A new species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Manipur State, northeast India, with a critical review highlighting extensive errors in literature covering bent-toed geckos of the Indo-Burma region

Figures & data

Table 1. List of Cyrtodactylus ND2 sequences used in this study (newly generated sequence in bold)

Species	Museum No.	Locality	GenBank Accession No.
C. annandalei Bauer, 2003	CAS 215722	Myanmar, Sagaing Div., Alaungdaw Kathapa N.P.	JX440524.1
C. arunachalensis Mirza, Bhosale, Ansari, Phansalkar, Sawant, Gowande & Patel, 2021	CES13/1465	India, Arunachal Pradesh State, Changlang Dist., Glow Lake.	KM255193.1
C. aunglini Grismer, Wood, Thura, Win, Grismer, Trueblood & Quah, 2018c	LSUHC 13948	Myanmar, Mandalay Reg., Pyin Oo Lwin Dist., Kyauk Nagar Cave.	MH764589.1
^aC. ayeyarwadyensis Bauer, 2003	CAS 212459	Myanmar, Ayeyarwady Div., vicinity of Mwe Hauk Village.	JX440526.1
C. ayeyarwadyensis Bauer, 2003	CAS 216446	Myanmar, Rakhine State, vicinity of Kantheya Beach.	GU550715.1
C. battalensis Khan, 1993	PMNH 2301	Pakistan, North-West Frontier Prov., Battagram City.	KC151983.1
C. bhupathyi Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018b	BNHS 2255	India, West Bengal State, Kalimpong Dist., near Lower Mongpong.	KM255204.1
C. brevidactylus Bauer, 2002	CAS 214105	Myanmar, Mandalay Reg., Popa Mountain Park.	GU550714.1
C. chamba Agarwal, Khandekar & Bauer, 2018c	CES11/1291	India, Himachal Pradesh State, Chamba Dist., near Chamba.	KM255191.1
C. chanhomeae Bauer, Sumontha & Pauwels, 2003	CUMZ 2003.62	Thailand, Saraburi Prov., Phra Phutthabat Dist., Thep Nimit Cave.	JX440529.1
C. chrysopylos Bauer, 2003	LSUHC 13937	Myanmar, Shan State, Taunggyi Dist., Ywnagan Township.	MH764604.1
^bC. chrysopylos Bauer, 2003	LSUHC 13126	Myanmar, Shan State, Taunggyi Dist., Ywnagan Township.	MH764598.1
C. fasciolatus (Blyth, 1861)	CES11/1337	India, Himachal Pradesh State, Shimla Dist., near Subathu.	KM255184.1
C. cf. fasciolatus 1	CES11/1345	India, Uttarakhand State, Almora Dist., Almora.	KM255169.1
C. cf. fasciolatus 2	CES09/1196	India, Uttarakhand State, Dehradun Dist., Mussoorie-Kempty Road.	KM255172.1
C. gansi Bauer, 2003	CAS 222412	Myanmar, Chin State, Min Dat Township.	JX440537.1
C. gubernatoris (Annandale, 1913)	BNHS 2207	India, Sikkim State, East Sikkim Dist., Singtam Town.	KM255181.1
C. guwahatiensis Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018c	BNHS 2146	India, Assam State, Kamrup Metropolitan Dist., Guwahati.	KM255194.1
C. himalayanus Duda & Sahi, 1978	CES11/1317	India, Jammu and Kashmir State, Kishtwar Dist., near Kishtwar.	KM255187.1
C. jaintiaensis Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018c	BNHS 2248	India, Meghalaya State, Jaintia Hills Dist., near Jowai.	KM255195.1
C. kazirangaensis Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018c	BNHS 2147	India, Assam State, Golaghat Dist., Kohora.	KM255170.1
C. khasiensis (Jerdon, 1870)	BNHS 2249	India, Meghalaya State, East Khasi Hills Dist., Cherrapunjee Resort.	KM255188.1
C. lawderanus (Stoliczka, 1871)	CES11/1343	India, Uttarakhand State, Almora Dist., Almora.	KM255189.1
C. meersi Grismer, Wood, Quah, Murdoch, Grismer, Herr, Espinoza, Brown & Lin, 2018b	LSUHC 13455	Myanmar, Bago Reg., Yangon (north) Dist., Taikkyi Township.	MH624104.1
