Ptychozoon cicakterbang, Grismer & Wood Jr & Grismer & Quah & Thy & Phimmachak & Sivongxay & Seateun & Stuart & Siler & Mulcahy & Anamza & Brown, 2019

Grismer, L. Lee, Wood Jr, Perry L., Grismer, Jesse L., Quah, Evan S. H., Thy, Neang, Phimmachak, Somphouthone, Sivongxay, Niane, Seateun, Sengvilay, Stuart, Bryan L., Siler, Cameron B., Mulcahy, Daniel G., Anamza, Tashitso & Brown, Rafe M., 2019, Geographic structure of genetic variation in the Parachute Gecko Ptychozoon lionotum Annandale, 1905 across Indochina and Sundaland with descriptions of three new species, Zootaxa 4638 (2), pp. 151-198 : 164-173

Ptychozoon cicakterbang sp. nov.

Malaysia Parachute Gecko

Figs. 7 , 8

Ptychozoon homalocephalum Cantor, 1847:626 (in part); Boulenger, 1912:52 (in part) Ptychozoon kuhli Smith, 1935:119 (in part) Ptychozoon lionotum Manthey & Grossmann, 1997:247 (in part); Cox et al., 1998:81 (in part); Das, Dattagupta, & Gayen, 1998:131 (in part); Chan-ard et al., 1999:132 (in part); Kluge, 2001:25 (in part); Leong et al., 2003:170; Grismer, 2011a:532, 2011b:139; Grismer et al., 2011:78 (in part); Grismer et al., 2018a:203 (in part). Ptychozoon lionatum (sic.) Grismer, Youmans, Wood, & Grismer 2006:161; Grismer & Pan, 2008:278.

Holotype. LSUHC 10648 adult female collected by Evan S. H. Quah, L. Lee Grismer, Anuar Shahrul, and Jesse L. Grismer on 26 June 2012 on the road to the top of Gunung Jerai , Kedah, Peninsular Malaysia (5.8099°N, 100.4367°E, 744 m above sea level).

Paratypes. LSUHC 5597 adult male collected by Jesse L. Grismer, L. Lee Grismer, and Perry L. Wood Jr. on 23 July 2003 from Pulau Sibui , Johor, Peninsular Malaysia (2.217539°N, 104.069966°E, 16 m above sea level) . LSUHC 5783 bears the same data as LUSHC 5597 except it was collected on 1 September 2003 . LSUHC 8709 adult male collected by L. Lee Grismer, Perry L. Wood Jr., and Jesse L. Grismer from Pulau Perhentian Besar , Terengganu, Peninsular Malaysia (5.901514°N, 102.746672°E, 123 m above sea level) . LSUHC 9059 adult female bearing the same collection data as LSUHC 8709 except being collected on 18 October 2007 . LSUHC 9447 adult female collected by Perry L. Wood Jr., Jesse L. Grismer, and L. Lee Grismer from Gunung Machinchang , Pulau Langkawi, Kedah, Peninsular Malaysia (6.386111°N, 99.661111°E, 634 m above sea level) . LSUHC 10978 adult female collected by Evan S. H. Quah and L. Lee Grismer on 7 September 2013 from Hutan Lipur Lata Belatan , Terengganu, Peninsular Malaysia (5.579567°N, 102.589028°E, 527 m above sea level) . LSUHC 11058 adult female collected by L. Lee Grismer and Shahrul Anuar on 26 June 2013 from Kem Baha , Gunung Stone, Kelantan, Peninsular Malaysia (5.340092°N, 101.966917°E, 511 m above sea level) . LSUHC 11418 adult male collected by Evan S. H. Quah and L. Lee Grismer on 26 June 2013 from Pulau Bidong , Terengganu, Peninsular Malaysia (5.620990°N, 103.058062°E, 25 m above sea level) .

