A ‘li le d agon’ om Kunming Ci y: a new g een pi ipe om
Yunnan P o ince, China (Squama a, Vipe idae, T ime esu us)
Yuhao Xu1, Jundong Deng1, Tie ui Zhang2, Tan Van Nguyen3,4, Shiyang Weng5, Nikolay A. Poya ko 6,
Ge no Vogel7, Fanyue Sun1, Chencan Liao8,9, Li ang Peng1
1 S a e Key Labo a o y o Pla eau Ecology and Ag icul u e, Qinghai Uni e si y, Xining 810016, Qinghai, China
2 The Anhui P o incial Key Labo a o y o Biodi e si y Conse a ion and Ecological Secu i y in he Yang ze Ri e Basin, College o Li e
Sciences, Anhui No mal Uni e si y, Wuhu 241000, Anhui, China
3 The School o Medicine & Pha macy, Duy Tan Uni e si y, Da Nang 550000, Vie nam
4 Cen e o En omology & Pa asi ology Resea ch, Duy Tan Uni e si y, Da Nang 550000, Vie nam
5 Ins i u e o Pla eau Biology o Xizang Au onomous Region, Lhasa 850008, Xizang, China
6 Depa men o Ve eb a e Zoology, Lomonoso Moscow S a e Uni e si y, Leninskiye Go y, GSP–1, Moscow 119234, Russia
7 Socie y o Sou heas Asian He pe ology, D-69115 Heidelbe g, Baden-Wü embe g, Ge many
8 Ailaoshan Sub opical Fo es Ecosys em Obse a ion and Resea ch S a ion o Yunnan P o ince, Pue 676200, Yunnan, China
9 Xi Shuang Ban Na T opical Bo anical Ga den, Chinese Academy o Sciences, Dai Au onomous P e ec u e o Xishuangbanna 666303, Yunnan, China
h ps://zoobank.o g/E3A021AB-E3A6-434C-89B6-5C5E163B34C4
Co esponding au ho s: Li ang Peng ([email p o ec ed]); Tan Van Nguyen ([email p o ec ed])
Academic edi o : Jus in Be ns ein ♦
Recei ed
23 Oc obe 2025 ♦
Accep ed
7 No embe 2025 ♦
Published
18 No embe 2025
Abs ac
A new species o he genus T ime esu us Lacépède is desc ibed om Kunming Ci y, Yunnan P o ince, China, based on an in e-
g a i e analysis o mo phological and molecula da a. The new species, T ime esu us loong sp. no ., is assigned o he subgenus
Vi ido ipe a Malho a & Tho pe and can be dis inguished om i s congene s by he ollowing combina ion o cha ac e s: he i s
sup alabial comple ely sepa a ed om he nasal scale; a sho and spinose hemipenis; mode a e adul body size; do sal scales in 19
(21 o 23)-19-15 ows; 150–151 en al scales in males, and 157–158 in emales; 67–68 subcaudals in males, and 57–64 in emales;
and dis inc body colo pa e ns. Molecula analyses based on he mi ochond ial 16S, cy b, and ND4 gene agmen s indica e ha
T ime esu us loong sp. no . is gene ically di e gen om all cu en ly ecognized congene s, showing unco ec ed p-dis ances
anging om 6.7–11.2% in cy b and 6.1–8.3% in ND4. The disco e y o his species b ings he o al numbe o ecognized T ime e-
su us species o 57, including nine eco ded om Yunnan P o ince, u he emphasizing he complex opog aphy and excep ional
le el o endemism.
Key Wo ds
Hemipenis, molecula phylogeny, mo phology, axonomy, T ime esu us loong sp. no ., Sou hwes e n China, sys ema ics
In oduc ion
The genus T ime esu us Lacépède, commonly known as
Asian g een pi ipe s, ep esen s one o he mos spe-
cies- ich and axonomically challenging g oups wi hin
he sub amily C o alinae Oppel. These enomous snakes
a e o conside able medical impo ance and a e widely
dis ibu ed ac oss opical and sub opical Asia, om he
Himalayas and sou he n China o mainland and insula
Sou heas Asia, ex ending as a as Timo and he Less-
e Sunda Islands (Gump ech e al. 2004; Malho a and
Tho pe 2004a; Da id e al. 2011, 2023; Poya ko e al.
2023; Mi za e al. 2023). Species o T ime esu us ex-
hibi ema kable ecological and mo phological di e si y,
Zoosys . E ol. 101 (4) 2025, 2267–2293|DOI 10.3897/zse.101.175879
Copy igh Xu, Y. e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed use, dis ibu-
ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2268
which, oge he wi h sexual dimo phism and geog aphic
a ia ion, ha e long complica ed hei axonomy. Ex en-
si e e isions du ing he pas wo decades ha e esul ed
in he ecogni ion o mul iple lineages and nume ous new
species, pa icula ly om Indochina and sou he n Chi-
na (Vogel e al. 2023; Idiia ullina e al. 2023, 2024a–c;
Pawangkhanan e al. 2025; Xu e al. 2025). A p esen ,
57 alid species a e ecognized in he genus (Mi za e al.
2023; Idiia ullina e al. 2023, 2024a–c; Pawangkhanan
e al. 2025; Ue z e al. 2025). Despi e nume ous axo-
nomic e isions in ecen yea s, species di e si y wi hin
T ime esu us emains incomple ely esol ed. High mo -
phological simila i y among species, coupled wi h p o-
nounced sexual dimo phism and geog aphic a ia ion,
o en obscu es diagnos ic di e ences and conceals c yp-
ic lineages (e.g., Malho a and Tho pe 2004a,b; Ra hee
e al. 2022; Nguyen e al. 2024; Idiia ullina e al. 2023,
2024a–c; Liang e al. 2025; Xu e al. 2025).
Acco ding o he classi ica ion p oposed by Mi za
e al. (2023), T ime esu us is cu en ly di ided in o six
subgene a: T ime esu us sensu s ic o, Pa ias G ay,
Popeia Malho a & Tho pe, Himalayophis Malho a
& Tho pe, Sino ipe a Guo & Wang, and Vi ido ipe a
Malho a & Tho pe (see also Idiia ullina e al. 2024b;
Pawangkhanan e al. 2025; Xu e al. 2025). Membe s
o Vi ido ipe a a e de ined by wo key mo phological
ea u es: he i s sup alabial scale is comple ely sepa-
a ed om he nasal scale, and he hemipenis is sho
and spinose (Malho a and Tho pe 2004a; Dawson e
al. 2008; Mi za e al. 2023). Cu en ly, eigh species a e
assigned o his subgenus, including T. mayaae Ra hee,
Pu kayas ha, Lal emsanga, Dalal, Biakzuala, Muansan-
ga & Mi za [ ype locali y: Champhai Dis ic , Mizo am
S a e, India], T. medoensis Zhao [ ype locali y: Mo uo
(Medog) Coun y, Xizang Au onomous Region, China],
T. nujiang Liang, Ding, Vogel, Chen & Wu [ ype lo-
cali y: Cikai Town, Gongshan Coun y, Nujiang Lisu
Au onomous P e ec u e, Yunnan P o ince, China], T.
p e iosus Xu, Nguyen, Wang, Zhang, Poya ko , Wei,
Vogel, Peng & Weng in Xu, Nguyen, Wang, Zhang, Po-
ya ko , Wei, Vogel, Li, Deng, Sun, Peng & Weng [ ype
locali y: Xiayadong Township, Yadong Coun y, Xigaze
Ci y, Xizang Au onomous Region, China], T. s ejnege i
Schmid [ ype locali y: Shaowu Ci y, Nanping P e ec-
u e, Fujian P o ince, China], T. uongsonensis O lo ,
Ryabo , Bui & Ho [ ype locali y: Phong Nha-Ke Bang
Na ional Pa k, Quang Binh P o ince (now Quang T i
P o ince), Vie nam], T. ogeli Da id, Vidal & Pau-
wels [ ype locali y: Khao Yai Na ional Pa k, Nakhon
Ra chasima P o ince, Thailand], and T. yunnanensis
Schmid [ ype locali y: Tengchong Ci y, Baoshan Ci y,
Yunnan P o ince, China] (see Ra hee e al. 2022; Liang
e al. 2025; Xu e al. 2025).
Yunnan P o ince, si ua ed a he junc ion be ween he
Palea c ic and O ien al Realms, is ecognized as one o
China’s majo biodi e si y ho spo s. I ha bo s a pa icu-
la ly ich pi ipe auna, including se e al na owly dis-
ibu ed T ime esu us species. Cu en ly, eigh species o
T ime esu us a e eco ded om Yunnan P o ince: h ee
o he subgenus T ime esu us (T. albolab is G ay, T. guoi
Chen, Shi, Vogel & Ding in Chen, Shi, Gao, Vogel, Song,
Ding & Dai, and T. caudo na us Chen, Ding, Vogel &
Shi in Chen, Yu, Vogel, Shi, Song, Tang, Yang, Ding &
Chen); wo o Popeia (T. popeio um Smi h and T. lan-
na Idiia ullina, Nguyen, Pawangkhanan , Suwannapoom,
Chanhome, Mi za, Da id, Vogel & Poya ko ); and h ee
o Vi ido ipe a (T. s ejnege i Schmid , T. yunnanensis
Schmid , and T. nujiang Liang, Ding, Vogel, Chen &
Wu) (e.g., Idiia ullina e al. 2024b; Nguyen e al. 2024;
Liang e al. 2025). Al hough Yunnan has been he ocus
o nume ous he pe ological s udies, ecen su eys ha e
e ealed ha many mon ane alleys and o es ed go g-
es in sou hwes e n China emain insu icien ly explo ed.
Du ing ieldwo k conduc ed in 2025 in Qinglong Town,
Anning Ci y, Kunming Ci y, Yunnan P o ince, China, we
disco e ed a popula ion o g een pi ipe s ha could no
be assigned o any known species. Mo phological exam-
ina ion and mi ochond ial DNA analyses (16S, cy b, and
ND4 genes) con i med ha his popula ion ep esen s an
undesc ibed species o T ime esu us. We he e o e de-
sc ibe i he ein as a new species.
Ma e ial and me hods
Sampling and specimen p ese a ion
Fieldwo k was conduc ed in o es ed a eas o Anning
Ci y, Kuming Ci y, Yunnan P o ince, China in May and
Sep embe 2025 (Fig. 1). Geog aphic coo dina es and
ele a ion we e eco ded using he TwoS ep Ou doo
Assis an 7.9.13 (Shenzhen 2bulu In o ma ion Tech-
nology Co., L d., China). Specimens we e loca ed using
snake hooks, pho og aphed in li e, and humanely eu h-
anized wi h a bu e ed MS-222 ( icaine me hanesul-
ona e) solu ion. Subsequen ly, specimens we e ixed in
10% o malin and ans e ed o 75% e hanol o long-
e m p ese a ion. Li e issue samples we e collec ed
esh, p ese ed in 95% e hanol, and s o ed a –20 °C
o molecula analyses. All ouche specimens we e
deposi ed in he he pe ological collec ion o Qinghai
Uni e si y, Qinghai P o ince, China (QHU). All p oce-
du es ollowed he egula ions o he Wildli e P o ec ion
Law o China and we e app o ed by he Ins i u ional
E hics Commi ee o Qinghai Uni e si y (P o ocol No.
PJ202501-89).
