A new species of Cyrtodactylus Gray (Squamata; Gekkonidae) from the Thai Highlands with a discussion on the evolution of habitat preference Author Grismer, L. Lee 0000-0001-8422-3698 lgrismer@lasierra.edu Author Rujirawan, Attapol 0000-0001-9179-6910 Department of Zoology, Faculty of Science, Kasetsart University, 10900 Thailand. & rujirawan _ bank @ yahoo. com; https: // orcid. org / 0000 - 0001 - 9179 - 6910 rujirawan_bank@yahoo.com Author Termprayoon, Korkhwan 0000-0003-1903-3040 Department of Zoology, Faculty of Science, Kasetsart University, 10900 Thailand. & termprayoon _ kk @ yahoo. com; https: // orcid. org / 0000 - 0003 - 1903 - 3040 termprayoon_kk@yahoo.com Author Ampai, Natee Department of Biology, Faculty of Science, Srinakharinwirot University, 10110 Thailand. Author Yodthong, Siriporn 0000-0001-6577-5358 Department of Zoology, Faculty of Science, Kasetsart University, 10900 Thailand. & siri _ yodthong @ yahoo. com; https: // orcid. org / 0000 - 0001 - 6577 - 5358 siri_yodthong@yahoo.com Author Wood, Perry L. 0000-0003-3767-5274 Department of Biological Sciences & Museum of Natural History, Auburn University, Auburn, Alabama 36849, USA. & perryleewoodjr @ gmail. com; https: // orcid. org / 0000 - 0003 - 3767 - 5274 perryleewoodjr@gmail.com Author Oaks, Jamie R. Department of Biological Sciences & Museum of Natural History, Auburn University, Auburn, Alabama 36849, USA. Author Aowphol, Anchalee 0000-0001-9504-4601 Department of Zoology, Faculty of Science, Kasetsart University, 10900 Thailand. & fsciacl @ ku. ac. th; https: // orcid. org / 0000 - 0001 - 9504 - 4601 fsciacl@ku.ac.th text Zootaxa 2020 2020-09-17 4852 4 401 427 journal article 8446 10.11646/zootaxa.4852.4.1 10d52e07-bc81-4176-adf1-9af558b27b33 1175-5326 4410011 03D97A75-F75E-4610-96F9-A641E76A3E1E Cyrtodactylus maelanoi sp. nov. Mae La Noi Bent-toed Gecko ( Figs. 5–7 ) Holotype . Adult male ZMKU R 00857 (field tag AA 03726) collected on 22 March 2017 at 2100 hrs by Piyawan Puanprapai, Attapol Rujirawan, Siriporn Yodthong, Natee Ampai, and Elyse S. Freitas from the Tha Pha Pum Subdistrict, Mae La Noi District , Mae Hong Son Province , Thailand ( 18.34223°N , 98.02317°E WGS; 991 m in elevation). Paratypes . The paratypes ZMKU R 00852–00856 (field tag AA 03721–25) bear the same data as the holotype. The remaining paratypes ZMKU R 00858–00860 (field tag AA 06195–97) bear the same locality data as the holotype but were collected by Attapol Rujirawan , Siriporn Yodthong , Korkwan Termprayoon , Natee Ampai on 13 June 2018 . FIGURE 5. Holotype of adult male Cyrtodactylus maelanoi sp. nov. (ZMKU R 00857) immediately after euthanasia. Above. Ventral view of pelvic region showing enlarged femoral and precloacal scale and pore arrangement. Middle. Dorsal view showing nuchal loop, dorsal band, and caudal patterns. Below. Ventral view showing belly and subcaudal coloration. FIGURE 6. Cyrtodactylus maelanoi sp. nov. type series. From left to right top panel: ZMKU R 00853 (field tag AA 03722), ZMKU R 00857 (field tag AA 03726), ZMKU R 00859 (field tag AA 06196) and ZMKU R 00860 (field tag AA 06197); bottom panel: ZMKU R 00856 (field tag AA 03725), ZMKU R 00854 (field tag AA 03723), ZMKU R 00858 (field tag AA 06195), ZMKU R 00852 (field tag AA 03721) and ZMKU R 00855 (field tag AA 03724). Diagnosis. Cyrtodactylus maelanoi sp. nov. differs from all species in the C. sinyineensis group by having the combination of 7–9 supralabials; six or seven infralabials; 29–37 paravertebral tubercles; 16–19 longitudinal rows of dorsal tubercles; 27–33 ventral scales ventral scales; 9–12 expanded subdigital lamellae on the fourth toe; 11–14 unmodified subdigital lamellae on the fourth toe; 22–24 total subdigital lamellae on the fourth toe; 24–28 enlarged femoral scales; a total of 8–13 pore-bearing femoral scales in males; 4–8 enlarged precloacal scales; four or five pore-bearing precloacal scales in males; three rows of enlarged post-precloacal scales; approximately five or six jagged dorsal body bands; 11 light-colored caudal bands (n=2); 11 dark-colored caudal bands (n=2); raised and strongly keeled dorsal tubercles that extend beyond base of tail; enlarged femoral and precloacal scales nearly the same size and continuous; pore-bearing femoral and precloacal scales not continuous; medial subcaudals two to three times wider than long and not extending onto lateral side of tail; iris reddish; nuchal loop lacking an anterior azygous notch, and bearing a jagged posterior border; dorsal bands not bearing paravertebral elements, wider than interspaces, bearing lightened centers, edged with white tubercles; dark markings in dorsal interspaces; dark caudal bands wider than light caudal bands; light caudal bands in adults bearing dark-colored markings; light-colored caudal bands not encircling tail; and mature regenerated tail not spotted ( Table 5 ). Description of holotype . Adult male SVL 75.2 mm ( Fig. 5 ); head moderate in length (HL/SVL 0.28), wide (HW/HL 0.73), flat (HD/HL 0.39), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region concave, canthus rostralis rounded; snout elongate (ES/HL 0.39), rounded in dorsal profile, broad in lateral profile; eye large (ED/HL 0.27); ear opening oval (EL/HL 0.11); eye to ear distance greater than diameter of eye; rostral rectangular, partially divided dorsally, bordered posteriorly by supranasals and one internasal, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by supranasals, posteriorly by two postnasals, and ventrally by first supralabials; 8(R,L) rectangular supralabials extending to below midpoint of eye; 6(R,L) infralabials tapering posteriorly to commissure of jaw; scales of rostrum and lores slightly raised, larger than granular scales on top of head and occiput; scales on top of head and occiput intermixed with small tubercles; dorsal superciliaries weakly pointed and directed