An integrative taxonomic review of the agamid genus Bronchocela (Kuhl, 1820) from Peninsular Malaysia with descriptions of new montane and insular endemics Author Grismer, L. Lee Author Wood, P. L. Author Lee, Cheol Haeng Author Quah, Evan S. H. Author Anuar, Shahrul Author Ngadi, Ehwan Author Sites, Jack W. text Zootaxa 2015 3948 1 1 23 journal article 10.11646/zootaxa.3948.1.1 e038824f-8500-4757-8aa8-d2a093bec8fe 1175-5326 288356 B596920C-270F-48B3-98B3-3FDFBF184576 Bronchocela shenlong sp. nov. Sheng Long Green-crested Lizard Figs. 5 , 6 , 7 Bronchocela cristatella Diong & Lim 1998 :347 ; Grismer et al. 2010 :149 ; Grismer 2011 :141 . Holotype . Adult male ( LSUHC 9017) collected on 16 June 2008 by L. Lee Grismer and Perry L. Wood Jr., at 2145 h at 1100 m a.s.l. from Bukit Larut, Perak, Peninsular Malaysia (04° 51.715 N , 100° 47.933 E ). Paratypes . LSUHC 9144 collected on 16 November 2008 by L. Lee Grismer, Jesse L. Grismer, Evan S. H. Quah, P. L. Wood Jr., and Chan K. O . between 2150 and 1350 hrs 1169 m a.s.l. from Bukit Larut, Perak, Peninsular Malaysia (04° 51.715 N , 100° 47.933 E ). LSUHC 10660 and 11301 bear the same collector data as the holotype but were collected on 30 June 2012 and 6 July 2013 , respectively, and LSUHC 10660 was collected at 1058 m a.s.l. Additional specimens examined from Bukit Larut. LSUHC 9416, 11587, 10271; ZRC 2.336, 2.491. Diagnosis (adults only). A moderately sized species reaching at least 106 mm SVL; tympanum small (DT/DO 0.38–0.44); head squarish in lateral profile (HD/HL 0.43–0.50), relatively narrow (HW/HL 0.54–0.63); forelimbs relatively long (FLL/SVL 0.51–0.59); foot not particularly short (FoL/HLL 0.39–0.41); hind limbs relatively long (HLL/SVL 0.84–0.95); 6–11 loreal scales; two or three postmentals; nuchal crest high, longer than diameter of orbit, spines lancolate; 71–92 midbody scales; 4–7 rows of paravertebral scales bearing keels pointing dorsoposteriorly; 1–5 rows of dorsal scales bearing keels pointing posteriorly; ventral scales less than five times the width of dorsal scales; 26–30 subdigital lamellae on the third finger; 27–36 subdigital lamellae on the fourth toe; third finger not longer than fourth finger; fourth finger not longer than fifth toe; no white patch between tympanum and orbit; tympanum green in adults; white infralabial stripe extending beyond tympanum; no red gular patch; no white ventrolateral line on body; postfemoral red line; and faint caudal bands. These characters are scored across all species in Tables 4 and 5 . Description of holotype . Adult male (SVL 106.0 mm); head disproportionately large, squarish in lateral profile (HD/HL 0.44); interorbital, frontal, and rostral regions sloped anteriorly; canthus rostralis sharp, extending to anterior margin of eye and continuous with enlarged, superciliary scales extending slightly beyond posterior border of eye; loreal region concave bearing seven canthal scales; rostral moderately sized, subrectangular, three times as wide as high, bordered laterally by first supralabials and posteriorly by four small scales; nasals protuberant, centrally located, external nares elevated above surface of rostrum, nasals bordered ventrally by two infranasals, posteriorly by two postnasal scales and dorsally by two elongate supranasals; scales on top of head small, slightly to moderately elongate, keeled to subspinose, those bordering medial margins of orbits enlarged; eyespot bearing parietal scale absent; single row of slightly enlarged, elongate, weekly keeled scales above supralabials; eye deeply set in orbit, surrounded by granular scales; tympanum naked; 11(R) 9(L) elongate, keeled supralabials, first scale of series smallest and square, remaining supralabials larger and rectangular; mental triangular, much larger than adjacent infralabials; three, elongate postmentals of nearly equal size; lateral postmentals contact first infralabial, first sublabial, and first chinshield; chinshields slightly enlarged, anterior two weakly keeled, decreasing in size posteriorly, and extending to a point below anterior margin of orbit; anterior chinshields separated from infralabials by one scale row of sublabials increasing to two and usually three rows posteriorly; 8(R,L) large, thin, rectangular, concave infralabials with upturned, laterally projecting, labial margins and strongly keeled, ventral sections; gulars sharply keeled; small, non-extensible dewlap. Body somewhat long, thin, triangular in cross-section; vertebral crest on nape composed of 11 greatly enlarged, lancolate scales; nuchal crest longer than diameter of orbit and abruptly tapers posteriorly to a single row of much smaller, keeled, triangular scales extending onto base of tail; lateral neck fold anterior to