C. mombergi Grismer, Wood, Quah, Thura, Herr & Lin, 2019b	LSUHC 14734	Myanmar, Kachin State, Mohnyin Township, Indawgyi W.S.	MN059875.1
C. cf. mombergi	LSUHC 14591	Myanmar, Kachin State, Mohnyin Township, Indawgyi W.S.	MN059869.1
C. montanus Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018c	BNHS 2231	India, Tripura State, North Tripura Dist., Phuldungsei Village.	KM255200.1
C. montanus Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018c	MZMU 1630	India, Mizoram State, Dampa Tiger Reserve, Pathlawi Lunglen Tlang.	MT250544.1
^cC. myaleiktaung Grismer, Wood, Thura, Win, Grismer, Trueblood & Quah, 2018c	LSUHC 13965	Myanmar, Mandalay Reg., Mandalay Dist., Mya Leik Taung.	Grismer pers. comm., 17/10/2020
C. myintkyawthurai Grismer, Wood, Quah, Murdoch, Grismer, Herr, Espinoza, Brown & Lin, 2018b	CAS 245200	Myanmar, Bago Reg., Central Bago Yoma.	MH624107.1
C. myintkyawthurai (erroneously as C. feae on GenBank)	USNM 559805	Myanmar, Mandalay Reg., Popa Mountain Park.	JX440536.1
C. nagalandensis Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018c	BNHS 2253	India, Nagaland State, Kohima Dist., Khonoma.	KM255199.1
C. namtiram sp. nov.	BNHS 2751	India, Manipur State, Tamenglong Dist., Namtiram.	MW367433.1
C. novaeguineae (Schlegel, 1837)	BPBM 23316	Papua New Guinea, West Sepik Prov., Parkop, Toricelli Mts.	JX440547.1
^dC. nyinyikyawi Grismer, Wood, Thura, Win & Quah, 2019a	CAS 226139	Myanmar, Magwe Reg., Min Bu Township, Shwe Settaw W.S.	MH624118.1
C. peguensis (Boulenger, 1893)	LSUHC 13454	Myanmar, Bago Reg., Myin Mo Shwe Taung Pagoda.	MH756190.1
C. philippinicus (Steindachner, 1867)	FMNH 236073	Philippines, Romblon Island.	JX440550.1
C. pyadalinensis Grismer, Wood, Thura, Win & Quah, 2019a	LSUHC 13932	Myanmar, Shan State, Ywangan Township, Panluang-Pyadalin Cave W.S.	MK488057.1
C. pyinyaungensis Grismer, Wood, Thura, Zin, Quah, Murdoch, Grismer, Lin, Kyaw & Lwin, 2018a	BYU 52234	Myanmar, Mandalay Reg., 5.3 km north of Pyinyaung Village at the Apache Cement factory mining site.	MF872307.1
C. russelli Bauer, 2003	CAS 226137	Myanmar, Sagaing Div., Htamanthi W.S.	JX440555.1
C. septentrionalis Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018c	BNHS 1989	India, Assam State, Bongaigaon Dist., near Abhayapuri.	MH971164.1
C. slowinskii Bauer, 2002	CAS 210205	Myanmar, Sagaing Div., Alaungdaw Kathapa N.P.	JX440559.1
C. tibetanus (Boulenger, 1905)	MVZ 233251	China, Tibet Autonomous Reg., Lhasa, 3 km from of Potala Palace.	JX440561.1
C. tripuraensis Agarwal, Mahony, Giri, Chaitanya & Bauer, 2018b	BNHS 2238	India, Tripura State, Sepahijala Dist., Sepahijala W.S.	KM255183.1
C. cf. tripuraensis	BNHS 2230	India, Tripura State, North Tripura Dist., Rowa W.S.	KM255201.1
C. urbanus Purkayastha, Das, Bohra, Bauer & Agarwal, 2020	VR/ERS/ZSI/688	India, Assam State, Kamrup Metropolitan Dist., Guwahati, Basishta Temple.	MN911174.1
C. sp. Changlang1	CES11/1349	India, Arunachal Pradesh State, Changlang Dist., Miao.	KM255179.1
C. sp. Changlang2	CES13/1459	India, Arunachal Pradesh State, Changlang Dist., Miao.	KM255192.1
C. sp. Khellong	CES13/1464	India, Arunachal Pradesh State, West Kameng Dist., Khellong.	KM255196.1
C. sp. Mizoram	CES13/1455	India, Mizoram State, near Aizawl Town.	KM255197.1