Additional specimens examined. LSUHC 5539 adult female collected by Jesse L. Grismer, L. Lee Grismer, Perry L. Wood Jr. on 23 July 2003 from Pulau Sibu , Johor, Peninsular Malaysia (2.21754°N, 104.06997°E, 16 m above sea level) ; LSUHC 6437 adult female collected by Jesse L. Grismer and Perry L. Wood Jr. on 2 July 2004 from the Tekek-Juara Trail , Pulau Tioman, Pahang, Peninsular Malaysia (2.82070°N, 104.168954°E, 110 m above sea level) ; LSUHC 9409 adult female collected by Evan S. H. Quah and L. Lee Grismer at the Reng Reef field station, Pulau Redang, Terengganu, Peninsular Malaysia (5.81212°N, 103.000749°E, 49 m above sea level) ; LSUHC 9891 adult female collected by Perry L. Wood Jr., L. Lee Grismer, and Jesse L. Grismer on 6 September 2007 from Lata Tembaka , Terengganu, Peninsular Malaysia (5.554063°N, 102.431311°E, 690 m above sea level) ; LSUHC 11439 adult female collected by Evan S. H. Quah and L. Lee Grismer on 26 June 2013 from Pulau Bidong , Terengganu, Peninsular Malaysia (5.620990°N, 103.058062°E, 25 m above sea level) . LSUHC 11234–36 collected by Evan S. H. Quah and L. Lee Grismer on 3 July 2013 from Hutan Lipur Lata Belatan , Terengganu, Peninsular Malaysia (5.57957°N, 102.58902°E, 527 m above sea level) .

Diagnosis. Ptychozoon cicakterbang sp. nov. differs from all other species of Ptychozoon by having the following unique combination of characters: a maximum SVL of 93.4 mm; supranasals not in contact; 9–13 supralabials; 11–15 infralabials; infra-auricular cutaneous flap present; prominent supra-auricular lobe; no dorsal or caudal tu- bercles; imbricate parachute support scales on dorsal surface of patagia; raised, prominent ridges on ventral surface of patagia; 79–105 midbody dorsal scales; 30–48 ventral scales; pre-antebrachial flap not inserting on the full length of digit V but leaving a space between them; no enlarged femoral scales; 17–25 pore-bearing precloacal scales in males; 18–28 enlarged precloacal scales; 5–8 rows of enlarged post-precloacal scales; 14–17 transverse subdigital lamellae on the fourth toe; 25–34 scales across the widest portion of the caudal flap; enlarged dorsal caudal scales forming intermittent whorls; distal caudal lobes fusing to form a narrow caudal flap; edges of caudal flap weakly crenulated; caudal lobes angled posteriorly; caudal lobes decrease posteriorly in size; no thick, dark, postorbital stripe; four dark body bands between limb insertions; no irregularly shaped, white, vertebral markings; and subcau- dal region banded in adults ( Tables 4 , 5 ).

Description of holotype. Adult female SVL 86.0 mm; head moderate (HL/SVL 0.27), wide (HW/HL 0.76), depressed (HD/HL 0.40), distinct from neck; snout rounded at tip in dorsal profile; prefrontal region concave; lores rounded; rostral scale large, rectangular, bearing an inverted Y-shaped, dorsomedial groove, in contact posteriorly with two supranasals, one postnasal, dorsolaterally with nostrils, and laterally with first supralabials; supralabials (10R,L) to mid-orbital position; infralabials (12R,14L); nostrils elliptical with long axes oriented dorsoventrally, occupying anterior portion of nasal scale, bordered anteriorly by rostral, dorsally by supranasal, posteriorly by four postnasals of varying sizes (upper largest), and ventrally by first and second supralabials; scales on rostrum granular slightly larger than granular scales on top of head and occiput; eyes large (ED/HL 0.22), less than snout length; pupil vertically elliptical, crenelated; supraciliaries elongate, posteriormost weakly pointed; auricular opening rounded, bearing a prominent supra-auricular lobe; tympanum deeply sunk; infra-auricular flap broad, rounded, extending from below corner of mouth to lateral margin of neck midway between posterior margin of ear opening and forelimb insertion, measuring 3.9 mm at its widest point; dorsal scales of infra-auricular flap large, subimbricate proximally, small juxtaposed distally, ventral flap scales minute and granular; mental triangular, slightly wider than deep, bordered laterally by first infralabials and posteriorly by paired, rectangular postmentals contacting medially for 80% of their length; one row of enlarged sublabials bordering infralabials, anteriormost largest; gular scales small, rounded, grading into larger, imbricating throat and subimbricate pectoral and ventral scales.