forelimb, posterior end forming a shallow pocket dorsal to forelimb insertion; dorsal scales of body small, keeled, spinose, 75 around midbody; seven rows of paravertebral scales bearing keels pointing dorsoposteriorly; one row of dorsal scales bearing keels pointing posteriorly; scales of flanks point ventroposteriorly; scales of belly and pectoral region similar in size, heavily keeled, spinose, arranged in semi-transverse rows more than twice the size of dorsal scales; forelimbs long (FLL/SVL 0.55), thin; dorsal and ventral scales of forelimbs keeled, spinose, nearly equal in size; five digits on manus; 26 subdigital lamellae on third finger; third finger not longer than fourth finger, fourth finger not longer than fifth toe; hind limbs long (HLL/SVL 0.90), thin; scales of hind limbs keeled, spinose; postfemoral scales slightly smaller, less keeled; five digits on pes; 27 subdigital lamellae on fourth toe; tail 3.04 times SVL, covered with longitudinal rows of keeled, spinose scales of similar size, Coloration (unstressed). Dorsal surfaces of head, body limbs and anterior portion of tail light green; broad, white stripe extends from anterior section of lower jaw through the infralabials past the angle of the jaw and below the tympanum to nearly the anterior edge of the shoulder region; loreal region and anterior supralabials bearing white flecks and faint striping; tympanum green, scales in orbit surrounding eye yellowish; six faint, transversely arranged, rows of diffuse, dull-white spots on dorsum between limb insertions that extend onto the tail to form faint, caudal bands; red postfemoral stripe extends from half way between the knee and the hind limb insertion on the body onto the anterior portion of the tail and bordered above by a diffuse, short, white stripe centered at the junction of the hind limb and the body; gular region yellowish with diffuse, broken, oblique, white stripes; all other ventral surfaces yellow; posterior three-fourths of tail brown, faintly banded. Variation (figs 5,6,7). The paratypes are remarkably similar to the holotype in all aspects of coloration and pattern although the white dorsal spots are more defined and less diffuse. The hatchling specimen (LSUHC 11587; SVL 43 mm ) has a less well-defined white infralabial stripe and the red postfemoral stripe has not yet appeared. These markings, however, are present in the juvenile specimen LSUHC 9416 (SVL 66 mm ) indicating that they are ontogenetically variable. Stressed individuals are usually gray (Figs. 6,7) but may also manifest a mosaic pattern of greens and grays. Females lack a dewlap. The specimens from Parit Falls may have relatively slightly wider heads than those from Bukit Larut (HW/HL 0.63 vs. 0.54–0.62). Meristic differences for all individuals are presented in Table 6. FIGURE 5. Bronchocela shenlong sp. nov. from Bukit Larut, Perak, Peninsular Malaysia. Upper left and right: adult female paratypes LSUHC 10660 (light color phase) and 9144 (dark color phase), respectively. Lower right: juvenile female paratype LSUHC 9416. Lower left: hatchling LSUHC 11587 sleeping on a fern at night. Photographs by L. L. Grismer. Distribution. Bronchocela shenlong sp. nov. is known only from Bukit Larut, Perak in the Bintang Mountain Range and from Parit Falls, Cameron Highlands, Pahang in the Titiwangsa Mountain Range ( Fig. 1 ). Natural history. Bronchocela shenlong sp. nov. occurs in both disturbed and undisturbed habitats in hill dipterocarp and montane forests ( Fig. 8 ). At Bukit Larut, specimens have been recorded from 942–1541 meters in elevation (ZRC 2.336 and ZRC 2.491, respectively). On Bukit Larut, we have observed lizards sleeping at night on forest-edge vegetation along a narrow road that winds through hill dipterocarp and montane forests. One specimen (LSUHC 9144) was collected during the day in a park while sitting on the top of an ornamental bush 1.5 meters above the ground. At Parit Falls, both specimens were collected at night at 1493 meters in elevation. LSUHC 12105 was shot out of a tree fern with a blowpipe while sleeping 10 meters above the ground in disturbed montane forest along a highly polluted river. LSUHC 12104 was collected by hand from the top of an ornamental bush in a highly disturbed area bordering an apartment complex along the same polluted river. At Bukit Larut, gravid females have been observed from June through November and hatchings and juveniles have been collected during September. LSUHC 12104 was a gravid female collected during September at Parit Falls. These data may indicate that the reproductive season for B. shenlong sp. nov. takes place during the earlier, drier part of the year. FIGURE 6. Upper and lower left and upper right: Bronchocela shenlong sp. nov. from Parit