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Tantilla excelsa sp. nov. (Figs. 6, 7) Tantilla taeniata: Wilson and Meyer 1971:32, in part; Wilson 1982:56, in part; Wilson and Meyer 1982:109, in part; Wilson and Meyer 1985:103, in part; Wilson and McCranie 1999:328, in part; McCranie 2011a:227, in part; McCranie 2011b:44, in part; McCranie et al. 2012:623; Wilson and Mata-Silva 2015:457, in part. Holotype. — USNM 579682, an adult male from Lancetilla (15°44 ′ N, 87°27 ′ W), 30 m elevation, department of Atlántida, Honduras, collected 26 February 2011 by Leonel Marineros. Paratypes (n ¼ 3). — UF 157566, an adult female from Compañia Agricola Paradise near Peña Blanca (14°58 ′ N, 88°01 ′ W), 700 m elevation, Cortés, Honduras; UMMZ 58417, an adult female that is dehydrated and has an incomplete tail, from El Progreso (15°24 ′ N, 87°48 ′ W), Yoro, Honduras, 30 m elevation; MCZ 22045, head and anterior portion of body of an adult, from Progreso District, Yoro, Honduras, about 30 m elevation. Diagnosis. — Tantilla excelsa (in preservative) is defined by the following combination of characters: (1) pale middorsal stripe tan, extending length of body and at least half of tail, stripe on vertebral row and adjacent halves of paravertebral rows throughout its length; (2) tan to white lateral stripe occupying adjacent halves of scale rows 3 and 4; (3) ventrolateral area similar in pale color to that of lateral stripe; (4) lower two-thirds anteriorly and about lower third posteriorly of scale row 1 white similar to that of ventrals; (5) lateral edges of ventrals white, no, or very few, dark spots present; (6) ventral and subcaudal surfaces yellow (shortly after death) and white; (7) two females with 161–178 ventrals, single male with 169 ventrals, subcaudals 61 in one female, 70 in male; (8) tail length 23% of total length in one female, 24% in male. Tantilla excelsa can be distinguished from T. taeniata by having 169 ventrals in single male, 161–178 in two females, and by having yellow ventral surfaces in life (vs. 141–152 ventrals in males and 150 in single known female, and ventral surfaces pale purplish pink grading to darker salmon color on posterior two-thirds of body and tail). Tantilla excelsa differs from T. impensa and T. stenigrammi by having the middorsal pale stripe involving all of the vertebral scale row and adjacent thirds of paravertebral scale rows throughout its length (vs. middorsal stripe confined to vertebral row in those two species [on at least anterior half of body in T. stenigrammi]) and having female ventrals numbering up to 178 (vs. to 172 in T. impensa and 159 in female T. stenigrammi). Tantilla excelsa differs from T. olŋmpia in having complete middorsal and lateral pale stripes, with the middorsal stripe extending onto lateral edges of paravertebral rows, and having 169 ventrals and 70 subcaudal scales in single male (vs. those stripes reduced to dashes and spots, respectively; pale middorsal dashes restricted to vertebral row; and 148 ventrals and 49 subcaudals in a male specimen). Tantilla excelsa differs from T. psittaca by having yellow ventral and subcaudal surfaces in life and white to pale yellow in preservative and in having 169 ventrals in single male and 161–178 in two females (vs. those surfaces pink and grading to red in life and pale brown medially and pale brown with pinkish tinge on posterior third of body and under tail in preservative, 153–163 ventrals in males and 154–161 in female specimens). Tantilla excelsa differs from T. tritaeniata in having 169 ventrals in single male and 161–178 ventrals in two females, and having a complete pale nuchal collar (vs. 157 ventrals in single male and 155–161 in females, and pale nuchal collar divided). Description of holotype. —An adult male; TOL 400 mm; SVL 305 mm; TAL 95 mm (23.8% of TOL); HL 9.8 mm; HW 5.5 mm (level of angle of mouth); head barely distinct from neck; snout broadly rounded in dorsal view; eye length 1.3 mm; snout length 3.0 mm, about 2.3 times longer than eye; pupil circular; rostral about 1.1 times wider than high (2.2 X 2.0 mm); internasal length about 0.8 times width (1.1 X 1.4 mm); prefrontal larger than internasal, wider than long (2.2 X 1.5 mm); median prefrontal suture 1.4 mm, about half as long as frontal; frontal with slight anterior extension, Vshaped posteriorly, about 1.0 times longer than wide (2.9 X 2.8 mm), about 1.0 times longer than distance from its anterior edge to tip of snout (2.9 X 2.8 mm); parietal about 1.5 times longer than wide (4.1 X 2.8 mm), median parietal suture length 3.0 mm, about 1.0 times longer than length of frontal (2.9 mm); parietals contacting five nuchal scales; supraocular about 1.6 times longer than wide (2.3 X 1.4 mm), bordering orbit, contacting parietal, upper postocular, preocular, frontal, and prefrontal. Nasal divided, anterior nasal contacting rostral, internasal, and supralabial 1, posterior nasal contacting internasal, prefrontal, preocular, and supralabials 1 and 2, nostril located in central portion of nasal (forming division of nasal); loreal absent; preocular single, about 1.4 times higher than long (0.7 m X 0.5 mm), lower edge contacting supralabials 2 and 3; postoculars 2, upper about 1.5 times larger than lower (upper height 0.6 mm, lower height 0.4 mm); temporals 1 þ 1, anterior temporal 1.5 times longer than high (1.9 X 1.3 mm), posterior temporal 1.8 times longer than high (1.6 X 0.9 mm); supralabials 7, 3 and 4 bordering orbit, 4 and 5 contacting lower postocular, 5 also contacting anterior temporal, 6 contacting anterior temporal, 7 contacting anterior and posterior temporals; infralabials 6– 7, first 2 contacting anterior pair of chinshields on side with 6 infralabials (left side), first 3 contacting anterior pair of chinshields on side with 7 infralabials (right side); infralabial 1 on each side separated medially by contact with anterior chinshield; mental about 1.6 times wider than long (1.4 X 0.9 mm), contacting first pair of infralabials and anterior pair of chinshields; anterior chinshields about 2.4 times longer than wide (2.6 X 1.1 mm), not extending laterally to border of lip; posterior pair of chinshields about 1.7 times longer than wide (1.7 X 1.0 mm), posterior tips separated from each other by two rows of gular scales; four preventral scales between gular 1 and ventral 1; dorsal scales in 15–15–15 transverse rows, smooth throughout, lacking apical pits and supracloacal tubercles; dorsal scales in six rows at level of subcaudal 10; ventrals 169; cloacal scute divided; subcaudals 70, paired; ventrals plus subcaudals 239. Color of holotype in life. —Other than noting the ventral and subcaudal surfaces were yellow (L. Marineros, personal communication), the collector of the holotype did not record color notes of the recently killed specimen. Color of holotype in preservative (Figs. 6, 7). —Dorsal surfaces brown with white nuchal collar, tan middorsal stripe, and white lateral stripe; tan middorsal stripe includes vertebral row and adjacent tips to a third of paravertebral scale rows; tan middorsal stripe narrowing anteriorly on first scale involved in stripe, extending onto tail, fading out at about midlength of tail; tan middorsal stripe beginning on scale 4 posterior to parietals; broader white lateral stripe present, covering adjacent halves of scale rows 3 and 4, bordered above by continuous, thin dark brown line; white lateral stripe beginning on scale 5 posterior to supralabial 7, extending well onto tail to nearly reaching tip; lower twothirds of scale row 1 white anteriorly, grading to white only reaching lower tips of scale row 1 posteriorly on body; about upper third of scale row 1, all of scale row 2, and lower third of scale row 3 same shade of brown as paravertebral rows on about anterior half of body, before reducing width posteriorly; pale nuchal collar complete, not involving posterior tips of parietals, but involving first two scale rows dorsally, pale nuchal collar also involving posterior half of ultimate supralabial plus two scale rows laterally; dorsal surface of head brown, other than dark brown line crossing posterior tips of parietals to connect with lateral dark head bar; white, with dirty white flecking lateral spot involving postnasal, supralabial 1, and anterior half of supralabial 2; larger white spot posterior to eye involving anterior half of anterior temporal, all of supralabial 5, and tips of supralabials 4 and 6; dark brown lateral bar involving posterior half of supralabial 7, adjacent two-thirds of supralabial 6, and all of posterior temporal to connect with dark brown parietal cross-line; lateral dark bar narrowing only slightly dorsally; anterior tip of mental scale and lateral tips of infralabials 1 and 2 with brown flecking; dark brown spot present on infralabial 4 and adjacent edge of infralabial 5, remainder of ventral surface of head white; ventral surfaces of body and tail white for full length, except for variation discussed above. Variation in paratypes. —Two adult female paratypes show the following variation (followed by mean in parentheses): TOL 326 mm in one (UF 157566; UMMZ 58417 has incomplete tail); SVL 251–276 (263.5). mm; TAL 23.0% of TOL in one; ventrals 161–178 (169.5); subcaudals 61 in one; ventrals plus subcaudals 222 in one. All paratypes (including MCZ R22045, a head and anterior portion of body only of an adult) have 7 supralabials on each side, with supralabials 3 and 4 bordering the eye. Available color pattern notes indicate those paratypes were similar overall to the color pattern of the holotype. Distribution and habitat. —Very little is known about the habitats of Tantilla excelsa. The holotype was killed by a worker at the Lancetilla Botanical Gardens during February. The UF specimen was taken after dark as it was crawling on the ground on a coffee farm during May. The UMMZ specimen was collected on 27 March 1923 and possibly came from an insect trap (G. Schneider, personal communication). No data are available for the MCZ paratype other than it was collected in 1925. The Lancetilla Botanical Gardens formerly contained Lowland Moist Forest and lies at 30 m elevation. The coffee farm is in an area formerly containing Premontane Wet Forest and lies at 700 m elevation. The remaining two localities are in Lowland Dry Forest, although more mesic corridors of gallery forest do occur in those areas. Thus, T. excelsa is known to occur at low and moderate elevations (30–700 m) on both sides of the Río Ulúa drainage, and from the vicinity of Lago de Yojoa, Cortés, to the north-northeast in the vicinity of Lancetilla, Atlántida (Fig. 5). Etymology. —The specific name excelsa is an adjective from the Latin excelsus, which in this case means high. The name refers to the relatively high number of ventral scales in this nominal form.

