Cnemapsis biocellata, Grismer, Lee, Onn, Chan Kin, Nasir, Nurolhuda & Sumontha, Montri, 2008
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publication ID |
https://doi.org/ 10.5281/zenodo.184071 |
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DOI |
https://doi.org/10.5281/zenodo.5611088 |
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persistent identifier |
https://treatment.plazi.org/id/6D3C87EA-C161-6708-1191-3DE2AD19FA4B |
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treatment provided by |
Plazi |
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scientific name |
Cnemapsis biocellata |
| status |
sp. nov. |
Cnemapsis biocellata sp. nov.
Figures 2 View FIGURE 2 , 3 View FIGURE 3
Cnemaspis siamensis Manthey & Grossmann 1997:215 ; Cox et al. 1998:91.
Holotype. Adult male ( ZRC 2.6693) collected on 3 March 2008 by Chan Kin Onn, L. Lee Grismer, and Rick Gregory at 2330 h at 37 m a.s.l. from Kuala Perlis, Perlis, Peninsular Malaysia (06° 24.437N 100° 08.564E).
Paratypes. Five paratypes ( ZRC 2.6694, 2.6696 adult females and ZRC 2.6695, 2.6697– 98 adult males) have the same collection data as the holotype. THNHM 8895 is from Khao Tohphayawang, Muang Satun District, Satun Province, Thailand collected in August 2003 by M. Sumontha.
Diagnosis. Cnemaspis biocellata differs from all other Southeast Asian Cnemaspis in having the unique combination of a maximum SVL of 40.1 mm; 6–10 supralabials; 5–7 infralabials; scales of anterior portion of forearm weakly keeled; ventral scales smooth; no femoral pores; 8–12 precloacal pores; no row of linearly arranged tubercles on flanks; paravertebral, longitudinal rows of caudal tubercles present but no lateral caudal rows; smooth subcaudals with an enlarged median row; one or two cloacal tubercles; no large, shield-like subtibial or submetatarsal scales; 29–37 subdigital lamellae on fourth toe; no distinct, large, dark spots on neck and back alternating with transverse, white markings; no white markings on flanks alternating with dark blotches; no dark bands encircling tail; two distinct, white, well defined occipital ocelli; black occipital band bordering a series of closely spaced, large, white to yellow spots which form a nuchal band extending from posterior margin of one eye to the other eye; small, black shoulder patch enclosing a single white to yellow ocellus; shoulder patches not meeting middorsally; posterior 25% of tail not white. These differences are summarized across all Southeast Asian species in Table 1 View TABLE 1 .
..... continued ..... continued No. of 4th toe lamellae 13 25–31 16-18 18–23 ..... continued ..... continued ..... continued Description of holotype. Adult male; snout-vent length 36.8 mm; head oval in dorsal profile, moderate in size (HL/SVL 0.23), somewhat narrow (HW/SVL 0.20), flattened (HD/HL 0.39), distinct from neck; snout short (ES/HL 0.48), slightly concave in lateral profile, longer than eye diameter; postnasal region constricted medially, concave; scales of rostrum weakly keeled, raised, slightly larger than those on occiput; low, rounded, supraorbital ridges; shallow frontorostral sulcus; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.22); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; eye to ear distance greater than diameter of eye; rostral scale concave, dorsal 75% divided by deep, longitudinal groove; rostral bordered posteriorly by two large supranasals and posterolaterally by one small, lower postnasal and first supralabial; supralabials 10 (R and L), decreasing in size posteriorly; seven (R and L) infralabials, decreasing in size posteriorly, first, second, and third infralabials nearly same size; nostrils elliptical, oriented dorsoposteriorly; three small, granular, postnasal scales (R and L) undifferentiated from lateral scales of rostrum; mental large, triangular, deeper than wide, concave medially, bordered posteriorly by three postmentals; lateral postmentals largest; postmentals bordered posteriorly by five smaller, rounded scales, lateralmost scales largest; enlarged sublabials absent; gular scales granular, slightly raised; throat scales smooth, imbricate.
pemaggilensis
perhemtianensis
Body slender, elongate; dorsal scales equal in size throughout body; dorsal tubercles more or less randomly distributed from occiput to base of tail; dorsal tubercles large with multiple keels; pectoral and abdominal scales smooth, flat, slightly elongate, imbricate, equal in size throughout; ventral scales slightly larger than dorsal scales; eight precloacal pores arranged in a chevron, not separated medially by intervening scales lacking pores; precloacal depression absent; femoral pores absent.