laterally; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals which contact medially for 50% of their length posterior to mental; one row of enlarged chinshields, outermost row bordering first four infralabials; gular and throat scales granular, grading posteriorly into larger, subimbricate pectoral and ventral scales. Body relatively short (AG/SVL 0.39) with well-defined ventrolateral folds; dorsal scales small, raised and interspersed with large, raised, semi-regularly arranged, strongly keeled tubercles; tubercles extend from top of head onto base of tail just beyond the postcloacal swelling; tubercles on nape smaller than those on body; 30 paravertebral tubercles; approximately 17 longitudinal rows of dorsal tubercles; 28 flat, subimbricate, ventral scales larger than dorsal scales; eight enlarged precloacal scales; five pore-bearing precloacal scales not separated on the midline by a poreless scale; three rows of large, post-precloacal scales; and no deep precloacal groove or depression. Forelimbs moderate in stature, relatively short (FL/SVL 0.17); slightly raised, juxtaposed scales of forearm larger than those on body, intermixed with large tubercles; palmar scales slightly raised, juxtaposed; digits welldeveloped, relatively long, inflected at basal, interphalangeal joints; digits narrow distal to inflections; widened proximal subdigital lamellae extend onto palm; slight webbing at base of digit; claws well-developed, sheathed by a dorsal and ventral scale at base; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.21), covered dorsally by small, raised, juxtaposed scales intermixed with large pointed tubercles and bearing flat, slightly larger imbricate scales anteriorly; ventral femoral scales flat, imbricate, much larger than dorsals; one row of 14(R)13(L) enlarged femoral scales and enlarged precloacal scales continuous; enlarged femoral scales nearly equal in size; small, postfemoral scales form an abrupt union with larger, flat ventral scales on posteroventral margin of thigh; 5,6(R,L) pore-bearing femoral scales not continuous with pore-bearing precloacal scales; subtibial scales flat, imbricate; plantar scales raised; digits relatively long, well-developed, inflected at basal, interphalangeal joints; 11(R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that do not extend onto sole, 13(R,L) unmodified subdigital lamellae distal to inflection; and claws well-developed, base of claw sheathed by a dorsal and ventral scale. Tail nearly complete, gracile in proportions, 115.1 mm in length, 8.1 mm in width at base, tapering to a point, TL/SVL (1.42); dorsal scales of tail flat, forming indistinct whorls; median row of transversely expanded subcaudal scales three times as wide as long, not extending onto lateral subcaudal region; three enlarged postcloacal tubercles at base of tail on hemipenal swellings; and postcloacal scales large, flat. Color pattern ( Figs. 5 , 6 ). Dorsal ground color of head, body, limbs, and tail yellowish-brown; top of head and rostrum nearly unicolor, bearing areas of slightly darker, diffuse irregularly shaped markings; nuchal loop smooth posteriorly with two posterior projections, not divided medially; approximately five dark jagged body bands bearing lightened centers, lacking paravertebral elements, edged with whitish tubercles extend from the shoulder to the presacral region; lighter colored interspaces between bands bear darker markings; whitish tubercles scattered on flanks; sacral and postsacral bands continue onto the tail to form five black caudal bands that are wider than the five light-colored caudal bands; light-colored caudal bands bear dark markings and do not encircle tail; limbs bear distinct, dark-colored irregularly shaped markings; posterior one-third of tail regenerated with dark mottled pattern; gular scales bearing only two or three black stipples; black stippling in throat, pectoral region, and anterior portion of belly more dense; subcaudal region darkly mottled, posterior one-third grey with faint mottling. Variation. The paratypes closely resemble the holotype in dorsal banding and nuchal loop pattern ( Fig. 6 ). The nape band are more prominent in ZMKU R 00858–00860 and ZMKU R 00855. Paratypes ZMKU R 00858–00860 , 00852 , 00855–00857 are darker overall. Paratypes ZMKU R 00853, 00859–00860 have regenerated tails. Paratype ZMKU R 00854 is missing the posterior one-thrid to one-half of the tail. Additional variation in meristic and mensural characters are presented in Table 6 . Distribution . Cyrtodactylus maelanoi sp. nov. is known only from the type locality from the Tha Pha Pum Subdistrict, Mae La Noi District, Mae Hong Son Province , Thailand ( 18.34223°N , 98.02317°E WGS; 991 m in elevation; Fig. 1 ). Etymology. The specific epithet “ maelanoi ” is a toponym of the type locality Mae La Noi. The recommended vernacular name in English is Mae La Noi Bent-toed Gecko . Natural history. All lizards observed were abroad at night ( Fig. 7 ) in the vicinity of a waterfall ( Fig. 8 ). Lizards were found on the ground, the walls of a building, and on granite rocks and within their cracks. Lizards were also seen on tree trunks as high 70–200 cm above the ground and on a twig approximately 160 cm above the ground. These observations clearly indicate that Cyrtodactylus maelanoi sp. nov. is a habitat generalist. No hatchlings of gravid females were observed indicating that at least the period of March through June is probably outside the reproductive season of this species. FIGURE 7. Uncatalogued specimen of Cyrtodactylus maelanoi sp. nov. photographed in situ at the type locality of Tha Pha Pum Subdistrict, Mae La Noi District, Mae Hong Son Province, Thailand. TABLE 5. Summary statistics and diagnostic characters of the species of the Cyrtodactylus sinyineensis species group. SD = standard deviation, n = sample size, and / indicates data unavailable. nov