Tantilla stenigrammi sp. nov. (Figs. 3, 4) Tantilla taeniata: McCranie 2011a:227, in part; McCranie 2011b:44, in part. Holotype. — University of Texas at Arlington (UTA) R-52591 (Field No. ENS 10897), a subadult female from Cuaca (15°23 ′ 01.75 ′′ N, 86°12 ′ 41.22 ′′ W;. datum ¼ WGS 84 in all cases), 895 m elevation, department of Olancho, Honduras, collected 3 February 2005 by Eric N Smith, Jorge A. FerrariCastro, Jorge Murillo, C. Chavez, Arturo Sosa, and John H. Malone. Paratype (n ¼ 1). — UNAH 3032, an adult male with an incomplete head and tail, from about 20 km E of Gualaco, Olancho, Honduras, at beginning of a foot trail (15°02 ′ N, 85°54 ′ W) leading to Cerro La Picucha, 1180 m elevation. Diagnosis. — Tantilla stenigrammi is defined, following Campbell (1998), by the following combination of characters: (1) middorsal stripe Warm Buff to Pale Horn Color (Color 118, 92), extending length of body and most of tail, confined to vertebral row on at least anterior half of body, extending onto adjacent edges of paravertebral scale rows posteriorly on body; (2) Buff-Yellow (Color 53) lateral stripe occupying adjacent thirds of scale rows 3 and 4; (3) ventrolateral area similar in pale color to that of lateral stripe; (4) lower half to two-thirds of scale row 1 colored similarly to that of ventrals; (5) lateral edges of ventrals immaculate white (in preservative), no dark spots present; (6) ventral and subcaudal surfaces Pearl Gray (in life; Color 81) to white (in preservative); (7) female with 159 ventrals, male with 164 ventrals; (8) tail incomplete in both. Tantilla stenigrammi can be distinguished from T. taeniata by having the pale middorsal stripe confined to the vertebral row on at least anterior half of body and in having pale gray ventral surfaces in life (vs. pale middorsal stripe also extending onto paravertebral rows throughout its length and ventral surfaces pale purplish pink grading to darker salmon color on posterior two-thirds of body and under tail in T. taeniata). It further differs from T. taeniata in having more ventral scales (164 vs. 141–152 in males, and 159 vs. 150 in females). Tantilla stenigrammi differs from T. psittaca and T. tritaeniata by having the middorsal pale stripe confined to the vertebral row on at least anterior half of body (vs. middorsal stripe involving all of vertebral scale. and adjacent third to half of paravertebral scale rows throughout its length in those two species); further from T psittaca by having pale gray ventral and subcaudal surfaces in life and white in preservative (vs. those surfaces being pink and grading to red in life and pale brown medially and pale brown with pinkish tinge on posterior third of body and under tail in preservative), and further from T. tritaeniata in having a complete nuchal collar and the pale lateral stripe covering adjacent thirds of scale rows 3 and 4 (vs. nuchal collar divided and pale lateral stripe including adjacent halves to two-thirds of scale rows 3 and 4). Tantilla stenigrammi differs from T. impensa in having 159 ventrals in the female, the pale middorsal stripe extending onto adjacent edges of paravertebral scale rows only posteriorly on the body, the pale brown lateral stripe covering adjacent thirds of scale rows 3 and 4, and by having a pale brown crossbar on the internasals that is bordered on the posterior edge by a dark brown cross-line (vs. 164–172 ventrals in females, the pale middorsal stripe confined to vertebral row throughout its length, pale lateral stripe including adjacent halves to twothirds of scale rows 3 and 4, and those head surfaces same ground color as remainder of head). Tantilla stenigrammi differs from T. olŋmpia in having complete middorsal and lateral pale stripes, in having a pale internasal crossbar, and having distinct pale large spots on the lateral surface of the head (vs. those body pale stripes reduced to dashes and spots, respectively, no internasal crossbar, and those lateral pale areas on head absent to indistinct). Description of holotype. —A subadult female; TOL 173 mm plus tail tip; SVL 137 mm; TAL 36 mm plus tail tip (20.8% plus tail tip of TOL); HL 4.6 mm; HW 3.4 mm (level of angle of mouth); head barely distinct from neck; snout broadly rounded in dorsal view; eye length 1.1 mm; snout length 2.2 mm, about 2.0 times longer than eye; pupil circular; rostral about 1.2 times wider than high (1.2 X 1.0 mm); internasal length about three-quarters of width (0.8 X 1.1 mm); prefrontal much larger than internasal, wider than long (1.3 X 0.9 mm); median prefrontal suture 0.9 mm, nearly half as long as frontal; frontal with slight anterior extension, V-shaped posteriorly, about 1.3 times longer than wide (2.1 X 1.6 mm), 1.3 times longer than distance from its anterior edge to tip of snout (2.1 X 1.6 mm); parietal about 1.6 times longer than wide (3.0 X 1.9 mm), median suture length 1.6 mm, about 0.8 times frontal length; parietals contacting seven nuchal scales; supraocular about 1.8 times longer than wide (1.6 X 0.9 mm), bordering orbit, contacting parietal, upper postocular, preocular, and prefrontal. Nasal divided, anterior nasal contacting rostral, internasal, and first supralabial, posterior nasal contacting internasal, prefrontal, preocular, and first and second supralabials, nostril located in central portion of nasal (forming division of nasal); loreal absent; preocular single, as high as long (both dimensions 0.5 mm), lower edge contacting supralabials 2 and 3; postoculars 2, upper about same size as lower (height of both 0.5 mm); temporals 1 þ 1, anterior temporal 1.1 times longer than high (1.0 X 0.9 mm), posterior temporal 1.6 times longer than high (1.3 X 0.8 mm); supralabials 7, 3rd and 4th bordering orbit, 4th and 5th contacting lower postocular, 5th and 6th contacting anterior temporal, and 7th contacting anterior and posterior temporals; infralabials 6, first 3 contacting anterior pair of chinshields, infralabial 1 on each side separated medially by contact with anterior chinshield; mental 1.7 times wider than long (1.2 X 0.7 mm), contacting first pair of infralabials and anterior pair of chinshields; anterior chinshields 2.1 times longer than wide (1.9 X 0.9 mm), not extending laterally to border of lip; posterior pair of chinshields 1.9 times longer than wide (1.3 X 0.7 mm), posterior tips separated from each other by one gular scale; three preventral scales present between gular and ventral 1; dorsal scales in 15–15–15 transverse rows, smooth throughout, lacking apical pits and supracloacal tubercles; dorsal scales in eight rows at level of subcaudal 10; ventrals 159; cloacal scute divided; subcaudals 64 plus tail tip (an estimated 5–10 subcaudal scales missing), paired; ventrals plus subcaudals 223 plus tail tip. Ventrals Subcaudals TAL/TOL n SpeciesMales/FemalesMalesFemalesMalesFemalesMalesFemalesMaximum TOL T. excelsa 3/1169161–178706124%23%400 mm T. gottei 6/1142–15814762–677024–26%26%367 mm T. impensa 9/12161–171164–17268–7264–7221–24%21–25% ~ 725 mm T. olŋmpia 1/0148—49—21%—338 mm T. psittaca 4/3153–163154–16163–73—24–25%—413 mm T. stenigrammi 1/1164159————173 mmþ T. taeniata 9/1141–15215060–705923–27%23%415 mm T. tritaeniata 1/2157155–161—59–65—23–24%273 mm Color of holotype in life (Fig. 3). —Coloration of freshly euthanized holotype (UTADC 8752–53): dorsum of head Olive-Brown (28), transforming to Sepia (219) along scale margins; anterior portion of rostral, supralabial 1, and nasals Sepia; posterior portion of rostral, internasal, and interparietal areas Grayish Horn Color (91); two Pale Horn Color (92) lateral spots on head, first occupying postnasal, lower edge of prefrontal, anterior tip of preocular, and most of supralabials 1 and 2, second lateral spot located posterior to eye and occupying most of lower postocular, lower borders of upper postocular and parietal, part of supralabials 4 and 6 and all of supralabial 5, and anterior half of anterior temporal; nuchal collar Warm Buff (118) dorsally to Pale Horn Color laterally, with Sepia anterior and posterior borders, collar about two dorsal scales long and occupying posterior tips of parietal and secondary temporal, and posterior third of supralabial 7; dorsum of body Hair Brown (119A), turning Sepia alongside all pale body stripes and toward nuchal collar; middorsal stripe Warm Buff, occupying only vertebral scale row anteriorly on body, extending onto adjacent edges of paravertebral scale rows posteriorly on body, beginning one and a half scales posterior to nuchal collar, stripe extending onto tail to reach tip, occupying adjacent halves of two most-middorsal scale rows; lateral body stripe on each side Buff-Yellow (53), occupying adjacent thirds of scale rows 3 and 4, beginning three scales posterior to nuchal collar, pale stripe on tail occupying second dorsal scale row and ending about 10 scales anterior to tip; underside of head mostly Pearl Gray (81), but Smoke Gray (44) toward mouth, and mental and buccal borders of infralabials 1–4 Sepia; venter of body, including anal plate, Pearl Gray, Smoke Gray toward sides, underside of tail Pearl Gray; ventral coloration extends to middle of dorsal scale row 1 on body, occupying all of dorsal scale row 1 on tail. Color of holotype in preservative (Fig. 4). —Dorsal surfaces brown with white nuchal collar and white middorsal and lateral stripes; pale middorsal stripe confined to vertebral row on at least anterior half of body, and extending onto adjacent edges of paravertebral rows posteriorly on body, with lateral tips of each vertebral scale dark brown; pale middorsal stripe not narrowing anteriorly on first scale involved in stripe; pale middorsal stripe extending onto tail, fading out at about a third of its length until becoming poorly visible at about midlength of tail; pale middorsal stripe beginning on 4th scale posterior to parietals; pale lateral stripe present on adjacent thirds of scale rows 3 and 4, bordered above by continuous, thin dark brown line; pale lateral stripe beginning on 5th scale posterior to ultimate (7th) supralabial, extending well onto tail; lower two-thirds of scale row 1 white; upper third of scale row 1, all of scale row 2, and lower third of scale row 3 same shade of brown as paravertebral rows; pale nuchal collar complete, involving posterior tips of parietals plus one and one-half scale rows dorsally, pale nuchal collar also involving posterior third of ultimate supralabial plus one scale row laterally; dorsal surface of head generally brown; off-white crossbar involving upper tip of rostral and both internasals, crossbar bordered posteriorly by dark brown cross-line involving posterior tips of internasal scales; lateral tips of each parietal scale white, contacting off-white line following parietal suture; anterior lateral white spot involving lower tips of prefrontals, postnasal, posterior two-thirds of supralabial 1, and all but posterior edge of supralabial 2; larger lateral white spot also present posterior to eye, involving all of supralabial 5 and anterior edge of supralabial 6, extending dorsally to cover about anterior half of anterior temporal; dark brown lateral bar covering posterior half of anterior temporal, all of posterior temporal, and posterior three-quarters and anterior two-thirds of supralabials 6 and 7, respectively; dark brown lateral bar extending dorsally to lateral third of parietal, not narrowing dorsally; about anterior half of mental and lateral tips of infralabials 1 and 2 dark brown, small portions of infralabials 3 and 4 with brown spots, remainder of ventral surface of head white; ventral surface of body white for full length, white pigment also extending onto lower tips of scale row 1 on body; subcaudal surface white, except upper tips of subcaudal scales dark brown. Variation in paratype. —The paratype is an adult male that lacks part of its head and most of its tail, thus no measurements were taken, nor was the color pattern recorded by JRM when examined. Data and photographs in preservative provided by J.E. Mérida (26 December 2016), however, indicate that it has 164 ventrals and a narrow vertebral stripe anteriorly on the body and extending onto adjacent edges of paravertebral scale rows posteriorly on body; thus, similar to that of the holotype. Distribution and habitat. —The holotype of Tantilla stenigrammi was found at 1630 h beside the school in Cuaca, Olancho (Fig. 5). The specimen was under a root mat on the buttress of a fallen tree about 1 m above ground, and that site located between a soccer field and secondary vegetation. The historic habitat of Cuaca was pine–oak forest (Premontane Moist Forest of Holdridge [1967]) along the flanks of Montaña de Botaderos in a transition zone between pine– oak forest and broadleaf rainforest (Premontane Wet Forest). The highest elevations of the Botaderos Mountains contain remnants of closed-canopy broadleaf rainforest (Lower Montane Wet Forest), usually referred to as ‘‘cloud forest.’’ The paratype was found dead during the day near the start of a foot trail in pine–oak forest (Premontane Moist Forest) along the lower flanks of the Sierra de Agalta. That trail also passes through Premontane Wet Forest, before ultimately entering Lower Montane Wet Forest as it gains altitude. Thus, both specimens of Tantilla stenigrammi were found in similar, formerly forested areas, and at similar elevations (895 and 1180 m) in north-central and central Olancho in east-central Honduras. Both localities are in low mountains on both sides of the upper drainage of the Río Sico Tinto (also called Río Negro) that forms the Agalta Valley (Fig. 5). Remarks. —A photograph of the holotype (UTA R-52591) of Tantilla stenigrammi was previously published in McCranie et al. (2006; as T. taeniata). Etymology. —The specific epithet, stenigrammi, is a Greek substantive used as an adjective and meaning narrow-lined, derived from the Greek adjective stenos (narrow) and the Greek noun grammi (line), in allusion to the narrow middorsal stripe that is characteristic of the species., Published as part of McCranie, James R. & Smith, Eric N., 2017, A Review of the Tantilla taeniata Species Group (Reptilia: Squamata: Colubridae: Colubrinae) in Honduras, with the Description of Three New Species, pp. 338-348 in Herpetologica 73 (4) on pages 339-342, DOI: 10.1655/Herpetologica-D-16-00080.1, http://zenodo.org/record/7712317, {"references":["McCranie, J. R. 2011 a. The Snakes of Honduras: Systematics, Distribution, and Conservation. Society for the Study of Amphibians and Reptiles, USA.","McCranie, J. R. 2011 b. A new species of Tantilla of the taeniata species group (Reptilia, Squamata, Colubridae, Colubrinae) from northeastern Honduras. Zootaxa 3037: 37 - 44.","Campbell, J. A. 1998. Comments on the identities of certain Tantilla (Squamata: Colubridae) from Guatemala, with the descriptions of two new species. Scientific Papers, Natural History Museum, University of Kansas 7: 1 - 14.","Holdridge, L. R. 1967. Life Zone Ecology, revised edition. Tropical Science Center, Costa Rica.","McCranie, J. R., J. H. Townsend, and L. D. Wilson. 2006. The Amphibians and Reptiles of the Honduran Mosquitia. Krieger Publishing Company, USA."]}

