A new ock-dwelling gecko o he subgenus Japonigekko (Squama a,
Gekkonidae, Gekko) om no hwes e n Sichuan P o ince, China
Yuhao Xu1, Shun Ma2,3, Bo Cai2, Shuo Qi4, Tomoya Ma sukoji5, Nikolay A. Poya ko 6, Fanyue Sun1,
Shiyang Weng7, Tianxuan Gu8, Kaidi Tian9, Di Zhang9, Jianping Jiang2, 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 China-C oa ia Bel and Road Join Labo a o y on Biodi e si y and Ecosys em Se ices, Chengdu Ins i u e o Biology, Chinese Academy o
Sciences, Chengdu 610213, Sichuan, China
3 Uni e si y o Chinese Academy o Science, Beijing 100049, China
4 School o Li e Sciences / School o Ecology, Sun Ya -sen Uni e si y, Guangzhou 510275, Guangdong, China
5 G adua e School o Science and Li e Technology, Uni e si y o Tsukuba, Iba aki 305-8572, Japan
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 Ins i u e o Pla eau Biology o Xizang Au onomous Region, Lhasa 850008, Xizang, China
8 Long o U Ci y Communi y, Chongqing 401331, China
9 Yellow Ri e Wa e Resou ces P o ec ion Ins i u e, Zhengzhou 450004, Henan, China
h ps://zoobank.o g/CCCD4D14-5256-40AC-B382-A16E665FD39A
Co esponding au ho s: Li ang Peng ([email p o ec ed]); Jianping Jiang ([email p o ec ed])
Academic edi o : Jus in Be ns ein ♦
Recei ed
17 Oc obe 2025 ♦
Accep ed
3 No embe 2025 ♦
Published
18 No embe 2025
Abs ac
We desc ibe a new ock-dwelling species o he genus Gekko (subgenus Japonigekko) om Heishui Coun y, Aba Tibe an and Qiang
Au onomous P e ec u e, Sichuan P o ince, China, based on bo h mo phological and molecula (1,574 bp om he mi ochond ial 16S
RNA and ND2 genes) e idence. Mo phologically, Gekko essela us sp. no . is cha ac e ized by i s mode a e body size; wo ( a ely
one) enla ged pos men als; la ened do sal ube cles ex ending om he pos e io head h ough he neck o he an e io po ion o
he ail, a anged in 12–15 ows a midbody; 130–157 en al scales be ween he men al and he cloacal sli ; 98–106 midbody scale
ows; 31–39 en al scale ows; subdigi al lamellae numbe ing 7–10 on inge I, 9–12 on inge IV, 7–9 on oe I, and 9–12 on oe IV;
absence o webbing; 6–8 p ecloacal po es in males and absence in emales; one ( a ely wo) pos cloacal ube cle on each side; and
a dis inc i e do sal colo a ion. Phylogene ically, he new species o ms a dis inc clade wi hin he subgenus Japonigekko, showing
unco ec ed 16S sequence di e gences o a leas 8.2% om i s closes ela i e, G. liboensis, and a leas 8.9% om all o he conge-
ne s, as well as ND2 di e gences o a leas 14.3% om G. engshanensis and a leas 14.9% om o he species in he subgenus. The
disco e y o his new species aises he numbe o Japonigekko species eco ded in China o 26 and in Sichuan P o ince o se en.
Key Wo ds
Gekko essela us sp. no ., molecula phylogeny, mo phological cha ac e s, axonomy
In oduc ion
The genus Gekko Lau en i, 1768, ep esen s a axonomical-
ly di e se g oup wi hin he amily Gekkonidae G ay, 1825.
Cu en ly, 93 alid species a e ecognized wi hin he genus,
and i s membe s a e widely dis ibu ed ac oss eas e n and
sou heas e n Asia, as well as pa s o no hwes e n Oceania
and Melanesia (Ma e al. 2024; Cao e al. 2025; Huang e
al. 2025; Pauwels e al. 2025; Ue z e al. 2025). Due o he
high mo phological simila i y among species, which com-
plica es diagnosis and hampe s he iden i ica ion o c yp ic
lineages, he axonomy o he genus Gekko emained un e-
Zoosys . E ol. 101 (4) 2025, 2221–2242|DOI 10.3897/zse.101.175246
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 Gekko species
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sol ed o long pe iod o ime (Zhao and Adle 1993; Zhao
e al. 1999; Kluge 1993, 2001; Rösle e al. 2005, 2011). In
ecen yea s, howe e , he use o in eg a i e axonomic ap-
p oaches ha combine de ailed mo phological assessmen s
wi h molecula phylogene ic analyses has cla i ied many
p e iously ambiguous species bounda ies and g adually e-
ealed he ue ex en o di e si y wi hin he genus (B own
e al. 2009; Rösle e al. 2011; Luu e al. 2015; Wood e al.
2020; Lyu e al. 2021; Xia e al. 2023; Huang e al. 2025).
Acco ding o he classi ica ion p oposed by Wood e al.
(2020), he genus is cu en ly di ided in o se en subgen-
e a: Gekko s. s ., Japonigekko Wood, Guo, T a e s, Su,
Olson, Baue , G isme , Sile , Moyle, Ande sen & B own,
P ychozoon Kuhl & Van Hassel , Rhacogekko Wood, Guo,
T a e s, Su, Olson, Baue , G isme , Sile , Moyle, Ande sen
& B own, Loma odac ylus an de Hoe en, Balawangek-
ko Wood, Guo, T a e s, Su, Olson, Baue , G isme , Sile ,
Moyle, Ande sen & B own, and A chipelagekko Wood,
Guo, T a e s, Su, Olson, Baue , G isme , Sile , Moyle,
Ande sen & B own. Among hese, Japonigekko is he
mos di e se g oup, comp ising 38 species, 25 o which
occu in China (Cao e al. 2025; Huang e al. 2025; Zhou e
al. 2025b). Membe s o Japonigekko a e cha ac e ized by
he ollowing diagnos ic ea u es: mode a e body size; na-
es usually in con ac wi h he os al; 2–4 nasals (includ-
ing naso os als, sup anasals, and pos nasals); 0–21 ows
o do sal ube cles; 0–32 p ecloacal po es; 1–4 pos cloacal
ube cles; and la e al olds lacking ube cles (Wood e al.
2020; Hou e al. 2021; Ma e al. 2024; Wang e al. 2024;
Zhou e al. 2025a; Ue z e al. 2025).
Loca ed in wes e n China, Sichuan P o ince encom-
passes se e al majo geomo phological egions, including
he Qinghai–Xizang Pla eau, he Hengduan Moun ains, he
Yunnan–Guizhou Pla eau, he Qinba Moun ains, and he
Sichuan Basin. I s complex geological s uc u e and highly
di e se opog aphy ha e gi en ise o a complex and a -
ied clima e. Toge he wi h signi ican geog aphical ba ie s
and a mosaic o mic ohabi a s, hese condi ions ha e d i -
en p onounced egional di e en ia ion o species. Con-
sequen ly, Sichuan suppo s excep ionally high le els o
biodi e si y and is ecognized as one o China’s mos im-
po an ese oi s o gene ic esou ces. Du ing a ield su -
ey conduc ed in he summe o 2025, we collec ed a se ies
o Gekko (Japonigekko) specimens om a d y-ho alley
in Heishui Coun y, Aba Tibe an and Qiang Au onomous
P e ec u e, Sichuan P o ince, China (Fig. 1). Howe e , de-
ailed mo phological compa isons and mi ochond ial DNA
analyses e ealed ha hese indi iduals di e ma kedly
om all known Gekko species. Based on his e idence, we
he ein desc ibe his popula ion as a new species.