characters	Clade 2	Clade 1
amphipetraeus aequalis		bayinnyiensis chaunghanak- waensis dammathetensis dattkyaikensis naungkayain- gensis maelanoi sp. inthanon sinyineensis taungwineensis welpyanensis doisuthep
Supralabial scales (SL)
mean (±SD) Range	9.0 (0.00) 9	8.1 (0.46) 7–9	8.0 (0.84) 7–9	9.4 (0.71) 8–11	9.0 (0.00) 9	8.3 (0.58) 8 or 9	8.3 (0.58) 8 or 9	8.1 (0.60) 7–9	11.0 (1.00) 10–12	9.7 (0.57) 9 or 10	9.1 (0.86) 7–10	8.3 (0.6) 8 or 9	11.0 (1.0) 10–12
n	9	37	5	24	3	3	3	9	3	3	14	3	3
Infralabial scales (IL)
Mean (±SD) Range	7.0 (0.00) 7	6.8 (0.53) 6–8	6.2 (0.44) 6 or 7	7.9 (0.50) 7–9	7.7 (0.58) 7 or 8	6.3 (0.57) 6 or 7	7.0 (0.00) 7	6.3 (0.50) 6 or 7	9.3 (0.58) 9 or 10	8 (0.00) 8	7.00 (0.39) 6–8	7.0 (0.00) 7	9.5 (1.5) 8–11
n	9	37	5	24	3	3	3	9	3	3	14	3	3
Paravertebral tubercles (PV)
Mean	34.7	32.0	26.0	32.5	31.7	34. 0	34.3	31.1	/	33.7	30.3	31.7	/
(±SD) Range	(0.50) 34 or 35	(1.64) 29–36	(0.71) 25–27	(0.98) 31–36	(1.15) 31–33	(0.00) 33–35	(0.58) 34 or 35	(2.37) 29–37	/ /	(0.00) 33–35	(0.84) 29–32	(1.5) 30–33	/
n	9	37	5	24	3	3	3	9	/	3	14	3	/
Longitudunal rows of body tubercles (LT)
Mean (±SD) Range	17.9 (0.78) 17–19	20.6 (1.42) 18–23	17.4 (01.14) 16–19	19.5 (1.47) 17–22	14.3 (1.15) 13–15	19.0 (1.00) 18–20	17.3 (1.54) 16–18	15.5 (0.87) 16–19	19. 0 (1.00) 18–20	15 (0.00) 15	18.7 (0.61) 18–20	16.0 (0.00) 16	19.7 (0.58) 19 or 20
n	9	37	5	24	3	3	3	9	3	3	14	3	3
Ventral scales (VS)
Mean (±SD) Range	28.3 (0.71) 28–30	24.5 (1.68) 22–31	25.6 (1.52) 24–28	25.4 (1.10) 23–27	26.7 (1.53) 25–28	25.3 (0.58) 25 or 26	27.0 (0.00) 27	28.4 (1.94) 27–33	30.7 (2.89) 29–34	28 (0.00) 27–29	32.3 (2.02) 30–36	29.3 (1.20) 28–30	32.0 (3.00) 29–35
n	9	37	5	24	3	3	3	9	3	3	14	3	3