Tantilla gottei sp. nov. (Figs. 8, 9) Tantilla taeniata: Wilson and Meyer 1971:32, in part; Wilson 1982:56, in part; Wilson and Meyer 1982:109, in part; Wilson and Meyer 1985:103, in part; Wilson and McCranie 1999:328, in part; McCranie 2011a:227, in part; McCranie 2011b:44, in part; Wilson and Mata-Silva 2015:457, in part. Holotype. — ROM 19996, an adult female from El Picacho (14°07 ′ N, 87°11 ′ W), a zoological park located near Tegucigalpa, 1280 m elevation, department of Francisco Morazán, Honduras, collected 12 December 1986 by Jorge Porras. Paratypes (n ¼ 4). — UNAH 5394, an adult male from Galeras (13°55 ′ N, 86°59 ′ W) about 5 km north of Güinope, 860 m elevation, El Paraíso, Honduras; MCZ R49886, a juvenile male from El Zamorano (14°00 ′ N, 87°01 ′ W), 800 m elevation, Francisco Morazán, Honduras; UNAH 3903, 4833, a subadult male and an adult male, respectively, from Tegucigalpa (centered at about 14°06 ′ N, 87°12 ′ W), 930– 1000 m elevation, Francisco Morazán, Honduras. Referred specimens (n ¼ 2). — UNAH 6141, a dehydrated specimen, from Tegucigalpa (centered at about 14°06 ′ N, 87°12 ′ W), 930–1000 m elevation, Francisco Morazán, Honduras; UNAH 1765 from Alauca (13°51 ′ N, 86°41 ′ W), 500 m elevation, El Paraíso, Honduras. Diagnosis. — Tantilla gottei is defined by the following combination of characters: (1) middorsal stripe pale cream (in preservative), extending length of body and most of tail, on vertebral row and adjacent third of paravertebral rows; (2) pale cream (in preservative) lateral stripe occupying adjacent third of scale row 3 and lower three-quarters of scale row 4; (3) ventrolateral area white as is that of lateral stripe (in preservative); (4) lower two-thirds of scale row 1 colored similarly to that of ventrals; (5) lateral edges of ventrals white, except some dark spots present (in preservative); (6) ventral and subcaudal surfaces yellow (in life) and white (in preservative); (7) female with 147 ventrals, male ventrals 142–158, subcaudals 70 in female, 62–67 in males; (8) female tail length 26% of total length, 24–26% in males. Tantilla gottei most closely resembles T. taeniata in having low numbers of ventral scales, but differs in having yellow ventral surfaces in life, and by having a pale brown internasal crossbar that is confluent with the pale lateral spot anterior to the eye, (vs. ventral surfaces pale purplish pink grading to darker salmon color on posterior two-thirds of body and tail in life, and in lacking a internasal crossbar). Tantilla gottei differs from T. impensa, T. olŋmpia, and T. stenigrammi by having the middorsal pale stripe involving all of vertebral scale row and adjacent third to half of paravertebral scale rows (vs. middorsal stripe confined to vertebral row in those three species, with that of T. olŋmpia also reduced to a series of dashes; but that stripe also extends onto adjacent edges of paravertebral scale rows posteriorly on body in T. stenigrammi). Tantilla gottei differs further from T. impensa and T. stenigrammi in having 142–152 ventrals in both sexes combined (vs. ventrals 161–172 in T. impensa, and 159 in one T. stenigrammi), and further from T. impensa in having 204–217 ventral plus subcaudal scales in both sexes combined (vs. 233–240). Tantilla gottei differs further from T. olŋmpia in having 62–70 subcaudals in both sexes combined and having a complete pale lateral stripe (vs. 49 subcaudals and pale lateral stripe reduced to dots in T. olŋmpia). Tantilla gottei differs from T. excelsa in having 142–152 ventrals and 204–217 ventral plus subcaudal scales in both sexes combined and having a pale brown internasal crossbar that is confluent with the pale lateral spot anterior to the eye (vs. 161–178 ventrals and 222–239 ventral plus subcaudal scales, and internasal crossbar absent). Tantilla gottei differs from T. psittaca by having yellow ventral and subcaudal surfaces in life and cream in preservative and in having 142–152 ventrals and 204–217 ventrals plus subcaudals in both sexes combined (vs. those surfaces pink and grading to red in life and pale brown medially and pale brown with pinkish tinge on posterior third of body and under tail in preservative, and 153–163 ventrals and 216–234 ventrals plus subcaudals). Tantilla gottei differs from T. tritaeniata in having 142–152 ventrals in both sexes combined and having a complete pale nuchal collar (vs. 155–161 ventrals and pale nuchal collar divided). Description of holotype. —A subadult female; TOL 367 mm; SVL 270 mm; TAL 97 mm (26.4% of TOL); HL 10.8 mm; HW 5.6 mm (between level of angle of mouth and eye); head barely distinct from neck; snout broadly rounded in dorsal view; eye length 1.4 mm; snout length 3.3 mm, about 2.4 times longer than eye; pupil circular; rostral about 1.4 times wider than high (2.2 X 1.6 mm); internasal length about 0.7 times of width (1.1 X 1.6 mm); prefrontal about 1.7 times larger than internasal (1.9 X 1.1 mm), as wide as long (both dimensions 1.9 mm); median prefrontal suture 1.6 mm, about 0.5 times as long as frontal (1.6 X 3.5 mm); frontal with slight anterior extension, V-shaped posteriorly, about 1.3 times longer than wide (3.5 X 2.7 mm), about 1.1 times longer than distance from its anterior edge to tip of snout (3.1 mm); parietal about 1.5 times longer than wide (4.4 X 3.0 mm), median suture length 2.8 mm, about 0.8 times length of frontal; parietals contacting six nuchal scales; supraocular about 1.9 times longer than wide (2.6 X 1.4 mm), bordering orbit, contacting parietal, upper postocular, preocular, and prefrontal. Nasal divided; anterior nasal contacting rostral, internasal, and supralabial 1; posterior nasal contacting internasal, prefrontal, preocular, and supralabials 1 and 2; nostril located in central portion of nasal (forming division of nasal); loreal absent; preocular single, not higher than long (1.3 X 1.4 mm), lower edge contacting supralabials 2 and 3; postoculars 2, upper about same size as lower (height of both scales 0.9 mm); temporals 1 þ 1, anterior temporal 1.3 times longer than high (1.8 X 1.4 mm), posterior temporal 1.3 times longer than high (1.9 X 1.5 mm); supralabials 7, 3 and 4 bordering orbit, 4 and 5 contacting lower postocular, 5 contacting anterior temporal and lower postocular, 6 contacting anterior temporal, and 7 contacting anterior and posterior temporals; infralabials 6, 1–4 contacting anterior pair of chinshields, first infralabial on each side separated medially by contact with anterior chinshield; mental about 0.9 times as wide as long (1.8 X 1.9 mm), contacting first pair of infralabials and anterior pair of chinshields; anterior chinshields about 1.6 times longer than wide (2.2 X 1.4 mm), not extending laterally to border of lip; posterior pair of chinshields about 1.7 times longer than wide (1.7 X 1.0 mm), posterior tips separated from each other by one gular scale; four preventral scales present between gular and first ventral; dorsal scales in 15–15–15 transverse rows, smooth throughout, lacking apical pits and supracloacal tubercles; dorsal scales in six rows at level of 10th subcaudal; ventrals 147; cloacal scute divided; subcaudals 70, paired; ventrals plus subcaudals 217. Color of holotype in life. —Nothing was recorded on color in life of the holotype, other than that the ventral and subcaudal surfaces were yellow (J. Porras, personal communication). Color of holotype in preservative (Figs. 8, 9). —Dorsal surfaces brown with pale cream nuchal collar and pale cream middorsal and white lateral stripes; pale middorsal stripe on vertebral row and adjacent third of paravertebral rows; pale middorsal stripe slightly narrowing anteriorly on first scale involved in stripe; pale middorsal stripe extending onto tail to nearly its tip; pale middorsal stripe beginning on fifth scale posterior to parietals; pale lateral stripe present on upper two-thirds of scale row 3 and lower three-quarters of scale row 4, bordered above by continuous, thin dark brown line on upper quarter of scale row four and below by incomplete dark brown line on lower third of scale row 3; pale lateral stripe beginning on scale 5 posterior to supralabial 7, extending well onto tail; lower two-thirds of scale row 1 cream; upper third of scale row 1 and all of scale row 2 same shade of brown as dorsal ground color; pale nuchal collar complete, involving posterior tips of parietals plus ~ 1.5 scale rows dorsally, pale nuchal collar also involving posterior third of ultimate supralabial plus one scale row laterally; dorsal surface of head generally brown, except distinct pale brown crossbar involving most of internasals and anterior third of prefrontals, pale crossbar confluent with pale cream lateral spot anterior to eye; lateral pale spot involving postnasal, supralabial 1, and anterior third of supralabial 2; larger pale cream spot also present posterior to eye, involving all of supralabial 5, anterior edge of supralabial 6, and posterior half of supralabial 4, pale spot extending dorsally to cover about anterior half of anterior temporal, and lower postocular; dark brown lateral bar covering posterior twothirds of anterior temporal, all of posterior temporal, and posterior three-quarters and anterior two-thirds of supralabials 6 and 7, respectively; dark brown lateral bar continuous with dorsal surface of head color; dark brown mottling and tiny spots present on mental and first infralabial, respectively; dark brown spot also present on anterior tips of infralabial four; ventral surface of head pale cream; ventral surface of body cream for full length, pale pigment also extending onto lower tips of scale row 1 on body; subcaudal surface cream, except upper tips of subcaudal scales dark brown. Variation in paratypes. —The four male paratypes show the following variation: TOL 165–343 mm (mean ± 1 SD, 265.8 ± 75.1 mm); SVL 125–254 mm (199.5 ± 54.9 mm); TAL 24–26% of TOL; ventrals 142–158 (147.3 ± 4.1); subcaudals 62–67 (64.0 ± 2.2); and ventrals plus subcaudals 204–215 (211.3 ± 5.0). Some of the paratypes were recorded to have slightly narrower pale lateral stripes than the holotype, with those variations involving the adjacent thirds of scale rows 3 and 4. Distribution and habitat. —Almost nothing is known about the habitats of Tantilla gottei, except the low montane habitats where this species was collected are in pine forest in the south-central portion of Honduras (Premontane Moist Forest of Holdridge 1967), with one exception. That exception is a single locality in Lowland Dry Forest (Holdridge 1967). The holotype was collected alive by a worker on the grounds of the El Picacho Zoo during December. One paratype (MCZ R49886) was collected during May. The known elevational range is 500–1280 m. The known geographical distribution of Tantilla gottei is from the upper Río Choluteca drainage, in the vicinity of the capital city of Tegucigalpa, Francisco Morazán, east- and southeast-ward to the south-central portion of the department of El Paraíso (Fig. 5) along the middle portion of the Río Choluteca. Anthropogenic alteration of forest habitats in recent times has further isolated these localities. In addition, most of those surviving pine forests are burned on an annual basis. Those repeated burnings serve to have a permanent drying effect on those burned areas. Therefore, T. gottei should be considered an endangered species. Etymology. —The name gottei is a patronymic noun honoring Steve W. Gotte, a long-time friend of the first author, who also made several field trips with JRM to Honduras. Steve has also been a long-term employee of the US Department of Interior at the National Museum of Natural History in Washington, DC, and most recently in Suitland, Maryland., Published as part of McCranie, James R. & Smith, Eric N., 2017, A Review of the Tantilla taeniata Species Group (Reptilia: Squamata: Colubridae: Colubrinae) in Honduras, with the Description of Three New Species, pp. 338-348 in Herpetologica 73 (4) on pages 344-346, DOI: 10.1655/Herpetologica-D-16-00080.1, http://zenodo.org/record/7712317, {"references":["Wilson, L. D., and J. R. Meyer. 1971. A revision of the taeniata group of the colubrid snake genus Tantilla. Herpetologica 27: 11 - 40.","Wilson, L. D., and J. R. Meyer 1982. The snakes of Honduras. Milwaukee Public Museum Contributions in Biology and Geology 6: 1 - 159.","Wilson, L. D., and J. R. Meyer. 1985. The Snakes of Honduras, 2 nd edition. Milwaukee Public Museum, USA.","Wilson, L. D., and J. R. McCranie. 1999. The systematic status of Honduran populations of the Tantilla taeniata group (Serpentes: Colubridae), with notes on other populations. Amphibia-Reptilia 20: 326 - 329.","McCranie, J. R. 2011 a. The Snakes of Honduras: Systematics, Distribution, and Conservation. Society for the Study of Amphibians and Reptiles, USA.","McCranie, J. R. 2011 b. A new species of Tantilla of the taeniata species group (Reptilia, Squamata, Colubridae, Colubrinae) from northeastern Honduras. Zootaxa 3037: 37 - 44.","Wilson, L. D., and V. Mata-Silva. 2015. A checklist and key to the snakes of the Tantilla clade (Squamata: Colubridae), with comments on taxonomy, distribution, and conservation. Mesoamerican Herpetology 2: 418 - 498.","Holdridge, L. R. 1967. Life Zone Ecology, revised edition. Tropical Science Center, Costa Rica."]}