Forelimbs moderately long, slender; dorsal scales of brachium flat, weakly keeled; scales of forearm same size as brachials, imbricate, those on anterior margin weakly keeled; ventral scales of brachium smooth, rounded, sharply juxtaposed; scales beneath forearm, smooth, flat; palmar scales smooth, juxtaposed, slightly raised; digits long with an inflected joint; claws slightly recurved; subdigital lamellae unnotched; lamellae beneath first phalanges widened; slight interdigital webbing; fingers increase in length from first to fourth with fifth slightly shorter than fourth; hind limbs slightly longer and thicker than forelimbs; dorsal scales of thigh keeled, slightly raised, juxtaposed to subimbricate; scales of anterior margin of thigh keeled; ventral scales of thigh flat, imbricate; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws slightly recurved; subdigital lamellae unnotched; lamellae beneath first phalanx widened; interdigital webbing weak; toes increase in length from first to fourth with fourth and fifth shorter than fourth; 29 subdigital lamellae on fourth toe (R, L).
Tail 1.37 times SVL, swollen at base; caudal scales arranged in segmented whorls, first segment at base of tail eight scales wide middorsally; anteriorly caudal scales raised, juxtaposed, weakly keeled becoming flatter, imbricate, and keeled posteriorly; shallow middorsal furrow; deeper, single lateral furrow; single median row of enlarged subcaudal scales; subcaudal scales smooth; a paravertebral and dorsolateral row of large, flattened tubercles on either side of midline; transverse tubercle rows do not completely encircle tail; two (R and L) enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail.
Coloration (in life; Fig. 2 View FIGURE 2 ). The holotype has a dorsal ground color of dull yellow which is overlain by five yellow, butterfly-shaped vertebral blotches extending from the shoulder region to the base of the tail. Small, faint yellow blotches occur on the flanks and limbs and tend to form caudal bands. The rostrum is grayish with faint, light markings highlighting the bright yellow, anterior, extrabrillar fringe. The interorbital region is yellow and two distinct, white, immaculate, well defined occipital ocelli are present. The ocelli are accentuated by a wide, black occipital band that forms the anterior border of a series of closely spaced, large, white to yellow spots that form a nuchal band extending from the posterior margin of one eye to the posterior margin of the other eye. Posterior to this, is a small, black shoulder patch enclosing a single white to yellow ocellus. The shoulder patches do not meet middorsally. The venter is beige and immaculate with the exception of faint, subcaudal mottling. In alcohol, the overall ground color dulls to gray but the essentials of the color pattern remain visible.
Variation. The male paratypes closely resemble the holotype in overall coloration and pattern ( Fig. 3 View FIGURE 3 ). The white, nuchal band in ZRC 2.6698 is more continuous than that of the holotype and the black occipital band is wider. The nuchal band in ZRC 2.6697 is more a series of closely spaced spots than a continuous band. The entire tail of ZRC 2.6695 is regenerated and uniform yellow. ZRC 2.6697 and 2.6698 have broken tails. This species has a marked sexual dimorphism with respect to coloration and pattern. The adult females (ZRC 2.6694 and 2.6696) have a base color of light brown and lack the black occipital and shoulder markings seen in the males ( Fig. 2 View FIGURE 2 ). Faint, straw colored ocelli, however, are present which are homologous to those in males. A series of poorly defined, straw colored, butterfly-shaped, vertebral markings extend from the nape to the base of the tail and tend to form poorly defined caudal bands. Small, irregularly shaped, faint spots occur on the flanks and limbs. The venter is beige and immaculate with the exception of faint, subcaudal mottling. A patch of skin is missing from the dorsum between the hind limb insertions and on the base of the tail in ZRC 2.6694. Morphometric variation and variation in scalation are presented in Table 2.
TABLE 2. Descriptive measurements and characteristics of the type series of Cnemaspis biocellata . All measurements in milimeters. PT = number of paravertebral tubercles; PP = number of precloacal pores; PS = number of postcloacal tubercles; and 4th toe = number of subdigital lamellae on 4th toe.