Ma e ial and me hods
Sampling
Field su eys we e conduc ed in he d y-ho alley o
Heishui Coun y, Sichuan P o ince, China, in he sum-
me o 2025. 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 Technology Co., L d.,
Shenzhen, China). A e being pho og aphed, all newly
collec ed specimens we e humanely eu hanized using a
le hal injec ion o 0.7% icaine me hanesul ona e (MS-
222) solu ion, hen ixed in 95% e hanol o one day and
ans e ed o 75% e hanol o long- e m p ese a ion.
F esh li e issue was ex ac ed, immedia ely p ese ed
in 95% e hanol, and s o ed a –20 °C. The specimens
we e deposi ed a Qinghai Uni e si y (QHU), he Muse-
um o Biology, Sun Ya -sen Uni e si y (SYS), and he
Chengdu Ins i u e o Biology (CIB), Chinese Academy o
Sciences. All sampling p ocedu es in ol ing li e geckos
complied wi h 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).
Molecula phylogeny
Since all specimens we e collec ed om he same locali y
and showed consis en mo phological ai s, we andom-
ly selec ed 10 indi iduals o DNA sequencing. To al ge-
nomic DNA was ex ac ed om p ese ed li e issue us-
ing he QIAamp DNA Mini Ki (QIAGEN, Changsheng
Bio echnology Co., L d.). Two mi ochond ial gene ag-
men s, 16S ibosomal RNA (16S) and NADH dehyd o-
genase subuni 2 (ND2), we e ampli ied ia polyme ase
chain eac ion (PCR) ollowing he p o ocols ou lined
by Wang e al. (2024). The ollowing p ime pai s we e
used: L3975 (5’-CGCCTGTTTACCAAAAACAT-3’)
and H4551 (5’-CCGGTCTGAACTCAGATCACGT-3’)
o 16S (Simon e al. 1994), and Me -3L (5’-ATAC-
CCCGACAATGTTGG-3’) and Ala-1H (5’-GCCT-
TAGCTTAATTAAAGTG-3’) o ND2 (Jonniaux and
Kumazawa 2008). The PCR p oduc s we e sequenced
by Shanghai Map Bio ech Co., L d. Raw sequences we e
assembled using SeqMan in he DNASTAR so wa e
package (Bu land 2000). Newly gene a ed sequences
ha e been submi ed o he DDBJ; accession numbe s a e
lis ed in Table 1.
Apa om he 20 newly ob ained sequences, we
included 80 addi ional sequences ep esen ing 48 indi-
iduals om 28 species o he subgenus Japonigekko
(Table 1). Gekko gecko (Linnaeus, 1758) and G. ee esii
(G ay, 1831) we e selec ed as ou g oups, ollowing he
app oach o Wang e al. (2024). A conca ena ed da ase
o he wo gene agmen s (16S + ND2, 1,574 bp in o-
al) was aligned using MUSCLE (Edga 2004) in MEGA
X (Kuma e al. 2018). Unco ec ed pai wise dis ances
(p-dis ances) be ween closely ela ed species we e also
calcula ed in MEGA X. The Maximum Likelihood (ML)
analysis was conduc ed in IQ-TREE 1.6.12 (Nguyen
e al. 2015), pa i ioned by gene, wi h bo h 16S and
ND2 agmen s independen ly selec ed unde he bes -
i model GTR+F+I+G4 by ModelFinde (Kalyaana-
moo hy e al. 2017) implemen ed in PhyloSui e 1.2.3
(Zhang e al. 2020), acco ding o he Bayesian In o -
ma ion C i e ion (BIC) (Kalyaanamoo hy e al. 2017).
Zoosys . E ol. 101 (4) 2025, 2221–2242
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Table 1. NCBI/DDBJ accession numbe s, locali ies, and ouche in o ma ion o all specimens used in his s udy. * indica es DDBJ
accession numbe .
No. Species Locali ies Vouche ID. 16S ND2 Re e ence
1G. essela us sp. no . Heishui, Sichuan, China CIB 119371 LC899308*LC899318* This s udy
2G. essela us sp. no . Heishui, Sichuan, China CIB 119372 LC899309*LC899319* This s udy
3G. essela us sp. no . Heishui, Sichuan, China CIB 119373 LC899310*LC899320* This s udy
4G. essela us sp. no . Heishui, Sichuan, China CIB 119374 LC899311*LC899321* This s udy
5G. essela us sp. no . Heishui, Sichuan, China CIB 119375 LC899312*LC899322* This s udy
6G. essela us sp. no . Heishui, Sichuan, China QHU R2025022 LC899313*LC899323* This s udy
7G. essela us sp. no . Heishui, Sichuan, China QHU R2025023 LC899314*LC899324* This s udy
8G. essela us sp. no . Heishui, Sichuan, China QHU R2025024 LC899315*LC899325* This s udy
9G. essela us sp. no . Heishui, Sichuan, China QHU R2025025 LC899316*LC899326* This s udy
10 G. essela us sp. no . Heishui, Sichuan, China QHU R2025026 LC899317*LC899327* This s udy
11 G. adle i Jingxi, Guangxi, China SYS 001400 MW451654 OR902178 Lyu e al. (2021); Wang e al.
(2024)
12 G. adle i Cao Bang, Vie am IEBR A.2012.24 KC700623 – Nguyen e al. (2013)
13 G. alpinus Ba ang, Sichuan, China CIB 121663 PQ255983 PQ303501 Ma e al. (2024)
14 G. alpinus Mangkang, Xizang, China CIB 121656 PQ255976 PQ303494 Ma e al. (2024)
15 G. au i e ucosus Yuncheng, Shanxi, China NNU Z20050801.004 – JN019062 Rösle e al. (2011)
16 G. bonkowskii Khammouane, Laos VFU R.2014.10 – KT266818 Luu e al. 2015
17 G. chinensis Hong Kong, China SYS 001211 MW451644 OR902183 Lyu e al. (2021); Wang e al.
(2024)
18 G. chinensis Shenzhen, Guangdong,
China
SYS 001085 MW451632 OR902184 Lyu e al. (2021); Wang e al.
(2024)
19 G. cib Hejiang, Sichuan, China SYS 001489 MW451655 OR902165 Lyu e al. (2021); Wang e al.
(2024)
20 G. cib Emeishan, Sichuan, China SYS 000708 MW451629 OR902166 Lyu e al. (2021); Wang e al.
(2024)
21 G. engshanensis Fengshan, Guangxi, China NHMG 240713 PV652773 PV657377 Huang e al. (2025)
22 G. engshanensis Fengshan, Guangxi, China NHMG 240714 PV652774 PV657378 Huang e al. (2025)
23 G. hokouensis Wuyishan, Fujian, China SYS 001290 MW451647 OR902173 Lyu e al. (2021); Wang e al.
(2024)
24 G. hokouensis Nanchang, Jiangxi, China SYS 001311 MW451648 OR902172 Lyu e al. (2021); Wang e al.
(2024)
25 G. ichangensis Yichang, Hubei, China SWU 0011602 PQ587503 – Cao e al. (2025)
26 G. ichangensis Yichang, Hubei, China SWU 0011603 PQ587504 – Cao e al. (2025)
27 G. japonicus Wuyishan, Fujian, China SYS 000672 MW451628 OR902176 Lyu e al. (2021); Wang e al.
(2024)
