%, n = 32) enlarged postmentals contacting infralabials; thereafter, sublabial scales separating enlarged postmentals from infralabials; first pair of postmentals in contact medially (38%, n = 8) or separated medially by single small gular (62%, n = 8); gulars keeled, 23–31 (28 ± 2, n = 16) from posterior tip of mental to preaxial margin of arm; gular pouch well developed, longitudinal; scales at distal end of gular pouch about one-fourth size of large gulars lateral to them. Nuchal crest consisting of 10–12 (11 ± 1, n = 16) elongate scales, longer scales separated from one another by low scales similar to paravertebral series; dorsal crest prominent in males, consisting of flattened, triangular scales (posterior to midbody, elongate scales of crest only slightly projecting in females), 24–34 (29 ± 3, n = 16) scales to posterior margin of thigh and some separated from one another by small scales of paravertebral series that meet one another medially; scales of dorsal crest separated from nuchal crest by gap of 4–8 (6 ± 1, n = 16) low, keeled dorsals; dorsals larger than ventrals, paravertebral two or three longitudinal rows pointing posteriorly and upward, remaining dorsals directed downward and posteriorly; orientation of scales on neck same as that between limbs; scales of flanks feebly keeled, homogenous except for single dorsolateral row of widely spaced, heavily keeled scales located in seventh row of dorsals below dorsal crest; dorsolateral keeled scales six (63%, n = 30) or seven (37%, n = 30) between limbs, separated from one another by 1–8 (usually 3–5) scales; dorsolateral series of heavily keeled scales extending in front of arm with about four large keeled scales separated from one another by 1–2 scales; heavily keeled scales absent from lower flanks; some specimens with single heavily keeled scale on lower neck in front of arm; scales around midbody 45–55 (49 ± 3, n = 16); no sharp transition to ventrals so that ventrals in a transverse row not countable; ventrals heavily keeled, noticeably smaller in area between arms, 45–54 (49 ± 3, n = 16) from preaxial edge of arm to preaxial edge of leg; preanal scales keeled, similar to, though somewhat smaller than ventrals, 7–10 (8 ± 1, n = 16) from preaxial edge of leg to vent; scales around vent unspecialized; scales around base of tail (counted five subcaudals behind vent) 15–20 (17 ± 2, n = 16); except for very short rictal fold, folds and pockets absent. Scales of brachium, antebrachium, thigh, and shank imbricate, keeled; some scales on postaxial border of antebrachium with more prominent mucrons than those on preaxial border; on thigh, enlarged scale with more developed, somewhat blade-like, keel positioned slightly postaxial to middorsal in center of thigh (some specimens with similar scale contacting this modified scale and positioned more postaxial and medial to it); this modified scale of thigh also much larger than all other scales on dorsal surface of thigh; palmar scales tiny, keeled with spinose mucrons; palmar subdigital lamellae multicarinate with spinose mucrons at base of fingers, bicarinate distally, 22–28 (25 ± 2, n = 16) 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, their keels similar except at base of Toe III where preaxial keels prominent and postaxial keels absent or greatly reduced; subdigital lamellae 26–32 (29 ± 2, n = 16) 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 lamella 4 on all fingers and toes; lamellae under all distal phalanges bicarinate; in relative length, Finger III> IV> II> V> I and Toe IV> III> V> II> I; when adpressed, tip of claw on Toe V closely approaching proximal phalangeal articulation of Toe IV but not passing it; leg relatively long, 46.4–53.7% (50.3 ± 2.3, n = 16) as long as SVL; length of foot accounting for 36.9–42.0% (39.6 ± 1.7, n = 15) 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, well developed on all scales of digits and present on most ventral surfaces of body, limbs, and tail, absent from dorsal body and head; at least some ventral scales with both types of sense organs on same scale; callous glands, pigmented generation glands (sensu Maderson & Chiu 1970 ; see also Harvey et al. 2014), and femoral and precloacal pores absent. 