THNHM ZRC ZRC ZRC ZRC ZRC ZRC
8895 2.6693 2.6694 2.6695 2.6696 2.6697 2.9998
paratype holotype paratype paratype paratype paratype paratype Distribution. Cnemaspis biocellata extends at least 40 km through the isolated karst mountain range of the Banjaran Nakawan from Thale Ban National Park, Satun Province, Thailand ( Manthey and Grossmann 1997) southward to Gua Kelam, Tasik Meranti, and finally Kuala Perlis in the state of Perlis, northern Peninsular Malaysia ( Fig. 3 View FIGURE 3 ). Given this species proclivities for existing on a karst substrate it is expected to range more widely in the nearby karst formations in both Malaysia and Thailand.
Natural history. All specimens of Cnemaspis biocellata were found on a karst substrate ( Fig. 4 View FIGURE 4 ) or in one case on a tree trunk growing within a karst formation. In Gua Kelam, the habitat was highly disturbed and continually frequented by visitors to the park yet C. biocellata were abundant. At Tasik Meranti, lizards were observed on large rocks in primary lowland dipterocarp forest. In Kuala Perlis, lizards were abundant on karst formations in an area with no immediate native vegetation in the vicinity of a housing community on the edge of a parking lot. These observations suggest the most important environmental parameter for the presence of this species is the karst substrate and the microhabitats it offers, regardless if the forest is disturbed or natural. Cnemaspis biocellata were observed abroad during the day on the shaded sides of karst rocks and boulders as well as within crevice microhabitats and beneath small rocks in piles on the ground. During the day, lizards were very wary and difficult to approach and retreated into nearby crevice shelters at our slightest advance. At night, lizards moved out further away from their retreats and were much easier to approach. This small species is amazingly quick and agile and effortlessly moves from one inclined surface to another. Upon capture, large sections of their skin are easily torn off in much the same manner as Gehyra mutilata .
Etymology. The specific epithet biocellata is derived from the Latin prefix bi- meaning “two” and the Latin ocellus meaning “a little eye” and refers to the two small occipital eyespots.
Comparisons to other species. Cnemaspis biocellata differs from C. affinis , C. argus , C. aurantiacopes , C. baueri , C. boulengeri , C. caudanivea , C. dringi , C. flavigaster , C. flavolineatus , C. kendallii , C. kumpoli , C. limi , C. nigridia , C. nuicamensis , C. pemanggilensis , and C. perhentianensis , C. tucdupensis in having a maximum SVL of less than 44.0 mm. It differs from C. dezwaani , C. jacobseni , C. modiglianii , C. phuketensis , and C. siamensis in being larger, having a maximun SVL greater than 39.7 mm. Cnemaspis biocellata differs from C. argus , C. baueri , C. dringi , C. flavigaster , C. limi , and C. nigridius in having less than 11 supralabials. Cnemaspis biocellata has less than eight infralabials which differentiates it from C. affinis , C. argus , C. aurantiacopes , C. baueri , C. caudanivea , C. chantaburiensis , C. dringi , C. flavigaster , C. flavolineatus , C. kendallii , C. kumpoli , C. limi , C. nigridius , C. pemanggilensis , C. siamensis , and C. tucdupensis but resembles all other species of Cnemaspis except C. affinis , C. argus , C. baueri , C. boulengeri , C. dringi , C. nigridius , C. pemanggilensis , and C. siamensis in having keeled scales on the forearm. All species of Southeast Asian Cnemaspis lack femoral pores except C. dezwaani , and C. modiglianii . Preanal pores occur in male C. affinis , C. argus , C. biocellata , C. flavigaster , C. nuicamensis , and C. perhentianensis but are lacking in male C. baueri , C. boulengeri , C. jacobseni , C. kendallii , C. limi , C. pemanggilensis , C. phuketensis , C. whittenorum , C. aurantiacopes , and C. tucdupensis . Cnemaspis biocellata is differentiated from C. affinis , C. caudanivea , C. dringi , C. modiglianii , C. flavolineatus , and C. nuicamensis in having more than six preanal pores. It is differentiated from C. nigridia in having less than 16 preanal pores. Cnemaspis biocellata is differentiated from C. boulengeri , C. aurantiacopes , and C. tucdupensis in that the ventralmost row of body tubercles are linearly arranged. All Southeast Asian species of Cnemaspis have caudal tubercles and only C. biocellata , C. boulengeri , C. nuicamensis , and C. tucdupensis lack a lateral row (this condition is unknown for C. dezwaani , C. jacobseni , and C. whittenorum ) and only C. biocellata , C. boulengeri , C. chanthaburiensis , C. kumpoli , C. modiglianii , C. caudanivea , and C. tucdupensis lack a ventrolateral row (this condition is unknown for C. dezwaani , C. dringi , C. jacobseni , C. siamensis , and C. whittenorum ). Only in C. boulengeri and C. tucdupensis are the caudal tubercles restricted to a single paravertebral row on either side of the midline. Cnemaspis biocellata is like all other Southeast Asia Cnemaspis except C. baueri , C. flavigaster , C. boulengeri , C. chanthaburiensis , C. kumpoli , C. limi , C. nigridius , C. phuketensis , C. whittenorum , C. caudanivea , C. aurantiacopes , C. nuicamensis , and C. tucdupensis in lacking, as opposed to having, keeled subcaudals although it lacks a single, median row of keeled subcaudals unlike C. dezwaani , C. jacobseni , C. kendallii , C. pemanggilensis , C. phuketensis , C. aurantiacopes , C. nuicamensis , and C. tucdupensis where this row is present (the condition in C. dringi is unknown). Cnemaspis biocellata is differentiated from all other Southeast Asian species except C. baueri , C. flavolineata , C. kendallii , C. pemanggilensis , and C. phuketensis in that the caudal tubercles do not encircle the tail. As in C. boulengeri , C. dezwaani , C. jacobseni , C. kendallii , C. limi , C. modiglianii , C. nigridius , C. pemanggilensis , C. phuketensis , C. siamensis , C. whittenorum , C. aurantiacopes , C. nuicamensis , and C. tucdupensis , C. biocellata has an enlarged, median subcaudal row of scales. Cnemaspis biocellata differs from C. dringi and C. whittenorum in having enlarged postcloacal tubercles and differs from all other Cnemaspis except C. affinis , C. baueri , C. boulengeri , C. chanthaburiensis , C. dezwaani , C. flavolineata , C. jacobseni , C. kendallii , C. limi , C. modiglianii , C. nigridius , C. pemanggilensis , C. phuketensis , C. caudanivea , C. aurantiacopes , and C. nuicamensis in having less than three postcloacal tubercles. Cnemaspis biocellata differs from C. boulengeri and C. caudanivea in lacking shield-like subtibial scales. Cnemaspis biocellata lacks keeled subtibial scales like C. argus , C. boulengeri , C. chanthaburiensis , C. modiglianii , C. pemanggilensis , C. caudanivea , C. nuicamensis , C. siamensis , and C. tucdupensis which have smooth subtibial scales (the condition in C. nigridia and C. whittenorum is unknown). Cnemaspis biocellata differs from C. boulengeri , C. dringi , C. caudanivea , C. aurantiacopes , and C. tucdupensis in lacking enlarged submetatarsal scales on the first toe. Cnemaspis biocellata differs from all other Southeast Asian Cnemaspis except C. flavigaster , C. flavolineatus , C. kumpoli , C. pemanggilensis , C. caudanivea , C. aurantiacopes , C. nuicamensis , and C. tucdupensis in having more than 27 subdigital lamellae on the fourth toe. Cnemaspis biocellata is similar to all other Southeast Asian species of Cnemaspis except C. boulengeri and C. baueri in lacking, as opposed to having, distinct, large, dark, markings on the neck. Cnemaspis biocellata differs from C. affinis , C. baueri , C. boulengeri , C. kendallii , C. limi , C. nigridius , C. phuketensis , C. aurantiacopes , and C. nuicamensis in lacking, as opposed to having, black bands on the tail. Cnemaspis biocellata differs from C. caudanivea in lacking a white tail tip. These character states are summarized in Table 1 View TABLE 1 .