Body dorsoventrally depressed, relatively stout (AXG/SVL 0.46); patagium 9.4 mm at midpoint of body, bearing enlarged subimbricate, rectangular scales dorsally, minute juxtaposed, subrectangular scales ventrally, ventral surface bearing raised ridges of granular scales extending from body to edge of flap; 86 minute, flat, round, juxtaposed midbody dorsal scales, largest mid-dorsally; four (L) and one (R) large, flat, isolated, dorsal scale(s) immediately anterior to the hind limb insertions; 42 transverse rows of large, smooth, flat, subimbricate ventrals, ventrals much larger than dorsals, decreasing in size laterally into granular scales at base of flap; 23 enlarged, precloacal scales; five rows of enlarged, post-precloacal scales; and scales immediately anterior to vent granular.

Limbs short, robust (FL/SVL 0.11; TBL/SVL 0.15); dorsal scales of forelimbs, flat, juxtaposed, larger than dorsal body scales; ventral forelimb scales subimbricate; anterior and posterior margins of forelimbs, and posterior margins of hind limbs bearing wide, cutaneous flaps; that of anterior margin of forearm (i.e. pre-antebrachial flap) emarginated distally and terminating low on base of digit I; scales of forelimb flap large, elongate, subimbricate; those of hind limb flap much smaller, rounded, subimbricate; palmar scales smooth, rounded; digits fully webbed, relatively short, dorsoventrally compressed; undivided transverse subdigital lamellae number 12 (I), 15 (II), 18 (III), 17 (IV), 18 (V), distalmost lamellae V-shaped; claws arise from within the dorsal surface of digital pads; claw of digit I replaced by an enlarged, flat scale; dorsal scales of hind limbs, flat, juxtaposed, larger than dorsal body scales; ventral scales of hind limbs flat, subimbricate, smaller than ventral scales of belly; flat scales of anterior margin of thigh subimbricate; plantar scales smooth, subimbricate; digits fully webbed; transverse subdigital lamellae number 10 (I), 13 (II), 17 (III), 17 (IV), 14 (V), distalmost lamellae V-shaped; claws arise from within the dorsal surface of digital pads; and claw of digit I replaced by an enlarged, flat scale.

Tail original, flattened, shorter than SVL (TL/SVL 0.88); two median rows of transversely widened, smooth subcaudals anteriorly becoming less regular and broken up posteriorly; postcloacal scales large, flat, imbricate; dorsal caudals flat, juxtaposed, larger than dorsal body scales; 6–10 smaller scales between much larger, transversely aligned scales composing whorls; tail width and caudal lobes decrease posteriorly; 23 caudal lobes on each side slightly angled posteriorly; and tail terminates in a short, narrow, flap (7.3 mm) bearing weakly crenulated edges.

Light phase coloration and pattern in life ( Fig. 7 ). Dorsal ground color of head, body, and tail beige; top of head essentially unicolor; labial scales lighter than body, delimited by thin, dark lines at their junctures; infra-auricular flap same lighter color as labial scales; four faint, thin, sinuous dorsal bands between limb insertions transitioning into approximately six darker caudal bands; terminal caudal band on caudal flap nearly black, edged posteriorly by narrow white band; subcaudal region mottled, weakly banded; iris deep-red; gular region, throat, ventral surfaces of forelimbs, pectoral region, and anterior portion of belly dull-white with stippled scales; and posterior margin of belly dull-white, immaculate; stippling on ventral surfaces of hind limbs dense.

Variation ( Fig. 8 ). Variation in coloration and pattern is extensive due to this species’ having very different dark and light phases and its ability to substrate-match (see Grismer 2011a). Color pattern variation in the paratypes described here is based on preserved material. LSUHC 9059, 9447, and 11418 have more boldly marked dorsal bands. Dorsal caudal bands in LSUHC 11418 are much more prominent than those of the holotype. Dark, subcaudal banding in LSUHC 9447 and 11418 is far more distinct than that of the holotype and in LSUHC 9059 and 10991 the banding is more defined than that of the holotype but not nearly as much as the former three specimens. The belly of LSUHC 11418 is much more densely stippled than other specimens of the type series. The parachute scales of the body patagia do not imbricate in LSUHC 5539, 5597, 6437, 8709, and 9059 whereas in the holotype and the other paratypes they are subimbricate. The tails of LSUHC 5783 and 11058 are regenerated and bear no caudal lobes. Males LSUHC 5597, 5783, 8709, and 11418 have 17–25 enlarged pore-bearing precloacal scales. Meristic differences amongst the type material and the additional specimens examined are presented in Table 6 .