DNA ex ac ion, ampli ica ion, and sequencing
To al genomic DNA was ex ac ed om e hanol-p ese ed
li e issues using he QIAamp DNA Mini Ki (QIA-
GEN, Changsheng Bio echnology Co. L d., Changchun,
China). Th ee mi ochond ial DNA (m DNA) agmen s,
including 16S ibosomal RNA (16S), cy och ome b (cy
b), and NADH dehyd ogenase subuni 4 (ND4), we e am-
Zoosys . E ol. 101 (4) 2025, 2267–2293
zse.penso .ne
2269
pli ied using polyme ase chain eac ion (PCR). The am-
pli ica ion o 16S was conduc ed wi h p ime s 16S1LM
(5′-CCGACTGTTGACCAAAAACAT-3′) and 16SH1
(5′-TCCGGTCTGAACTCAGATCACGTAGG-3′), ol-
lowing he p o ocol o Nguyen e al. (2020). Fo cy
b, we used p ime s L14910 (5′-GACCTGTGATMT-
GAAAACCAYCGTTGT-3′) and H16064 (5′-CTTTG-
GTTTACAAGAACAATGCTTTA-3′), as desc ibed by
Bu b ink e al. (2000). The ND4 agmen was ampli ied
using T im-ND4F (5′-CACCTATGACTACCAAAAGCT-
CATGTAGAGC-3′) and T im-ND4LEUR (5′-CAT-
TACTTTTACTTGGATTTGCACCA-3′), ollowing Sal i
e al. (2013). The PCR he mal cycling p o ile was as ol-
lows: ini ial dena u a ion a 95 °C o 3 min; ollowed by
36 cycles o dena u a ion a 95 °C o 30 s, annealing a
48 °C o 45 s, and ex ension a 72 °C o 1 min; wi h a
inal ex ension a 72 °C o 10 min; and a inal hold a 4 °C.
PCR p oduc s we e sequenced by Shanghai Map Bio ech
Co., L d. (Shanghai, China). Raw sequences we e assem-
bled using SeqMan (DNASTAR; Bu land 2000), and new-
ly gene a ed sequences we e submi ed o DDBJ (Table 1).
Phylogene ic analysis
Fo phylogene ic analysis, a o al o 235 sequences we e
included (Table 1), o which 21 we e newly gene a ed in
his s udy, while 214 we e e ie ed om wo public da-
abases: GenBank and GenBase (Na ional Genomics Da a
Cen e (NGDC), Beijing Ins i u e o Genomics, Chinese
Academy o Sciences / China Na ional Cen e o Bioin-
o ma ion; Bu e al. 2024; CNCB-NGDC Membe s and
Pa ne s 2025). These included ep esen a i es om 34
species o T ime esu us and wo ou g oup axa: C aspe-
docephalus puniceus (Boie) and Pel opelo malaba icus
(Je don). Sequences we e aligned in MEGA X (Kuma e
al. 2018). The Maximum Likelihood (ML) analysis was
conduc ed in IQ-TREE 1.6.12 (Nguyen e al. 2015) using
he bes - i model GTR + F + I + G4 o all h ee agmen s
(16S, cy b, and ND4), as de e mined by ModelFinde
o IQ-T ee in PhyloSui e 1.2.3 acco ding o Akaike in-
o ma ion c i e ion (AIC) (Akaike 1973; Akaike 1974;
Kalyaanamoo hy e al. 2017; Zhang e al. 2020). Node
suppo was assessed using bo h he Ul a as Boo s ap
Figu e 1. Map showing he ype locali ies o he new species and o he T ime esu us species o he subgenus Vi ido ipe a. Red
s a : T ime esu us loong sp. no . om Anning, Kunming, Yunnan, China; b own ci cle: T. c . loong om Huili, Sichuan, China;
yellow ci cle: T. mayaae om Champhai, Mizo am, India; blue ci cle: T. medoensis om Mo uo (Medog), Xizang, China; pink
ci cle: T. nujiang om Gongshan, Yunnan, China; g een ci cle: T. p e iosus om Xiayadong, Yadong, Xizang, China; pu ple
ci cle: T. s ejnege i om Shaowu, Fujian, China; o ange ci cle: T ime esu us sp. om Chengdu, Sichuan, China; black ci cle: T.
uongsonensis om Phong Nha-Ke Bang NP, Quang T i, Vie nam; whi e ci cle: T. ogeli om Khao Yai NP, Nakhon Ra chasima,
Thailand; and g ey ci cle: T. yunnanensis om Tengchong, Yunnan, China.
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2270
Table 1. NCBI / DDBJ / NGDC accession numbe s, locali ies, and ouche in o ma ion o all specimens used in his s udy. * =
DDBJ Accession Numbe , and # = NGDC Accession Numbe .
NO. Species name Locali y Vouche NO. 16S cy b ND4 Re e ences
Genus T ime esu us
Subgenus Vi ido ipe a
1T ime esu us loong
sp. no .
Anning, Kunming,
Yunnan, China
QHU R2025027 LC899328*LC899334*LC899341* This s udy
2T ime esu us loong
sp. no .
Anning, Kunming,
Yunnan, China
QHU R2025028 LC899329*LC899335*LC899342* This s udy
3T ime esu us loong
sp. no .
Anning, Kunming,
Yunnan, China
QHU R2025029 LC899330*LC899336*LC899343* This s udy
4T ime esu us loong
sp. no .
Anning, Kunming,
Yunnan, China
QHU R2025030 LC899331*LC899337*LC899344* This s udy
5T ime esu us loong
sp. no .
Anning, Kunming,
Yunnan, China
QHU R2025031 LC899332*LC899338*LC899345* This s udy
6T ime esu us loong
sp. no .
Anning, Kunming,
Yunnan, China
QHU R2025032 LC899333*LC899339*LC899346* This s udy
7T. c . loong Huili, Sichuan, China GP 37 EU443812 EF597522 EF597527 Dawson e al. (2008)
8T. c . loong Huili, Sichuan, China GP 38 EU443814 EF597523 EF597528 Dawson e al. (2008)
9T ime esu us sp. M . Emei, Sichuan,
China
CIB CB2120 C_AA122998.1#C_AA122989.1#C_AA123004.1#Unpublished
10 T ime esu us sp. M . Yuping, Sichuan,
China
CIB CB2121 C_AA122999.1#C_AA122990.1#C_AA123005.1#Unpublished
11 T ime esu us sp. M . Emei, Sichuan,
China
CIB CB2127 C_AA123000.1#C_AA122991.1#C_AA123006.1#Unpublished
12 T. mayaae Champhai, Mizo am,
India
NCBS NRC-AA-0012 – OM966859 – Ra hee e al. (2022)
13 T. mayaae Ri-Bhoi, Meghalaya,
India
BNHS 3658 – OM966860 – Ra hee e al. (2022)
14 T. mayaae Ri-Bhoi, Meghalaya,
India
VR/ERS/ZSI/833 – OM966862 – Ra hee e al. (2022)
15 T. mayaae Manipu , India MZMU 2970 – OQ968476 – Elangbam e al.
(2023)
16 T. mayaae Manipu , India MZMU 2971 PP566116 – – Elangbam e al.
(2023)
17 T. medoensis Mo uo, Xizang, China SYS 001831 / CHS
824
MK194252 MK201553 – Li e al. (2020)
18 T. medoensis Mo uo, Xizang, China KIZ YPX46122 MW020095 MW111479 – Che e al. (2020)
19 T. medoensis Mo uo, Xizang, China KIZ YPX46123 MW020331 MW133479 – Che e al. (2020)
20 T. medoensis Mo uo, Xizang, China ANU ZR24025 – PX068371 PX094008 This s udy
21 T. medoensis Mo uo, Xizang, China ANU ZR24026 – PX068372 PX094007 This s udy
22 T. medoensis Mo uo, Xizang, China ANU ZR24072 – PX068373 – Xu e al. (2025)
23 T. c . medoensis Pu ao, Kachin,
Myanma
AM B416 / CAS
221528
AY352735 AY352765 AY352831 Malho a and Tho pe
(2004a)
24 T. c . medoensis No heas India V18 MG995794 MG995819 MG995834 Unpublished
25 T. nujiang Fugong, Yunnan,
China
ANU ZR24133 PX061893 PX068374 PX094009 Xu e al. (2025)
26 T. nujiang Gongshan, Yunnan,
China
CIB DLR353 PV994716 PX021391 PX021401 Liang e al. (2025)
27 T. nujiang Gongshan, Yunnan,
China
CIB DLR365 PX021397 PX021407 PV994722 Liang e al. (2025)
28 T. nujiang Gongshan, Yunnan,
China
CIB DLR375 PX021398 PX021408 PV994723 Liang e al. (2025)
29 T. nujiang Gongshan, Yunnan,
China
CIB DLR380 PX021399 PX021409 PV994724 Liang e al. (2025)
30 T. p e iosus Yadong, Xizang, China QHU R2025019 PX061894 PX068368 PX094011 Xu e al. (2025)
31 T. p e iosus Yadong, Xizang, China QHU R2025020 PX061895 PX068369 PX094010 Xu e al. (2025)
32 T. p e iosus Yadong, Xizang, China QHU R2025021 PX061896 PX068370 PX094012 Xu e al. (2025)
33 T. s ejnege i M . Wuyi, Fujian, China GP 816 – KX019140 KX019294 Guo e al. (2016)
34 T. s ejnege i M . Jinggang, Jiangxi,
China
GP 662 – KX019120 KX019274 Guo e al. (2016)
35 T. s ejnege i Qimen, Anhui, China GP 470 – KX019094 KX019248 Guo e al. (2016)
36 T. s ejnege i Jinhua, Zhejiang,
China
GP 633 – KX019111 KX019265 Guo e al. (2016)
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NO. Species name Locali y Vouche NO. 16S cy b ND4 Re e ences
37 T. s ejnege i Youxi, Fujian, China GP 755 – KX019131 KX019285 Guo e al. (2016)
38 T. s ejnege i Dehua, Fujian, China GP 2435 – KX019054 KX019209 Guo e al. (2016)
39 T. s ejnege i Taichung, Taiwan,
China
AM TST23 – AF277689 EU443799 C ee e al. (2003);
Dawson e al. (2008)
40 T. s ejnege i M . Diaoluo, Hainan,
china
GP 45 – KX019088 KX019242 Guo e al. (2016)
41 T. s ejnege i Qiongzhong, Hainan,
China
GP 46 – KX019089 KX019243 Guo e al. (2016)
42 T. s ejnege i Qiongzhong, Hainan,
China
GP 48 – KX019104 KX019258 Guo e al. (2016)
43 T. s ejnege i Ca Ba NP, Hai Phong,
Vie nam
GP 881 – KX019145 KX019299 Guo e al. (2016)
44 T. s ejnege i Phia Oac-Phia Den NP,
Cao Bang, Vie nam
ROM 35321 – KT216408 KT216457 Guo e al. (2016)
45 T. s ejnege i Nonggang, Guangxi,
China
GP 2461 – KX019056 KX019211 Guo e al. (2016)
46 T. s ejnege i Nanchuan, Chongqing,
China
GP 1227 – KX019007 KX019162 Guo e al. (2016)
47 T. s ejnege i Leishan, Guizhou,
China
GP 1887 – KX019034 KX019189 Guo e al. (2016)
48 T. s ejnege i Luoding, Guangdong,
China
KIZ 09939 – KX019158 KX019312 Guo e al. (2016)
49 T. s ejnege i Enshi, Hubei, China GP 2011 – KX019040 KX019195 Guo e al. (2016)
50 T. uongsonensis Phong Nha-Ke Bang
NP, Quang T i Vie nam
AM B659 / VNUH
190606
EU443818 EU443815 EU443816 Dawson e al. (2008)
51 T. uongsonensis Khammouane, Laos NAP-09115 – PX068365 PX094015 Xu e al. (2025)
52 T. ogeli Nakhon Ra chasima,
Thailand
AM B97 AY059562 AY059574 AY059596 Malho a and Tho pe
(2004a)
53 T. ogeli Nakhon Ra chasima,
Thailand
NAP-08227 – PX068366 PX094013 Xu e al. (2025)
54 T. ogeli Nakhon Ra chasima,
Thailand
NAP-08228 – PX068367 PX094014 Xu e al. (2025)
55 T. yunnanensis Tengchong, Yunnan,
China
QHU R2025033 LC899641*LC899340*LC899347* This s udy
56 T. yunnanensis Loei, Thailand AM A164 AF517181 AY352766 AF517224 Malho a and Tho pe
(2004a); C ee e al.