......continued on the next page TABLE 5. (Continued)

characters	Clade 2	Clade 1
amphipetraeus aequalis		bayinnyiensis chaunghanak- waensis dammathetensis dattkyaikensis naungkayain- gensis maelanoi sp. nov inthanon							sinyineensis taungwineensis welpyanensis doisuthep
Expanded 4th toe lamellae (ET4)
Mean	7.0	8.5	9.0	7.1	8.7	9.3	7. 0	10.4	/	9	7.4	8.0	/
(±SD) Range	(0.00) 7	(0.77) 7–10	(0.00) 9	(0.61) 6–9	(0.58) 8 or 9	(0.58) 9 or 10	(0.00) 7	(1.01) 9–12	/	(0.00) 8–10	(0.50) 7 or 8	(0.00) 8	/
n	9	37	5	24	3	3	3	9	/	3	14	3	/
Unmodified 4th toe lamellae (UT4)
Mean (±SD)	11.2 (0.44)	14.4 (1.01)	13.3 (0.50)	12.4 (0.92)	12.3 (0.58)	13.0 (0.00)	11.3 (0.58)	12.9 (0.93)	/	11.3 (0.58)	12.1 (0.62)	12.0 (1.00)	/
Range	11 or 12	13–17	13 or 14	11–14	12 or 13	13	11 or 12	11–14	/	11 or 12	11–13	11–13	/
n	9	37	5	24	3	3	3	9	/	3	14	3	/
Total 4th toe lamellae (TT4)
Mean	18.2	22.9	22.3	19.5	21.0	23.3	18.3	23.3	/	20.3	19.4	20.0	/
(±SD) Range	(0.44) 18 or 19	(0.94) 21–25	(0.50) 22 or 23	(1.06) 18–21	(0.00) 21	(0.58) 22 or 23	(0.58) 18 or 19	(0.70) 22–24	/	(1.6) 19–21	(0.85) 18–21	(1.0) 19–21	/
n	9	37	5	24	3	3	3	9	/	3	14	3	/
Enlarged femoral scales (FS)
Mean	31.7	25.4	26.8	28.0	33.0	26.0	31.7	26.9	30.3	26.3	24.1	30.3	/
(±SD)	(2.06)	(1.85)	(1.79)	(1.06)	(2.65)	(1.70)	(2.89)	(1.17)	(0.61) (0.58)		(3.83)	(0.58)
Range	29–34	22–30	24–28	27–32	31–36	25–28	30–35	24–28	29–32 26 or 27		18–30	30–31	/
n	9	37	5	24	3	3	3	9	3	3	14	3	/
Femoral pores (FP)
Mean (±SD) Range	11.4 (0.89) 10–12	14.4 (2.94) 10–19	15.0 (0.00) 15	28.6 (1.51) 27–32	36.0 (0.00) 36	25.0 (0.00) 25	11.0 (0.00) 11	10.5 (0.2.08) 8–13	5.0 (1.00) 4–6	16 (0.00) 16	16.3 (3.15) 13–22	20.0 (0.00) 20	5.0 (0.00) 0
n	5	7	1	9	1	2	2	4	3	1	7	2	1