Sigalegalephrŋnus minangkabauensis sp. nov. Holotype. — Museum Zoologicum Bogoriense Amphibian Collection, MZB 25738 (field number ENS 16028), an adult male (Fig. 7) from Gunung Kunyit, Kecamatan Panyabungan Selatan, Kabupaten Kerinci, Provinsi Jambi, Indonesia, 2.26013°S, 101.49512°E at 1402 m, collected by E. Wostl, E.N. Smith, W. Trilaksono, and G. Barraza on 24 June 2013. Diagnosis and comparison. —The following combination of characters is unique to Sigalegalephrŋnus minangkabauensis: (1) A small (19.32 mm SVL) and slender toad without parotoid glands. (2) Fingertips I and II are rounded and not expanded. (3) Fingertips III and IV are rounded and expanded. (4) The toe tips are rounded but not expanded. (5) The webbing is rudimentary in the hands and moderate in the feet. (6) The dorsum is light greenish-brown with a middorsal pinstripe extending from the tip of the snout to the vent. (7) The flanks have a single stroke of dark brown extending from the posterior end of the orbit to the inguinal region. (8) The dorsal surface is moderately tuberculate. (9) The ventral surface is smooth with scattered black spots. Sigalegalephrŋnus minangkabauensis can be distinguished from S. mandailinguensis (characters in parentheses) based on the following differences. The tympanum is barely discernible (tympanum distinct). The finger-pads are moderately defined (finger-pads prominent). The fingertips are rounded but not expanded (distinctly spatulate on tips III to IV). The hands lack subarticular tubercles (distinct subarticular tubercles under Fingers III and IV). The webbing of the foot is more extensive, extending to the last phalanx on Toes I and II (last phalanx free of webbing). The pads on toes are moderately defined (pads prominent). The feet lack subarticular tubercles (distinct subarticular tubercles under Toes IV and V). The overall texture is glossy with fewer tubercles on the dorsum and flanks (overall texture rugose, body and flanks extensively tuberculate). Taking into account the uncorrected genetic distance between the two species of Sigalegalephrŋnus (Table 2), the aforementioned comparisons provide adequate diagnostic characters to warrant S. minangkabauensis as a species distinct from S. mandailinguensis. Description of holotype. —The holotype (sex indistinguishable) has SVL of 19.32 mm; head length 6.94 mm; head width 6.57 mm; snout length 2.7 mm; eye length 2.2 mm; eye–nares length 1.6 mm; distance between nares to tip of snout 0.6 mm; internarial distance 1.8 mm; intercanthal distance 3.6 mm; forearm length 6.0 mm; hand length 5.5 mm; femur length 8.5 mm; tibia length 8.71 mm; tarsal length 5.1 mm; foot length 7.5 mm; width of fingertip pad of Finger III 1.6 mm; Finger IV 1.4 mm. Body slender, head little longer than wide; head length 36.1% SVL, head width 34.0% SVL; snout length 14% SVL; canthus rostralis concave; loreal area without tubercules and concave; snout truncated and slightly sloping back toward mouth; snout mucronate and with prominent median keel in dorsal view; eye length 11.4% SVL; pupil horizontal; upper eyelid granular; tympanum barely visible, with no supratympanic fold; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval-shaped, longer than wide; skin of dorsal surfaces rough to finely shagreen with few large, scattered tubercles; tubercles small, rounded, and almost without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smooth with few fine warts; forearm length 31.1% SVL; hand length 28.5% SVL; relative lengths of Fingers I Color in life. —Edges of lore and head golden with black shades; area below eyes with prominent white marking with yellowish tint; dorsum light greenish-brown with light brown hourglass figure extending from posterior of orbit to top of the sacroiliac joint; hourglass shape ends with distinct horizontal black bean color mark on each side; yellowishgreen marking on each shoulder; flanks black with red tubercles, maculated with greenish-yellow blotches, and possessing very prominent dark brown stripe starting from posterior end of orbit to inguinal region; inguinal areas greenish with golden tint; sacroiliac joint to inguinal region of flanks, golden yellowish-green; dorsal sides of limbs light brown; forearm, femur, tibia–fibula, and tarsus with distinct dark spot encircled with golden-yellowish-green color; venter opaque; throat golden-yellow; abdomen, ventral side of arms and legs pinkish with scattered yellow and black blotches; webbing in hand and foot translucent. Color in preservative. —In preservative, the animal appears dull because it has lost its golden-yellowish and greenish colors. The hourglass pattern has turned gray. The venter has lost all of its pinkish and golden-yellowish shades and turned a greenish-white. Etymology. —The specific epithet refers to the Minangkabau or Minang ethnic group inhabiting the region where the new species was found. Common name. —Minangkabau Puppet Toads. Province, Sumatra (MZB 25738). A color version of this figure is available online. Distribution and natural history. — Sigalegalephrŋnus minangkabauensis is known only from Gunung Kunyit from an elevation of 1428 m (Fig. 4). The holotype was found perched on a leaf ~ 1.25 m above ground, by the edge of a forest stream at 2015 h. Before collecting it, ENS watched the specimen move in reverse toward the edge of the leaf on which it was perched, where it defecated (outside of the surface of the leaf), and then return to its original position. The holotype weighed 0.5 g., Published as part of Smart, Utpal, Sarker, Goutam C., Arifin, Umilaela, Harvey, Michael B., Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2017, A New Genus and Two New Species of Arboreal Toads from the Highlands of Sumatra with a Phylogeny of Sundaland Toad Genera, pp. 63-75 in Herpetologica 73 (1) on pages 63-75, DOI: 10.1655/Herpetologica-D-16-00041, http://zenodo.org/record/7716435

Sigalegalephrynus gen. nov. Type species. — Sigalegalephrŋnus mandailinguensis by present designation. Diagnosis and comparisons. —The genus can be diagnosed based on the following ecological attributes and morphology: medium-sized (Ansonia (> 40 mm SVL), commonly called Stream toads, is typically found on low vegetation near watercourses. The generally diminutive members of Pelophrŋne (Leptophrŋne (> 40 mm SVL) occur on shrubs. The only other genus of toads that has true arboreal habits is Rentapia, whose members typically tend to be stockier (> 70 mm SVL) than Sigalegalephrŋnus. The genus Phrŋnoidis is represent-ed by two large (> 70 mm SVL) semiaquatic toads, usually found on rocks along streams and rivers; the members of the Duttaphrŋnus (> 40 mm SVL) and Ingerophrŋnus (> 40 mm SVL) display terrestrial or somewhat riparian habits. The monotypic genus Pseudobufo is represented by a large (> 75 mm SVL) and aquatic species with completely webbed feet that inhabits the peat swamps of the Malay Peninsula, Borneo, and Eastern Sumatra. Like all other Sundaland toad genera (characters in parentheses), the new genus possesses a visible or slightly visible tympanum. Sigalegalephrŋnus most closely resembles Ansonia; however, the former lacks mandibular spines (mandibular spines present) and possesses combined femur and tibia lengths smaller than its SVL (SVL Pelophrŋne, in Sigalegalephrŋnus Finger I projects beyond the webbing by two phalanges (reduced Finger I, with one or no phalanges projecting beyond webbing), and males possess nuptial excrescences with well-keratinized spicules (poorly spiculated with only slight keratinization or not keratinized at all). Unlike Rentapia, Sigalegalephrŋnus lacks paratoid glands (paratoids prominent). The new genus can be told apart from Ingerophrŋnus by the lack of well-defined parallel crests between the eyes (parallel crests prominent). Unlike Leptophrŋne, Sigalegalephrŋnus lacks enlarged tubercles at the base of each toe, between at the articulation of the first phalanx and metacarpus (large tubercles present). Additionally, males of L. cruentata are unique among Southeast Asian toads in having nuptial excrescences that are white and swollen on the first and second fingers; males of Sigalegalephrŋnus (and all other genera) have nuptial excrescences with no white and swollen tissue. Unlike Phrŋnoidis, Sigalegalephrŋnus has slender limbs (limbs robust) and toes that are less than half webbed (toes fully webbed, with the exception of the fourth). Unlike Pseudobufo —the only Sundaland toad with fully webbed toes— Sigalegalephrŋnus has toes that are less than half webbed. Males of Sigalegalephrŋnus can be distinguished from all other toads in the region by the presence of an elongate inner metacarpal-thenar tubercle, which is as distinct and large as the outer metacarpal tubercle, and is located medially (Fig. 2). Males of Leptophrŋne cruentata have an elongate and medially located inner metacarpal tubercle, but this is less distinct and noticeably smaller than the outer metacarpal tubercle, whereas Rentapia and Pelophrŋne lack the inner metacarpal tubercle altogether. Fingertips three and four of the new genus are truncated, reflecting arboreality as in Pelophrŋne, Sabahphrŋnus, Rentapia, and some species of Ansonia. Etymology. —The generic name is derived from the name given by the indigenous Batak people of the Toba region in Sumatera Utara to life-sized wooden puppets called Sigale Gale. These puppets are used during the papurpur sepata funerary festivals to placate the spirits of the dead who have left no children behind. The suffix is derived from the masculine and Latinized Greek noun for toad, phrŋnos. The new genus, with a relatively large size compared with most arboreal toads in the region, lanky hands, and a wood-brown complexion, is evocative of the Sigale Gale. Common name. —Puppet Toads. Content. —Our phylogenetic analyses indicate the presence of two species within the new genus: S. mandailinguensis Smart et al.; and S. minangkabauensis Smart et al., Published as part of Smart, Utpal, Sarker, Goutam C., Arifin, Umilaela, Harvey, Michael B., Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2017, A New Genus and Two New Species of Arboreal Toads from the Highlands of Sumatra with a Phylogeny of Sundaland Toad Genera, pp. 63-75 in Herpetologica 73 (1) on pages 63-75, DOI: 10.1655/Herpetologica-D-16-00041, http://zenodo.org/record/7716435

Harvey, Michael B., O'Connell, Kyle A., Wostl, Elijah, Riyanto, Awal, Kurniawan, Nia, Smith, Eric N., Grismer, L. Lee (2016): Redescription Cyrtodactylus lateralis (Werner) (Squamata: Gekkonidae) and Phylogeny of the Prehensile-tailed Cyrtodactylus. Zootaxa 4107 (4): 517-540, DOI: http://doi.org/10.11646/zootaxa.4107.4.3

Cyrtodactylus lateralis (Werner) Gymnodactylus lateralis Werner 1896: 11. Holotype female (ZMB 12029, Fig. 1, 2, examined) from ���Sumatra.��� [herein traced to Binjai, Sumatera Utara, Indonesia]. Gymnodactylus lateralis Werner: Werner 1900: 499; Barbour 1912: 80; De Rooij 1915: 7. Cyrtodactylus lateralis (Werner): Kluge 1993: 8; Manthey and Grossmann 1997: 223; R��sler 2000: 66; Teyni�� et al. 2010: 33. Referred specimens examined (19). Seven males (MZB 13171, MZB 13173; UTA 62916, UTA 62919 ���62921) and six females (MZB 13172, MZB 13174 ���13176, MZB 18330, UTA 62917 ���62918) from trail up SE slope of Gunung Seulawah Agam, Aceh Besar Regency, Aceh Province, Sumatra, Indonesia, 685���1065 m collected 30 July and 1 August 2015 by M. B. Harvey and E. Wostl and one female (UTA 62915) from Gunung Batee Meucica, Aceh Besar Regency, Aceh Province, Sumatra, Indonesia, 5.26157 �� N, 95.54176 �� E, 498 m collected 1 August 2015 by G. C. Sarker and I. Sidik; two males (ZMB 65273 ���65274) and two females (ZMB 65271 ���65272) from Gurah, Aceh Province, Sumatra, Indonesia, collected January 1997 by C. Sch��fer and W. Grossmann (traced to a village of the same name in Kecamatan Peukan Bada, Kabupaten Aceh Besar). One female (ZMB 49125) from Ketambe, Aceh Province, Sumatra, Indonesia, collected March 1990 by W. Denzer, C. Steiof and U. Manthey. Diagnosis. A large species of Cyrtodactylus reaching 100 mm SVL (LSUHC 12579) and distinguished from all congeners by the following combination of characters: (1) body robust; limbs and digits moderate in length; (2) tail prehensile; intact tails 0.97���1.22 times longer than body; (3) tubercles extending from frontal region and supraorbital skin to cover most of tail, 16���20 irregular longitudinal rows of tubercles at midbody, 21���28 irregular transverse rows between limbs; (4) dorsal antebrachium and brachium tuberculate; (5) tubercles on tail extending to 90 % of its length; (6) 51���66 ventrals in a transverse row at midbody; (7) conical, spinose tubercles in ventrolateral fold; (8) subcaudals not transversely enlarged; (9) moderate longitudinal sulcus in precloacal region of males; (10) 9���13 precloacal pores in males (0���15) pore primordia in females), femoral pores absent, precloacal pores sunken into precloacal sulcus; pore secretions red; (11) greatly enlarged precloacal pore-bearing scales present; (12) ventral surface of thighs entirely granular or granular except for 2���13 (count combined for both sides) widely spaced enlarged femoral scales; (13) 18���24 lamellae under fourth toe; (14) cloacal tubercles 1���2 on each side, usually in contact with one another; (15) dorsum gray to brown with dark brown markings; venter pale pink, immaculate or with diffuse, darkly pigmented scales forming no obvious pattern; (16) labials pale, contrasting with darker facial band; occiput with few large blotches; postocular stripe brown, edged dorsally by thin black line then by pale pigmented scales; (17) 5���6 transverse, usually broken bands on body; 7���9 bands on tail; caudal bands complete ventrally though faint. Comparisons. Cyrtodactylus lateralis is very similar to C. durio. Cyrtodactylus lateralis has rounded to subpyramidal tubercles in the temporal region, on the occiput, and on the neck, whereas tubercles in these areas are noticeably larger and distinctly spinose in C. durio. We scored three new specimens of C. durio for the same characters used in the description of C. lateralis. Ranges of meristic and mensural characters of C. durio overlap those of C. lateralis, however extremes of three characters were slightly higher or lower than those of C. lateralis. The three specimens of C. durio (data for KU 173086 followed by KU 173087 then LSUHC 12579 vs. ranges [mean �� standard deviation, sample size] for C. lateralis) had slightly fewer tubercles at midbody (17, 15, and 15 vs 16���20 [18 �� 1, n = 13]), slightly fewer lamellae under finger IV (15, 17, and 17 vs. 16���20 [18 �� 1, n = 14]), and a slightly larger orbit (diameter of orbit 27.8 %, 28.7 %, and 26.5 % of head length vs. 20.6���27.6 % [25.4 �� 1.6, n = 16]) than the series of C. lateralis. All specimens of C. durio lack enlarged subfemoral scales, whereas 70 % of C. lateralis had these scales. Cyrtodactylus lateralis, C. brevipalmatus, C. durio, C. elok, and C. stresemanni are the only Sunda Shelf species with a spinose, prehensile tail. Unlike C. stresemanni (characters in parentheses), C. lateralis has 16���20 tubercles across midbody (13), a transversely banded dorsum (wide, dark brown chevrons on flanks separated by undulating pale vertebral stripe), and small irregular blotches on the occiput (large, dark triangular occipital mark). Unlike C. brevipalmatus and C. elok, C. lateralis has a precloacal sulcus (absent), spinose tubercles in the ventrolateral fold (tubercles absent from fold), 51���66 ventrals (35���49), and caudal whorls of six tubercles separated by granular scales ventrolaterally (caudal whorls of four tubercles; ventrolateral tubercles