Aside from the differences in SVL and scalation between Cnemaspis siamensis and C. biocellata ( Table 1 View TABLE 1 ), these two species also differ in various aspects of adult male color pattern ( Fig. 2 View FIGURE 2 ). Cnemaspis biocellata has a pair of white occipital ocelli, a white to yellow nuchal band, and a black shoulder patch whereas C. siamensis lacks these characters but has a series of white vertebral spots and a tripartite, dark, occipital marking which are absent in C. biocellata .
Discussion. The need for a phylogeny of Southeast Asian Cnemaspis can not be understated. With the presence of at least 16 species of this genus on the Malay Peninsula and its associated islands and a total of at least 23 in Southeast Asia (Grismer et al. in prep.), Cnemaspis comprises a significant portion of the lizard fauna of the Indo-Malayan Region and until these relationships are resolved, little else can be said about the evolutionary biology of this large, widespread group. An interesting feature occurring in C. biocellata and C. kumpoli which are sympatric, but not in C. siamensis with which the former two species are likely sympatric ( Taylor 1963), is the presence of black shoulder patches enclosing a yellow ocelli, a condition further exaggerated in C. biocellata on the nape and occiput ( Fig. 2 View FIGURE 2 ). Whether or not this character indicates a close phylogenetic relationship between these species or is the result of independent, parallel evolution stemming from similar selection pressures will have to await the acquisition of a phylogeny (Bauer et al. in prep.).
TABLE 1. — Diagnostic characters of Southeast Asian species of Cnemaspis. / = character not applicable;? = character could not be evaluated from specimens or literature.
| affinis | argus | aurantiacopes | baueri | |
|---|---|---|---|---|
| Maximum SVL (mm) | 49.0 | 65.8 | 56.6 | 64.9 |
| Supralabials Infralabials | 8–11 8–10 | 8–10 10–12 | 9–11 8–10 | 11–13 10–12 |
| Forearm scales keeled (1) or not (0) | 1 | 0 | 1 | 0 |
| Ventral scales keeled (1) or not (0) | 1 | 1 | 0 | 1 |
| Femoral pores present (1) or absent (0) | 0 | 0 | 0 | 0 |
| No. of femoral pores | / | / | / | / |
| Precloacal pores present (1) or absent (0) No. of precloacal pores | 1 5–6 | 0 / | 0 / | 0 / |
| Tubercles of ventralmost row on flank linearly arranged and in contact or nearly so (1) or tubercles on flank widely spaced and more randomly distributed (0) | 0 | 0 | 1 | 0 |
| Caudal tubercles present (1) or absent (0) | 1 | 1 | 1 | 1 |
| Lateral caudal tubercles present (1) or not (0) | 1 | 1 | 1 | 1 |
| Ventrolateral caudal tubercles anteriorly (1) or not (0) | 1 | 1 | 1 | 1 |
| Caudal tubercles restricted to a single Paravertebral row on each side (1) or not (0) | 0 | 0 | 0 | 0 |
| Subcaudals keeled (1) or not (0) | 1 | 1 | 0 | 0 |
| Single median row of keeled subcaudals (1) or not (0) | 0 | 0 | 1 | 0 |
| Caudal tubercles encircle tail (1) or not (0) | 1 | 0 | 0 | 1 |
| Enlarged median subcaudal scale row (1) or not (0) | 0 | 0 | 1 | 0 |
| Postcloacal tubercles present (1) or absent (0) | 1 | 1 | 1 | 1 |
| Shield-like subtibial scales present (1) or absent (0) | 0 | 0 | 0 | 0 |
| Subtibial scales keeled (1) or not (0) Enlarged submetatarsal scales on 1st toe (1) or not (0) | 1 0 | 0 0 | 1 1 | 1 0 |
| No. of 4th toe lamellae | 25–32 | 23–24 | 28–30 | 26–27 |
| White markings on flanks (1) or not (0) | 1 | 0 | 0 | 0 |
| Distinct, large, dark spots on neck (1) or not (0) | 0 | 0 | 0 | 1 |
| Dark caudal bands present (1) or absent (0) | 1 | 1 | 0 | 0 |
| Posterior 25% of tail white (1) or not (0) | 0 | 0 | 0 | 0 |
| ZRC |
Zoological Reference Collection, National University of Singapore |
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