......continued on the next page TABLE 5. (Continued)

characters	Clade 2	Clade 1
amphipetraeus aequalis		bayinnyiensis chaunghanak- waensis dammathetensis dattkyaikensis naungkayain- gensis maelanoi sp. nov inthanon							sinyineensis taungwineensis welpyanensis doisuthep
Enlarged precloacal scales (PS)
Mean	10.0	9.1	8.2	10.2	8.3	8.3	9.7	6.4	/	11	8.6	12.0	/
(±SD) Range	(1.22) 8–11	(0.77) 7–10	(0.45) 8 or 9	(0.70) 9–12	(0.58) 8 or 9	(0.58) 8 or 9	(2.08) 8–12	(1.24) 4–8	/	(1.0) 10–12	(0.84) 7–10	(1.0) 11–13	/
n	9	37	5	24	3	3	3	9	/	3	14	3	/
Precloacal pores (PP)
Mean (±SD) Range	8.0 (1.0) 7–9	7.4 (2.07) 5–10	9.0 (0.00) 9	9.9 (0.60) 9–11	9.0 (0.00) 9	7.0 (1.41) 6–8	3.5 (0.5) 3 or 4	4.5 (0.58) 4 or 5	5.0 (0.00) 5	5 (0.00) 5	6.6 (0.79) 6–8	7.5 (0.5) 7 or 8	6.0 (0.00) 6
n	5	7	1	9	1	2	2	4	1	1	7	2	1
Post-precloacal scale rows (PPS)
Mean (±SD) Range	3.0 (0.00) 3	3.0 (0.00) 3	2.4 (0.54) 2 or 3	3.0 (0.21) 2 or 3	3.0 (0.00) 3	3.0 (0.00) 3	3. 0 (0.00) 3	3.0 (0.00) 3	3.0 (0.00) 3	3 (0.00) 3	3.0 (0.00) 3	3.0 (0.00) 3	3.0 (0.00) 3
n	9	37	5	24	3	3	3	9	3	3	3	3	3
Body bands (BB)
Mean (±SD)	5.5 (0.94)	5.3 (1.00)	5.2 (0.83)	5.92 (0.41)	5.0 (0.00)	6.0 (0.00)	4.0 (0.00)	5.4 (0.49)	6.0 (0.00)	5.5 (0.71)	4.1 (0.26)	6.0 (0.00)	8.7 (1.53)
Range	4–7	5–7	4–6	5–7	5	6	4	5 or 6	6	5 or 6	4 or 5	6	7–10
n	9	12	5	24	3	3	3	9	3	2	14	3	3
Light caudal bands (LCB)
Mean (±SD) Range	11.0 (1.00) 10–12	9.6 9.3 (0.52) (2.08) 9 to 10 7–11		12.3 (1.37) 10–14	10.3 (0.71) 10 or 11	8.0 (1.41) 7–9	11.0 (0.00) 11	11.0 (0.00) 11	12.0 (0.00) 12	9 (0.00) 9	12.1 (0.60) 11–13	9.0 (0.00) 9	12.0 (0.00) 12
n	2	8	3	6	3	2	1	2	2	1	9	1	1