28 G. japonicus Lushan, Jiangxi, China SYS 001317 MW451649 OR902177 Lyu e al. (2021); Wang e al.
(2024)
29 G. jinjiangensis Deqin, Yunan, China CIB 5334220088 PQ255987 MT449432 Hou e al. (2021); Ma e al.
(2024)
30 G. jinjiangensis De ong, Sichuan, China CIB 5133380017 – MT449437 Hou e al. (2021)
31 G. kaiyai Xinxian, Henan, China AHUXXBH01 OQ780318 – Zhang e al. (2023)
32 G. kaiyai Shangcheng, Henan, China AHUJGTBH01 OQ780322 – Zhang e al. (2023)
33 G. khunkhamensis Khammouane, Laos VNUF R.2021.23 – OL416111 Si hi ong e al. (2021)
34 G. kwangsiensis Wuming, Guangxi, China SYS 001195 MW451642 OR902175 Lyu e al. (2021); Wang e al.
(2024)
35 G. kwangsiensis Wuming, Guangxi, China SYS 001194 MW451641 OR902174 Lyu e al. (2021); Wang e al.
(2024)
36 G. liboensis Libo, Guizhou, China SYS 002876 PV652777 PV657381 Huang e al. (2025)
37 G. liboensis Libo, Guizhou, China SYS 002877 PV652778 PV657382 Huang e al. (2025)
38 G. melli Dongguan, Guangdong,
China
SYS 001742 MW451661 OR902169 Lyu e al. (2021); Wang e al.
(2024)
39 G. melli Puning, Guangdong, China SYS 001702 MW451660 OR902170 Lyu e al. (2021); Wang e al.
(2024)
40 G. nadenensis Khammouane, Laos ZFMK 98741 – KY421618 Luu e al. (2017)
41 G. palma us Zhaoqing, Guangdong, China SYS 002797 OR903156 OR902179 Wang e al. (2024)
42 G. palma us Napo, Guangxi, China SYS 001185 MW451637 OR902182 Lyu e al. (2021); Wang e al.
(2024)
43 G. pauci ube cula us Baise, Guangxi, China SYS 002806 OR903154 OR902163 Wang e al. (2024)
44 G. pauci ube cula us Baise, Guangxi, China SYS 002806 OR903155 OR902164 Wang e al. (2024)
45 G. p ep Changjiang, Hainan, China HN2024R038 PV368865 PV389916 Zhou e al. (2025a)
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Xu, Y. e al.: A new Gekko species
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No. Species Locali ies Vouche ID. 16S ND2 Re e ence
46 G. p ep Changjiang, Hainan, China HN2024R039 PV368866 PV389915 Zhou e al. (2025a)
47 G. scien iad en u a Quang Binh, Vie nam IEBR A.2014.7 – KP205392 Luu e al. (2014)
48 G. sengchan ha ongi Khammouane, Laos VFU R2014.14 – KT266816 Luu e al. (2015)
49 G. similignum Wuzhishan, Hainan, China SYS 001597 MW451658 OR902185 Lyu e al. (2021); Wang e al.
(2024)
50 G. similignum Wuzhishan, Hainan, China SYS 001598 MW451659 OR902186 Lyu e al. (2021); Wang e al.
(2024)
51 G. scab idus Yanbian, Sichuan, China CIB YN201909199 PQ255992 MT449429 Hou e al. (2021); Ma e al.
(2024)
52 G. scab idus Yanbian, Sichuan, China CIB YN201909200 – MT449430 Hou e al. (2021)
53 G. subpalma us Fenghua, Zhejiang, China SYS 001762 MW451662 OR902167 Lyu e al. (2021); Wang e al.
(2024)
54 G. subpalma us Fenghua, Zhejiang, China SYS 001767 MW451663 OR902168 Lyu e al. (2021); Wang e al.
(2024)
55 G. swinhonis Zunhua, Hebei, China SYS 001814 MW451666 OR902171 Lyu e al. (2021); Wang e al.
(2024)
56 G. swinhonis Zunhua, Hebei, China SYS 001815 MW451667 – Lyu e al. (2021)
57 G. hakhekensis Thakhek, Khammouane,
Laos
IEBR A.2014.6 – KP205396 Luu e al. (2014)
58 G. uongi Khanh Hoa, Vie nam IEBR A.2011.1 – KP205398 Luu e al. (2014)
Ou g oup
59 G. gecko Nanning, Guangxi, China N/A AY282753 AY282753 Zhou e al. (2006)
60 G. ee esii M . Yinping, Guangdong,
China
SYS 000796 MW451630 OR902187 Lyu e al. (2021); Wang e al.
(2024)
Nodes we e ega ded as well suppo ed when ul a as
boo s ap alues (UFB) ≥ 95% and SH-like app oxima e
likelihood a io es alues (SH-aLRT) ≥ 80% (S ephane
e al. 2010; Hoang e al. 2018). The phylogene ic ee
was isualized using FigT ee 1.4.4 (Rambau 2018).
Mo phological examina ion
Compa isons wi h o he congene s o he subgenus Ja-
ponigekko we e based on a ailable li e a u e: S ejnege
(1907); Zhou e al. (1982); Song (1985); O a e al. (1995);
Zhao e al. (1999); Go is and Maeda (2004); Rösle e al.
(2005, 2011); Toda e al. (2008); Zhou and Wang (2008);
Phung and Ziegle (2011); Nguyen e al. (2013); Luu e al.
(2014, 2015, 2017); Jono e al. (2015); Ngo e al. (2015);
Yang (2015); Lin and Yao (2016); Hou e al. (2021); Lyu
e al. (2021); Si hi ong e al. (2021); Zhang e al. (2023);
Ma e al. (2024); Wang e al. (2024); Cao e al. (2025);
Huang e al. (2025); Zhou e al. (2025a, b).
Mo phological desc ip ions ollowed he me hodology
ou lined in Lyu e al. (2021) and G isme e al. (2022), wi h
ce ain abb e ia ions e ised acco ding o he s anda ds se
ollowing he abb e ia ions lis in Da ko e al. (2022). Th ee
mo phome ic cha ac e s we e measu ed wi h Deli digi al
calipe s (No. 90150B) o he nea es 0.1 mm: SVL = snou –
en leng h, measu ed om he ip o he snou o he pos e-
io edge o he en ; TAL = ail leng h, measu ed om he
pos e io ma gin o he en o he ip o he ail; AGD = ax-
illa–g oin dis ance, de ined as he dis ance be ween he pos-
e io edge o he o elimb inse ion and he an e io edge o
he hindlimb inse ion, wi h bo h limbs inse ed pe pendic-
ula ly o he body wall. All o he measu emen s we e aken
using Mi u oyo digi al calipe s (CD-15AX) o he nea es
0.01 mm unde a Leica s e eomic oscope (EZ4): HL = head
leng h, measu ed om he ip o he snou o he pos e io
ma gin o he ea opening; HW = head wid h, measu ed a
he angle o he jaws; HH = head heigh , measu ed om
he op o he head pos e io o he eyes o he bo om o
he lowe jaw; ESD = eye- o-snou dis ance/snou h leng h,
measu ed om he ip o he snou o he an e io co ne o
he eye; ED = eye diame e , measu ed as he g ea es dis-
ance om he an e io o he pos e io co ne o he eye; EL
= maximum ea opening diame e ; RW = maximum os al
wid h; RH = maximum os al heigh ; MW = maximum
men al wid h; and ML = maximum men al leng h.