(2003)
57 T. yunnanensis Pu Ma NP, Nghe An,
Vie nam
AM B174 AY059563 AY059573 AY059595, Malho a and Tho pe
(2004b)
58 T. yunnanensis Jingdong, Yunnan,
China
GP 851 – KT216392 KT216441 Guo e al. (2015)
59 T. yunnanensis Lincang, Yunnan,
China
GP 3507 KT216339 KT216384 KT216433 Guo e al. (2015)
60 T. yunnanensis Lincang, Yunnan,
China
GP 3509 KT216341 KT216386 KT216435 Guo e al. (2015)
61 T. yunnanensis Longling, Yunnan,
China
KIZ 05089 KT216352 KT216399 KT216448 Guo e al. (2015)
62 T. yunnanensis Jingdong, Yunnan,
China
KIZ 047083 KT216351 KT216398 KT216447 Guo e al. (2015)
63 T. c . yunnanensis Honghe, Yunnan,
China
GP 2532 KT216331 KT216376 KT216425 Guo e al. (2015)
64 T. c . yunnanensis Pingbian, Yunnan,
China
GP 3273 KT216334 KT216379 KT216428 Guo e al. (2015)
65 T. c . yunnanensis Pingbian, Yunnan,
China
GP 3275 KT216336 KT216381 KT216430 Guo e al. (2015)
66 T. c . yunnanensis Mengzi, Yunnan, China GP 3564 KT216344 KT216389 KT216438 Guo e al. (2015)
67 T. c . yunnanensis Mengzi, Yunnan, China GP 3589 KT216345 KT216390 KT216439 Guo e al. (2015)
Subgenus Himalayophis
68 T. a unachalensis No heas India APF/SFRI-1871 MK722155 MK720609 – Cap ain e al. (2019)
69 T. ibe anus Nepal AM B258 AY352715 AY352749 AY352810 Malho a and Tho pe
(2004a)
Subgenus Pa ias
70 T. la omacula us Mindanao, Philippines AM B4 AY352734 AY352764 AY352830 Malho a and Tho pe
(2004a)
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2272
NO. Species name Locali y Vouche NO. 16S cy b ND4 Re e ences
71 T. hageni Songhkla, Thailand AM B33 AY059552 AY059567 AY059585
72 T. malcolmi M . Kinabalu, Sabah,
Malaysia
AM B349 AY371786 AY371832 AY371861 Malho a and Tho pe
(2004a)
73 T. mcg ego i Ba an Is., Philippines AM B289 AY371795 AY371831 AY371858 Malho a and Tho pe
(2004a)
74 T. schul zei Palawan, Philippines AM B210 AY352725 AY352756 AY352819 Malho a and Tho pe
(2004a)
75 T. suma anus Bengkulu, Suma a,
Indonesia
AM B367 AY371791 AY371824 AY371864 Malho a and Tho pe
(2004a)
Subgenus Sino ipe a
76 T. sichuanensis Sichuan, China GP7 / YBU030116 HQ850449 HQ850447 HQ850446 Guo and Wang (2011)
Subgenus Popeia
77 T. lanna Doi In hanon NP,
Chiangmai, Thailand
AUP-00061 OR471637 OR470571 OR470534 Idiia ullina e al.
(2024b)
78 T. nebula is Came on Highlands,
Pahang, Malaysia
AM B345 AY371775 AY371811 AY371849 Sande s e al. (2006)
79 T. phuke ensis Phang Nga, Thailand AM B467 AY371781 AY371807 AY371851 Sande s e al. (2006)
80 T. popeio um Yingjiang, Yunnan,
China
DL2017070101 MH779887 MH779875 MH779879 Chen e al. (2019)
81 T. sabahi sabahi M . Kinabalu, Sabah,
Malaysia
AM B344 AY371771 AY371815 AY371842 Malho a and Tho pe
(2004a)
82 T. enasse imensis Suan Phueng,
Ra chabu i, Thailand
ZMMU Re-17669 PP032802 OR999089 PP032781 Idiia ullina e al.
(2024b)
Subgenus T ime esu us
83 T. albolab is Shek Kwu Chan, Hong
Kong, China
AM A157 AY352744 AF171884 AY352839 Malho a and Tho pe
(2000, 2004a)
84 T. can o i Nicoba Is., India AM A85 AY352741 AF171889 AY352836 Malho a and Tho pe
(2000, 2004a)
85 T. caudo na us Yingjiang,
Yunnan,China
AR1238 MK575042 MK575036 MK575038 Chen e al. (2020)
86 T. cilia is Thum Khao Ting,
T ang, Thailand
ZMMU Re-17661 OR471621 OR470557 OR470538 Idiia ullina e al.
(2023)
87 T. e y h ochlo is Tham Si Va Ca e,
Klong Ha , Sa Kaeo,
Thailand
RIM-0079 PQ654052 PQ658816 PQ658818 Pawangkhanan e al.
(2025)
88 T. ayeya wadyensis Rangoon, Myanma AM A209 AF517174 AF171900 AF517217 C ee e al. (2003)
89 T. kanbu iensis Khao Yai NP,
Kanchanabu i,
Thailand
ZMMU Re-17667 – OR470579 OR470553 Idiia ullina e al.
(2023); Idiia ullina e
al. (2024a)
90 T. mac ops Bangkok, Thailand ZMMU Re-17856 – PP766219 PP779475 Idiia ullina e al.
(2024c)
Ou g oup
91 C aspedocephalus
puniceus
Indonesia AM B213 AF517177 AF517192 AF517220 C ee e al. (2003)
92 Pel opelo
malaba icus
Tamil Nadu, India AM A218 AY059564 AY059569,AY059587 Malho a and Tho pe
(2004a)
App oxima ion (UFB) and he SH-like app oxima e like-
lihood a io es (SH). UFB alues we e calcula ed wi h
5000 boo s ap eplica es, wi h alues ≥ 95% conside ed
s ong suppo ; and SH was conduc ed wi h 1000 epli-
ca es, and alues ≥ 80% we e ega ded as well suppo ed
(S ephane e al. 2010; Hoang e al. 2018). The Bayes-
ian In e ence (BI) analysis was conduc ed ia M Bayes
3.2.7a (Ronquis e al. 2012) unde he bes - i model
GTR + F + I + G4 o all h ee agmen s (16S, cy b, and
ND4), which was calcula ed acco ding o BIC as well by
ModelFinde o M -Bayes in PhyloSui e 1.2.3 (Zhang e
al. 2020). In he BI analysis, h ee independen uns we e
conduc ed wi h 1 × 107 gene a ions and sampled e e y
1000 gene a ions, wi h he i s 25% o samples disca ded
as bu n-in. Nodes wi h Bayesian pos e io p obabili ies
(PP) ≥ 0.95 we e conside ed s ongly suppo ed (Huelsen-
beck e al. 2001). The phylogene ic ee was isualized
using FigT ee .1.4.4 (Rambau 2018). Unco ec ed pai -
wise gene ic dis ances (p-dis ances) among species we e
also calcula ed using MEGA X (Kuma e al. 2018).
Mo phological and hemipenial examina ion
A o al o 31 mo phological cha ac e s we e eco ded
o each specimen ( ollowing Vogel e al. 2023; Xu e al.
2025). Measu emen s we e aken wi h a Mi u oyo digi-
al calipe s (CD-15AX) o he nea es 0.1 mm, excep o
body and ail leng hs, which we e measu ed o he nea es
millime e wi h a measu ing ape. The numbe o en al
scales was coun ed acco ding o Dowling (1951). Hal
en als we e coun ed as one. The i s enla ged shield
Zoosys . E ol. 101 (4) 2025, 2267–2293
zse.penso .ne
2273
an e io o he en als was ega ded as a p e en al and
was p esen in all examined specimens. The i s scale
unde he ail mee ing i s opposi e was ega ded as he
i s subcaudal, and he e minal scu e was no included
in he numbe o subcaudals. The do sal scale ows we e
coun ed a one head leng h behind he head, a mid-body,
and a one head leng h be o e he en . In he numbe
o sup alabials ouching he subocula , hose only ouch-
ing he p esubocula we e no included. In alabials we e
conside ed o be hose shields ha we e comple ely be-
low a sup alabial and bo de ing he mou h gap. The i s
sublabial was de ined as he scale ha s a s be ween he
pos e io chin shield and he in alabials and ha bo de s
he in alabials. Values o pai ed head cha ac e s we e
eco ded on bo h sides o he head and we e epo ed in
a le - igh o de . In addi ion, we paid special a en ion
o diagnos ic colo pa e n cha ac e s, such as eye colo ,
pos ocula s eak, en ola e al s ipe, ail colo a ion, and
o e all body colo a ion. The sex was de e mined by dis-
sec ion o he en al ail base.
Compa a i e mo phological da a we e compiled om
he li e a u e, including Schmid (1925,1927), Maki (1931),
Pope (1935), Zhao e al. (1998), Da id e al. (2001a, b,
2002), Ao e al. (2004), O lo e al. (2004), Da id and
Ma hew (2005), Zhao (2006), Guo e al. (2009), Teynié
and Da id (2014), Nguyen e al. (2018), Che e al. (2020),
Ra hee e al. (2022), Elangbam e al. (2023), Nguyen e al.
(2025), Liang e al. (2025), and Xu e al. (2025).
Hemipenial mo phology
The p epa a ion, measu emen , and desc ip ion o hemi-
penes ollowed he me hodologies o Zhang e al. (1984),
Mye s and Cadle (2003), Jiang (2010), and Ren e al.
(2022, 2025). Hemipenial ma e ial was ob ained om he
le side o esh o p ese ed adul male specimens. E e -
ed hemipenes we e ein la ed using colo ed pe oleum jel-
ly, and images we e cap u ed using a digi al came a.
The measu emen cha ac e s o he hemipenis a e as
ollows: HTL = hemipenial o al leng h; HTW = hemip-
enial o al wid h; HCL = hemipenial uncus leng h; and
SPBD = sulcus spe ma icus bi u ca ion dis ance, mea-
su ed om he base o he hemipenis o he bi u ca ion
poin o he sulcus spe ma icus in he e ical di ec ion.
The ollowing a ios we e used o compa a i e pu poses:
HCL/HTL = a io o uncus leng h o o al leng h; and
SPBD/HCL = a io o sulcus bi u ca ion dis ance o un-
cus leng h, o e alua e he ela i e posi ion o he sulcus
bi u ca ion poin along he hemipenial uncus.
Specimen in o ma ion o Vi ido ipe a species used in
hemipenial compa isons conduc ed in his s udy a e p o-
ided below:
T ime esu us medoensis (N = 1): QHU R2025040 (adul
male), collec ed on June 2018, by Jundong Deng, om
Beibeng Township, Mo uo (Medog) Coun y, Nyingc-
hi Ci y, Xizang Au onomous Region, China (app ox.
29.23°N, 95.18°E).
T ime esu us p e iosus (N = 2): QHU R2025019 (adul
male), collec ed on 22 June 2025 by Zhenqi Wang, Yu-
hao Xu, Fanyue Sun, and Li ang Peng, om Xiayadong
Township, Yadong Coun y, Xigaze Ci y, Xizang Au-
onomous Region, China (27.262°N, 89.016°E; ele a-
ion 1824 m asl.); QHU R2025020 (adul male), col-
lec ed on 22 June 2025, sha es he same locali y and
collec o as QHU R2025019.
T ime esu us s ejnege i (N = 1): QHU R2025037 (adul
male), collec ed on Sep embe 2025 by Yuhao Xu,
om Liandu Dis ic , Lishui Ci y, Zhejiang P o ince,
China (28.499°N, 119.922°E; ele a ion 526 m asl.).
T ime esu us yunnanensis (N = 1): QHU R2025035
(adul male), collec ed on July 2025 by Jundong Deng,
om Lincang Ci y, Yunnan P o ince, China (app ox.