Comparisons ( Tables 4 , 5 ; Figs. 3 , 5 , 6 ). Ptychozoon cicakterbang sp. nov. differs from all other species of Ptychozoon in having a prominent supra-auricular lobe as opposed to a small ridge or no enlargement at all. It differs further from P. intermedium Taylor, 1915 , P. kuhli ( Stejneger, 1902) , and P. trinotaterra in lacking as opposed to having caudal tubercles. From P. intermedium , P. rhacophorus ( Boulenger, 1899) , P. trinotaterra , and P. kaengkrachanense it differs in having four body bands as opposed to 0–3. Ptychozoon cicakterbang sp. nov. differs from P. bannaense Wang, Wang , & Liu, 2016, P. horsfieldii , P. intermedium , P. kuhli , P. nicobarense , P. rhacophorus , P. trinotaterra , and P. kaengkrachanense in having an emargination between the pre-antebrachial flap and digit I as opposed to no emargination. From P. popaense it differs by having a maximum SVL of 93.4 mm versus 86.2 mm, 14–17 subdigital lamellae on the fourth toe versus 13 or 14, the absence versus the presence of a thick, dark postorbital stripe; and the absence versus the presence of large, irregularly shaped, white, vertebral markings. Ptychozoon cicakterbang sp. nov. differs futher from P. lionotum , P. kabkaebin sp. nov., and P. tokehos sp. nov. by having a significantly higher mean number of infralabials and a prominent supra-auricular lobe versus a slightly raised ridge. From C. tokehos sp. nov. it differs further by having a significantly higher mean number of subdigital lamellae on the fourth toe, a higher mean number of ventral scales, and a significantly longer axilla-groin length. From P. lionotum it differs even further by having, as opposed to lacking, prominent ridges on the ventral surface of the patagia and weakly crenulated as opposed to smooth caudal flap edges. From C. kabkaebin sp. nov. and P. lionotum it differs further by having a significantly higher mean number of supralabial scales and midbody scales and caudal whorls composed of enlarged scales. From C. kabkaebin sp. nov. it differs further by lacking, as opposed to having, a thick, dark, postorbital stripe. From C. lionotum it differs further by having a significantly narrower head. Ptychozoon cicakterbang sp. nov. is well-separated from other species previously recognized as P. lionotum in the PCA and DAPC and occupies a significantly different position along PC1 from that of P. kabkaebin sp. nov. and P. tokehos sp. nov. and along PC2, it occupies a significantly different positon from that of P. lionotum . From these three species, it is further separated by an uncorrected pairwise sequence divergence of 4.1–14.4%. Combinations of other characters differentiating P. cicakterbang sp. nov. from other more distantly related species are presented in Table 5 .

Distribution ( Fig. 1 ). Ptychozoon cicakterbang sp. nov. ranges throughout Peninsular Malaysia and its associated east and west coast islands and most probably extends into extreme southern Thailand. A population has also been reported from Natuna Besar Island, Indonesia that lies between Peninsular Malaysia and Borneo ( Leong et al. 2003).

Etymology. The specific epithet cicakterbang is derived from the Malay word for flying lizards. The word “cicak” means lizard and “terbang” means flight. The term is used for both Ptychozoon and Draco .

Natural history. Ptychozoon cicakterbang sp. nov. is found on trees of varying sizes in primary and old secondary, lowland dipterocarp forests ( Fig. 9 ) up to approximately 800 meters in elevation ( Dring 1979; Grismer 2011a,b; Grismer et al. 2011). On Pulau Sibu, it occurs at sea level on small trees within coastal forest adjacent to mangrove swamps ( Grismer 2011a) and can often be found sleeping on the smaller branches of small trees and large shrubs during the day and night less than two meters above the ground ( Grismer 2011a; Grismer et al. 2011). When alarmed, it may jump to the ground to escape. On Pulau Lang Tengah, Terengganu, specimens were observed on the ground eating winged termites on a rainy evening during a storm (Evan S. H. Quah; pers. obs.). Ptychozoon cicakterbang sp. nov. is adept at substrate matching and can vary considerably in coloration and pattern from one area to the next ( Fig. 8 ).