Scala ion ea u es and hei abb e ia ions we e as ol-
lows: NS = numbe o nasals, including naso os als, su-
p anasals, and pos nasals; INS = in e nasals, numbe o
scales be ween sup anasals, in con ac wi h he os al; SL
= sup alabials; IL = in alabials; IOS = in e o bi als, num-
be o scales in a line be ween he an e io co ne s o he
eyes; PO = p eo bi als, numbe o scales in a line om he
nos il o he an e io ma gin o he eye; PM = pos men-
als, numbe o scales bo de ing he men al; GP = gula s
bo de ing he pos men als; MBSR = midbody scale ows;
VS = en al scale ows a midbody; SMC = scales in a
line om he men al o he on o he cloacal sli ; DTR
= do sal ube cle ows a midbody; GSDT = g anules su -
ounding do sal ube cles, coun ed by andomly selec ing
10 ube cles on he unk and eco ding he numbe o
g anules su ounding each; LF1 = numbe o he subdigi-
al lamellae benea h he en i e i s inge ; LF4 = numbe
o he subdigi al lamellae benea h he en i e ou h inge ;
LT1 = numbe o he subdigi al lamellae benea h he en-
i e i s oe; LT4 = numbe o he subdigi al lamellae be-
nea h he en i e ou h oe; PP = numbe o he p ecloacal
po es; PAT = numbe o he pos cloacal ube cles.
Zoosys . E ol. 101 (4) 2025, 2221–2242
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Figu e 1. Map showing he locali y o Gekko essela us sp. no . The s a indica es he ype locali y, and ci cles deno e o he known
dis ibu ion si es.
In addi ion, we eco ded he p esence o absence o
webbing (Web); he p esence o ube cles on he o e-
limbs (Fo e ube cles), hindlimbs (Hind ube cles), and
ail (Tail ube cles); he p esence o enla ged subcaudal
scales; and he ypes o do sal colo a ion pa e ns.
O he abb e ia ions a e as ollows: Is.: Island; M .:
Moun ain; M s.: Moun ains; NP: Na ional Pa k; NR: Na-
u e Rese e; WS: Wildli e Sanc ua y; asl.: abo e sea le el.
Resul s
The leng hs o he 16S and ND2 agmen s in he inal
alignmen comp ised 563 base pai s (bp) and 1,011 bp,
espec i ely. In he ML analysis (Fig. 2), all Gekko (Japo-
nigekko) samples included in his s udy o med a s ong-
ly suppo ed monophyle ic lineage (SH = 100 / UFB =
100). Howe e , ela ionships among species wi hin he
subgenus emain incomple ely esol ed due o low nodal
suppo a in e speci ic le els. The newly collec ed spec-
imens om Heishui Coun y, Sichuan P o ince, China,
cons i u ed a well-suppo ed monophyle ic lineage wi h-
in Japonigekko (SH = 100 / UFB = 100), which was e-
co e ed as sis e o a clade comp ising G. engshanensis
Huang, Wang, Qi, Song, Huang, Wang & Mo, 2025, G. li-
boensis Zhou, Liu & Li, 1982, and G. kwangsiensis Yang,
2015, al hough wi h ela i ely low suppo (SH = 80 /
UFB = 65). Finally, hese ou axa, along wi h G. pau-
ci ube cula us Wang, Qi, Zhou & Wang, 2024, o med
a well-suppo ed clade (SH = 100 / UFB = 100), whose
membe s exhibi specialized ecological adap a ions o
ocky en i onmen s.
Unco ec ed p-dis ances o he 16S and ND2 gene
agmen s a e summa ized in Tables 2 and 3, espec i e-
ly. In e speci ic gene ic dis ances wi hin Japonigekko
anged om 3.5% be ween G. chinensis (G ay, 1842) and
G. similignum Smi h, 1923 o 16.8% be ween G. adle i
Nguyen, Wang, Yang, Lehmann, Le, Ziegle & Bonkow-
ski, 2013 and G. swinhonis Gün he , 1864 o he 16S
gene (Table 2); and om 5.4% be ween G. chinensis and
G. similignum o 26.7% be ween G. p ep Zhou, Chen,
Liu, Li, Qi, Li & Peng, 2025 and G. swinhonis Gün he ,
1864 o he ND2 gene (Table 3). The Heishui popula ion
displayed subs an ial gene ic di e gence om all conge-
ne s, anging om 8.2% ( s. G. liboensis) o 16.0% ( s.
G. swinhonis) o he 16S gene and om 14.3% ( s. G.
engshanensis) o 25.2% ( s. G. melli (Vog , 1922)) o
he ND2 gene. In ligh o he well-suppo ed monophyly,
subs an ial gene ic di e gence, and dis inc mo pholog-
ical cha ac e s ha consis en ly di e en ia e he Gekko
popula ion om Sichuan P o ince om all known con-
gene s (see Compa isons below), we he ein desc ibe he
Gekko sp. om Heishui Coun y, Sichuan P o ince, Chi-
na, as a new species.
zse.penso .ne
Xu, Y. e al.: A new Gekko species
2226
Table 2. Unco ec ed p-dis ances (%) o he 16S gene among species o Japonigekko used in his s udy.
NO Species 1 2 3 4 5 6 7 8 9 10 11
1G. essela us sp. no . 0.0–0.8
2G. adle i 13.6–14.2 0.7
3G. alpinus 10.7–11.4 13.8–14.8 0.5
4G. chinensis 12.8–13.5 5.6–5.8 11.8–12.6 0.2
5G. cib 11.9–12.6 14.9 9.4–9.5 13.5–13.7 0.0
6G. engshanensis 9.1–9.7 14.6–15.5 11.2–11.9 14.6–14.9 13.7–14.0 0.6
7G. hokouensis 11.3–11.9 14.2–14.4 12.0–13.1 12.8–13.2 13.2–13.4 12.6–13.1 0.4
8G. ichangensis 12.1–13.0 15.4 9.6–10.5 15.1–15.3 12.4–12.5 14.8–15.0 14.0–14.3 0.0
9G. japonicus 13.4–13.8 15.1–15.6 10.5–11.4 14.4–14.8 13.6–14.0 14.3–14.6 13.7–14.1 7.7–7.8 0.2
10 G. jinjiangensis 9.5–9.8 13.0–13.4 4.0–4.4 11.3–11.5 10.2 12.0–12.2 10.0–10.2 9.9–10.0 10.8 –
11 G. kaiyai 12.7–13.6 14.1–15.4 12.4–13.5 13.1–14.2 12.6–13.6 13.9–14.0 8.8–9.5 13.7–14.9 14.2–15.2 10.9–11.2 1.2
12 G. kwangsiensis 10.2–10.6 14.7 11.5–12.5 13.4–13.7 14.9–15.1 10.3–10.7 13.2–13.7 15.3–15.5 15.3–15.6 11.4–11.5 13.2–13.5
13 G. liboensis 8.2–9.2 14.2–15.2 11.3–12.4 13.8–14.2 13.3–13.5 9.0–9.4 14.6–14.7 14.4–14.6 13.3–13.7 12.4–12.6 14.3–15.2