23.91°N, 100.15°E).
Abb e ia ions
Mo phological desc ip ions and mo phome y ollow
s anda dized abb e ia ions e ised acco ding o Da ko e
al. (2022). The mo phome ic cha ac e s a e as ollows:
SVL = snou - en leng h; TAL = ail leng h; TL = o al
leng h; TAL/TL = a io o ail leng h o o al leng h; HL
= head leng h; HW = head wid h; ESD = eye-snou dis-
ance, measu ed om he ip o he snou o he an e io
edge o eye; ED = eye diame e ; EN = eye o nos il dis-
ance, measu ed om he an e io ma gin o he eye o he
pos e io ma gin o he nos il; SOL = subocula leng h;
and SOW = subocula wid h.
The scala ion cha ac e s a e as ollows: VS = numbe
o en al scales; SC = numbe o subcaudal scales; CP =
cloacal pla e; SL = sup alabials; IL = in alabials; DSR
= do sal scale ows; ASR = an e io do sal scale ows;
MSR = do sal scale ows a midbody; PSR = pos e io
do sal scale ows; PRO = p eocula s; PO = pos ocula s;
SO = sup aocu-la s; SBO = subocula s; IOS = in e o -
bi al scales, numbe o scales a he na owes poin be-
ween he o bi s; SpOC = numbe o do sal head scales
su ounding he sup aocula ; NS = Nasals; INS = in e na-
sals; NINN = numbe o scales sepa a ing he in e nasals;
KTEM = keeling o he empo al scales, and KOCC =
keeling o he occipi al scales.
O he abb e ia ions: AR = Au onomous egion; NP =
Na ional Pa k; NR = Na u e Rese e; M = Moun ains;
asl. = abo e sea le el; WS = Wildli e Sanc ua y.
Resul
Phylogene ic ela ionships
The conca ena ed sequence alignmen was 2,314 base
pai s (bp) in leng h (16S = 508 bp; cy b = 1,034 bp; and
ND4 = 772 bp). Bo h ML and BI analyses yielded cong u-
en opologies (Fig. 2). Acco ding o ou m DNA-based
phylogeny, all pu a i e T ime esu us species clus e in o a
single clade. Howe e , ela ionships among species wi hin
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2274
GP 662
GP 755
GP 470
GP 633
GP 2435
AM TST23
GP 816
GP 45
GP 46
GP 48
GP 1227
GP 2011
GP 1887
KIZ 09939
GP 881
GP 2461
ROM 35321
GP 3509
GP 3507
KIZ 05089
QHU R2025033
KIZ 47083
GP 851
AM A164
AM B174
GP 2532
GP 3564
GP 3589
GP 3275
GP 3273
ANU ZR24133
CIB DLR353
CIB DLR380
CIB DLR365
CIB DLR365
AM B659
ZMMU NAP-09115
AM B97
ZMMU NAP-08227
ZMMU NAP-08228
ANU ZR24025
ANU ZR24026
ANU ZR24072
KIZ YPX46122
KIZ YPX46123
SYS 001831
NCBS NRC-AA-0012
MZMU 2970
MZMU 2971
BNHS 3658
VR/ERS/ZSI/833
AM B416
V18
QHU R2025020
QHU R2025021
QHU R2025019
QHU R2025028
QHU R2025032
QHU R2025027
QHU R2025029
QHU R2025030
QHU R2025031
GP 37
GP 38
CIB CB2127
CIB CB2120
CIB CB2121 AM A85 T. can o i
AM A209 T. ayeya wadyensis
AM A157 T. albolab is
AR 1238 T. caudo na us
RIM 0079 T. e y h ochlo is
ZMMU Re-17856 T. mac ops
ZMMU Re-17667 T. kanbu iensis
ZMMU Re-17661 T. cilia is
GP 7 T. sichuanensis
AM B4 T. la omacula us
AM B289 T. mcg ego i
AM B210 T. schul zei
AM B367 T. suma anus
AM B349 T. malcolmi
AM B33 T. hageni
AUP-00061 T. lanna
AM B467 T. phuke ensis
AM B345 T. nebula is
DL2017070101 T. popeio um
AM B344 T. sabahi sabahi
ZMMU Re-17669 T. enasse imensis
APF/SFRI-1871 T. a unachalensis
AM B258 T. ibe anus AM B213 C aspedocephalus puniceus
AM A218 Pel opelo malaba icus
0.03
86/92/0.72
99/100/1.0
79/90/0.99
99/99/1.0
100/100/1.0
75/95/0.79
100/100/1.0
97/99/1.0
80/94/0.81
100/100/1.0
100/100/1.0
87/95/0.93
100/100/0.97
–/62/0.65 100/100/1.0
89/98/0.95
96/99/1.0
95/98/0.96
99/95/0.95
100/100/1.0
100/100/1.0
100/100/1.0
82/74/0.75
83/92/0.97
100/100/1.0
85/84/0.50 100/100/1.0
51/70/0.74
95/99/0.95
68/97/0.76
100/100/1.0
76/97/0.68
93/100/0.78
100/100/1.0
–/64/0.79
84/77/1.0
100/100/1.0
69/83/0.74
83/86/0.96
99/100/1.0
99/100/1.0
–/53/–
75/89/–
72/86/0.52
73/73/0.93
80/78/0.97
100/100/1.0
82/67/0.75
87/72/0.78
99/100/1.0
Genus
T ime esu us
Vi ido ipe a
T ime esu us
Sino ipe a
Pa ias
Popeia
Himalayophis
T. s ejnege i
T. yunnanensis
T. c . yunnanensis
T.
T. nujiang
uongsonensis
T. ogeli
T. medoensis
T. mayaae
T. c . medoensis
T. p e iosus
T. loong sp. no .
T. c . loong
T ime esu us sp.
Figu e 2. Phylog am o he genus T ime esu us in e ed om h ee mi ochond ial (16S/cy b/ND4) agmen s. The b anch suppo
alues a e p esen ed wi h he SH-like app oxima e likelihood a io es (SH)/Ul a as Boo s ap App oxima ion (UFB)/Bayesian
pos e io p obabili ies (PP); he ones lowe han 50 o 0.5 a e displayed as “–”.
he genus emain poo ly esol ed. The monophyly o he
subgene a Pa ias and Popeia was s ongly suppo ed (SH
99/ UFB 100 / PP 1.0; SH 100 / UFB 100 / PP 1.0 espec-
i ely), while subgenus Himalayophis and Vi ido ipe a
ecei ed mode a e suppo (SH 87 / UFB 72 / PP 0.78;
SH 82 / UFB 74 / PP 0.75, espec i ely). The subgenus
T ime esu us is no eco e ed as monophyle ic in analysis
and is clea ly di ided in o wo deeply di e gen clades.
Wi hin he subgenus Vi ido ipe a, phylogene ic e-
la ionships among species a e gene ally well esol ed
(Fig. 2). Two specimens AM B416 (= CAS 221528) om
Kachin S a e, Myanma and V18 om No heas India,
p e iously iden i ied as T. medoensis, o m a dis inc
clade ha is sis e o he clade comp ising T. medoensis
and T. mayaae. The e o e, we en a i ely e e o hese
specimens as T. c . medoensis. In addi ion, i e specimens
Zoosys . E ol. 101 (4) 2025, 2267–2293
zse.penso .ne
2275
om sou heas e n Yunnan P o ince, China (GP 2532,
GP 3273, GP 3275, GP 3564, and GP 3589), p e iously
iden i ied as T. gump ech i, o m a sis e clade o T. yun-
nanensis wi h mode a e suppo (SH = 75 / UFB = 97 /
PP = 0.79). The unco ec ed p-dis ances be ween he wo
clades ange om 4.0–6.2% o cy b and 2.9–5.1% o
ND4. As mo phological da a o his popula ion a e no
ye a ailable, we en a i ely e e o hese specimens as T.
c . yunnanensis. The six uniden i ied T ime esu us speci-
mens om Anning, Kunming Ci y, Yunnan P o ince o m
a well-suppo ed monophyle ic clade (SH = 100 / UFB =
100 / PP = 1.0). This clade is sis e o wo specimens om
Huili Ci y, Sichuan P o ince (GP 37 and GP 38), which
had long been misiden i ied as T. yunnanensis. Toge he ,
hey o m a dis inc subclade wi h h ee addi ional speci-
mens om wes e n Sichuan P o ince (CIB CB2120, CIB
CB2121, and CIB CB2127), wi h mode a e suppo in he
ML analysis (SH = 85 / UFB = 84), bu low suppo in he
BI analysis (PP = 0.5). This en i e assemblage clus e s a
he ou e ma gin o he clade o med by all o he cu en ly
ecognized species o Vi ido ipe a.
Unco ec ed p-dis ances o he cy b and ND4 gene
agmen s a e summa ized in Table 2 and Table 3, espec-
i ely. Among cu en ly ecognized Vi ido ipe a species,
in e speci ic gene ic dis ances anged om 4.9% (T. s e-
jnege i and T. yunnanensis) o 12.8% (T. ogeli and T.
yunnanensis) o he cy b gene (Table 2), and om 3.1%
(T. s ejnege i and T. yunnanensis) o 11.0% (T. p e iosus
and T. ogeli) o he ND4 gene (Table 3). The Anning
popula ion exhibi ed subs an ial gene ic di e gence om
all o he ecognized congene s, wi h unco ec ed p-dis-
ances anging om 6.7% ( s. T. mayaae) o 11.2% ( s. T.
s ejnege i) o he cy b gene, and om 6.1% ( s. T. yun-
nanensis) o 8.3% ( s. T. medoensis) o he ND4 gene.
In addi ion, specimens om Huili Ci y, Sichuan P o -
ince showed cy b di e gence o 4.8–4.9% and ND4 di-
e gence o 4.1–5.2% om he specimens om Anning,
Yunnan P o ince. These alues app oach he minimum
in e speci ic di e gence obse ed wi hin Vi ido ipe a
o cy b, and exceed he minimum obse ed o ND4.
Meanwhile, specimens om wes e n Sichuan P o ince
di e ed om he Anning popula ion by 7.0–7.1% in cy
b and 5.9–6.1% in ND4, clea ly exceeding he minimum
in e speci ic dis ances wi hin he subgenus.
Liang e al. (2025) assessed he axonomic s a us o
T ime esu us gump ech i Da id, Vogel, Pauwels & Vidal
[ ype locali y: Phu Luang WS, Loei P o ince, Thailand],
based on ou mi ochond ial gene agmen s (12S, 16S,
cy b, and ND4) oge he wi h mo phological da a. Thei
analyses e ealed ha he opo ypes o T. gump ech i
clus e ed wi h he opo ypes o T. yunnanensis, and ha
he gene ic di e gence be ween he wo (0.0–1.6% in
cy b) was ma kedly lowe han ypical in e speci ic di-
e gence wi hin he subgenus Vi ido ipe a. Consequen -
ly, T. gump ech i was ea ed as a junio synonym o T.
yunnanensis. Ou phylogene ic analyses yielded simila
esul s in his ega d. Howe e , he phylogene ic opolo-
gy p esen ed by Liang e al. (2025) placed he subgenus
T ime esu us nes ed wi hin wha was p e iously consid-
e ed he subgenus Vi ido ipe a, o ming a sis e g oup
o T. medoensis, T. mayaae, and T ime esu us sp. (Hui-
li Ci y, Sichuan P o ince, China). Based on his esul ,
hey anno a ed Vi ido ipe a on hei phylogene ic ee
as comp ising only he clade including T. s ejnege i, T.
yunnanensis, T. ogeli, T. uongsonensis, and T. nujiang,
he eby excluding T. medoensis, T. mayaae, and T ime e-
su us sp. (Huili Ci y, Sichuan P o ince, China) om he
subgenus. This a angemen is incong uen wi h bo h p e-
ious s udies and ou own mo phological and molecula
e idence (C ee e al. 2003; Malho a and Tho pe 2004a,
b; Dawson e al. 2008; Guo e al. 2015, 2016; Ra hee e
al. 2022; Xu e al. 2025; his s udy). A likely explana ion
o his inconsis ency is ha Liang e al. (2025) included
only h ee species o he subgenus T ime esu us, namely
T. sep en ionalis K ame , 1977, T. guoi, and T. pu pu eo-
macula us (G ay, 1832), and he phylogene ic esolu ion
in his pa o he ee is a he weak. The e o e, he o e all
phylogene ic amewo k p oposed by Liang e al. (2025)
wa an s addi ional e i ica ion wi h expanded sampling.