Figures

FIGURE 1. Distribution of Ptychozoon lionotum, P. cicakterbang sp. nov., P. kabkaebin sp. nov., and P. tokehos sp. nov. Diamond designated localities are non-specimen based and considered tentative. Their sources are: (1) Pulau Natuna Besar, Indonesia (Leong et al. 2003); (14) Hat Yai, Songkla, Thailand (LSUDPC 10946); (18) Ko Samui, Surat Thani, Thailand (LSUDPC 10948); (22) Dong Paya Fei Mounatains, Nakhon Ratchasima, Thailand (Smith 1935); (23) Chantabun, Chanthaburi, Thailand (Smith 1935); (24) Phnom Samkos, Pursat, Cambodia (LSUDPC 10949–53); (28) Phu Quoc Island, Kiem Giang Province, Vietnam (Nguyen et al. 2009); (29) Trang Bom, Dong Nai Province, Vietnam (Nguyen et al. 2009) however this specimen may be P. trinotaterra (see Distribution section for P. tokehos sp. nov.); (30) Cat Tien, Dong Nai Province, Vietnam (Nguyen et al. 2009); and (41) Siracha, Chanthaburi, Thailand (Smith 1935). Numbered locations for the specimen-based localities are listed in Tables 1 and 6–9 and on the tree in Figure 2.

FIGURE 3. Uncorrected pairwise sequence divergence between species of rhe Ptychzoon lionotum group (boxplots) and between individuals of the P. lionotum group (matrix). Colored squares represent the 50% quartile. Light-blue circles are means.

FIGURE 5. Boxplot analyses of the characters bearing statistically significant mean values between various combinations of species pairs. The species pairs are listed on the plot. Colored squares represent the 50% quartile. Orange circles are the means.

FIGURE 6. Multivariate analyses of P. lionotum, P. cicakterbang sp. nov., P. kabkaebin sp. nov., and P. tokehos sp. nov. A. PCA ordination along PC 1 and PC2. B. DAPC showing complete separation of the 95% confidence ellipses. C. PCA ordination along PC1, PC2, and PC3.

FIGURE 7. Holotype of Ptychozoon cicakterbang sp. nov. LSUHC 10648 from Gunung Jerai, Kedah, Peninsular Malaysia. Photograph by L. L. Grismer.

FIGURE 8. Color pattern variation in Ptychozoon cicakterbang sp. nov. A. LSUHC 9059 from Pulau Perhentian Besar, Tereng- ganu, Peninsular Malaysia. B. LSUHC 8709 from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. C. LSUDPC 3207 from Butterfly Park at Cameron Highlands, Pahang, Peninsular Malaysia. D. LSUHC 5539 from Pulau Sibu, Johor, Peninsular Malaysia.

FIGURE 9. Habitat of Ptychozoon cicakterbang sp. nov. A. Forest at the type locality of Gunung Jerai, Kedah, Peninsular Malaysia. B. Microhabitat along the Tekek-Juara Trail, Pulau Tioman, Peninsular Malaysia.

Tables

TABLE 4. Summary statistics and diagnostic characters among Ptychozoon lionotum, P. cicakterbang sp. nov., P. kabkaebin sp. nov., and P. tokehos sp. nov. SD = standard deviation and N = sample size.