separated by lanceolate scales forming ventrolateral fringe). Description. Characteristics of the holotype placed in brackets follow those of all specimens examined. Large Cyrtodactylus reaching at least 93 mm SVL in females [77 mm], 82 mm in males; smallest specimen subadult female, 62.1 mm SVL; body robust, accounting for 38���49 % (45.6 �� 3.7, n = 7) of SVL in males, 43.0��� 53.9 % (48.2 �� 3.8, n = 8) [49.4 %] of SVL in females; tail 0.99���1.14 (n = 8) [0.97] times longer than body and accounting for 49.3���55.1 % (51.8 �� 2.3, n = 8) [49.3 %] of total length; head triangular in dorsal view, much wider than body and distinct from neck; head 28.9 ���34.0% (30.8 �� 1.7, n = 7) of SVL, 63.9���78.9 % (70.2 �� 6.1, n = 7) as wide as long, and 36.2���43.2 % (38.8 �� 2.6 n = 7) as deep as wide in males and 28.5���33.4 % (31.0 �� 1.7, n = 9) [30.1 %] of SVL, 64.3���71.6 % (69.4 �� 2.7, n = 9) [64.3 %] as wide as long, and 38.0��� 46.2 % (41.6 �� 2.8, n = 9) [41.1 %] as deep as wide in females. Rostral wider than or about as wide as mental (96.8���126.81 %, 106.8 �� 8.5, n = 17, of width of mental) [103.2 %], posteriorly forming border of nostril; dorsal margin of rostral notched medially by shallow groove; rostral groove bifurcating slightly ventral to center of nostril; rostral bordered posteriorly by four (50 %, n = 16), five (31 %), six (6 %), or seven (13 %) postrostrals [rostral irregular in holotype; rostral partially fragmented into two large plates just anterior to straight row of 5 postrostrals; short, bifurcating groove below these large irregular plates; rostral of ZMB 49125 also irregular, postrostral series fused to rostral, large diamond-shaped plate partially fractured within rostral in front of fused postrostrals]; internarial region narrow, 10.4���18.8 % (15.7 �� 2.0, n = 17) [17.6 %] as wide as head, 55.6���70.3 % (63.8 �� 4.5, n = 17) [60.9 %] as wide as narrowest point between orbits; narrowest point between orbits 18.7���28.9 % (24.5 �� 2.7, n = 17) [28.9 %] as wide as head; deep depression occupying most of prefrontal and frontal areas; prefrontal-frontal depression bordered laterally by raised canthal and circumorbital ridges, bordered posteriorly by slightly arched, transverse ridge located just behind orbits; central parietal region slightly depressed; scales on dorsal snout granular, rounded, smooth, subequal in size, about as large as tubercles in parietal regions and distinctly smaller than granules between eyes; tubercles extending to frontal region and supraorbital skin; tubercles on dorsal surface of head subpyrimidal and smooth, much smaller than those on body, those in occipital region surrounded by seven or eight granular scales; few tubercles around back of head subconical; dorsal body tubercles unicarinate, circular, slightly projecting from dorsum, about ten times as large as dorsal granules, 16���20 (18 �� 1, n = 13) [18] longitudinal rows at midbody including tubercles in ventrolateral folds, 21���28 (23 �� 2, n = 13) [27] irregular transverse rows between limbs; dorsal tubercles on flanks like those middorsally or attenuated into conical tubercles; dorsal granules smooth. Nostril oval, directed postero-laterally and somewhat dorsally; bordered by rostral, postrostral, supranasal, postnasals, and narrow antero-dorsal process of first supralabial; supranasal rounded to rhomboidal, about four times as large as adjacent postnasals; postnasals granular, 5���7 (6 �� 1, n = 19) [5 / 5]; snout length 36.7���41.2 % (39.3 �� 1.4, n = 16)[39.4 %] of head length and distance from orbit to nostril accounting for 71.2���78.4 % (74.1 �� 2.0, n = 17) [75.3 %] of snout length; loreal region sloping, concave just behind nostril; lips flared; loreals granular, juxtaposed; 19���25 (21.9 �� 2, n = 21) [21 / 21] loreals between nasal and anterior border of orbit; lorilabials in front of orbit flat, plate-like, enlarged relative to other loreals (1.5���2 times as large as loreals above them, but much smaller than supralabials); eye separated from supralabials by one or two longitudinal rows of tiny granules; diameter of orbit 20.6���27.6 % (25.4 �� 1.6, n = 16) [26.3 %] of head length; temporal region with granular scales separating rounded to subpyrimidal, smooth or weakly keeled tubercles; distance from posterior border of orbit to anterior border of auditory meatus 23.6���31.3 % (29.0 �� 1.9, n = 16) [30.9 %] of head length; skin fold overlapping dorsal edge of auditory meatus; auditory meatus elliptical, it���s greatest diameter vertical or directed posterodorsally less than 20 �� to vertical, positioned at posterior end of head and with its dorsal margin at same level as rictus; greatest diameter of auditory meatus 17.7���39.6 % (25.6 �� 5.8, n = 17) [25.5 %] of diameter of orbit; smallest diameter of auditory meatus 60���70 % smaller than greatest diameter; external ear canal deep; supralabials 8���12 (10 �� 1, n = 21) [9 / 10] to center of orbit; total enlarged supralabials 10���14 (12 �� 1, n = 21) [11 / 12], last separated by 2���5 small granular scales from rictus (13���17 total supralabials to rictus) [15 / 16]; first supralabial with anterodorsal extension reaching nostril and separating lorilabials from rostral; remaining supralabials subrectangular with rounded dorsal edges, reducing in size to anterior border of orbit where one or two supralabials somewhat longer than rest. Mental triangular; infralabials 7���9 to center of orbit (8 �� 1, n = 22) [7 / 7], 9���13 (10 �� 1, n = 22) [10 / 10] total enlarged infralabials to rictus; first pair of postmentals in contact medially, length of their contact about half as long as mental; second pair of postmentals as long as first pair but extending medially only one-half to two-thirds as far, separated from one another medially by 2���7 (6 �� 1, n = 17) [5] small granular scales bordering first pair of postmentals; sublabial row decreasing in size posteriorly, not differentiated from adjacent gulars at level of eye; gulars small, smooth, granular, nonimbricate, much smaller than ventrals; in front of pectoral girdle, gulars grading into progressively larger, imbricate scales; scale of chest like ventrals; ventrals smooth, imbricate, flat, circular, about three-fourths as wide as tubercles on dorsum at midbody; ventrals in transverse row at midbody 51���66 (55 �� 3) [59]; ventrolateral skin fold distinct, containing long spinose tubercles separated by granular scales intermixed with smaller spinose tubercles, subpyrimidal tubercles, and smooth rounded tubercles (Fig. 2); tubercles of ventrolateral fold lacking keels. Tail cylindrical, though somewhat flattened dorsally and ventrally, with distinct lateral sulcus proximally for about two-thirds its length; caudal tubercles 2���3 times as large as those on body, heavily keeled, and projecting, separated from one another by granular scales; caudal tubercles on about 90 % of intact tails, disposed in six longitudinal rows and in regular whorls, one tubercle of each whorl below lateral sulcus on each side; 2���3 tubercles in irregular transverse row between successive whorls on proximal one-third to one-half of tail; tubercles between whorls same shape but only one-half to one-third as large as tubercles forming whorls; size and distribution of tubercles at base of tail like posterior body; subcaudals smooth, flat, homogenous, imbricate, 2���3 times as large as granules on sides of tail; each subcaudal about as long as wide; transversely enlarged subcaudal plates absent; cloacal tubercles rounded, smooth, one (7 %, n = 27) or two (93 %) on each side, contacting one another (80 %, n = 25 paired tubercles) or separated from one another by a single granular scale (20 %) [2 / 2 cloacal tubercles contacting one another on each side of holotype]. Tubercles of limbs keeled, subpyramidal, largest same size as those on dorsum; tuberculation of brachium variable, almost entirely granular to as tuberculate as antebrachium [brachium of holotype tuberculate, but not as heavily tuberculate as antebrachium]; antebrachium tuberculate; antebrachial tubercles about six times as large as granular scales between them; granules and tubercles grading to dorsal patch of about seven rows of large juxtaposed scales covering distal antebrachium and proximal hand; ventral surfaces of brachium covered in smooth, slightly imbricate, homogenous granules; tubercles extending across postaxial and ventral surfaces of antebrachium to approximate position of radius, then transitioning to granular scales on preaxial side of antebrachium; scales on palm juxtaposed to slightly imbricate; lamellae under finger I 8���9 [8], under finger II 12���16 [13], under finger III 15���19 [16], under finger IV 16���20 [17], under finger V 13���16 [14]; one or two scales widened and similar to lamellae extending onto palm at base of digits. Thigh, shank, and postaxial half of dorsal foot (at base of toes IV���V) tuberculate; tubercles of leg subpyrimidal, unicarinate, largest about same size as those on dorsum, about six times as large as small granular scales between them; tubercles on thigh decreasing in size postaxially, grading to tiny granular scales on postaxial edge of thigh (no sharp transition along postaxial edge of thigh); ventral thigh entirely granular (30 %, n = 23) or granular except for row of 1���7 (3���13 total count including both sides; 70 % of the time; Fig. 3) flat enlarged subfemoral scales [ventral thigh entirely granular in holotype, lacking enlarged subfemoral scales]; enlarged subfemoral scales separated from one another by gaps occupied by granular scales; gaps between subfemorals increasing in width distally; sole covered in flat, juxtaposed to weakly imbricate, rounded scales; lamellae under toe I 8���10 [10], under toe II 11���16 [13], under toe III 16���19 [16]; under toe IV 18���24 [19], under toe V 17���20 [18]; one or two scales widened and similar to lamellae extending onto sole at base of toes II���V, three or four extending onto sole at base of toe I. Scales of precloacal region forming patch (Fig. 2, 3) of enlarged, smooth, imbricate scales grading to smaller ventrals anteriorly and small imbricate scales edging vent posteriorly; males with parallel series of 9���13 total precloacal pores; pore-bearing series of males sunken into shallow longitudinal sulcus; same scales of females with 0���15 pore primordia lacking secretions [no detectable pore or pore primordia in holotype]; females lacking longitudinal sulcus; scales flanking pore-bearing scales smaller than pore-bearing scales anteriorly, subequal to them posteriorly; both sexes with pair of post-cloacal slits; males with post-cloacal ossicle in wall of each slit. Coloration. In life (based on field notes of M.B. Harvey and photos of all specimens collected by the summer 2015 expedition to Aceh), Cyrtodactylus lateralis resembles tree bark. This species has a gray, light brown, or reddish brown ground coloration overlain by diffuse, fine darker speckling in addition to bold, dark brown blotches and pale areas largely devoid of speckling (Fig. 4). Dorsal tubercles are the same color as adjacent scales. Dorsally, a few, small, dark brown blotches mark the parietal and occipital regions; however, the rest of the dorsal head is usually devoid of solid dark brown blotches [holotype faded to cream with light brown markings]. The supraciliaries are suffused with orange; they may be almost entirely orange in young specimens, whereas brown pigment obscures the orange supraciliary pigment in older individuals. A wide band extends from the rostral to the eye and continues as a postocular stripe to the arm. The facial band is always weakly defined and may be quite diffuse in some specimens [loreal region of holotype brownish]. Behind the eye, weak, thin, and often broken black pinstripes edge the postocular band dorsally and ventrally [postocular band with black pinstripes extending above and slightly behind arm]. Several rows of temporals above the black pinstripe are cream, pale gray, or pale yellow. These pale temporals form a band behind the eye that breaks up into subcircular spots at the back of the head and on the neck. The labials are also pale, almost immaculate to heavily mottled but always contrasting with the facial band and gular skin [bands of melanophores covering supralabials 5���6, anterior portion of 7, and 8���9 on left/ 1���2 and 9���10 on right; chin diffusely speckled with melanophores]. In specimens with mottled labials, subcircular cream, pale gray, or pale yellow spots, each about as large as a supralabial, mark the labial series. The iris is grayish tan or tan, heavily reticulated with black lines. The ciliary body is only slightly lighter than the iris and indistinct. The tongue is white and lacks the black ventral patch described for some congeners (Harvey et al. 2015); the buccal epithelium is pink. Cyrtodactylus lateralis has a broken V-shaped mark on the nape, one narrow band on the neck, 5���6 [5] bands on the body, and 7���9 (8 �� 1) [7] bands on the tail. Bands on the neck and body usually break up into oval blotches, and pale areas separate the bands middorsally and on the upper flanks. Bands on the body usually have a ���butterflylike��� shape [Fig. 1, 4, see also plate 1, figure 4 of Werner, 1896]. The lower flanks are darkly pigmented and frequently sharply contrast with a pale ventrolateral fold or a fold of the same color with pale yellow or cream spinose tubercles [fold of holotype like flanks, covered in melanophores but with large spinose tubercles of fold white and lacking melanophores]. Caudal bands are square to rectangular, narrower than the pale intersp, Published as part of Harvey, Michael B., O'Connell, Kyle A., Wostl, Elijah, Riyanto, Awal, Kurniawan, Nia, Smith, Eric N. & Grismer, L. Lee, 2016, Redescription Cyrtodactylus lateralis (Werner) (Squamata: Gekkonidae) and Phylogeny of the Prehensile-tailed Cyrtodactylus, pp. 517-540 in Zootaxa 4107 (4) on pages 520-527, DOI: 10.11646/zootaxa.4107.4.3, http://zenodo.org/record/256578, {"references":["Werner, F. (1896) Zweiter Beitrag zur Herpetologie der indo-orientalischen Region. Verhandlungen der Zoologische- Botanischen Gesellschaft in Wien, 46, 6 - 24. http: // dx. doi. org / 10.5962 / bhl. part. 19752","Werner, F. (1900) Reptilien und Batrachier aus Sumatra. Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Tiere, 13, 479 - 508.","Barbour, T. (1912) A contribution to the zoogeography of the East Indian islands. Memoirs of the Museum of Comparative Zoology, 44, 1 - 203, pls. 1 - 8.","De Rooij, N. de (1915) The Reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria. E. J. Brill, Leiden, xiv + 384 pp.","Kluge, A. G. (1993) Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, 245 pp.","Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien Sudostasiens. Natur und Tier Verlag, Munster, 512 pp.","Rosler, H. (2000) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Teynie, A., David, P. & Ohler, A. (2010) Notes on a collection of amphibians and reptiles from Western Sumatra (Indonesia), with the description of a new species of the genus Bufo. Zootaxa, 2416, 1 - 43.","Harvey, M. B., O'Connell, K. A., Barraza, G., Riyanto, A., Kurniawan, N. & Smith, E. N. (2015) Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the southern Bukit Barisan Range of Sumatra and an estimation of their phylogeny. Zootaxa, 4020, 495 - 516. http: // dx. doi. org / 10.11646 / zootaxa. 4020.3.5","Cameron, P. (1906) A new genus of Cryptinae (Ichneumonidae) from Sumatra. The Entomologist, 39, 196 - 197. http: // dx. doi. org / 10.5962 / bhl. part. 1560","Grismer, L. L. (2008) On the distribution and identification of Cyrtodactylus brevipalmatus Smith, 1923 and Cyrtodactylus elok Dring, 1979. The Raffles Bulletin of Zoology, 65, 177 - 179.","Grismer, L. L. (2011) Lizards of Peninsular Malaysia, Singapore and their Adjacent Archipelagos. Edition Chimaira, Frankfurt, 728 pp.","Grismer, L. L., Anuar, S., Quah, E., Muin, M. A., Onn, C. K., Grismer, J. L. & Ahmad, N. (2010) A new spiny, prehensile-tailed species of Cyrtodactylus (Squamata: Gekkonidae) from Peninsular Malaysia with a preliminary hypothesis of relationships based on morphology. Zootaxa, 2625, 40 - 52.","Linkem, C. W., McGuire, J. A., Hayden, C. J., Mohammed, I. S., Bickford, D. P. & Brown, R. M. (2008) A new species of bent-toe gecko (Gekkonidae: Cyrtodactylus) from Sulawesi Island, eastern Indonesia. Herpetologica, 64, 224 - 234. http: // dx. doi. org / 10.1655 / 07 - 040.1"]}