Museum abbreviations are defined and original citations for sequences given in the ‘Materials and methods’ section. Dist. District; Subdist. Subdistrict; Prov. Province; Div. Division; Reg. Region; N.P. National Park; W.S. Wildlife Sanctuary; Mts. Mountains.

^aSpecimen number erroneously as ‘CAS 216459’ and locality erroneously as ‘Myanmar, Rakhine State, Than Dawe Dist., Gwa Township’ in original publication, see point 22 in ‘Errors in literature’ section.

^bSee point 19 in ‘Errors in literature’ section.

^cSee point 19 in ‘Errors in literature’ section.

^dThe specimen number for the sequence MH624118.1 was erroneously published as LSUHC 226139 in Grismer et al. (2018d: table 1), but correctly given as ‘CAS 226139’ elsewhere in that paper, in Grismer et al. (2019a) and on the GenBank record. The error was copied in some subsequent papers (e.g. Purkayastha et al. 2020: table 1).

Figure 1. Topographic map of northeast India and surrounding countries showing the type locality (red star) of Cyrtodactylus namtiram sp. nov. and all known localities for the other members of the gansi group from the following literature: Agarwal et al. (2018b); Bauer (2002, 2003); Grismer et al. (2018c); Muansanga et al. (2020). Circle with an x, type localities; without an x, other published localities: C. aunglini (blue), C. brevidactylus (brown), C. chrysopylos (grey), C. gansi (turquoise), C. jaintiaensis (light green), C. montanus (pink), C. nagalandensis (dark green), C. myaleiktaung (orange), Mizoram sp. (black). Country borders black; state borders brown; major rivers blue. Inset: Map of northeast India showing country borders in black, and green box representing the area depicted in the main map; abbreviations: In India, Np Nepal, Bd Bangladesh, Bt Bhutan, Cn China, Mm Myanmar, Tl Thailand, L Laos.

Figure 2. Maximum Likelihood phylogeny of the Indo-Burma radiation of Cyrtodactylus based on the complete ND2, tRNA-Trp, tRNA-Ala genes and partial tRNA-Asn gene alignment. Coloured portion of the tree denotes the Indo-Burma radiation, with species clades and species groups coloured as follows: outgroups (black); fasciolatus clade (light green); slowinskii clade (brown); peguensis clade comprising the gubernatoris group (light blue) and peguensis group (red); khasiensis clade comprising the arunachalensis group (pink), mombergi group (orange), khasiensis group (dark blue), and gansi group (dark green). Bootstrap support values ≥95 represented by a black spot, values <95 are given next to nodes.

Figure 3. Cyrtodactylus namtiram sp. nov. adult male holotype (BNHS 2751) in preservation: (A). dorsal view; (B). ventral view; (C). details of dorsal pholidosis; (D). dorsal view of head; (E). ventral view of head; (F). lateral view of head. (A and B) to scale; (D–F) to scale. Scale bars = 10 mm.

Figure 4. Cyrtodactylus namtiram sp. nov. adult male holotype (BNHS 2751) in life from Tamenglong District, Nagaland State, northeast India: (A). dorsolateral view; (B). close-up of head, left side; (C). ventral view while under anaesthesia, scale bar = 10 mm. Photos taken ex-situ. (D). habitat at the type locality.