......continued on the next page TABLE 5. (Continued)

characters	Clade 2		Clade 1
amphipetraeus		aequalis	bayinnyiensis chaunghanak- waensis dammathetensis dattkyaikensis naungkayain- gensis maelanoi sp. nov						inthanon	sinyineensis taungwineensis welpyanensis			doisuthep
Dark caudal bands (DCB)
Mean (±SD)	12.0 (1.0)	9.5 (0.76)	10.3 (2.08)	12.7 (1.37)	10.7 (0.71)	8.5 (0.71)	10.0 (0.00)	11.0 11.0 9 12.0 (0.00) (0.00) (0.00) (0.71)				10.0 (0.00)	11.0 (0.00)
Range	11–13 9–11		8–12	11–15	10 or 11	8 or 9	10	11	11	9	11–13	10	11
n	2	8	3	6	3	2	1	2	2	1	9	1	1
Morphology
Body tubercles low, weakly keeled	no	no	yes/no	yes	no	no	no	no	no	no	no	no	no
Body tubercles raised, moderately to strongly keeled	yes	yes	yes/no	no	yes	yes	yes	yes	yes	yes	yes	yes	yes
Tubercles extend beyond base of tail	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	no	yes
Enlarged femoral and pre- yes cloacal scales continuous		yes	yes	yes	yes	yes	yes	yes	yes	yes	variable	yes	yes
Pore-bearing femoral and precloacal scales continuous	no	no	no	yes	yes	no	no	no	yes	no	no	no	yes
Enlarged proximal femoral scales ~1/2 size of distal femorals	no	no	no	no	yes	no	no	no	no	no	no	no	no
Medial subcaudals 2 or 3 times wider than long	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes
Medial subcaudals extend no onto lateral surface of tail Color Pattern		no	yes	yes	yes	no	yes	no	no	no	no	no	no
Color of iris	green	reddish reddish		green	green	reddish	reddish	reddish green		reddish reddish		green	reddish
Nuchal loop divided medi- variable variable no ally				no	variable	yes	no	no	no	no	no	no	no

......continued on the next page TABLE 5. (Continued)

characters	Clade 2	Clade 1
amphipetraeus aequalis			bayinnyiensis chaunghanak- waensis dammathetensis dattkyaikensis naungkayain- gensis maelanoi sp. nov						inthanon	sinyineensis taungwineensis welpyanensis			doisuthep
Nuchal loop with anterior	no	no	no	no	no	no	no	no	no	no	no	no	no
azygous notch
Posterior border of nuchal jagged		smooth jagged		jagged	jagged	protracted jagged		smooth jagged jagged			protracted	jagged jagged
loop
Band on nape	variable	yes	yes	yes	yes	no	variable	yes	yes	yes	no	yes	yes
Dorsal banding with para- yes		yes	variable	yes	yes	no	yes	no	no	no	no	no	no
vertebrtal elements
Dorsal bands wider than	usually	equal to yes/no		yes	yes	yes	no	yes	yes	yes	yes	yes	yes
interspaces	wider
Dorsal bands bearing	yes	no	yes/no	variable	no	yes	yes	yes	yes	weak	yes	no	anterior
lightened centers	edge
Dorsal bands edged with	yes	partly	no	variable	no	partly	no	yes	yes	yes	partly	no	posteriorly
light-colored tubercles
Shape of dorsal bands	broken to	highly vari- jagged		jagged	jagged	zig-zag	jagged	jagged zig-zag jagged			zig-zag	jagged straight
hour glass able
Dark markings in dorsal	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes
interspaces in adults
Ventrolateral body fold	no	no	no	no	faintly	no	no	no	no	no	no	yes	no
whitish
Top of head diffusely	variable	mottled mottled		mottled	mottled	mottled	mottled	patternless mottled mottled			mottled	mottled no
mottled, blotched, or pat-	(adult)
ternless
Light reticulum on top of	no	no	no	no	no	no	no	no	no	no	no	no	yes
head
Anterodorsal margin of	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes
thighs darkly pigmented
Anterodorsal margin of	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes	yes
brachia darkly pigmented
Light caudal bands bearing yes		no	no	yes	no	yes	yes	yes	yes	yes	yes	yes	yes
dark markings in adults

......continued on the next page TABLE 5. (Continued)

characters Clade 2		Clade 1
amphipetraeus	aequalis	bayinnyiensis chaunghanak- waensis dammathetensis dattkyaikensis naungkayain- gensis maelanoi sp. nov	inthanon	sinyineensis taungwineensis welpyanensis	doisuthep
Light caudal bands encir- no	variable no no no no no no		no	no no no	no
cle tail
Dark caudal bands wider yes	yes	yes yes yes yes yes yes	yes	yes yes yes	yes
than light caudal bands
Mature regenerated tail no	no	yes no / no no no	no	no no no	no
spotted
Maximum SVL (mm) 93.4	87.0	84.1 76.3 69.3 83.0 66.9 92.5	87.3	91.6 82.0 70.6	90.5

TABLE 6. Meristic, mensural, and color pattern data from the type series of Cyrtodactylus maelanoi sp. nov. R = right, L = left, and / = data unobtainable or not applicable.