	P. lionotum	P. cicakterbang	P. kabkaebin	P. tokehos
		sp. nov.	sp. nov.	sp. nov.
supralabials to middle of eyeball (SU)
mean (±SD)	8.5 (±1.00)	10.3 (±1.08)	8.7 (±1.15)	9.5 (±0.87)
range	7–9	9–13	8–10	8–11
N	4	18	3	19
infralabials (IL)
mean (±SD)	10.3 (±0.96)	12.7 (±1.23)	10.7 (±1.15)	10.9 (±0.60)
range	9–11	11–15	10–12	10–12
N	4	18	3	17
transverse midbody dorsal scales (MB)
mean (±SD)	75.8 (±3.78)	89.8 (±7.43)	80.3 (±2.08)	89.0 (±4.04)
range	73–81	79–105	78–82	82–95
N	4	18	3	19
ventral scales (VS)
mean (±SD)	35.8 (±1.50)	38.2 (±4.95)	35.3 (±2.51)	33.6 (±2.35)
range	35–38	30–48	33–38	30–37
N	4	18	3	19
enlarged pore-bearing precloacal scales in males (PP)
mean (±SD)	21.0 (±1.0)	21.0 (±3.25)	21.0 (±0.00)	21.4 (±1.51)
range	20–22	17–25	21	20–24
N	2	4	1	8
enlarged precloacal scales (PS)
mean (±SD)	23.8 (±0.50)	22.8 (±2.65)	22.0 (±0.82)	21.8 (±1.91)
range	23 or 24	18–28	22–23	18–25
N	4	18	3	19
enlarged post-precloacal scale rows (PPS)
mean (±SD)	5.0 (±0.00)	5.9 (±0.99)	6.0 (±0.82)	5.28 (±0.75)
range	5	5–8	5–7	4–7
N	4	18	3	19
transversely expanded 4th toe lamellae (TL4)
mean (±SD)	16.5 (±0.58)	16.2 (±0.86)	14.6 (±2.08)	14.6 (±1.41)
range	16 or 17	14–17	13–17	13–18
N	4	18	3	19
scales across widest portion of caudal flap (CF)
mean (±SD)	31.5 (±2.5)	29.1 (±3.27)	28.3 (±1.53)	31.6 (±2.59)
range	29–34	25–34	27–30	28–34
N	2	14	3	11
supra-auricular lobe	weak ridge	prominent lobe	weak ridge	weak ridge
prominent ridges on ventral surface of body flap	no	yes	no	yes
edges of caudal flap	smooth	weakly crenu-	weakly crenu-	smooth
caudal whorls bearing enlarged scales	weak to	lated present	lated absent	present
thick dark, postorbital stripe	absent variable	absent	present	variable
irregularly shaped, white vertebral markings	variable	absent	variable	variable
maximum SVL (mm)	95.9	93.4	89.1	97.5

TABLE 5. Character states differentiating species of Ptychozoon from one another. Data in part come from Annandale (1905), Brown et al. (1997), Brown (1999), Sumontha et al. (2012) and Wang et al. (2016). / = data unavailable or the character does not exist.

cicakterbang sp. nov.		kabkaebin sp. nov. tokehos sp. nov. popaense bannaense				horsfieldii intermedium kuhli lionotum nicobarense					rhacophorus	trinotaterra		kaengkracha- nense
max SVL (mm)	93.4	95.4	97.5	86.2	87.5	73.9	98.8	107.8	95.9	100.3	64.5		71.3	86
supranasals in contact	no	no	no	no	no	no	yes, no	yes, no	no	yes	no		yes	yes
infra-auricular cutaneous flap	present	present	present	present	present	present	present	present	present	present	absent		present	present
prominent infra-auricular	present	absent	absent	absent	absent	absent	absent	absent	absent	absent	absent		absent	absent
cutaneous lobe
disposition of dorsal tubercles	absent	absent	absent	absent	absent or	absent	0–10 irregu-	2–6	absent	absent	6–10 rows,		0 or 1 mid- absent
					only on oc-		lar rows	straight			scattered		dorsal row
					ciput			rows
dorsal tubercle shape	absent	absent	absent	absent	flat when absent		flat to	convex or	absent	absent	spiculate to		flat	absent
					present		convex	spiculate			highly spinose
enlarged, dorsal parachute	present	present	present	present	present	present	present	present	present	present	absent		present	present
support scales
space between pre-antebrachi-	present	present	present	present	absent	absent	absent	absent	present	absent	absent		absent	absent
al flap and digit I
enlarged femoral scales	absent	absent	absent	absent	absent	8–11	12–19	absent	absent	absent	absent		absent	absent
enlarged pore-bearing precloa-	17–25	21	20–24	21	17	10–11	8–12	14–32	20–22	14–22	12–18		19–21	14–19
cals in males
fourth toe lamellae	14–17	13–17	13–18	13 or 14	16 or 17	11–13	9–14	12–16	16 or	13–21	11–13		12–14	15–17
									17
scales across widest portion of	25–34	27–30	28–34	28 or 29	15	9–21	14–18	42–51	29–34	20–29	/		26	/
caudal flap
distal lobes fused into narrow	narrow	narrow	narrow	narrow	narrow	narrow	narrow	wide	narrow	wide	narrow		narrow	narrow
or wide caudal flap
caudal tubercles continue to	absent	absent	absent	absent	absent	absent	variably	yes	absent	absent	absent		yes	absent
tail terminus
caudal lobes angled poste-	slight	slight	slight	slight	extreme	extreme	extreme	slight	slight	slight	extreme		slight	slight
riorly
tail and lobes bearing distal	minimal minimal		mini-	minimal	extensive extensive		extensive	minimal	mini-	minimal extensive			minimal	extensive
reduction			mal						mal
thick postorbital stripe	absent	present	vari-	present	present	present	present	present	variable	absent	absent		present	absent
			able
bands between limb insertions	4	4	4	4	4	4	3	4	4	3	0		3	3
irregularly shaped, white	absent	variable	vari-	present	absent	absent	absent	absent	variable	present	present		present	absent
vertebral markings			able