Phylogeny of Cyrtodactylus lateralis and other Prehensile-tailed Cyrtodactylus Currently, the Reptile Database lists 206 species of Cyrtodactylus (Uetz & Ho��ek 2015). Systematists face considerable challenges when trying to identify derived morphological characters in this diverse gekkonid radiation. On the basis of morphology, Grismer et al. (2010) proposed a phylogeny of Cyrtodactylus with spinose prehensile tails. These authors constructed their tree using parsimony and polarized a small matrix of characters by treating all other congeners as outgroups. In their phylogeny, Cyrtodactylus brevipalmatus and C. elok form a clade and share digital webbing, a square tail, and a caudal fringe. Cyrtodactylus spinosus, C. stresemanni, and C. durio form a second clade and share spinose tubercles in the ventrolateral fold on the body. Within the second clade, Cyrtodactylus durio and C. stresemanni are sister species and share enlarged, ventrolateral caudal tubercles. The authors argued that a prehensile tail is a synapomorphy of these five species. Using their methods (Table 1), we add Cyrtodactylus lateralis (Werner), C. interdigitalis Ulber, C. nuaulu Oliver et al., and C. serratus Kraus to this morphological phylogeny (Fig. 6). Ulber (1993) compared C. interdigitalis to C. brevipalmatus and C. elok and noted that, unlike these two species, C. interdigitalis has a round rather than square tail. All three species share digital webbing and a ventrolateral fringe. The ventrolateral fringe consists of a distinctive row of enlarged conical tubercles separated by smaller lanceolate tubercles. The enlarged tubercles are the ventrolateral components of caudal whorls, whereas the smaller conical tubercles are the only enlarged scales separating the whorls dorsally. All three species lack spinose tubercles in the ventrolateral fold on the body. Ulber (1993) did not mention whether C. interdigitalis coils its tail at rest, however we have seen photos of specimens from Laos exhibiting this behavior. Based on this combination of traits, we add C. interdigitalis to the morphological phylogeny of Grismer et al. (2010) and place C. interdigitalis as the sister species to a clade containing C. elok and C. brevipalmatus (Fig. 6). We refer to this group of three species as the ���mainland SE Asian clade��� of prehensile-tailed Cyrtodactylus, because these species occur in Laos, Myanmar, Peninsular Malaysia, and Thailand. In their phylogeny, Grismer et al. (2010) placed Cyrtodactylus durio as the sister species to C. stresemanni with C. spinosus sister to the C. durio + C. stresemanni clade. They note that C. spinosus lacks ventrolateral caudal tubercles, but did not explain how they scored this character. We count tubercles in the middle of the tail: C. lateralis, C. durio, and C. stresemanni have six spinose tubercles per caudal whorl throughout most of the tail (the count will be higher close to the body and lower distally), whereas C. spinosus has only 4 (also noted by Linkem et al. 2008). We add C. lateralis to the C. durio + C. stresemanni Clade, because C. durio and C. lateralis are identical for the characters used to construct the morphological phylogeny (Table 1). Cyrtodactylus nuaulu and C. serratus, both from east of Wallace���s Line, have conical tubercles in the ventrolateral fold, ventrolateral caudal tubercles, and tails with whorls of six spinose tubercles extending for most of the length of the tail. On the basis of morphology, we infer that the tails of these two species are prehensile, even though neither description mentions whether the species coils its tail in a manner similar to C. spinosus and the Sunda Shelf species. The unique holotype of C. serratus was collected in the Star Mountains of New Guinea in 1987 and Kraus (2007) provided no information about its behavior or natural history. All adults of the type series of C. nuaulu were collected on trees and vines, 50���200 cm aboveground on Seram (Oliver et al. 2009). Based on these observations, we add C. nuaulu and C. serratus to the clade containing C. stresemanni, C. durio, and C. lateralis. We refer to this clade as the ���insular clade��� of prehensile-tailed geckos, because most of the species occur on islands. Of the five species of the insular clade, only C. durio occurs on the mainland. We have shown that some distinctive characters originally suggesting a close relationship between Cyrtodactylus durio and C. stresemanni are also found in C. serratus and C. nuaulu. Although the specimen has faded with age, the dorsal pattern of the holotype of C. stresemanni remains well-defined (R��sler & Glaw 2008, their figure 7). The specimen has two pairs of elongate blotches on the flanks separated by a wide, pale vertebral stripe. This dorsal pattern is unlike that of any other Sunda Shelf Cyrtodactylus, however it resembles patterns in the type series of C. nuaulu (compare especially to MZB 2325 in Oliver et al. 2009, their figure 2), C. spinosus (see especially MZB 7029 with ���less complete��� chevrons, Linkem et al. 2008: 228), and C. serratus (Kraus 2007, his figure 1 A). All four species share a wide pale vertebral stripe and few (2���3 between limbs) large, dark-brown chevron-shaped blotches. The vertebral stripe may be interrupted by the chevrons medially or divide the chevrons as it does in the holotypes of C. serratus and C. stresemanni and some paratypes of C. spinosus (MZB 7029) and C. nuaulu (MZB 2329). Although the dark triangular blotch on the nape of C. stresemanni might at first appear to ally it with C. elok and C. brevipalmatus, C. nuaulu, C. serratus, and C. spinosus have triangular blotches on their napes as well. Cyrtodactylus durio and C. lateralis unequivocally lack the distinctive color pattern of C. stresemanni and the three other species. Like C. brevipalmatus, C. elok, and C. interdigitalis, C. durio and C. lateralis are banded species lacking a pale vertebral stripe. To determine polarity of the distinctive color pattern of the Wallacean and Papuan species, we reviewed recent descriptions and revisions of Cyrtodactylus from across the distribution of the genus. We were unable to find other species with a pale vertebral stripe and 2���3 dark brown chevrons. C. zugi Oliver has two large brown chevrons, but lacks a vertebral stripe. The pattern of the Wallacean and Papuan prehensile-tailed Cyrtodactylus is most likely derived, however, its distribution is not congruent with the morphological phylogeny regardless of polarity. If derived, the pattern is a synapomorphy of the insular clade, with a reversal to the banded pattern in C. durio and C. lateralis. If the pattern is primitive, a banded pattern would have to evolve twice: once leading to the mainland SE Asia clade and once leading to a clade containing C. durio and C. lateralis. Prior to this study, molecular data were available for only two of the eight species of prehensile-tailed Cyrtodactylus. Wood et al. (2012) confirmed the suspicions of Ulber (1993) that C. elok is closely related to C. interdigitalis, and these two species are sister taxa in Clade F of their phylogeny. We obtained ND 2 and COI sequences for four of our new specimens of C. lateralis and an ND 2 sequence of the holotype of C. durio. Our maximum likelihood and Bayesian analyses recovered similar topologies with minor differences in support for each gene. Relatively few sequences of the COI gene were available on GenBank (Appendix 1), however we were able to place Cyrtodactylus elok and C. lateralis within a phylogeny containing 34 additional congeners and Gekko gekko as an outgroup. Our Maximum Likelihood tree contained few mainland species and had low support for most nodes. Nonetheless, we found no evidence of a close relationship between C. elok and C. lateralis using the COI sequences. In this small matrix of species, C. elok and C. lateralis were placed near the base of the phylogeny and had uncorrected genetic distances greater than 19 % to other species in the phylogeny (Table 2). Our ND 2 matrix included Cyrtodactylus durio, C. elok, C. interdigitalis, C. lateralis, 83 additional congeners, 26 gecko species as near outgroups, and Anolis carolinensis as a distant outgroup. We only discuss the region of the tree containing prehensile-tailed Cyrtodactylus and have collapsed several clades in figure 6 to make the phylogeny easier to read. The base of the tree, including C. angularis, C. tibetanus, outgroups, and a clade of the Myanmar species, is not shown. Overall, our ND 2 phylogenies had high bootstrap support and posterior probabilities; it showed substantial congruence with the phylogeny of Wood et al. (2012). As for the COI gene, we found no evidence of a close relationship between Cyrtodactylus lateralis and C. elok using the ND 2 gene (Fig. 6). We found that C. elok and C. interdigitalis are closely related and belong to a clade containing the C. pulchellus Group, a large radiation of 18 Australo-Papuan species, and the mainland species C. oldhami, C. tygroides, C. hontreensis, C. intermedius, and C. peguensis. This result agrees with that of Wood et al. (2012), although they found the C. pulchellus Group to be sister to the C. elok + C. interdigitalis clade, whereas we found the mainland species to be sister to this pair. Nonetheless, C. elok and C. interdigitalis have relatively long branch lengths and an uncorrected genetic distance of 22.9 % from one another (Table 2). In contrast, Cyrtodactylus lateralis belongs to a large clade of mostly Sunda Shelf and Philippine species. Cyrtodactylus lateralis is sister to C. durio and their ND 2 sequences have diverged by 11.7 %. These two species have yet to show any clear affinitiess to three clades in the crown clade of figure 6, and we show all four lineages as an unresolved polytomy. The distant relationship of C. lateralis to the C. elok + C. interdigitalis clade is also reflected in high uncorrected genetic distances between C. lateralis and C. elok (27.6 %) and C. lateralis and C. interdigitalis (26.7 %)., Published as part of Harvey, Michael B., O'Connell, Kyle A., Wostl, Elijah, Riyanto, Awal, Kurniawan, Nia, Smith, Eric N. & Grismer, L. Lee, 2016, Redescription Cyrtodactylus lateralis (Werner) (Squamata: Gekkonidae) and Phylogeny of the Prehensile-tailed Cyrtodactylus, pp. 517-540 in Zootaxa 4107 (4) on pages 527-531, DOI: 10.11646/zootaxa.4107.4.3, http://zenodo.org/record/256578, {"references":["Uetz, P. & Hosek, J. (Eds.) (2015) The Reptile Database. Available from: http: // www. reptile-database. org (accessed 1 October 2015)","Grismer, L. L., Anuar, S., Quah, E., Muin, M. A., Onn, C. K., Grismer, J. L. & Ahmad, N. (2010) A new spiny, prehensile-tailed species of Cyrtodactylus (Squamata: Gekkonidae) from Peninsular Malaysia with a preliminary hypothesis of relationships based on morphology. Zootaxa, 2625, 40 - 52.","Ulber, T. (1993) Bemerkungen uber cyrtodactyline Geckos aus Thailand nebst Beschreibungen von zwei neuen Arten (Reptilia: Gekkonidae). Mitteilungen aus dem Museum fur Naturkunde in Berlin. Zoologisches Museum und Institut fur Spezielle Zoologie, 69, 187 - 200. http: // dx. doi. org / 10.1002 / mmnz. 19930690202","Linkem, C. W., McGuire, J. A., Hayden, C. J., Mohammed, I. S., Bickford, D. P. & Brown, R. M. (2008) A new species of bent-toe gecko (Gekkonidae: Cyrtodactylus) from Sulawesi Island, eastern Indonesia. Herpetologica, 64, 224 - 234. http: // dx. doi. org / 10.1655 / 07 - 040.1","Kraus, F. (2007) A new species of Cyrtodactylus (Squamata: Gekkonidae) from western Papua New Guinea. Zootaxa, 1425 (1), 63 - 68. http: // dx. doi. org / 10.11646 / zootaxa. 1425.1.8","Oliver, P., Edgar, P., Mumpuni, Iskandar, D. T. & Lilley, R. (2009) A new species of bent-toed gecko (Cyrtodactylus: Gekkonidae) from Seram Island, Indonesia. Zootaxa, 2115, 47 - 55.","Rosler, H. & Glaw, F. (2008) A new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Malaysia including a literature survey of mensural and meristic data in the genus. Zootaxa, 1729, 8 - 22.","Wood, P. L. Jr., Heinicke, M. P., Jackman, T. R. & Bauer, A. M. (2012) Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversification. Molecular Phylogenetics and Evolution, 65, 992 - 1003. http: // dx. doi. org / 10.1016 / j. ympev. 2012.08.025"]}