Falls, Cameron Highlands, Pahang, Peninsular Malaysia. Upper and lower left: adult male paratype LSUHC 12105 in light and dark color phase, respectively. Upper right: adult female paratype LSUHC 12104 in light color phase. Lower right: Bronchocela cristatella (LSUHC 12103) from 12 km S Parit Falls. Photographs by L. L. Grismer. Etymology. The specific epithet shenlong is used here to indicate this species’ morphological and ecological similarity to the “Shen Long” or “Spirit Dragon” of Chinese mythology. Shen Long is a wingless, five-toed, azurecolored reptile that resides in mountains and controls the wind, thunder, rain and clouds. As the epithet is used here, shenlong refers to this species’ upland, cloud forest habitat as well as its general dragon-like appearance. Comparisons. Bronchocela shenlong sp. nov. can be differentiated from B. danieli , B. smaragdina , B. vietnamensis , B. celebesensis , B. jubata , B. orlovi , B. rubrigularis and B. rayaensis sp. nov. by having 71–92 midbody scales as opposed to 43–71 collectively. It differs further from B. danieli , B. smaragdina , B. cristatella , B. marmorata Gray, B. hayeki , B. jubata , B. orlovi , and B. rubrigularis by the tympanum being less than 50% of the diameter of the eye. Having a high nuchal crest separates it from B. danieli , B. smaragdina , B. vietnamensis , B. marmorata , and B. rayaensis sp. nov. Bronchocela shenlong sp. nov. is also differentiated from various combinations of all other species on the basis of numerous color pattern characteristics (Tables 4,5). Bronchocela shenlong sp. nov. most closely resembles B. cristatella from Peninsular Malaysia but can be differentiated from it by having a relatively smaller tympanum (DT/DO 0.38–0.44 vs. 0.42–0.53); a relatively thicker head (HD/HL 0.43–0.50 vs. 0.40–0.45); a green as opposed to a black tympanum; a broad, white infralabial stripe as opposed to its absence; and a postfemoral red stripe extending onto the tail. For additional characters further differentiating B. shenlong sp. nov. from B. rayaensis sp. nov. see the comparisons section for the latter. These character states are summarized in Table 5 . FIGURE 7. Type series of Bronchocela shenlong sp. nov. From left to right holotype LSUHC 9017 and paratypes LSUHC 11301, 9144, and 10660 from Bukit Larut, Perak, Peninsular Malaysia and paratypes LSUHC 12105 and 12104 from Parit Falls, Cameron Highlands, Pahang, Peninsular Malaysia. Photographs by L. L. Grismer. FIGURE 8. Habitat of Bronchocela shenlong sp. nov. Left: disturbed habitat near Parit Falls, Cameron Highlands, Pahang, Peninsular Malaysia (Photograph by L. L. Grismer). Upper right: disturbed habitat at Bukit Larut, Perak, Peninsular Malaysia (Photograph by E. S. H. Quah). Lower right: undisturbed microhabitat at Bukit Larut (Photograph by L. L. Grismer). FIGURE 9. Color pattern variation in Bronchocela cristatella . Upper left: adult male from Taiping, Perak, Peninsular Malaysia (LSUHC 12102). Upper right: adult male from Hutan Lipur Sekayu, Terengganu, Peninsular Malaysia (LSUHC 11992). Lower left: adult male from Pulau Natuna Besar, Indonesia (LSUHC 11616). Photographs by L. L. Grismer. Lower right: adult male (LSUDPC 4651) from Kanowit, Sarawak, East Malaysia (Borneo). Photograph by Chan, K. O. Remarks. Grismer and Quah (2015) noted the exceptionally high degree of endemism and co-occurrence of closely related species in Cameron Highlands with respect to other montane areas in Peninsular Malaysia , which they attributed to the extensive plateau-like physiography of this upland region. Although Bronchocela shenlong sp. nov. is not endemic to Cameron Highlands it does occur in probable sympatry with B. cristatella . We collected a specimen of B. cristatella (LSUHC 12103; Fig. 9 ) in a small, Orang Asli village at 809 meters in elevation 12 km due south of Parit Falls near Ringlet on the western edge of the Cameron Highlands Plateau. The contact zone between B. shenlong sp. nov. and B. cristatella on the western slopes of the Bintang Mountain Range is unknown. Based on a specimen of B. cristatella (LSUHC 12102; Fig. 9 ) we collected along the base of the mountain at Taiping immediately below Bukit Larut at 76 meters in elevation, the two species thus far come to within 2.3 km of one another. The extent to which B. shenlong sp. nov. ranges north and south of Cameron Highlands and Bukit Larut in their respective upland corridors is unknown. South of Cameron Highlands in the Titiwangsa Mountain Range, Grismer (2011) lists an unconfirmed report from B. cristatella from Fraser’s Hill, Pahang ( Fig. 1 ). A specimen confirmed as B. cristatella from even farther south at Genting Highlands (LSUHC 5097) taken at 885 meters was used in this analysis (Figs. 1,2).