Fu he mo e, de ailed mo phological examina ions con-
i med ha he specimens om Anning, Kunming, Yunnan
a e diagnosably dis inc om all known Vi ido ipe a spe-
cies. Howe e , due o he lack o a ailable mo phological
da a o he Huili and Chengdu in Sichuan P o ince spec-
imens, we a e cu en ly unable o de e mine hei exac
axonomic s a us. Based on he combined molecula and
mo phological e idence, we desc ibe he popula ion om
Anning Ci y, Kunming Ci y, Yunnan P o ince, China as
a new species, and he Huili popula ion is en a i ely e-
assigned o his new species, while he wes e n Sichuan
popula ion is he ein ea ed as T ime esu us sp.
Taxonomy
T ime esu us loong Xu, Deng, Zhang, Nguyen,
Poya ko , Vogel & Peng, sp. no .
h ps://zoobank.o g/B557FA3A-3B3A-47C2-A9F8-239824C7DE63
Tables 4, 5; Figs 3, 4A, B, 5–8, 9B, C, 10A, B, 11A1, A2
Type ma e ial. Holo ype • QHU R2025027, adul male,
collec ed om Qinglongxia, Anning Ci y, Kunming Ci y,
Yunnan P o ince, China (25.0590°N, 102.3561°E; ele a-
ion 1,822 m asl.), on 15 Sep embe 2025 by J.D. Deng
and Y.H. Xu.
Pa a ypes (n = 5) • QHU R2025031 (adul emale),
wi h same da a as he holo ype. • QHU R2025028 (adul
male), collec ed on 22 Sep embe 2025 by J.D. Deng and
Y.H. Xu; • QHU R2025029 (subadul male) and • QHU
R2025032 (adul emale), collec ed on 27 May 2025 by
J.D. Deng; • QHU R2025030 (subadul male), collec ed
on 21 Sep embe 2025 by J.D. Deng and Y.H. Xu; all
om he same locali y as he holo ype.
Diagnosis. The new species T ime esu us loong sp.
no . is dis inguished om all o i s congene s by a combi-
na ion o he ollowing mo phological cha ac e s: (1) i s
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2282
Table 4. Main measu emen s and me is ic cha ac e s o he ype se ies o T ime esu us loong sp. no .
Specimen
ouche
QHU R2025027 QHU R2025028 QHU R2025029 QHU R2025030 QHU
R2025031
QHU
R2025032
Type Holo ype Pa a ype
Sex Male Male Subadul male Subadul male Female Female
SVL (mm) 517 601 402 335 657 632
TAL (mm) 129 147 96 83 135 146
TL (mm) 646 748 498 418 792 778
TAL/TL 0.20 0.20 0.19 0.20 0.17 0.19
HL (mm) 25.8 29.0 20.7 19.3 36.6 34.6
HW (mm) 17.7 20.9 14.4 13.9 27.6 20.9
HW/HL 0.69 0.72 0.70 0.72 0.75 0.60
ESD (mm) 8.6 9.5 6.6 5.7 11.4 8.9
EN (mm) 5.4 6.6 4.5 4.0 7.7 6.7
ED (mm) 3.3 3.6 2.4 2.7 4.2 4.3
ED/ESD 0.38 0.38 0.36 0.47 0.37 0.48
ED/HL 0.13 0.12 0.12 0.14 0.11 0.12
NS 1, undi ided 1, undi ided 1, undi ided 1, undi ided 1, undi ided 1, undi ided
INS Sligh ly ena ged Sligh ly ena ged Sligh ly ena ged Sligh ly ena ged Sligh ly ena ged Sligh ly ena ged
NINN 2 small + 1 la ge 2 small + 1 la ge 2 2 2 2
PRO 3 3 3 3 3 3
PO 2/2 4/4 2/2 3/3 2/2 3/3
SO 1/1 3/2 1/1 1/1 1/1 1/1
SOL 4.3 – 3.6 3.4 5.4 4.3
SOW 1.9 – 1.3 1.2 2.1 2.1
SBO 1/1 1/1 1/1 1/1 1/1 1/1
IOS 9 10 7 9 10 11
SpOC 8/8 8/7 7/8 9/8 6/5 6/7
KTEM Smoo h Smoo h Smoo h Smoo h Smoo h Smoo h
KOCC Sligh ly keeled Smoo h Sligh ly keeled Sligh ly keeled Sligh ly keeled Smoo h
SL 10/10 11/10 10/10 10/10 11/11 10/10
IL 12/11 12/11 12/12 11/12 13/14 12/12
DSR 19-19-15 21-19-15 23-19-15 19-19-15 21-19-15 19-19-15
VS 150+2 151+2 150+3 150+2 158+3 157+2
SC 67, pai ed 68, pai ed 67, pai ed 67, pai ed 57, pai ed 64, pai ed
VS+SC 217 219 217 217 215 221
CP 1 1 1 1 1 1
Eye colo Ambe wi h a ain
eddish hue
Ambe wi h a ain
eddish hue
Ambe Ambe Yellowish Yellowish
Pos ocula
s eak
Thin and whi e, wi h a ew
scale ma gins inged wi h
yellow
Whi e Thin and whi e, wi h he
uppe ma gins o a ew
scales inged wi h eddish-
b own
Thin and whi e, wi h a ew
scale ma gins inged wi h
yellow
Absen Absen
Ven ola e al
s ipe
Yellow+whi e+ ed+b own Thin, whi e and inged
wi h yellowish ma gins
Yellow+whi e+ ed+b own Yellow+whi e+ ed+b own Whi e Absen
while only limi ed in o ma ion is a ailable o emales
due o he small sample size. This species exhibi s sex-
ual dich oma ism. In males, he eyes a e ambe in colo .
Among he wo adul specimens (QHU R2025027 and
QHU R2025028), he i ises exhibi a sligh eddish hue,
whe eas in he wo subadul specimens (QHU R2025029
and QHU R2025030), he eyes a e pu ely ambe . The
pos ocula s eak is na ow and whi e, wi h a ew scale
ma gins sligh ly inged wi h yellow o eddish-b own.
The en ola e al s ipe uns along he i s do sal scale
ow and is a iable in colo a ion among males, occu ing
in wo o ms. In mos male specimens, i consis s o a
na ow whi e band bo de ed do sally by a ain yellowish
ma gin and en ally by a hin, ed line, wi h a di use
b ownish inge below he ed line. Howe e , in QHU
R2025028, he en ola e al s ipe is only a na ow whi e
band bo de ed on bo h sides by ain yellowish ma gins.
In some indi iduals, he uppe and lowe ma gins o he
i s do sal scale ow emain g een.
In emales, he eyes a e yellowish. The pos ocula
s eak is absen in bo h specimens. The en ola e al s ipe
is absen in QHU R2025032, while in QHU R2025031 i
is na ow, whi e, and es ic ed o he middle po ion o
he i s do sal scale ow.
In all indi iduals, he do sal su aces o he head and
body a e g ass g een. In males, he la e al body colo a ion
Zoosys . E ol. 101 (4) 2025, 2267–2293
zse.penso .ne
2283
Figu e 7. Pa a ypes o T ime esu us loong sp. no . A. Do sal iew o he head; B. La e al iew o he head; C. Ven al iew o he
head; D. La e al iew o he body; E. Ven al iew o he body. A1–E1. QHU R2025028, adul male; A2–E2. QHU R2025029,
subadul male; A3–E3. QHU R2025030, subadul male; A4–E4. QHU R2025031, adul emale; A5–E5. QHU R2025032, adul
emale. Pho og aphs by Y.H. Xu.
is sligh ly pale han he mid-do sal egion, and he la -
e al do sal scales a e a iably speckled wi h ine c eam-
ish-blue o whi e spo s. In con as , emales exhibi a uni-
o mly g ass-g een colo a ion ac oss he en i e do sum.
The do sal su ace o he ail is dis inc ly bicolo ed: g een
an e io ly, g adually ansi ioning o nea ly uni o m o -
ange o ligh o ange- ed pos e io ly, wi h he ail ip being
blackish b own (Fig. 6B–E). The en al su ace o he
head and he cen al po ion o he an e io en al body
a e no iceably pale , appea ing pale whi ish-g een o
c eamish-yellow. Pos e io o midbody, he en al colo -
a ion becomes uni o mly yellowish-g een o ligh g een.
The an e io po ion o en al su ace o he ail is pale
g een, while he pos e io po ion is ligh o ange- ed.
Main cha ac e s o he pa e n in p ese a i e. The
desc ip ion is based on a subadul male specimen (QHU
R2025029) and a emale specimen (QHU R2025032),
bo h collec ed in May 2025. A e app oxima ely i e
mon hs in alcohol, he do sal su ace o he head had
u ned oli e-yellow and ha o he body yellow in bo h
specimens. The eyes had become pale g ay, and he su-
p alabials as well as he en al su ace o he head had
aded o milky whi e. The la e al po ions o he en al
body we e pale yellow, while he en i e cen al en al
su ace had u ned c eamish-yellow. In he male speci-
men, he pos e io po ion o he en al body was sligh ly
yellowish-g een. The o ange- ed a ea o he ail emained
ela i ely well de ined. In he male specimen, he en o-
la e al s ipe had changed o whi e do sally and b ownish
wi h a sligh eddish hue en ally.
Dis ibu ion and na u al his o y. Cu en ly, T ime e-
su us loong sp. no . is ce ainly known only om he
ype locali y, Qinglongxia, Anning Ci y, Kunming Ci y,
Yunnan P o ince, China, a an ele a ion o app oxima e-
ly 1,822 m asl. All specimens we e encoun e ed du ing
he i s hal o he nigh , coiled wi hin sh ubs on hill-
sides nea s eams in well-p ese ed b oad-lea ed o es
(Fig. 10A–C), when nigh ime empe a u es anged 15 °C
~ 18 °C. The emale specimen QHU R2025031 de eca -
ed immedia ely when dis u bed du ing cap u e. I s eces
con ained nume ous bi d ea he s, sugges ing ha small
bi ds may cons i u e pa o he adul die o his species
(Fig. 10D).
Compa isons. T ime esu us loong sp. no . is assigned
o he subgenus Vi ido ipe a based on a combina ion o
diagnos ic cha ac e s, including i s phylogene ic posi ion,
he comple e sepa a ion o he i s sup alabial om he
nasal scale, and a sho spinose hemipenis, all o which
a e cha ac e is ic ea u es o Vi ido ipe a (e.g., Nguyen
e al. 2025; Xu e al. 2025). Acco dingly, mo phological
compa isons a e es ic ed o congene s wi hin Vi ido i-
pe a, which cu en ly comp ises nine ecognized species
and ep esen s he mos ele an g oup o di e en ial di-
agnosis. The p incipal cha ac e s dis inguishing T ime e-
su us loong sp. no . om hese congene s a e summa-
ized in Table 5 and illus a ed in Fig. 10.
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2284
Figu e 8. Colo a ion o he pa a ypes o T ime esu us loong sp. no . in li e. A. La e al iew o he head; B. Do sola e al iew;
C. Ven al iew. A1–C1. QHU R2025028, adul male; A2–C2. QHU R2025029, subadul male; A3–C3. QHU R2025030, subadul
male; A4–C4. QHU R2025031, adul emale. Pho og aphs by Y.H. Xu.