14 G. melli 11.0–11.8 13.6–14.2 8.6–8.9 13.1–13.5 7.8–8.0 12.6–13.3 13.4–13.6 12.6 14.5–14.7 9.6–9.8 13.4–14.0
15 G. palma us 13.1–13.5 2.5–2.9 13.2–14.0 3.8–4.4 13.7–14.1 15.3–15.6 12.6–13.2 15.4–15.8 13.7–14.1 12.3–12.7 13.6–14.0
16 G. pauci ube cula us 8.9–9.3 14.0 10.5–11.2 12.5–12.7 12.9 10.7–10.9 11.7 16.0–16.1 15.4–15.5 10.2 12.9
17 G. p ep 13.0–13.8 6.0–6.8 12.0–12.4 3.5–3.7 15.0–15.1 15.6–15.7 14.6–14.8 15.3–15.5 14.6–14.8 11.9–12.0 13.1–13.8
18 G. similignum 12.6–13.2 6.4 12.2–13.0 1.9–2.0 15.2 15.4–15.5 13.4–13.6 16.0–16.1 14.8–15.0 11.3 13.9–14.8
19 G. scab idus 11.0–11.2 13.5–13.9 4.7–4.9 12.2–12.4 9.7 11.5 11.1–11.3 11.5 11.1 5.4 11.2–11.5
20 G. subpalma us 14.3–14.9 16.0 9.8–9.9 15.0–15.1 9.6 13.9–14.4 14.9–15.3 14.0 14.8–15.0 12.0 14.6–15.0
21 G. swinhonis 15.8–16.0 16.2–16.8 13.0–14.4 15.0–15.4 12.4–13.0 16.1–16.8 14.3–15.1 14.9–15.5 15.7–16.1 14.1–14.5 15.4–16.4
NO. Species 12 13 14 15 16 17 18 19 20 21
12 G. kwangsiensis 0.4
13 G. liboensis 11.9–12.1 0.2
14 G. melli 14.4–14.8 13.5–13.9 0.9
15 G. palma us 13.5–13.7 14.1–14.3 12.4–13.0 0.4
16 G. pauci ube cula us 12.0–12.3 9.6–9.8 13.1–13.3 12.0–12.2 0.0
17 G. p ep 13.6–13.8 15.0–15.2 13.7–13.9 4.2–4.6 14.0 0.2
18 G. similignum 14.2–14.3 14.2–14.4 13.7–14.3 4.6–5.0 14.0 4.3 0.0
19 G. scab idus 12.1–12.4 12.2 10.0–10.2 12.6–13.1 10.6 12.6–12.7 12.4 –
20 G. subpalma us 13.4–13.6 15.5–15.7 8.8–9.0 14.1–14.5 15.1 14.8 15.7 12.0 –
21 G. swinhonis 16.0–16.3 15.1–15.5 14.3–14.8 14.6–15.0 15.8–16.2 15.4–16.0 15.4–15.9 14.9–15.1 13.9–14.4 1.1
Zoosys . E ol. 101 (4) 2025, 2221–2242
zse.penso .ne
2227
Table 3. Unco ec ed p-dis ances (%) o he ND2 gene among species o Japonigekko used in his s udy.
NO. Species 1 2 3 4 5 6 7 8 9 10 11 12 13
1G. essela us sp. no . 0.0–1.0
2G. adle i 23.8–25.0 –
3G. alpinus 19.6–20.9 23.4–23.6 1.6
4G. au i e ucosus 19.6–20.5 24.8 21.5–22.2 –
5G. bonkowskii 20.3–20.7 24.6 19.0–19.4 21.5 –
6G. chinensis 22.5–24.5 14.4–14.6 21.9–22.7 24.3–24.5 21.2–21.4 0.4
7G. cib 22.1–22.7 26.1 22.2–22.8 20.7 19.4 24.0–24.3 0.0
8G. engshanensis 14.3–14.9 23.4 20.0–20.3 22.1 19.6 23.7 21.7 0.0
9G. hokouensis 19.7–21.2 24.8–25.2 19.9–21.6 21.0–21.5 19.0–20.3 24.2–25.0 23.0–23.6 20.6–20.9 1.3
10 G. japonicus 20.3–21.8 26.1 19.6–20.6 20.9 19.6 24.9–25.2 22.9 21.4 22.5–22.7 0.0
11 G. jinjiangensis 16.9–17.2 20.7–21.3 7.1–8.4 19.9–20.5 17.9–19.0 21.1–22.4 20.6–20.7 17.0–18.3 18.0–20.4 16.9–17.2 2.4
12 G. khunkhamensis 22.4–22.8 26.0 21.3–21.7 24.3 15.2 24.6–24.8 21.1 22.2 23.8–24.2 22.6 20.9–21.7 –
13 G. kwangsiensis 16.6–17.4 24.1–24.2 21.8–22.7 22.4–22.5 20.9–21.0 23.0–23.3 20.7–20.9 15.9–16.1 21.4–21.8 22.7–22.8 19.1–19.4 21.5–21.7 0.4
14 G. liboensis 16.4–16.7 23.4–23.5 20.8–21.7 22.1–22.2 19.4–19.6 24.7–24.9 21.4–21.5 17.0–17.1 19.6–20.2 22.8–22.9 19.3–20.0 21.5–21.7 18.1–18.4
15 G. melli 23.4–25.2 24.5–25.1 21.9–23.0 22.6–23.5 21.2–21.8 24.7–25.2 18.9–19.0 24.1–24.3 23.6–24.6 23.6–23.8 19.3–21.1 23.4 23.3–23.8
16 G. nadenensis 21.6–22.0 23.3 18.6–19.4 20.2 6.9 21.2–21.8 20.3 20.5 21.2–22.2 19.9 16.9–18.1 13.8 20.5–20.7
17 G. palma us 22.3–24.2 6.5 22.4–22.6 23.6–23.8 23.1–23.6 14.3–14.4 25.9–26.1 23.1–23.2 23.7–24.4 24.9–25.0 20.7–21.1 25.6–26.2 23.4–23.5
18 G. pauci ube cula us 14.9–16.6 25.3–25.4 19.4–20.4 21.2–21.3 19.2 25.4–25.5 21.0–21.1 17.4–17.5 21.0–21.8 21.7–21.8 17.2–17.6 21.5 19.0–19.3
19 G. p ep 23.1–24.7 15.9–16.0 24.6–25.6 26.1–26.3 21.6–22.2 7.4–8.0 24.8–24.9 25.0 25.6–26.0 26.4–26.7 22.6–23.5 25.4–25.6 24.2–24.4
20 G. scien iad en u a 20.1–20.5 24.4 18.4–18.8 20.4 13.8 22.5 20.5 19.6 21.4–21.6 21.0 17.7 14.8 20.9
21 G. sengchan ha ongi 19.4 23.8 20.0–20.2 20.0 14.2 22.2–22.7 21.6 20.1 22.0–22.9 20.9 19.0–19.2 15.7 21.4
22 G. similignum 22.5–24.0 15.0–15.1 22.6–23.3 25.3–25.4 22.9–23.1 5.4–5.5 25.3–25.4 24.0–24.1 24.6–25.1 25.0–25.1 21.3–22.2 24.8–25.0 23.5–23.6
23 G. scab idus 17.4–17.8 18.3–19.1 11.5–12.6 19.2–19.7 19.9–20.3 20.2–20.7 21.1–21.7 16.7 17.0–17.4 17.2–17.6 9.6–11.5 21.7–22.6 20.2–20.9
24 G. subpalma us 21.3–22.4 25.0 21.7–22.2 21.6 20.5 24.1–24.4 17.9 21.7 23.3–23.4 23.1 19.1–20.2 22.6 22.2
25 G. swinhonis 20.2–20.9 25.1 22.6–22.7 20.7 21.2 25.7–25.8 21.9 20.7 22.0–22.5 21.9 20.4 23.6 21.8–21.9
26 G. hakhekensis 20.9–21.2 21.8 19.0–19.6 20.6 6.7 19.9–20.5 20.3 19.4 20.5–20.9 19.0 18.1–19.2 15.9 20.1–20.3
27 G. uongi 22.9 20.5 20.2–20.3 24.3 22.0 20.9–21.2 21.8 23.6 22.3 21.8 19.6–19.9 22.2 22.7–22.9
NO. Species 14 15 16 17 18 19 20 21 22 23 24 25 26
14 G. liboensis 0.1
15 G. melli 23.9–24.5 2.7
16 G. nadenensis 18.8–19.0 20.1–21.4 –
17 G. palma us 23.2–23.4 25.2–25.9 22.5–23.2 1.2
18 G. pauci ube cula us 17.8–18.0 25.0–25.3 18.6 24.4–24.5 0.1
19 G. p ep 26.2–26.3 26.4–27.0 22.3–22.9 15.2–15.4 26.4–26.5 0.4
20 G. scien iad en u a 20.3–20.5 21.2–21.6 13.4 22.9–23.5 18.6 23.8–24.4 –
21 G. sengchan ha ongi 19.7–19.9 21.8–22.5 11.9 23.5 19.0 22.7–23.3 10.4 –
22 G. similignum 24.5–24.7 25.8–26.4 22.5–22.7 13.9–14.5 24.9–25.0 8.5–9.0 22.9–23.1 22.5–22.7 0.1
23 G. scab idus 16.9–18.3 19.3–21.3 19.2–19.6 19.7–20.2 17.8 21.9–22.4 17.1–18.8 18.4–19.2 20.2–21.1 1.5