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. Coloration: In life [based on color notes taken in field by MBH from an adult male (UTA 60536) and an adult female (MZB 9796), supplemented by color photos of most specimens of type series], roof of mouth and throat black, tongue and skin medial to labials yellow-orange except for flesh-colored epithelium adjacent to teeth; iris yellow-gold; ciliary ring pale yellow, thin, indistinct; skin between scales of gular pouch pale grayish blue; scales of gular pouch green distally and grayish blue proximally so that pouch appearing green when retracted and grayish blue with oblique green lines when extended; dorsolateral row of heavily keeled scales pale grayish blue, contrasting with green scales adjacent to them; distinct light blue-green postfemoral stripe on proximal half of thigh, extending onto base of tail; venter brighter shade of green than dorsum from tip of snout onto base of tail, but broken by pale blue V-shaped mark anterior to chest; tail green proximally grading to brown distally or with alternating light and dark green bands basally of light bluish green alternating with green or bluish green, grading to brownish green and dark brown distally; when present, proximal bands only about one-fourth as long as distal band; darker caudal bands 10–12 (11 ± 1, n = 5), however difficult to count in most specimens due to fading, absence of bands on base of tail, or fusion of bands distally. Postfemoral stripe of female edged in black; female lacking blue pigment in gular region and pale blue V-shaped mark in front of chest. FIGURE 8. Adult Pseudocalotes guttalineatus (A, UTA 60501, SVL 73.6 mm) from montane forest on Gunung Pesagai, Sumatera Selatan, and juvenile (B, MZB 9652, SVL 39.4 mm) from montane forest above Ngarip, Lampung, Sumatra. In preservative, the green pigment has faded to dull grayish blue and the sky blue pigment of the dorsolateral series and modified scales of the neck has faded to cream. All specimens in the type series have blackened flanks. Etymology. The new name guttalineatus is an adjective derived from the Latin nouns gutta, meaning drop or spot, and lineatus, meaning lined. The new name refers to the dashed, bluish gray line on the neck and upper flanks of Pseudocalotes guttalineatus . Standard English Common Name. Dash-lined False Garden Lizard. Distribution and Natural History. Pseudocalotes guttalineatus occurs in sympatry with P. cybelidermus in humid montane forests of the Bukit Barisan Range of Lampung and Sumatera Selatan, Sumatra. This lizard is locally abundant on the mountains above Ngarip and at Maura Dua. Specimens were sleeping in low vegetation (below 2 m above the ground). Although we found all specimens between 1341–1625 m , coffee plantations replaced forest below this elevational range, and we did not sample areas above it. Although normally green in life, Pseudocalotes guttalineatus apparently can turn to various shades of brown, presumably in order to match its environment or temperament. One juvenile (MZB 9652, SVL 39.4 mm , Fig. 8 ) was brown when photographed alive, then turned green after being sacrificed for fixation. We did not record this specimen’s coloration upon capture, although, as we recall, all specimens of this species found sleeping in the forest were green and most remained this color after capture. Similarly, UTA 60501 lacked distinct banding on the tail, body, and limbs at the beginning of a photo session. However, as the session progressed, dark green bands appeared on the body, limbs, and tail. The blackened flanks of all specimens in the type series may not be an artifact of preservation. While handling specimens in the field, the flank color changed to black in some specimens, suggesting that P. guttalineatus undergoes this color change in response to stress. When threatened, P. guttalineatus opens its mouth in a threat display accentuated by its sharply contrasting black throat and bright yellow-orange tongue. By dissecting the right side only, we studied reproductive condition of two females in our sample. The right oviduct of UTA 60501 contained two large, oval-shaped eggs (dimensions 14.6 X 6.9 mm and 14.2 X 7.1 mm ; clutch size of four) already with tough fibrous shells, but no grossly evident calcification of the shell. In addition, this specimen’s right ovary contained two relatively large, yellow (diameters 4.7 mm ) and 5 small, cream-colored (diameters 1–1.5 mm ) eggs. The right oviduct of UTA 60540 was empty and highly convoluted, but not distended, suggesting that the specimen had not recently ovoposited. Its ovary contained one large, yellow, ovoid (4.8 X 3.7 mm ) egg, a smaller circular egg (diameter 2.7), and 7 small cream-colored eggs ( 0.7–1.5 mm ).