Mo phologically, T ime esu us loong sp. no . is mos
simila o T. yunnanensis (dis ibu ed in cen al o sou h-
wes e n Yunnan P o ince o China, he Mandalay Region
and Shan S a e o Myanma , no he n Laos, no he n and
cen al Thailand, and no he n o cen al Vie nam), bu i
can be clea ly dis inguished om he la e by he ollow-
ing combina ion o cha ac e s: (1) smalle maximum SVL
in emales (657 mm s. 804 mm); (2) lowe o al num-
be o VS+SC in males (217–219 [mean 217.5 ± 1.0] s.
221–231 [mean 225.9 ± 3.9]); (3) i ises ambe in males s.
b igh o deep ed i ises obse ed in males o T. yunnan-
ensis (Fig. 10A, B, K); (4) he en ola e al s ipe in he
males o he new species is slende and limi ed o he i s
do sal scale ow occu ing in wo o ms, ei he a na ow
whi e band bo de ed do sally by a ain yellowish ma gin
and en ally by a ed line wi h a di use b ownish inge
below he ed line, o a na ow whi e band bo de ed by
ain yellowish ma gins on bo h sides, while in he males o
T. yunnanensis he s ipe is b oade and mo e i id, occu-
pying he en i e i s do sal scale ow and he lowe ma gin
o he second ow, e en ex ending on o he la e al edges
o he en als, consis ing o a hick ed lowe bo de and
a whi e uppe zone wi h sha ply sepa a ed colo s isible
om a dis ance; (5) he new species has a an e io ly g een
colo a ion o ail, pos e io ly g adually ansi ioning o a
uni o mly o ange o ligh o ange- ed. In con as , T. yun-
nanensis shows da k ed blo ches o ming la ge, i egula
pa ches wi h une en ma gins on he an e io pa o he ail
(Fig. 11). In hemipenial mo phology, T ime esu us loong
sp. no . can be dis inguished om T. yunnanensis by sho -
e hemipenial uncus and a highe deg ee o bi u ca ion
(HCL/HTL a io 0.62–0.66 s. 0.69), a mo e dis al sulcus
spe ma icus bi u ca ion poin (SPBD/HCL a io 0.43–0.49
s. 0.30), and a g ea e numbe o enla ged spines (14–19
pe side s. 9–10, excluding he smalle spines lining he
ma gin o he sulcus spe ma icus) (Fig. 4A, B, F).
Zoosys . E ol. 101 (4) 2025, 2267–2293
zse.penso .ne
2285
Figu e 9. Habi a and ield obse a ions o T ime esu us loong sp. no . A. The mic ohabi a o he new species in Anning, Kunming,
China; B. Holo ype QHU R2025027 in li e, in si u; C. Pa a ype QHU R2025028 in li e, in si u; D. Feces o he pa a ype QHU
R2025031. Pho og aphs by Y.H. Xu (A–C) and J.D. Deng (D).
T ime esu us loong sp. no . di e s om T. mayaae
(dis ibu ed ac oss he Indian S a es o Manipu , Megha-
laya, Mizo am, Assam, Nagaland, Wes Bengal, Sikkim,
Bhu an, and Chin S a e o Myanma ) by he ollowing
combina ion o cha ac e s: la ge maximum SVL in e-
males (657 mm s. 590 mm); lowe numbe o VS in
males (150–151 [mean 150.3 ± 0.5] s. 153–163 [mean
158.1 ± 3.0]); highe numbe o SC in emales (57–64
[mean 60.5 ± 4.9] s. 53–55 [mean 54.0 ± 1.0]); and high-
e numbe o VS+SC in emales (215–221 [mean 218.0 ±
4.2] s. 205–208 [mean 206.7 ± 1.5]). In addi ion, emales
o T ime esu us loong sp. no . ha e yellowish i ises, dis-
inc ly di e ing om he g eenish eyes o T. mayaae.
T ime esu us loong sp. no . di e s om T. medoensis
(dis ibu ed in he Xizang AR o sou hwes e n China, and
possibly in no he n Myanma and no heas e n India)
by he ollowing combina ion o cha ac e s: sligh la ge
maximum SVL in bo h sexes (601 mm in males, 657 mm
in emales s. 553 mm in males, 624 mm in emales);
highe numbe o VS in bo h sexes (150–151 [mean 150.3
± 0.5] in males, 157–158 [mean 157.5 ± 0.7] in emales
s. 146–151 [mean 148.6 ± 1.8] in males, 145–147 [mean
146.0 ± 1.0] in emales); g ea e numbe o SC in males
(67–68 [mean 67.3 ± 0.5] s. 55–59 [mean 57.0 ± 1.4]);
and highe numbe o VS+SC in bo h sexes (217–219
[mean 217.5 ± 1.0] in males, 215–221 [mean 218.0 ±
4.2] in emales s. 201–208 [mean 205.6 ± 2.9] in males,
204–206 [mean 205.0 ± 1.0] in emales). The new species
also possesses a g ea e numbe o SL (10–11 s. 8–9)
and IL (11–14 s. 8–10). I u he di e s in ha ing mo e
do sal scale ows a midbody (19 s. 17) and pos e io ly
(15 s. 13, a ely 11). Eye colo a ion also di e s: males o
new species ha e ambe i ises wi h a sligh eddish hue,
and emales yellowish, whe eas bo h sexes o T. medoen-
sis possess g een o yellowish-g een eyes (Fig. 10A, B,
E). In addi ion, he en ola e al s ipe in emales o new
species is absen o uni o mly whi e, con as ing wi h he
ed-and-whi e s ipe o T. medoensis. In hemipenial mo -
phology, T ime esu us loong sp. no . can be dis inguished
om T. medoensis by longe hemipenial uncus and a
lowe deg ee o bi u ca ion (HCL/HTL a io 0.62–0.66
s. 0.53), and a mo e dis al sulcus spe ma icus bi u ca ion
poin (SPBD/HCL a io 0.43–0.49 s. 0.21) (Fig. 4A–C).
T ime esu us loong sp. no . di e s om T. nujiang (dis-
ibu ed ac oss se e al locali ies wi hin he Nujiang Ri e
Basin o no hwes e n Yunnan P o ince, China, including
Gongshan, Fugong, and Lushui Con ies) by he ollowing
combina ion o cha ac e s: sligh ly smalle maximum SVL
in bo h sexes (601 mm in males, 657 mm in emales s.
694 mm in males, 682 mm in emales); lowe numbe o
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2286
Table 5. Summa y o mo phological cha ac e s in membe s o he subgenus Vi ido ipe a. No es: Diagnos ic cha ac e s dis inguishing T ime esu us loong sp. no . a e indica ed in bold.
T ime esu us Sou ces Max SVL (mm) VS SC VS+SC ASR MSR PSR SL IL
♂♀♂ ♀ ♂ ♀ ♂ ♀
T. loong sp. no . (19) 601 657 150–151
[150.3 ± 0.5]
157–158
[157.5 ± 0.7]
67–68 [67.3 ± 0.5] 57–64 [60.5 ± 4.9] 217–219 [217.5
± 1.0]
215–221
[218.0 ± 4.2]
19–23 19 15 10 o 11 11–14
T. mayaae (7), (8), (9),
(14), (15), (16)
610 590 153–163
[158.1 ± 3.0]
152–153
[152.7 ± 0.6]
54–69 [61.1 ± 4.5] 53–55 [54.0 ±
1.0]
211–231 [218.9
± 6.1]
205–208
[206.7 ± 1.5]
19–28 19–21 15–17 8–10 10–13
T. medoensis (4), (13), (16) 553 624 146–151
[148.6 ± 1.8]
145–147
[146.0 ± 1.0]
55–59 [57.0 ±
1.4]
58–60 [59.0 ± 1.0] 201–208
[205.6 ± 2.9]
204–206
[205.0 ± 1.0]
17 ( a ely
19)
17 13 ( a ely
11)
8 o 9 8–10
T. nujiang (17) 694 682 164–173
[168.7 ± 3.6]
165–168
[166.6 ± 1.3]
59–68 [62.8 ± 3.9] 57–60 [58.2 ± 1.1] 226–241
[231.3 ± 6.8]
222–226
[224.8 ± 1.8]
19–21
( a ely 22)
19 15 9 o 10 11–13
T. p e iosus (18) 516 512 140–143
[141.5 ± 2.1]
142 56–58 [57.0 ±
1.4]
54 198–199
[198.5 ± 0.7]
196 19 19 15 8 o 9 10 o
11
T. s ejnege i (1), (3), (16) 635 670 154–178
[164.5 ± 4.8]
155–173
[163.9 ± 3.5]
60–80 [70.3 ± 4.2] 58–70
[63.4 ± 2.9]
218–256
[234.9 ± 7.7]
218–237
[227.5 ± 4.5]
21–25 21–23 15 9–12 10–14
T. uongsonensis (8), (11), (16) 521 488 170–190
[179.8 ± 8.7]
165–167
[166.0 ± 1.0]
65–71 [67.4 ± 2.5] 61–70 [66.7 ± 4.9] 235–259
[247.2 ± 10.7]
228–235
[232.7 ± 4.0]
21 21 15 9–11 11–13
T. ogeli (5), (16) 661 947 157–169
[162.8 ± 3.7]
157–173
[164.2 ± 5.8]
63–71 [67.0 ± 2.3] 59–70 [63.0 ± 3.8] 221–238
[229.8 ± 4.9]
218–238
[227.7 ± 6.3]
21–23 21 ( a ely
20)
15 9–11 11–16
T. yunnanensis (2), (3), (6),
(10), (12), (17)
602 804 151–164
[158.2 ± 3.9]
150–164
[157.9 ± 3.6]
61–71 [66.1 ± 2.9] 52–65 [57.8 ± 3.6] 221–231
[225.9 ± 3.9]
211–223
[216.4 ± 3.7]
19–21 19–21 15–17 9–11 10–13
T ime esu us Eye colo in li e Pos ocula s eak in li e Ven ola e al s ipe in li e Tail ed Tail ed pa e n Body colou a ion in li e
♂ ♀ ♂ ♀ ♂ ♀
T. loong sp. no . ambe ,
some imes wi h
a sligh eddish
hue
yellowish hin and whi e,
wi h a ew
scale ma gins
sligh ly inged
wi h yellow o
eddish-b own
absen yellow + whi e +
ed + b own o hin,
whi e wi h yellowish
ma gins
absen o whi e,
hin
yes o ange o ligh o ange- ed, uni o m,
blackish b own a he ip
uni o mly g ass-g een
T. mayaae us y o
g eenish
g eenish none o ed +
whi e
absen ed + whi e, wide pale yellow + whi e,
hin
yes us y ed, o ming blo ches an e io ly uni o mly g ass-g een
T. medoensis g een o
yellowish
g een
g een o
yellowish
g een
none o ain
whi e
absen ed + whi e, wide ed + whi e yes ed o da k us y ed, some imes
o ming blo ches an e io ly
uni o mly g ass-g een
T. nujiang golden yellow golden yellow none absen da k ed + whi e ain whi e yes eddish-b own wi h da k b own a he ip uni o mly g ass-g een
T. p e iosus eddish-b own o ange-yellow none o e y
ain , hin whi e
absen ed + whi e, wide whi e, hin yes b ick- ed, some imes o ming blo ches
an e io ly
uni o mly g ass-g een
T. s ejnege i b igh ed o
o ange- ed
( a ely yellow)
yellow o
o ange
ed + whi e o
whi e
absen o whi e,
hin
ed + whi e, wide ed + whi e o whi e yes ligh o ange- ed o ed, uni o m o o ming
blo ches an e io ly
uni o mly g ass-g een
T. uongsonensis g eenish-
yellow
g eenish-
yellow
none absen ed + b own, wide ed + b own no – g eenish blue wi h b own
b oad bands
T. ogeli ligh o ange ligh o ange none o ain
whi e
absen ed + whi e, wide pale yellow +
whi e, hin
no – uni o mly g ass-g een
T. yunnanensis b igh o deep
ed
golden yellow ed + whi e o
whi e o absen
absen o whi e,
hin
ed + whi e, wide absen o whi e o
pale g een, hin
yes da k ed, wi h la ge an e io blo ches
and i egula ly shaped ma gins
uni o mly g ass-g een
Sou ces: (1) = Maki 1931; (2) = Pope 1935; (3) = Zhao e al. 1998; (4) = Da id e al. 2001a; (5) = Da id e al. 2001b; (6) = Da id e al. 2002; (7) = Ao e al. 2004; (8) = O lo e al. 2004; (9) = Da id and Ma hew 2005; (10) = Guo e al. 2009; (11) =
Teynié and Da id 2004; (12) = Nguyen e al. 2018; (13) = Che e al. 2020; (14) = Ra hee e al. 2022; (15) = Elangbam e al. 2023; (16) = Nguyen e al. 2025; (17) = Liang e al. 2025; (18) = Xu e al. 2025; (19) = his s udy.