Cyrtodactylus psarops sp. nov. Harvey, O���Connell, Barraza, Riyanto, Kurniawan & Smith Holotype. An adult female (MZB 9699; field number ENS 14612, Fig. 1) from Gunung Tanggamus, above Gisting, 5.424 �� S, 104.692 �� N, 1163 m, Kabupaten Tanggamus, Lampung Province, Sumatra, Indonesia, collected June 11, 2013, by M. B. Harvey, E. N. Smith, and field parties from BC, MZB, UB, and UTA. Paratypes (14). Four males (MZB 9686���9687, MZB 9700, UTA R- 61589) and one unsexed juvenile (UTA R- 61579) from forest above Ngarip, 5.280��� 5.284 �� S, 104.557��� 104.558 ��E, 1318���1419 m; four females (UTA R- 61580���61581, MZB 9690���9691) and one male (UTA R- 61590) from forest NE of Ngarip, 5.298- 5.314 �� S, 104.528 ���104.546, 950��� 1065 m; one male (MZB 9697) and two females (UTA R- 61582���61583) from Gunung Tanggamus, above Gisting, 5.422��� 5.424 �� S, 104.692��� 104.695 �� N, 1016���1163 m, Lampung, Sumatra, Indonesia; and one female (MZB 9700) from Gunung Pesagi (known locally as ���Masagi���), Remanan Jaya, Muara dua, Sumatera Selatan Province, Sumatra, Indonesia. M. B. Harvey, E. N. Smith, and field parties from BC, MZB, UB, and UTA collected the paratypes on 9���17 June 2013. Diagnosis. A medium-size species of Cyrtodactylus reaching 82 mm SVL and distinguished from all congeners by the following combination of characters: (1) body moderately robust; limbs and digits relatively long; (2) tail 1.06���1.35 times longer than body; (3) subpyramidal, weakly keeled tubercles extending from frontal region and supraorbital skin to base of tail, 23���26 irregular longitudinal rows of tubercles at midbody, 28���38 irregular transverse rows between limbs; (4) dorsal antebrachium and often brachium tuberculate; (5) tubercles on tail restricted to basal third; (6) 38���49 ventrals in a transverse row at midbody; (7) subcaudals not transversely enlarged; (8) deep, subtriangular depression in precloacal region of males; (9) 28���32 femoral and precloacal pores in males, arrayed in a continuous series (fewer pore primordia usually present in females), precloacal pores sunken into precloacal depression; (10) scale at anterior apex of pore-bearing scales greatly enlarged, sunken into precloacal depression in males, superficial in females; (11) pore-bearing series sharply transitioning to tiny granular scales of postaxial surface of thigh; (12) 18���22 lamellae under fourth toe; (13) cloacal tubercles usually single; (14) dorsal pattern of brown blotches edged first in dark brown to black then in pale gray or brown; at least some tubercles on flanks yellow to pale cream; (15) labials charcoal or dark brown with yellow or cream spots; occiput with few large blotches; long postocular stripe extending beyond arm onto flank; (16) 7���9 transverse, irregular or broken bands on body; 10���13 regular bands on tail, mostly complete ventrally. Description. Medium-size Cyrtodactylus reaching at least 82 mm SVL in females, 74 mm in males; smallest juvenile 42 mm SVL; body moderately robust, accounting for 64.9���74.3 % (n = 6) of SVL in males, 70.1���74.3 % (n = 8) of SVL in females; tail 1.06���1.35 (1.21 �� 0.10, n = 6) times longer than body and accounting for 51.6���57.4 % (54.7 �� 2.2, n = 6) of total length; head triangular in dorsal view; head 25.7���35.1 % (29.1 �� 3.2, n = 6) of SVL, 57.8���63.2 % (61.2 �� 2.2, n = 6) as wide as long, and 33.7 ���44.0% (37.7 �� 4.5, n = 6) as deep as wide in males and 27.3���29.8 % (28.7 �� 0.8, n = 8) of SVL, 54.5���62.4 % (58.4 �� 2.8, n = 8) as wide as long, and 31.7���40.6 % (36.6 �� 3.1, n = 8) as deep as wide in females. Rostral narrower than mental to as wide as mental (67.6 ���100.0%, 85.2 �� 8.7, n = 15, of width of mental), posteriorly forming border of nostril; dorsal margin of rostral deeply notched medially by shallow groove; rostral groove bifurcating slightly ventral to center of nostril; rostral bordered posteriorly by three (60 %, n = 15), four (27 %, n = 15), or five (13 %, n = 15) postrostrals; postrostral at midline projecting into posterior margin of rostral, except in UTA R- 61582 where posterior margin of rostral straight; internarial region narrow, 16.8���22.6 % (19.3 �� 1.8, n = 15) as wide as head; distance between anterior margins of orbit 30.3���38.6 % (35.3 �� 2.5, n = 15) as wide as head; shallow, key-hole shaped depression occupying most of prefrontal, frontal, and parietal areas (depression bordered laterally by slightly raised canthal, circumorbital, and supratemporal ridges and thickened skin at occiput); scales on dorsal snout granular, subpyrimidal, smooth, subequal in size, about as large as tubercles in parietal regions and distinctly smaller than granules between eyes; tubercles extending to frontal region and supraorbital skin; tubercles on dorsal surface of head subpyrimidal and smooth, much smaller than those on body, each surrounded by seven or eight granular scales; dorsal body tubercles unicarinate, oval, slightly projecting from dorsum, about five times as large as dorsal granules, 23���26 (23 �� 2, n = 14) longitudinal rows at midbody including scales in ventrolateral folds, 28���38 (33 �� 3, n = 14) irregular transverse rows between limbs; dorsal granules smooth. Nostril oval, directed postero-laterally and somewhat dorsally; bordered by rostral, postrostral, supranasal, nasal, and narrow antero-dorsal process of first supralabial; supranasal rounded to rhomboidal, about three times as large as adjacent postnasals; postnasals 5���7 (6 �� 1, n = 14); snout length 34.5���42.6 % (38.1 �� 2.1, n = 15) of head length and distance from orbit to nostril accounting for 73.4���80.3 % (77.0 �� 2.0, n = 15) of snout length; loreal region sloping; lips somewhat flared; 16���22 (20 �� 2, n = 15) loreals between nasal and anterior border of orbit; lorilabials in front of orbit flat, plate-like, enlarged relative to other loreals (1.5���3 times as large as loreals above them, but much smaller than supralabials); eye separated from supralabials by one or two longitudinal rows of tiny granules; diameter of orbit 19.0��� 25.7 % (23.3 �� 2.0, n = 15) of head length; temporal region with granular scales separating large, rounded, smooth or weakly keeled tubercles; distance from posterior border of orbit to anterior border of auditory meatus 24.1���33.5 % (29.3 �� 2.4, n = 15) of head length; auditory meatus deep, elliptical, directed postero-dorsally at about 40 �� to vertical, positioned at posterior end of head and with its ventral margin at same level as rictus; smallest diameter of auditory meatus 60���70 % smaller than greatest diameter; enlarged supralabials 9���12 (11 �� 1, n = 16), last separated by 1���5 small granular scales from rictus (12���16 total supralabials; 8���10 from center of orbit to rostral); first supralabial with antero-dorsal extension reaching nostril and separating lorilabials from rostral; remaining supralabials subrectangular with rounded dorsal edges, reducing in size to rictus. Mental triangular; infralabials 8���11 (10 �� 1, n = 16; 6���8 from center of orbit); pair of postmentals in contact medially; second pair of postmentals oval to squarish, about half as wide as first pair, separated from one another medially by 6���11 (9 �� 1, n = 15) small granular scales bordering first pair of postmentals; sublabial row decreasing in size posteriorly, not differentiated from adjacent gulars at level of eye; gulars small, smooth, granular, nonimbricate, much smaller than ventrals; in front of pectoral girdle, gulars grading into progressively larger, imbricate scales; scale of chest like ventrals; ventrals smooth, imbricate, flat, circular, about as wide as tubercles on dorsum; ventrals in transverse row at midbody 38���49 (44 �� 4, n = 13); ventrolateral skin fold distinct, containing tubercles. Tail subcylindrical, though somewhat flattened dorsally and ventrally; tubercles restricted to 16.0��� 24.9 % (19.1 �� 3.1, n = 6) of intact tails; distribution of tubercles at base of tail like dorsal body, distally tubercles not projecting and arrayed in 3���6 verticils; verticils separated by 6���9 small scales along dorsal midline; tubercles on tail not extending to ventral side, although some scales of verticils may be slightly enlarged relative to other subcaudals; subcaudals flat, homogenous, 1.5���2 times as large as scales on sides of tail; each subcaudal about as long as wide; transversely enlarged subcaudal plates absent; cloacal tubercles rounded, smooth, single, slightly smaller than dorsal tubercles; tip of tail capped by granular scales. Brachium covered in small, homogenous, weakly keeled scales; brachial tubercles usually present (67 %, n = 15); antebrachium tuberculate; antebrachial tubercles subpyrimidal, unicarinate, about four times as large as granular scales between them, slightly smaller than those on dorsum; ventral surfaces of brachium and antebrachium covered in smooth, slightly imbricate, homogenous scales; scales on palm and dorsal surface of hand smooth, imbricate; scales under finger I 8���11 (n = 15 for all digits), under finger II 13���15, under finger III 14���18, under finger IV 16���19, under finger V 13���16 (all counts excluding ungual scale); one or two scales widened and similar to lamellae extending onto palm at base of digits. Thigh, shank, and postaxial half of dorsal foot (at base of toes IV���V) tuberculate; tubercles of leg subpyrimidal, unicarinate, about same size as those on dorsum, about five times as large as small granular scales between them; preaxial and ventral surface of thigh covered in rounded, flat, imbricate scales; ventral thigh with enlarged row of pore-bearing scales and similarly enlarged row of scales anterior to pore-bearing row; this doubled row of scales arranged in an inverted V with its apex at midline of belly; abrupt transition to tiny granules on postaxial surface of thigh; postfemoral granules only one-fifth to one-sixth size of pore-bearing scales; sole covered in flat, imbricate, rounded scales; scales under toe I 8���11 (n = 15), under toe II 13���15, under toe III 16���20; under toe IV 18���22, under toe V 17���21; one or two scales widened and similar to lamellae extending onto sole at base of digits. Arrangement of pore-bearing scales sexually dimorphic; both sexes with single, greatly enlarged scale at apex of pore-bearing series (Fig. 2); apical scale twice as long as wide, with single pore or pore primordium at its posterior end; apical scale superficial in females and juveniles, sunken into precloacal depression of males; total pores 28���32 (30 �� 2, n = 6) in males; pore rows of males continuous, extending onto thighs; pore primordia present (75 %, n = 8) or absent (25 %) in females; when present, total pore primordia 13���29 (20 �� 6, n = 6); in two specimens distal primordia separated by gaps of 2 / 4 and 3 / 3 (right/left) scales on thigh; in specimens without gaps, primordia extending for short distance onto thigh (50 %, n = 6) or restricted to body (50 %). Coloration. In life (based on field notes of M.B. Harvey and photos of all specimens in the type series), Cyrtodactylus psarops has a gray, greenish gray, or greenish brown ground coloration with brown markings widely edged in black. On the flanks, tubercles edging the brown dorsal markings are frequently bright yellow, sharply contrasting with the adjacent color. Anastomosing brown and pale yellow markings cover the dorsal snout. The supraciliaries are also yellow with some black areas. The rostral, mental, narial region, and labials are mostly charcoal to black, and the labials are marked by widely spaced, distinctive yellow spots. Dorsally, several large subcircular blotches cover the parietal area. A wide postocular stripe extends beyond the arm onto the flank. A transverse nuchal band may partially fuse to the postocular stripe, although the two markings are usually separate from one another. The iris is reddish brown, edged in black and with a black reticulum. The ciliary body is only slightly lighter than the iris and indistict. In Cyrtodactylus psarops, the distal third of the tongue is black above and below, whereas the rest of the tongue is pink. Cyrtodactylus psarops (Fig. 3) has one band on the neck, 7���9 (8 �� 1, n = 15) bands on the body, and 10���13 (11 �� 1, n = 6) bands on the tail. Bands on the neck and body frequently break up into subcircular blotches. Bands on the tail are narrower than the pale interspaces at the base of the tail but longer than the interspaces distally. Distally, the caudal interspaces become increasingly more pale. Ventrally the caudal bands are continuous (i.e., the tail is ringed). Regenerated tails are entirely tan above and below. The brachium, antebrachium, thigh, and shank each bears three to four bands. Banding extends onto the hand and foot, although the digits are mostly charcoal or dark brown. Except for the tail, the venter is pale pink to purple. Palms and soles are heavily melanic, whereas the porebearing scales are distinctly paler than scales around them. Adult males have a distinctive chalky white, semilunar area anterior and lateral to the precloacal depression. This chalky area extends to the groin. Etymology. This species��� name psarops is a noun in apposition derived from the Greek words psaros, meaning speckled, and ops, meaning face. The new name alludes to the characteristic yellow speckling of the labials and snout in this species. Standard English name. Speckle-faced Bent-toed Geckos. Standard Indonesian name. Cicak Jari Lengkung Muka Bintik. Distribution and natural history. We collected most of the type specimens from humid forest, coffee plantations, and adjacent secondary growth on Gunung Tanggamus and mountains surrounding Ngarip, a small town in the southern Bukit Barisan Range, Lampung, Sumatra (Fig. 4). Cyrtodactylus psarops extends into adjacent Sumatera Selatan at Gunung Pesagi. We encountered most specimens on the forest floor at night (19:00��� 21: 10 hrs), however this species readily ascends into low vegetation. We found UTA R- 61581 about 2 m aboveground on the stem of a plant at the forest edge and MZB 9702 about 1 m aboveground on a dead tree in a coffee plantation. Cyrtodactylus psarops occurs between 950���1419 m, however, its elevational range may be broader. We did not visit sites at higher or lower elevations in the vicinity of the type locality., Published as part of Harvey, Michael B., O'Connell, Kyle A., Barraza, Gabriel, Riyanto, Awal, Kurniawan, Nia & Smith, Eric N., 2015, Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the Southern Bukit Barisan Range of Sumatra and an estimation of their phylogeny, pp. 495-516 in Zootaxa 4020 (3) on pages 498-503, DOI: 10.11646/zootaxa.4020.3.5, http://zenodo.org/record/240877