24 G. subpalma us 21.4–21.5 17.8–18.4 18.6 25.3–25.5 22.4–22.5 25.5–25.6 20.3 20.5 25.0–25.1 20.0–21.1 0.0
25 G. swinhonis 21.9–22.0 23.1–23.4 21.4 24.0–24.3 23.7–23.8 26.5–26.7 22.5 22.2 25.5–25.6 19.6–20.2 21.8 –
26 G. hakhekensis 18.6–18.8 19.7–20.7 6.7 22.2–22.9 18.1 21.6–22.2 13.0 13.0 20.9–21.0 19.0–19.4 19.9 23.1 –
27 G. uongi 21.8–22.0 20.7–21.4 22.0 21.6–21.8 21.2 22.2–22.7 22.2 21.8 21.6–21.8 18.2–19.7 21.4 24.0 20.3
zse.penso .ne
Xu, Y. e al.: A new Gekko species
2228
Taxonomic accoun
Family Gekkonidae Oppel, 1811
Genus Gekko Lau en i, 1768
Subgenus Japonigekko Wood, Guo, T a e s, Su, Olson,
Baue , G isme , Sile , Moyle, Ande sen & B own, 2020
Gekko essela us Xu, Ma, Cai, Qi, Ma sukoji,
Poya ko , Sun, Jiang & Peng, sp. no .
h ps://zoobank.o g/6373BF5E-F05B-4248-8B7D-0E50B1E10BE3
Figs 3–10, Tables 4–6
Type ma e ial. Holo ype • CIB 119371 (Field no. LFR
2025142), adul male, om Se’e gu Town, Heishui Coun-
y, Aba Tibe an and Qiang Au onomous P e ec u e, Sich-
uan P o ince, China (31.9531°N, 103.3971°E; 1,847 m
asl.), collec ed by Yuhao Xu, Fanyue Sun, and Jiaxiang
Wu on 3 Augus 2025.
Pa a ypes (N = 13, all om Sichuan P o ince, Chi-
na) • QHU R2025022 (Field no. LFR 2025143, adul e-
male) • QHU R2025023 (Field no. LFR 2025144, adul
male) • QHU R2025024 (Field no. LFR 2025145, adul
emale), wi h he same collec ing in o ma ion as he ho-
lo ype • QHU R2025025 (Field no. LFR 2025146, adul
male), om Longba Town, Heishui Coun y (31.9855°N,
103.3503°E; 2,047 m asl.), collec ed by Yuhao Xu, Fa-
nyue Sun and Jiaxiang Wu on Augus 04, 2025 • QHU
R2025026 (Field no. LFR 2025147, subadul male), om
Se’e gu Town, Heishui Coun y (31.9290°N, 103.4211°E;
1,827 m asl.), collec ed by Yuhao Xu, Fanyue Sun and
Jiaxiang Wu on Augus 4, 2025 • SYS 003029 (Field no.
LFR 2025148, adul male) • SYS 003030 (Field no. LFR
2025149, adul emale), om Se’e gu Town, Heishui
Coun y (31.9260°N, 103.44324°E; 1,886 m asl.), collec -
ed by Yuhao Xu, Fanyue Sun and Jiaxiang Wu on Augus
4, 2025 • SYS 003031 (Field no. LFR 2025150, adul
emale) • SYS 003032 (Field no. LFR 2025151, subadul
emale), om he same locali y as he holo ype, collec -
ed by Yuhao Xu, Fanyue Sun and Jiaxiang Wu on Au-
gus 4, 2025 • CIB 119372 (adul emale) • CIB 119373
(adul emale) • CIB 119374 (adul emale) • CIB 119375
(subadul emale), om Se’e gu Town, Heishui Coun y
(31.9650°N, 103.3966°E; 1,938 m asl.), collec ed by Bo
Cai in July, 2025.
E ymology. The speci ic name “ essela us” is a La i-
nized adjec i e in he nomina i e singula (masculine
gende ), de i ed om “ essella” (a small squa e ile), and
means “ e icula ed” o “checke boa d-like.” I e e s o
he cha ac e is ic do sal pa e n o he new species, which
consis s o a se ies o i egula da k ans e se bands ha
a e ain ly in e up ed medially by a na ow e eb al line
and, la e ally, a e pa ly agmen ed and pa ly in e con-
nec ed, o ming an i egula , checke boa d-like e icula -
ed pa e n ac oss he do sum. Fo he common names, we
sugges “Checke ed Gecko” in English and “Bān Wén Bì
Hǔ” (斑纹壁虎) in Chinese.
Diagnosis. Gekko essela us sp. no . can be diagnosed
om o he Japonigekko species by he ollowing unique
combina ion o cha ac e s: (1) a mode a e body size (SVL
eaches up o 64.9 mm in males and 72.9 mm in emales);
(2) na es in con ac wi h he os al, in e nasal 0 o 1; (3)
wo enla ged pos men als, a ely one; (4) la ened do sal
ube cles p esen om he pos e io head h ough he neck
o he an e io po ion o he ail, a anged in 12–15 ows
CIB 119371
QHU R2025023
QHU R2025025
QHU R2025022
QHU R2025024
QHU R2025026
CIB 119374
CIB 119373
CIB 119375
CIB 119372
G. engshanensis NHMG 240713
G. engshanensis NHMG 240714
G. liboensis SYS 002876
G. melli SYS 001742
G. kwangsiensis SYS 001194
G. subpalma us SYS 001762
G. alpinus CIB 121663
G. pauci ube cula us SYS 002806
G. hokouensis SYS 001209
G. kaiyai AHUXXBH01
G. liboensis SYS 002877
G. kwangsiensis SYS 001195
G. pauci ube cula us SYS 002807
G. hokouensis SYS 001311
G. kaiyai AHUJGTBH01
G. melli SYS 001702
G. subpalma us SYS 001767
G. cib SYS 001489
G. swinhonis SYS 001814
G. au i e ucosus NNU Z20050801.004
G. cib SYS 000708
G. swinhonis SYS 001815
G. alpinus CIB 121656
G. jinjiangensis CIB 5334220088
G. jinjiangensis CIB 5133380017
G. japonicus SYS 000672
G. khunkhamensis VNUF R.2021.23
G. nadenensis ZFMK 98741
G. bonkowskii VFU R.2014.10
G. scab idus CIB YN201909199
G. ichangensis SWU 0011602
G. hakhekensis IEBR A.2014.6
G. scab idus CIB YN201909200
G. ichangensis SWU 0011603
G. japonicus SYS 001317
G. scien iad en u a IEBR A.2014.7
G. sengchan ha ongi VNUF R.2014.14
G. chinensis SYS 001211
G. similignum SYS 001597
G. p ep HN2024R038
G. adle i SYS 001400
G. chinensis SYS 001085
G. similignum SYS 001598
G. p ep HN2024R039
G. adle i IEBR A.2012.24
G. gecko N/A
G. palma us SYS 002797
G. uongi IEBR A.2011.1
G. ee esii SYS 000796
G. palma us SYS 001185
100/100
80/69 55/60
77/64
100/100
95/96
98/100
89/98
100/100
–/76
76/74
76/74
83/77
84/82
84/82
98/100
100/100
99/100
89/97
87/90
–/81
100/100
91/97
–/–
100/100
100/100
99/100
100/100
66/84
0.06
Gekko essela us sp. no .