Zoosys . E ol. 101 (4) 2025, 2267–2293
zse.penso .ne
2287
Figu e 10. Compa ison o he la e al head iew o males o species in he subgenus Vi ido ipe a in li e. A. T ime esu us loong sp.
no ., QHU R2025027, holo ype; B. T ime esu us loong sp. no ., QHU R2025028, pa a ype; C. T ime esu us sp. om Chengdu,
Sichuan, China; D. T. mayaae om Kangpokpi, Manipu , India; E. T. medoensis, QHU R2025040, om Mo uo (Medog), Xizang,
China; F. T. nujiang om Gongshan, Yunnan, China; G. T. p e iosus, QHU R2025020, om Yadong, Xizang AR, China; H. T. s ej-
nege i, QHU R2025037, om Lishui, Zhejiang, China; I. T. uongsonensis om Phong Nha-Ke Bang NP, Quang T i, Vie nam; J. T.
ogeli om Phong Nha-Ke Bang NP, Quang T i, Vie nam; K. T. yunnanensis om Jingdong, Yunnan, China; L. T. c . yunnanensis
om M . Dawei, Honghe, Yunnan, China. Pho og aphs by Y.H. Xu (A–C, E–L) and P. Shinde (D).
VS in bo h sexes (150–151 [mean 150.3 ± 0.5] in males,
157–158 [mean 157.5 ± 0.7] in emales s. 164–173 [mean
168.7 ± 3.6] in males, 165–168 [mean 166.6 ± 1.3] in e-
males); and lowe numbe o VS+SC in bo h sexes (217–
219 [mean 217.5 ± 1.0] in males, 215–221 [mean 218.0 ±
4.2] in emales s. 226–241 [mean 231.3 ± 6.8] in males,
222–226 [mean 224.8 ± 1.8] in emales). Eye colo a ion
also di e s, wi h males o he new species ha ing ambe
i ises wi h a sligh eddish hue s. yellow o mo led g ay
and lesh-colo ed in T. nujiang (Fig. 10A, B, F). In addi-
ion, a pos ocula s eak is p esen in males o new species
bu absen o e y ain in T. nujiang.
T ime esu us loong sp. no . di e s om T. p e iosus
(dis ibu ed in Yadong Coun y, Xigaze Ci y, Xizang AR,
China) by he ollowing combina ion o cha ac e s: ha ing
a la ge maximum SVL in bo h sexes (601 mm in males,
657 mm in emales s. 516 mm in males, 512 mm in e-
males); a highe numbe o VS (150–151 [mean 150.3 ±
0.5] in males, 157–158 [mean 157.5 ± 0.7] in emales s.
140–143 [mean 141.5 ± 2.1] in males, 142 in emale); a
g ea e numbe o SC (67–68 [mean 67.3 ± 0.5] in males,
57–64 [mean 60.5 ± 4.9] in emales s. 56–58 [mean 57.0 ±
1.4] in males, 54 in emale); a highe numbe o VS+SC
(217–219 [mean 217.5 ± 1.0] in males, 215–221 [mean
218.0 ± 4.2] in emales s. 198–199 [mean 198.5 ± 0.7]
in males, 196 in emale); a g ea e numbe o sup alabials
(10 o 11 s. 8 o 9); in possessing all subcaudal scales
pai ed ( s. pa ly single); and eye colo a ion in males is
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2288
Figu e 11. Compa ison o male colo a ion be ween T ime esu us loong sp. no . and T. yunnanensis. A1. T ime esu us loong sp.
no ., QHU R2025028; A2. T ime esu us loong sp. no ., QHU R2025029; B1. T. yunnanensis, QHU R2025035, om Lincang,
Yunnan, China; B2. T. yunnanensis, un ouche ed indi idual om Jingdong, Yunnan, China. The whi e a ows indica e he do sal
su ace o he ail, and blue a ows indica e he en ola e al s ipe. Pho og aphs by Y.H. Xu.
ambe wi h a sligh eddish hue ( s. b ick ed) (Fig. 10A,
B, G). In hemipenial mo phology, T ime esu us loong
sp. no . can be dis inguished om T. p e iosus by lon-
ge hemipenial uncus and a lowe deg ee o bi u ca ion
(HCL/HTL a io 0.62–0.66 s. 0.47–0.53), and a mo e
dis al sulcus spe ma icus bi u ca ion poin (SPBD/HCL
a io 0.43–0.49 s. 0.28–0.36) (Fig. 4A, B, D).
T ime esu us loong sp. no . di e s om T. s ejnege i
( es ic ed o eas e n and sou he n China, including Tai-
wan P o ince, no he n Vie nam, and no heas e n Laos)
by he ollowing combina ion o cha ac e s: lowe VS in
males (150–151 [mean 150.3 ± 0.5] s. 154–178 [mean
164.5 ± 4.8]); lowe numbe o VS+SC in males (217–
219 [mean 217.5 ± 1.0] s. 218–256 [mean 234.9 ± 7.7]);
and ewe do sal scale ows a midbody (19 s. 21–23). In
males, he i ises a e ambe wi h a sligh eddish hue, and
he en ola e al s ipe is slende and con ined o he i s
do sal scale ow, occu ing in wo o ms: ei he a na ow
whi e band bo de ed do sally by a ain yellowish ma -
gin and en ally by a ed line wi h a di use b ownish
inge below he ed line, o a na ow whi e band bo de ed
by ain yellowish ma gins on bo h sides in T ime esu us
loong sp. no . In con as , in T. s ejnege i, he i ises a e
o ange- ed o ed (Fig. 10A, B, H), and he en ola e al
s ipe consis s o a ed lowe bo de and a whi e uppe
zone, wi h he wo colo s sha ply sepa a ed and clea ly
isible om a dis ance. In emales, he en ola e al s ipe
is uni o mly whi e, whe eas in T. s ejnege i i is whi e o
ed-and-whi e. In hemipenial mo phology, T ime esu us
loong sp. no . can be dis inguished om T. s ejnege i by
sho e hemipenial uncus and a highe deg ee o bi u -
ca ion (HCL/HTL a io 0.62–0.66 s. 0.69), and a mo e
dis al sulcus spe ma icus bi u ca ion poin (SPBD/HCL
a io 0.43–0.49 s. 0.28) (Fig. 4A, B, E).
T ime esu us loong sp. no . di e s om T. uongson-
ensis ( es ic ed o cen al Vie nam and cen al Laos) by
he ollowing combina ion o cha ac e s: a la ge maxi-
mum SVL in bo h sexes (601 mm in males, 657 mm in e-
males s. 521 mm in males, 488 mm in emales); a lowe
numbe VS (150–151 [mean 150.3 ± 0.5] in males, 157–
158 [mean 157.5 ± 0.7] in emales s. 170–190 [mean
179.8 ± 8.7] in males, 165–167 [mean 166.0 ± 1.0] in e-
males); a g ea e numbe o SC (67–68 [mean 67.3 ± 0.5]
in males, 57–64 [mean 60.5 ± 4.9] in emales s. 56–58
[mean 57.0 ± 1.4] in males, 54 in he emale); and conse-
quen ly a lowe numbe o VS+SC (217–219 [mean 217.5
± 1.0] in males, 215–221 [mean 218.0 ± 4.2] in emales s.
235–259 [mean 247.2 ± 10.7] in males, 228–235 [mean
232.7 ± 4.0] in emales); and ewe do sal scale ows a
midbody (MSR 19 s. 21). Eye colo a ion is ambe wi h
a sligh eddish hue in males and yellowish in emales,
con as ing wi h he g eenish-yellow eyes o bo h sexes in
Zoosys . E ol. 101 (4) 2025, 2267–2293
zse.penso .ne
2289
he la e species (Fig. 10A, B, I). The body is uni o mly
b igh g ass-g een, whe eas T. uongsonensis exhibi s a
g eenish-blue do sum wi h b oad b own c ossbands.
Finally, T ime esu us loong sp. no . di e s om T.
ogeli (dis ibu ed in he sou heas e n pa o cen al
Thailand, cen al and sou he n Laos, Cambodia, and cen-
al and sou he n Vie nam) by he ollowing combina-
ion o cha ac e s: smalle maximum SVL in bo h sexes
(601 mm in males, 657 mm in emales s. 661 mm in
males, 947 mm in emales); lowe numbe o VS in males
(150–151 [mean 150.3 ± 0.5] s. 157–169 [mean 162.8 ±
3.7]); lowe numbe o VS+SC in males (217–219 [mean
217.5 ± 1.0] s. 221–238 [mean 229.8 ± 4.9]); ewe do -
sal scale ows a midbody (MSR 19 s. 21, a ely 20); and
eye colo a ion in males is ambe wi h a sligh eddish hue
( s. ligh o ange o lesh-colo ed) (Fig. 10A, B, J).
Discussion
Pholidosis (i.e., he pa e n and a angemen o scales) has
long been conside ed one o he mos impo an diagnos-
ic ea u es o species iden i ica ion and classi ica ion in
snakes. Howe e , in he genus T ime esu us, scale coun s
a e o en highly conse ed, making axonomic delimi a-
ion pa icula ly challenging (Guo e al. 2009; Da id e
al. 2011; Sumon ha e al. 2021; Idiia ullina e al. 2024b;
Nguyen e al. 2025; Pawangkhanan e al. 2025). Many
closely ela ed species a e nea ly indis inguishable based
on scala ion cha ac e s alone, especially wi hin he sub-
genus Vi ido ipe a, whe e se e al axonomic uni s had
long been misiden i ied as T. s ejnege i o T. yunnanensis
(Schmid 1927; Pope 1935; Smi h 1943; Zhao e al. 1998;
C ee e al. 2003; Malho a and Tho pe 2004a, b; Zhao
2006; Guo e al. 2009; Ra hee e al. 2022; Wu e al. 2023;
Liang e al. 2025). Fo example, T. mayaae, T. nujiang,
and he newly desc ibed T ime esu us loong sp. no . we e
p e iously ega ded as geog aphically dis inc popula-
ions o T. yunnanensis (Zhao e al. 1998; Malho a and
Tho pe 2004a, b; Zhao 2006; Ra hee e al. 2022; Wu e al.