TABLE 6. Meristic, mensural (in mm), and diagnostic characters of Ptychozoon cicakterbang sp. nov. Gn = Gunung (mountain) and P = Pulau (island). See Materials and methods for character abbreviations. Numbers in parentheses refer to numbered localities in Figure 1. / = data unobtainable; r = tail partially or completely regenerated.

	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC
	10648 (12)	5783 (2)	5597 (2)	9059 (8)	9447 (13)	8709 (8)	11418 (6)	11058 (11)	10978 (5)
	Gn. Jerai	P. Sibu	P. Sibu	P. Perhentian	P. Langkawi	P. Perhentian	P. Bidong	Gn. Stong	Lata Belatan
				Besar		Besar
	holotype	paratype	paratype	paratype	paratype	paratype	paratype	paratype	paratype
sex	f	m	m	f	f	m	m	f	f
supralabials to middle of eyeball (SU)	10	11	10	11	9	11	11	10	11
infralabials (IL)	14	15	14	15	12	13	14	13	12
supra-auricular lobe	yes	yes	yes	yes	yes	yes	yes	yes	yes
midbody dorsal scales (MB)	86	84	85	98	91	100	95	86	79
midbody ventral scales (VS)	42	36	34	44	32	32	44	36	30
enlarged pore-bearing precloacal scales in	/	17	21	/	/	21	25	/	/
males (PP)
enlarged precloacal scales (PS)	23	18	23	25	22	23	26	24	20
enlarged post-precloacal scale rows (PPS)	5	5	5	7	6	6	7	8	5
transversely expanded 4th toe lamellae	17	16	17	16	17	16	16	15	16
(TL4)
enlarged dorsal caudal scales forming	yes	yes	yes	yes	yes	yes	yes	yes	yes
whorls
scales across widest portion of caudal flap	34	28	26	28	30	25	29	/	27
(CF)
edges of caudal flap smooth or crenulated	crenulated	crenulated	crenulated	crenulated	crenulated	crenulated	crenulated	crenulated	crenulated
thick postorbital stripe	no	no	no	no	no	no	no	no	no
irregularly shaped, white vertebral markings	absent	absent	absent	absent	absent	absent	absent	absent	absent
SVL	86.0	81.9	74.7	84.6	82.0	80.1	84.2	93.4	91.3
TL	76.4	69.8r	77.2	88.3	86.7	80.5	79.1	65.6r	81.5
TW	7.9	6.7	6.3	7.2	7.1	7.0	7.3	6.8	7.4
HL	23.5	22.7	21.9	24	24.0	22.7	22.5	25	23.2
HW	17.8	16.7	15.8	17.1	17.4	16.7	17,4	19.4	16.3
HD	9.3	8.9	8.8	9.2	8.7	9.7	9.1	10.3	9.2
SNL	10.6	10.6	10.2	11.1	10.4	10.1	9.7	11.5	10.7
ED	5.2	4.7	4.5	3.7	4.5	5.6	5.6	6.7	6.0
TD	2.7	1.6	1.3	2.0	2.1	1.5	2.2	2.2	2.8
IN	3.4	2.6	2.7	2.9	2.4	2.8	3.0	3.4	3.1
IO	2.2	2.9	2.9	2.1	2.2	2.1	2.3	3.5	2.3
AXG	39.7	41.2	40.5	43.9	42.7	38	40.1	44.4	45.4
FL	9.6	10.6	9.5	9.9	10.6	8.8	10.0	11.4	11.0
TBL	13.2	14.6	13.8	13.7	14.6	12.9	13.7	14.1	16.0