ZMKU R 00858 (field tag AA 06195)	ZMKU R 00852 (field tag AA 03721)	ZMKU R 00853 (field tag AA 03722)	ZMKU R 00854 (field tag AA 03723)	ZMKU R 00855 (field tag AA 03724)	ZMKU R 00856 (field tag AA 03725)	ZMKU R 00857 (field tag AA 03726)	ZMKU R 00859 (field tag AA 06196)	ZMKU R 00860 (field tag AA 06197)
Mae Hong Son	paratype	paratype	paratype	paratype	paratype	paratype	holotype	paratype	paratype
Sex	M	F	M	F	F	F	M	M	F
Supralabials	8	8	7	9	8	8	8	9	8
Infralabials	6	6	6	7	7	6	6	7	6
Body tubercles low, weakly keeled	no	no	no	no	no	no	no	no	no
Body tubercles raised, moderately to strongly keeled	yes	yes	yes	yes	yes	yes	yes	yes	yes
Paravertebral tubercles	30	29	30	31	30	32	30	37	31
Longitudinal rows of body tubercles	18	17	18	16	17	18	17	23	19
Tubercles extend beyond base of tail	yes	yes	yes	yes	yes	yes	yes	yes	yes
Ventral scales	28	30	28	28	27	27	28	33	27
Modified subdigital lamellae on 4th toe	10	9	11	11	9	12	11	11	10
Unmodified subdigital lamellae on 4th toe	13	14	12	13	13	11	13	13	14
Total subdigital lamellae on 4th toe	23	23	23	24	22	23	24	24	24
Enlarged femoral scales (R/L)	14/13	13/14	12/12	14/13	14/13	14/13	15/13	14/13	14/14
Total femoral scales	27	27	24	27	27	27	28	27	28
Femoral pores (R/L)	5/6	/	4/4	/	/	/	5/5	7/6	/
Total femoral pores in males	11	/	8	/	/	/	10	13	/
Enlarged precolacal scales	8	4	5	7	6	7	7	7	7
Precloacal pores	5	/	5	/	/	/	4	4	/
Post-precloacal scales rows	3	3	3	3	3	3	3	3	3
Enlarged femoral and precloacal scales continuous	yes	yes	yes	yes	yes	yes	yes	yes	yes

......continued on the next page TABLE 6. (Continued)

ZMKU R 00858 (field tag AA 06195)	ZMKU R 00852 (field tag AA 03721)	ZMKU R 00853 (field tag AA 03722)	ZMKU R 00854 (field tag AA 03723)	ZMKU R 00855 (field tag AA 03724)	ZMKU R 00856 (field tag AA 03725)	ZMKU R 00857 (field tag AA 03726)	ZMKU R 00859 (field tag AA 06196)	ZMKU R 00860 (field tag AA 06197)
Pore-bearing femoral and precloacal scales continuous	no	/	no	/	/	/	no	no	no
Enlarged proximal femoral scales ~1/2 size of distal femorals	no	no	no	no	no	no	no	no	no
Medial subcaudals 2 or 3 times wider than long	yes	yes	yes	yes	yes	yes	yes	yes	yes
Medial subcaudals extend onto lateral surface of tail	no	no	no	no	no	no	no	no	no
Nuchal loop divided medially	no	no	no	no	no	no	no	no	no
2 posterior projections from nuchal loop	yes	yes	yes	yes	yes	yes	yes	yes	yes
Nuchal loop with anterior azygous notch	yes	no	no	no	no	no	no	no	no
Triangular marking anterior to nuchal loop	no	no	no	no	no	no	no	no	no
Posterior border of nuchal loop	smooth	smooth	smooth	smooth	smooth	smooth	smooth	smooth	smooth
Band on nape	yes	yes	yes	yes	yes	yes	yes	yes	yes
Dorsal banding with paravertebral elements	no	no	no	no	no	no	no	no	no
Number of body bands	5	5	5	5	5	6	5	6	5
Dorsal body bands wider than interspaces	yes	yes	yes	yes	yes	yes	yes	yes	yes
Dorsal body bands with lightened centers	yes	yes	yes	yes	yes	yes	yes	yes	yes
Dorsal bands edged with white tubercles	yes	yes	yes	yes	yes	yes	yes	yes	yes
Shape of dorsal bands	jagged	jagged	jagged	jagged	jagged	jagged	jagged	jagged	jagged
Dark markings in dorsal interspaces	yes	yes	yes	yes	yes	yes	yes	yes	yes