2023; Liang e al. 2025). In ecen yea s, he inco po a ion
o molecula phylogene ic da a has g adually unco e ed
he ue di e si y wi hin Vi ido ipe a, e ealing ha his
subgenus ha bo s a g ea e numbe o c yp ic species han
p e iously ecognized. None heless, due o signi ican
o e lap in pholidosis among many axa, he inclusion o
mo e di e se mo phological cha ac e s is essen ial o ac-
cu a e species iden i ica ion and classi ica ion. One such
p omising cha ac e se is colo a ion pa e n, which has
shown po en ial in esol ing axonomic ambigui ies in his
g oup. Sexual dich oma ism is common in Vi ido ipe a,
wi h males ypically exhibi ing mo e i id and dis inc i e
colo a ion pa e ns ha a e o en s able wi hin species. As
such, compa isons o male colo a ion ai s can p o ide
impo an diagnos ic clues o species delimi a ion. Th ee
ecen ly desc ibed species, T. mayaae, T. p e iosus, and
T. nujiang in Vi ido ipe a ha e also employed colo a ion
ea u es as key diagnos ic cha ac e s (Ra hee e al. 2022;
Liang e al. 2025; Xu e al. 2025). T ime esu us loong
sp. no . is likewise dis inguished by i s unique colo a ion
ai s. Al hough i was once misiden i ied as T. yunnanen-
sis, he wo species can be eadily dis inguished in males
by di e ences in eye colo a ion, as well as in he colo
pa e n o he en ola e al s ipe and do sal ail.
In addi ion, hemipenial mo phology se es as ano he
e ec i e diagnos ic ea u e. Al hough he hemipenes o
Vi ido ipe a species ha e been desc ibed in p e ious s ud-
ies (Pope 1935; Malho a and Tho pe 2004a; O lo e al.
2004; Dawson e al. 2008; Guo e al. 2009; Ra hee e al.
2022; Liang e al. 2025; Xu e al. 2025), mos accoun s
we e based on incomple ely e e ed o poo ly p ese ed
specimens, he eby limi ing hei diagnos ic alue. A com-
p ehensi e in e speci ic compa ison has emained lacking.
In his s udy, we compa ed he hemipenes o i e species
(Fig. 4) wi hin he subgenus Vi ido ipe a and ound ha
he p ima y di e ences in ol e he deg ee o bi u ca ion,
he ela i e posi ion o he sulcus spe ma icus bi u ca ion
poin , and he numbe o enla ged spines. While Guo e al.
(2009) desc ibed a ia ion in he zona ion o he spine and
calyx egions among se e al species, ou esul s indica e
ha hese ea u es a e, in ac , ema kably simila ac oss
axa. The p e iously epo ed disc epancies may esul om
me hodological di e ences in hemipenis p epa a ion, in-
cluding he use o in la ion ma e ials and p ese a ion s a e,
which can dis o he appa en shape (Mye s and Cadle
2003). Mo ing o wa d, hemipenial mo phology should be
documen ed ac oss a b oade ange o species wi hin Vi -
ido ipe a, and he e minology and p epa a ion echniques
should be s anda dized o ensu e consis en axonomic in-
e p e a ions. Taken oge he , bo h mo phological and mo-
lecula e idence consis en ly suppo he dis inc i eness o
T ime esu us loong sp. no . wi hin Vi ido ipe a.
The new species T ime esu us loong sp. no . ep esen s
a dis inc e olu iona y lineage wi hin he subgenus Vi ido i-
pe a. Phylogene ic analyses based on mi ochond ial DNA
(16S, cy b, and ND4 genes) eco e ed T. loong sp. no . as
sis e o T. c . loong om Huili Ci y, Sichuan P o ince, Chi-
na, wi h mode a e s a is ical suppo (SH = 83; UFB = 92;
PP = 0.97). Toge he , hey o m a clade wi h an uniden i ied
T ime esu us lineage om wes e n Sichuan P o ince, Chi-
na. This en i e assemblage occupies a pe iphe al posi ion
ela i e o he emaining ecognized species o Vi ido ipe a,
and T ime esu us loong sp. no . exhibi s subs an ial gene ic
di e gence om all cu en ly ecognized congene s, wi h
unco ec ed p-dis ances o a leas 6.7% (cy b) and 6.1%
(ND4). This le el o di e gence, combined wi h consis en
mo phological di e en ia ion, suppo s i s ecogni ion as a
alid species. Mo eo e , he specimens p e iously eco d-
ed as T. yunnanensis by Schmid (1927) and Pope (1935)
om “Yunnan- u” (now Kunming Ci y, Yunnan P o ince;
specimen MCZ 14671, a pa a ype o T. yunnanensis), hose
lis ed by Zhao e al. (1998) om Kunming (specimens KIZ
72001, KIZ 73008, and KIZ 85702), as well as specimen
NHMUK 1904.11.29.29 om “Yunnan- u” housed a he
Na u al His o y Museum, London, may likewise ep esen
T ime esu us loong sp. no . (Table 6). Howe e , due o he
zse.penso .ne
Xu, Y. e al.: A new T ime esu us species om Yunnan, China2290
lack o di ec examina ion, he colo a ion o hese speci-
mens om Kunming Ci y and i s su ounding a eas emains
unce ain. Fu he eassessmen is he e o e necessa y o de-
e mine whe he hey indeed ep esen T ime esu us loong
sp. no ., o whe he he wo species exhibi pa ially o e -
lapping dis ibu ions.
Liang e al. (2025) ecen ly synonymized T. gump ech-
i wi h T. yunnanensis on he basis o minimal gene ic
di e gence and o e lapping mo phological a ia ion.
Ne e heless, some specimens included in hei com-
pa a i e da ase appea o ha e been assigned o di e -
en axa. Fo ins ance, ZFMK 92790 and ZFMK 92791
om Phuoc Binh Na ional Pa k, Ninh Thuan P o ince
(cu en ly Khanh Hoa P o ince), sou he n Vie nam,
we e labeled as T. gump ech i bu seem mo phologically
consis en wi h T. ogeli (T.V. Nguyen, pe s. obs.). Like-
wise, CAS 241182 om Moenyin Township, Myi kyina
Dis ic , Kachin S a e, no he n Myanma , appea s e e -
able o T. popeio um (see Idiia ullina e al. 2024b), and
ZFMK 92793 om Chu Yang Sin Na ional Pa k, Dak Lak
P o ince, sou he n Vie nam, ini ially iden i ied as T. s e-
jnege i, also co esponds o T. ogeli. Such e o s may
ha e a ec ed hei axonomic in e p e a ion and highligh
he need o a ho ough e-examina ion o compa a i e
ma e ial used in hei s udy. Conside ing he ecen de-
sc ip ions o T. nujiang and T. p e iosus, i is e iden ha
sou hwes e n China ha bo s mul iple c yp ic lineages o
he subgenus Vi ido ipe a wi h es ic ed dis ibu ions.
The disco e y o T ime esu us loong sp. no . om Kun-
ming Ci y, Yunnan P o ince, u he suppo s he iew
ha T. yunnanensis sensu la o ep esen s a complex o
closely ela ed species ha wa an s comp ehensi e e-
ision using bo h genomic and mo phological da a. The
disco e y o T ime esu us loong sp. no . unde sco es he
axonomic complexi y o he T. yunnanensis g oup and
he impo ance o in eg a i e app oaches in esol ing
c yp ic di e si y among Asian g een pi ipe s.
Biogeog aphically, he occu ence o T ime esu us
loong sp. no . in he Anning Ci y, Kunming Ci y adds an
impo an elemen o ou unde s anding o Vi ido ipe a
di e si ica ion in Yunnan P o ince. This a ea ep esen s
a ansi ional zone be ween he Red Ri e d ainage and
he Yunnan-Guizhou Pla eau, cha ac e ized by dissec ed
mon ane e ain and b oad-lea ed o es s o coni e ous
o es s. The es ic ed ange o T ime esu us loong sp.
no . con as s wi h he mo e wes e nly species T. nujiang
and he Himalayan occu ence o T. p e iosus. Toge he ,
hese species e eal a clea geog aphic and ele a ional
g adien o di e si ica ion wi hin he Vi ido ipe a, like-
ly in luenced by complex o ogeny, clima ic oscilla ions,
and ecological isola ion. The na ow en ola e al s ipe,
ligh o ange ed ail, and ambe i is o T ime esu us loong
sp. no . may ep esen local adap i e ea u es associa ed
wi h i s mid-mon ane o es en i onmen . Wi h he dis-
co e y o T ime esu us loong sp. no ., he o al numbe
o T ime esu us species eco ded om Yunnan P o ince
now ises o nine, including wo species endemic o Yun-
nan, namely T ime esu us loong sp. no . and T. nujiang.
These endemics u he emphasize Yunnan’s excep ional
biogeog aphic he e ogenei y and i s ole as a co e di e -
si ica ion cen e o Asian g een pi ipe s.
F om a conse a ion pe spec i e, T ime esu us loong
sp. no . is cu en ly known only om i s ype locali y in
Qinglongxia, Anning Ci y, Kunming Ci y, Yunnan P o -
ince, China, a an ele a ion o app oxima ely 1,822 m asl.
Gi en i s highly es ic ed known dis ibu ion and he ab-
sence o popula ion da a, he species should be p o ision-
ally classi ied as Da a De icien (DD) unde IUCN Red
Lis c i e ia. Ta ge ed ield su eys a e u gen ly needed o
e alua e i s ac ual dis ibu ion, popula ion size, and eco-
logical equi emen s. Because g een pi ipe s a e medi-
cally impo an , accu a e axonomic ecogni ion o he T.
yunnanensis complex is also c i ical o clinical iden i i-
ca ion and enom managemen .
In conclusion, T ime esu us loong sp. no . ep esen s
ano he lineage wi hin he apidly di e si ying subge-
nus Vi ido ipe a adia ion o sou hwes e n China. The
disco e y o his species, oge he wi h T. nujiang and
T. p e iosus, highligh s Yunnan and adjacen Himalayan
egions as key cen e s o specia ion o Asian g een pi
ipe s. In eg a i e axonomic app oaches ha combine
mo phological, molecula , and ecological da a will be es-
sen ial o ully esol e he complex e olu iona y his o y
o his medically signi ican g oup. This inding u he
ein o ces he impo ance o sou hwes e n China as a
ho spo o pi ipe di e si ica ion and highligh s he need
o con inued in eg a i e su eys in mon ane ecosys ems.
Acknowledgemen s
We a e deeply g a e ul o D . Bo Cai (CIB, China) and
Pa ag Shinde (India) o p o iding aluable in o ma-
ion and pho og aphs o T ime esu us spp. We since e-
ly hank D . Jinlong Ren (CIB, China) o his in alu-
able assis ance in he p epa a ion and mo phological
desc ip ion o he hemipenes. T.V. Nguyen hanks D .
Pa ick Da id (MNHN, F ance) o sha ing axonomic
Table 6. Specimens om Kunming, Yunnan, China p e iously assigned o T ime esu us yunnanensis. Rema k: N/a = no a ailable.
Collec ion numbe Sex SVL (mm) TAL (mm) ASR MSR PSR VS SC SL IL Sou ce
KIZ 72001 ♂417 100 20 19 15 157 71 9/10 11/11 Zhao e al. (1998)
KIZ 73008 ♀457 99 19 19 15 150 57 10/11 12/11 Zhao e al. (1998)
KIZ 85702 ♀617 141 21 19 15 158 65 10/10 11/11 Zhao e al. (1998)
MCZ 14671 ♂N/a N/a 21 19 15 160 68 N/a N/a Schmid (1927); Pope (1935)
NHMUK 1904.11.29.29 ♀652 137 21 20 15 156 55 10/10 12/12 This s udy
Zoosys . E ol. 101 (4) 2025, 2267–2293
zse.penso .ne
2291
in o ma ion and help ul discussions ega ding T ime e-
su us species. N.A. Poya ko hanks S.S. Idiia ullina
(MSU, Russia) o help and assis ance. We also hank
Jus in M. Be ns ein (AMNH, USA) and he anonymous
e iewe o cons uc i e commen s on an ea lie e -
sion o he manusc ip . This esea ch was unded by
he Na ional Na u al Science Founda ion o China
[32301325], he Open P ojec o S a e Key Labo a o y
o Pla eau Ecology and Ag icul u e, Qinghai Uni e si-
y [2025-KF-02], and he Russian Science Founda ion
(RSF g an Nº 22-14-00037-P, o suppo ing he wo k
o D . Nikolay A. Poya ko ).
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