Gekko (Japonigekko
G. essela us sp. no .
)
Gekko (Gekko)100/100
Figu e 2. Maximum likelihood (ML) ee o Japonigekko in e ed om 16S and ND2 genes. Node suppo alues a e p esen ed
as SH-like app oxima e likelihood a io es (SH)/ul a as boo s ap app oxima ion (UFB); hose lowe han 50 a e shown as “–”.
Pho og aph on humbnail by YHX.
Zoosys . E ol. 101 (4) 2025, 2221–2242
zse.penso .ne
2229
a midbody; (5) 130–157 en al scales be ween he men al
and he cloacal sli ; (6) 98–106 midbody scale ows; (7)
31–39 en al scale ows; (8) subdigi al lamellae 7–10 on
inge I, 9–12 on inge IV, 7–9 on oe I, and 9–12 on oe
IV; (9) webbing absen ; (10) 6–8 p ecloacal po es in males,
absen in emales; (11) one pos cloacal ube cle on each
side, a ely wo; (12) in li e, he do sum is p edominan ly
g eyish b own, wi h a se ies o i egula da k ans e se
bands ha a e ain ly in e up ed medially by a na ow e -
eb al s ipe and la e ally agmen ed and in e connec ed,
o ming an i egula , checke boa d-like e icula ed pa e n.
Desc ip ion o he holo ype. (Fig. 3). An adul male
specimen wi h o iginal ail. Size medium, SVL 61.4 mm;
body slende and unk ela i ely elonga e, AGD 29.0 mm,
AGD/SVL a io 0.47; ail sligh ly longe han body, dis-
inc ly swollen a base, o al in sec ion, TAL 72.3 mm,
TAL/SVL a io 1.19.
Head dep essed, no iceably longe han wide, and
clea ly sepa a ed om he neck, HL 17.09 mm, HW
12.31 mm, HH 5.00 mm, HH/HL a io 0.29, HL/HW a io
1.39. Snou ob use, ound an e io ly, ESD 7.14 mm, ESD/
HL a io 0.42. Eye la ge, ED 3.89 mm, ED/HL a io 0.23;
pupil e ical, ea u ing c enula ed edges. Ea opening
small, app oxima ely ellip ical, obliquely o ien ed, EL
1.12 mm, EL/ED a io 0.29.
Ros al app oxima ely ec angula in shape, wide han
he high, RW 2.31 mm, RH 1.46 mm, RW/RH a io 1.58.
Na es o al, ounded by he os al, 1s sup alabial, one dis-
inc ly enla ged sup anasal, and h ee sligh ly enla ged na-
sals pos e io ly; in e nasals absen . P eo bi als 18/17, p e-
o bi al egion deeply conca e; in e o bi al scales be ween
he an e io co ne s o eyes 24; sup alabials 10/10; in al-
abials 9/9. Men al pen agonal, wide han long, MW 1.80,
ML 1.42, MW/ML a io 1.27; men al na owe han he
os al, RW/MW a io 1.28; pos men als wo, enla ged,
wice as long as wide, ouching he men al and he i s
in alabial on bo h sides and h ee gula scales pos e io ly.
Tube cles p esen on he do sal su ace o he head behind
he eyes; g anula scales on he an e odo sal egion o he
head a e la ge han hose on he pos e io egion.
Do sal scales smoo h, ound o o al, g anula , and
jux aposed; ube cles la ened, i egula ly a anged, ex-
ending om he pos e io pa o he head o he an e io
one- hi d o he ail, o ming app oxima ely 14 ows a
midbody, su ounded by 8–9 do sal scales. Ven ola e -
al old p esen , wi hou ube cles; en als sligh ly la g-
e han do sals, smoo h, imb ica e, and la ges in middle
o belly; en al scale ows a midbody 36; scale ows
a ound midbody 106; en al scales in a ow be ween
men al and cloacal sli 138; p ecloacal scales enla ged,
bu no enla ged scales on highs; p ecloacal po es 6 (Fig.
4A), si ua ed a he dis al o each scale, he ips o he wo
cen al scales a e sligh ly inclined la e ally, sepa a ing
he p ecloacal po es a he midline in o wo g oups (3/3);
pos cloacal ube cle 1/1 (Fig. 5A), la ge, su ounded by
se e al smalle ube cles delinea ed by shallow g oo es.
Do sal scales o ail la and smoo h, i egula in size, wi h
a ew ube cles p esen on he an e io one- hi d o he ail;
subcaudals small a he base, dis inc ly enla ged beyond
he swollen po ion, and a anged in a longi udinal ow.
Fo elimbs and hindlimbs well de eloped; do sal su -
ace o o elimbs wi hou ube cles, bu do sal su ace
o hindlimbs bea ing ube cles; no webbing be ween he
inge s and oes; digi s mode a ely dila ed; inge s and
oes II–IV clawed; claws la e ally comp essed, ex ending
beyond he e minal lamellae; no webbing be ween in-
ge s and oes; subdigi al lamellae undi ided, unde ma-
nus 7-9-10-11-9 (le ) / 7-8-9-11-9 ( igh ), and unde pes
8-8-10-11-8 (le ) / 8-8-11-11-10 ( igh ); ela i e leng h
o inge s and oes I < II < V < III < IV.
Colo a ion o he holo ype in li e. (Fig. 6). In li e, he
do sal g ound colo o he head and body is ligh g ey-
ish-b own, sca e ed wi h small da k b own o yellow-
ish-b own spo s. A ain W-shaped da k ma king ex end-
ing an e io ly om he occipi al egion o he pos e io
co ne s o he eyes. On he la e al sides o he head, a
da k longi udinal s ipe unning om he pos e o en al
co ne o he eye o he ea opening. Eigh i egula da k
ans e se bands be ween he nape and he sac um. The
da k bands ain ly di ided along he midline by a na -
ow, ligh -colo ed e eb al line unning in e mi en ly
om he nape o he ail ip. La e ally, he bands pa -
ly agmen ed and in e connec ed, o ming an i egula
e icula ed pa e n eminiscen o a checke boa d. All
da k ma kings and bands consis o a mix u e o yellow-
ish-b own and da k b own spo s. The do sal su aces o
he limbs bea ing i egula da k ans e se o e icula ed
ma kings. The ail ligh g eyish-b own a he base, g ad-
ually becoming pale pos e io ly, and bea ing 12 da k
ans e se bands; he ail ip da k b own. The head and
en al su aces o he body uni o mly c eamy whi e,
while he en al su ace o he ail pale yellowish-whi e.