Figure 5. Cyrtodactylus montanus in life showing variation of dorsal markings and colouration. Photos taken in-situ of six different uncollected adult individuals from the Jampui Hills, North District, Tripura State, northeast India: (A). adult female from Phuldungsei Village, nearby the type locality of C. montanus; (B–f). from the vicinity of Vanghmun Village: (B and C). adult females; (D–F). adult males.

Table

Species	Locality	Latitude (N)	Longitude (E)	Reference
C. aunglini	‘Kyauk Nagar Cave, 11 km southwest of Pyin Oo Lwin, Pyin Oo Lwin Township, Pyin Oo Lwin District, Mandalay Region, Myanmar (20.93087°N, 95.22580°E [error]; 715 m)’. GPS coordinates for the type locality estimated from Google Maps. Refer to point no. 3 in ‘Errors in literature’ section for C. aunglini type locality.	21.978445	96.37001	Grismer et al. (2018c); Google Maps
C. brevidactylus	‘Popa Mountain Park, Kyauk Pan Tawn Township, Mandalay Division, Myanmar (20°57’56.8”N; 95°14’18.8”E)’	20.96578	95.23856	Bauer (2002)
C. chrysopylos	‘Panlaung-Pyadalin Cave Wildlife Sanctuary, Ywa Ngan Township, Shan State, Myanmar (21°07’58.4”N, 96°20’25.0”E, 319 m.)’. Note: original coordinates erroneously given as ‘(21°0’58.4”N, 96°20’25.0”E)’ by Grismer et al. (2018c) who also did not give the type locality in its original form despite implying so with the use of quotation marks.	21.13289	96.340	Bauer (2003)
*C. chrysopylos	‘5.1 km south of Pyadalin Cave, Panlaung-Pyadalin Cave Wildlife Sanctuary, Ywnagan Township, Taunggyi District, Shan State, Myanmar in Nwalabo Mountains (21.02783°N, 96.41283°E; 951 m in elevation)’.	21.02783	96.41283	Grismer et al. (2018c)
C. gansi	‘Che Stream, Min Dat Township, Min Dat District, Chin State [Myanmar] (21°21’15.5”N, 93° 56’13.3”E, 780 m.)’	21.35431	93.93703	Bauer (2003)
C. jaintiaensis	‘vicinity of Tyrshi Falls near Jowai town, West Jaintia Hills district, Meghalaya state, India (25.46956°N, 92.18313°E, 1350 m asl.)’	25.46956	92.18313	Agarwal et al. (2018b)
C. montanus	‘near Phuldungsei village in the southern Jampui Hills, Kanchanpur subdivision, North Tripura district, Tripura state, India (23.82257°N, 92.26034°E, 850 m asl.)’	23.82257	92.26034	Agarwal et al. (2018b)
*C. montanus	‘Dampa Tiger Reserve …. alongside a trail up a hill called Pathlawi Lunglen Tlang (23.7051°N, 92.4075°E; elev. 553 m asl)’, Mizoram State, India	23.7051	92.4075	Muansanga et al. (2020)
*C. montanus	Below Eden Lodge, Vanghmun Village, Kanchanpur Subdivision, North Tripura District, Tripura State, India.	24.00758	92.27738	Present study
C. myaleiktaung	‘Mya Leik Taung, 25.3 km southeast of Mandalay, Aungmyethazan Township, Mandalay District, Mandalay Region, Myanmar (21.77722 N, 96.25138 E; 250 m)’	21.77722	96.25138	Grismer et al. (2018c)
C. nagalandensis	‘Near Khonoma village, Kohima district, Nagaland state, India (25.65818°N, 94.02142°E, 1450 m asl.)’	25.65818	94.02142	Agarwal et al. (2018b)
C. namtiram sp. nov.	Vicinity of a waterfall (…, 770 m asl.), Namtiram Village, Tamenglong West Subdivision, Tousem Tribal Development Block, Tamenglong District, Manipur State, India.	25.046381	93.429483	Present study
Mizoram sp.	‘Near Aizawl town, Mizoram, India’. Estimated coordinates for Aizawl from Google Maps.	23.7307	92.7173	Agarwal et al. (2014); Google Maps

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