......continued on the next page TABLE 6. (Continued)

ZMKU R 00858 (field tag AA 06195)	ZMKU R 00852 (field tag AA 03721)	ZMKU R 00853 (field tag AA 03722)	ZMKU R 00854 (field tag AA 03723)	ZMKU R 00855 (field tag AA 03724)	ZMKU R 00856 (field tag AA 03725)	ZMKU R 00857 (field tag AA 03726)	ZMKU R 00859 (field tag AA 06196)	ZMKU R 00860 (field tag AA 06197)
Ventrolateral fold whitish	no	no	no	no	no	no	no	no	no
Top of head diffusely mottled, blotched, or patternless	patternless	patternless	patternless	patternless	patternless	patternless	patternless	patternless	patternless
Light-colored reticulum on top of head	no	no	no	no	no	no	no	no	no
Anterodorsal margin of thighs darkly pigmented	yes	yes	yes	yes	yes	yes	yes	yes	yes
Anterodorsal margin of brachia darkly pigmented	yes	yes	yes	yes	yes	yes	yes	yes	yes
White caudal bands with dark mark- ings	yes	yes	/	yes	yes	yes	yes	/	/
White caudal bands encircle tail	no	no	/	no	no	no	no	/	/
Number of light caudal bands	/	/	/	/	11	11	/	/	/
Number of dark caudal bands	/	/	/	/	11	11	/	/	/
Dark caudal bands wider than light caudal bands	yes	yes	/	yes	yes	yes	yes	/	/
Mature regenerated tail spotted	/	/	no	no	/	/	no	no	no
SVL	75.2	73.5	92.5	75.9	68.5	74.	90.0	88.8	91.5
TW	8.1	6.6	10.6	6.6	6.5	6.7	9.3	9.5	9.0
FL	12.9	12.3	14.2	12.9	11.9	12.2	14.1	15.0	14.2
TBL	15.9	14.9	17.2	15.1	14.1	15.1	17.2	17.8	17.4
AG	29.6	31.3	40.2	33.1	30.2	33.3	39.6	36.5	38.7
HL	21.1	21.1	26.7	20.9	19.2	21.2	24.8	25.1	24.9
HW	15.3	14.2	18.9	14.7	13.6	14.2	17.2	17.1	16.9
HD	8.3	8.5	10.9	8.7	8.4	8.1	10.0	10.6	10.3
ED	5.6	5.4	6.5	4.5	5.4	4.8	5.5	5.7	5.2
EE	6.3	6.3	7.8	6.3	6.3	5.8	7.7	7.7	7.3
ES	8.3	9.4	10.5	9.1	8.7	8.5	10.1	10.8	10.6

......continued on the next page Comparisons. Cyrtodactylus maelanoi sp. nov. (n=9) differs from all other species of the C. sinyineensis based on various combinations of character states ( Table 5 ). In clade 1, it differs from C. inthanon , C. sinyineensis , and C. taungwineensis in having significantly fewer supralabials and from these species plus C. welpyanensis in having significantly fewer infralabials ( Fig. 4 ; Table 4 ). It differs further from C. sinyineensis , C. taungwineensis , and C. welpyanensis having significantly fewer precloacal scales; from C. sinyineensis it differs significantly in having fewer enlarged femoral scales; it differs further from C. taungwineensis by having significantly fewer ventral scales ( Fig. 4 ; Table 4 ). Cyrtodactylus maelanoi sp. nov. may differ further from its sister species C. inthanon in having 24–28 enlarged femoral scales versus 29–32 in C. inthanon and they plot completely separate in the PCA and DAPC analyses ( Fig. 3 ). Increases in sample sizes may indicate that some of these character differences are not statistically significant whereas other differences may be statistically significant. Differences in color pattern among all species of the C. sinyineensis group are listed in Table 5 .