Cyrtodactylus semicinctus sp. nov. Harvey, O���Connell, Barraza, Riyanto, Kurniawan & Smith Holotype. An adult male (MZB 9703, collector���s tag ENS 14958, Fig. 5) from montane forest on a trail above Kayu Aro, leading to the top of Mt. Kerinci, Kabupaten Kerinci, Jambi Province, Sumatra, Indonesia, 1905 m, 1.74159 �� S, 101.26015 ��E, collected on 24 June 2013 by M. B. Harvey and K. A. O���Connell. Paratypes (7). Two males (MZB 9695, 9696), four females (MZB 9694, UTA R- 61576, 61577, 61588), and one unsexed juvenile (MZB 9701) collected from the same trail as the holotype, 1752���2006 m, on 21 June 2014 by M. B. Harvey, E. N. Smith, and other members of the summer 2013 field party. Diagnosis. A medium size species of Cyrtodactylus reaching 89 mm SVL and distinguished from all congeners by the following combination of characters: (1) body moderately robust; limbs and digits relatively long; (2) tail 1.05���1.19 times longer than body; (3) subpyramidal, weakly keeled tubercles extending from parietal region and supraorbital skin to base of tail, 24���27 irregular longitudinal rows of tubercles at midbody, 29���35 irregular transverse rows between limbs; (4) dorsal antebrachium tuberculate, brachium lacking tubercles; (5) tubercles on tail restricted to basal third; (6) 33���44 ventrals in a transverse row at midbody; (7) subcaudals not transversely enlarged; (8) deep, subtriangular precloacal depression in males; (9) 36���38 femoral and precloacal pores in males, arrayed in a continuous series (fewer pore primordia usually present in females), sunken into precloacal depression; (10) scale at anterior apex of pore-bearing scales greatly enlarged, sunken into precloacal depression in males, superficial in females; (11) pore-bearing series sharply transitioning to tiny granular scales of postaxial surface of thigh; (12) 19���22 lamellae under fourth toe; (13) cloacal tubercles 2 / 2 or 3 / 3 contacting one another; (14) dorsal pattern of brown blotches widely edged in black; tubercles on flanks yellow to pale cream; (15) labials mostly yellow; wide U-shaped band around occiput to loreal region; (16) 5���7 transverse, irregular or broken bands on body; 9���11 regular bands on tail, incomplete ventrally, except near distal tip of some specimens. Description. Medium-size Cyrtodactylus reaching at least 89 mm SVL in females, 75 mm in males; smallest juvenile 36 mm SVL; body moderately robust, accounting for 69.9���72.9 % (n = 3) of SVL in males, 70.1���74.3 % (n = 4) of SVL in females; tail 1.05���1.19 (1.10 �� 0.05, n = 5) times longer than body and accounting for 51.4���54.4 % (52.4 �� 1.2, n = 5) of total length; head triangular in dorsal view; head 27.1���30.1 % of SVL, 62.4���71.1 % as wide as long, and 37.6���40.3 % as deep as wide in males (n = 3) and 25.7���29.9 % of SVL, 58.7���72.3 % as wide as long, and 32.7���37.1 % as deep as wide in females (n = 4). Rostral narrower than mental (76.7���99.5 %, 89.5 �� 8.6, n = 8, of width of mental), posteriorly forming border of nostril; dorsal margin of rostral deeply notched medially by shallow groove; rostral bordered posteriorly by three (50 %, n = 8), four (38 %, n = 8), or five (12 %, n = 8) postrostrals; postrostral at midline projecting into posterior margin of rostral; internarial region narrow, 17.2���21.1 % (19.5 �� 1.4, n = 8) as wide as head; distance between anterior margins of orbit 29.2���39.6 % (34.8 �� 3.9, n = 8) as wide as head; shallow, key-hole shaped depression occupying most of prefrontal, frontal, and parietal areas (depression bordered laterally by slightly raised canthal, circumorbital, and supratemporal ridges and thickened skin at occiput); scales on dorsal snout granular, subpyrimidal, smooth, subequal in size, and slightly larger than granules of frontal and parietal regions; tubercles extending to parietal region and on supraorbital skin but absent from snout and depressed areas between eyes; tubercles on dorsal surface of head subpyrimidal and smooth, much smaller than those on body, each surrounded by seven or eight granular scales; dorsal body tubercles unicarinate, oval, slightly projecting from dorsum, about five times as large as dorsal granules, 24���27 (25 �� 1, n = 8) longitudinal rows at midbody including scales in ventrolateral folds, 29���35 (32 �� 2, n = 8) irregular transverse rows between limbs; dorsal granules smooth. Nostril oval, directed postero-laterally and somewhat dorsally; bordered by rostral, postrostral, supranasal, nasal, and narrow antero-dorsal process of first supralabial; supranasal rounded, about twice as large as adjacent postnasals; postnasals 4���6 (5 �� 0, n = 16); snout length 37.3���44.7 % (40.8 �� 2.1, n = 8) of head length and distance from orbit to nostril accounting for 68.1���77.6 % (72.7 �� 2.8, n = 8) of snout length; loreal region sloping; lips flared; 16���23 (18 �� 2, n = 8) loreals between nasal and anterior border of orbit; lorilabials in front of orbit flat, plate-like, enlarged relative to other loreals (1.5���2 times as large as loreals above them, but much smaller than supralabials); eye separated from supralabials by one or two longitudinal rows of tiny granules; diameter of orbit 20.8���26.6 % (23.4 �� 2.0, n = 8) of head length; temporal region with granular scales separating large, subpyrimidal, and keeled tubercles; distance from posterior border of orbit to anterior border of auditory meatus 27.0��� 33.3 % (29.7 �� 2.3, n = 8) of head length; auditory meatus deep, narrowly elliptical, directed postero-dorsally at about 30 �� to vertical, positioned at posterior end of head and with its ventral margin at same level as rictus; smallest diameter of auditory meatus 60���70 % smaller than greatest diameter; enlarged supralabials 8���11 (10 �� 1, n = 15), last separated by 2���4 small granular scales from rictus (10���15 total supralabials; 7���9 from center of orbit to rostral); first supralabial with antero-dorsal extension reaching nostril and separating lorilabials from rostral; remaining supralabials subrectangular with rounded dorsal edges, reducing in size to rictus. Mental triangular; infralabials 8���10 (9 �� 1, n = 15; 6���7 from center of orbit); pair of postmentals in contact medially; second pair of postmentals squarish, about half as wide as first pair, separated from one another medially by 6���8 (7 �� 1, n = 8) small granular scales bordering first pair of postmentals; sublabial row decreasing in size posteriorly; gulars small, smooth, granular, nonimbricate, much smaller than ventrals; gulars grading into progressively larger, imbricate scales just in front of pectoral girdle; scale of chest like ventrals; ventrals smooth, imbricate, flat, circular, slightly smaller than tubercles on dorsum, but much smaller than dorsal granules; ventrals in transverse row at midbody 33���44 (38 �� 5, n = 6); ventrolateral skin fold distinct, containing tubercles. Tail subcylindrical, though somewhat flattened dorsally and ventrally; tubercles restricted to 17.2���23.3 % of intact tails (n = 2); distribution of tubercles at base of tail like dorsal body, distally tubercles not projecting and arrayed in 1���5 verticils; verticiles separated by 5���7 small scales along dorsal midline; tubercles on tail not extending to ventral side, although some scales of verticils may be slightly enlarged relative to other subcaudals; subcaudals flat, homogenous, two or three times as large as scales on sides of tail; each subcaudal about as long as wide; transversely enlarged subcaudal plates absent; in specimens with regenerated tails, subcaudals just proximal to regenerated portion enlarged: about half again as long as scale row proximal to them; cloacal tubercles rounded, smooth, two (88 %, n = 16) or three (12 %, n = 16, both counts of three in one specimen: UTA R- 61577), in contact with one another, half again as large as dorsal tubercles; tip of tail capped by granular scales. Brachium covered in small, homogenous, weakly keeled scales; brachial tubercles absent; antebrachium tuberculate; antebrachial tubercles subpyrimidal, unicarinate, about four times as large as granular scales between them, slightly smaller than those on dorsum; ventral surfaces of brachium and antebrachium covered in smooth, slightly imbricate, homogenous scales; scales on palm and dorsal surface of hand granular, smooth, weakly (ventrally) to moderately (dorsal surface) imbricate; scales under finger I 8���10 (n = 8 for all digits), under finger II 12���15, under finger III 15���17, under finger IV 16���18, under finger V 12���15 (all counts excluding ungual scale); one or two scales widened and similar to lamellae extending onto palm at base of digits. Thigh, shank, and postaxial half of dorsal foot (at base of toes IV���V) tuberculate; tubercles of leg subpyrimidal, unicarinate, about same size as those on dorsum, about five times as large as small granular scales between them; preaxial and ventral surface of thigh covered in rounded, flat, imbricate scales; ventral thigh with enlarged row of pore-bearing scales and similarly enlarged row of scales anterior to pore-bearing row; this doubled row of scales arranged in an inverted V with its apex at midline of belly; abrupt transition to tiny granules on postaxial surface of thigh; postfemoral granules only one-fifth to one-sixth size of pore-bearing scales; sole covered in flat, imbricate, rounded scales; scales under toe I 8���11, under toe II 12���15, under toe III 17���19; under toe IV 19��� 22, under toe V 17���20; one or two scales widened and similar to lamellae extending onto sole at base of digits. Arrangement of pore-bearing scales sexually dimorphic; both sexes with single, greatly enlarged scale at apex of pore-bearing series; apical scale twice as long as wide, with single pore or pore primordium at its posterior end; apical scale superficial in females and juveniles, sunken into precloacal depression in males; total pores 36 and 38 in two adult males (SVL 75 mm and 68 mm, respectively), only 14 in small male (SVL 55 mm) perhaps due to ontogenetic differences; two females (SVL 51 mm and 89 mm, respectively) with 17 and 19 pore primordia, one lacking primordia (SVL 83 mm); in both sexes pores and/or primordia extending onto thigh. Coloration. In life, Cyrtodactylus semicinctus has a light gray, greenish tan, or dull orange ground coloration with brown to reddish brown markings widely edged in black (MZB 9695 was in the process of shedding and its ground coloration was gray rather than orange or greenish tan). Narrow markings are entirely black. The dorsal tubercles are a paler shade than the dorsal ground color. On the flanks, even some of the tubercles within dark markings are this pale color, so that the flanks appear to be densely speckled. A yellow wash covers the labials, rostral, and supraciliary scales. These yellowish scales are mostly immaculate, although small, irregular black blotches mark the narial and internarial region and sometimes extend ventrally onto the rostral and or first supralabial. Dorsally, the head usually has one to three poorly defined blotches in the parietal region and a sharp black line edging the gray to yellow supraciliaries medially. This species has a distinctive U-shaped marking extending from the loreal regions around the occiput. The marking begins a short distance behind the nasal just above the enlarged series of lorilabials and widens just in front of the eye. Behind the eye and around the occiput, a slightly more narrow pale band edges the U-shaped band dorsally and accentuates it. The iris is reddish brown to greenish bronze, edged in black and with a black reticulum. The ciliary body is only slightly lighter than the iris and indistict. Cyrtodactylus semicinctus (Fig. 6) has one band on the neck, 5���7 (6 �� 1, n = 8) bands on the body, and 9���11 (10 �� 1, n = 5) bands on the tail. Bands on the neck and body are jagged, irregular, and frequently broken on the flanks. When broken, blotches on the flanks may fuse with one another, sometimes to produce long sinuous markings. In particular, the band on the neck usually breaks up into blotches. However, the band on the neck does not fuse to the U-shaped nuchal band. Bands on the tail are slightly longer than the pale interspaces. Except near the distal tip of the tail of some specimens, the bands are not continuous ventrally (i.e., the tail is banded, not ringed; all specimens have at least the basal one-third of the tail not regenerated and none have ringed tails). Distally, the caudal interspaces become increasingly more pale. Regenerated tails are entirely black or mostly black with few scattered cream scales. The brachium, antebrachium, thigh, and shank each bears two or three jagged, irregular bands. Except for the tail, the venter is mostly rose pink to grayish purple. Palms and soles are a darker shade of the same ground color, whereas the pore-bearing series are distinctly paler than scales around them. Some scales at the edges of the venter are the same color as the dorsal ground coloration. They form a diffuse reticulum against the rose pink to grayish purple ground color. This diffuse, greenish tan to orange reticulum is restricted to the lateral edges of the belly and pre- and postaxial margins of the ventral limbs, but extends across the entire gular region. The cloacal tubercles are greenish tan or orange. Ventrally, intact tails are tessellated with charcoal and elements of the dorsal ground coloration. Etymology. This species��� name semicinctus is a masculine adjective derived from the Latin noun cinctum, meaning girdle or belt, and prefix semi, meaning partial or incomplete. The name alludes to the incomplete caudal rings of this species. Standard English name. Kerinci Bent-toed Geckos. Standard Indonesian name. Cicak Jari Lengkung Kerinci. Distribution and natural history. This species is known only from the type locality. We collected the entire type series between 1752���2006 m along a trail above the tiny hamlet of Kayu Aro. The trail extends through apparently pristine forest to the top of Mt. Kerinci and is frequented by hikers. Field parties collected Cyrtodactylus semicinctus on the forest floor at night (1950���2340 hrs) and we found most specimens around tree trunks. In field notes, Harvey noted that UTA R- 61576 emitted a faint ���baaap��� when first grasped and a louder ���raaank��� when struggling as the senior author collected color notes., Published as part of Harvey, Michael B., O'Connell, Kyle A., Barraza, Gabriel, Riyanto, Awal, Kurniawan, Nia & Smith, Eric N., 2015, Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the Southern Bukit Barisan Range of Sumatra and an estimation of their phylogeny, pp. 495-516 in Zootaxa 4020 (3) on pages 503-507, DOI: 10.11646/zootaxa.4020.3.5, http://zenodo.org/record/240877

Harvey, Michael B., O'Connell, Kyle A., Barraza, Gabriel, Riyanto, Awal, Kurniawan, Nia, Smith, Eric N. (2015): Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the Southern Bukit Barisan Range of Sumatra and an estimation of their phylogeny. Zootaxa 4020 (3): 495-516, DOI: http://dx.doi.org/10.11646/zootaxa.4020.3.5

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). 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 2.14–2.41 X (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. 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. 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, U