A e cap u e, he o e all body colo becomes pale ,
u ning yellowish-b own, and he da k ma kings on he
do sum become blu ed. Howe e , a e being kep in cap-
i i y o se e al days, he colo pa e n g adually e u ns
o i s o iginal s a e, e en when housed in a whi e con aine .
Colo a ion o he holo ype in p ese a ion. (Fig.
3). In p ese a i e, he do sal g ound colo o he head,
body, and limbs u ns g ey, wi h he yellowish-b own
spo s on he do sum disappea ing, while he da k b own
ma kings emained dis inc ; he en al su ace aded o
g eyish-whi e.
Va ia ion. The main mo phological cha ac e s o Gek-
ko essela us sp. no . a e summa ized in Table 4. The spe-
cies exhibi s no e iden sexual dimo phism in mo phome -
ic a ios, scale coun s, o colo a ion. The longes known
male measu es SVL 64.9 mm, wi h TAL 72.9 mm (QHU
R2025025); and he emale measu es SVL 65.3 mm, wi h
TAL 65.1+ mm (QHU R2025022); TAL/SVL a io 1.07–
1.23. Head dep essed, no iceably longe han wide, and
clea ly sepa a ed om he neck, HL/HW a io 1.16–1.47.
Snou ob use, ound an e io ly, ESD/HL a io 0.38–0.43.
Eye la ge; pupil e ical, ea u ing c enula ed edges. Ea
opening small, app oxima ely ellip ical, obliquely o ien -
ed, EL/ED a io 0.19–0.40, and EL/HL a io 0.04–0.08.
zse.penso .ne
Xu, Y. e al.: A new Gekko species
2236
Figu e 8. P ese ed specimen o he emale pa a ype o Gekko essela us sp. no . (QHU R2025024). A. Do sal iew; B. Ven al
iew; C. do sal iew o head; D. La e al iew o head; E. Ven al iew o head; F. Ven al iew o hand; G. Ven al iew o oo ;
H. Do sal iew o midbody. Pho og aphs by YHX. Scale ba s: 5 mm.
mo e, i can be dis inguished om G. pauci ube cula us
by (1) MBSR 98–106 ( s. 136–142), (2) VS 31–39 ( s.
42–44), (3) SMC 130–157 ( s. 189–192), (4) DTR 12–15
( s. 4), (5) PP 6–8 ( s. 12), and (6) IOS 24–28 ( s. 37).
The main cha ac e is ics ha dis inguish Gekko es-
sela us sp. no . om o he species o he subgenus Ja-
ponigekko a e summa ized in Table 6. By ha ing 6–8
p ecloacal po es in males, he new species can be easily
dis inguished om G. aa onbaue i T i, Thai, Phim o-
han, Da id & Teynié, 2015, G. adle i Nguyen, Wang,
Yang, Lehmann, Le, Ziegle & Bonkowski, 2013, G.
canhi canhi Rösle , Nguyen, Van Doan, Ho, Nguyen &
Ziegle , 2010, G. chinensis, G. jinjiangensis Hou, Shi,
Wang, Shu, Zheng, Qi, Liu, Jiang & Xie, 2021, G. kai-
yai Zhang, Wu & Zhang, 2023, G. khunkhamensis Si -
hi ong, Lo, Nguyen, Ngo, Kho pa hoom, Le, Ziegle
& Luu, 2021, G. liui Zhou, Zhou, Wang, Li, Zhang, Li,
Shen, Liu & Rao, 2025, G. melli, G. palma us Boulenge ,
Zoosys . E ol. 101 (4) 2025, 2221–2242
zse.penso .ne
2237
Figu e 9. Habi a and ield obse a ions o Gekko essela us sp. no . A. Mac ohabi a o he new species in Se’e gu Town, Heishui
Coun y, Sichuan, China; B. Mic ohabi a o he new species; C. Gekko essela us sp. no . in li e, in si u; D. The sympa ic Lycodon
mul izona us. Pho og aphs by YHX (A–C) and TXG (D).
1907, G. p ep, G. scab idus Liu & Zhou, 1982, G. sci-
en iad en u a Rösle , Ziegle , Vu, He mann & Böhme,
2004, G. sengchan ha ongi Luu, Calame, Nguyen, Le
& Ziegle , 2015, G. shiba ai Toda, Sengoku, Hikida &
O a, 2008, G. similignum, G. awaensis Okada, 1956, G.
hakhekensis Luu, Calame, Nguyen, Le, Bonkowski &
Ziegle , 2014; G. uongi Phung & Ziegle , 2011; and
G. e eb alis Toda, Sengoku, Hikida & O a, 2008 ( s.
3–4 in G. aa onbaue i, 17–21 in G. adle i, 5 in G. canhi,
17–27 in G. chinensis, 4–5 in G. jinjiangensis, 9–12 in
G. kaiyai, 0 in G. khunkhamensis, 20–23 in G. liui, 9–11
in G. melli, 23–30 in G. palma us, 24 in G. p ep, 10–15
in G. scab idus, 23–30 in G. scien iad en u a, 4–5 in G.
sengchan ha ongi, 0–3 in G. shiba ai, 17 in G. similig-
num, 0 in G. awaensis, 1–5 in G. hakhekensis, 10–11 in
G. uongi, and 0–1 in G. e eb alis).
By ha ing MBSR 98–106, Gekko essela us sp. no .
can be dis inguished om G. bonkowskii Luu, Calame,
Nguyen, Le & Ziegle , 2015; G. cib Lyu, Lin, Ren, Jiang,
Zhang, Qi & Wang, 2021; G. hokouensis Pope, 1928; G.
ichangensis Cao, Sucha i akul, Tie, Suwannapoom, Yan
& Chomdej, 2025; G. japonicus (Dumé il & Bib on,
1836); G. nadenensis Luu, Nguyen, Le, Bonkowski &
Ziegle , 2017; and G. subpalma us (Gün he , 1864) ( s.
117 in G. bonkowskii, 128–149 in G. cib, 119–130 in
G. hokouensis, 127–145 in G. ichangensis, 130–144 in
G. japonicus, and 129–156 in G. nadenensis).
By ha ing DTR 12–15, Gekko essela us sp. no . can
be dis inguished om G. au i e ucosus Zhou & Liu,
1982, G. guishanicus Lin & Yao, 2016, G. swinhonis, and
G. wenxianensis Zhou & Wang, 2008 ( s. 16–20 in G.
au i e ucosus, 0 in G. guishanicus, 6–8 in G. swinhonis,
and 10 in G. wenxianensis).
Mo eo e , by ha ing LT4 9–12, Gekko essela us sp.
no . can be dis inguished om G. alpinus Ma, Shi, Shen,
Chang & Jiang, 2024, G. aibaiensis Song, 1985, and G.
yakuensis Ma sui & Okada, 1968 ( s. 13–15 in G. alpi-
nus, 7–8 in G. aibaiensis, and 15 in G. yakuensis).
Discussion
Molecula phylogene ic analyses eco e ed Gekko es-
sela us sp. no . as a s ongly suppo ed monophyle ic
lineage (SH = 100 / UFB = 100) wi hin he subgenus
Japonigekko. This lineage is nes ed wi hin a well-sup-
po ed clade (SH = 100 / UFB = 100) ha also includes
G. engshanensis, G. kwangsiensis, G. liboensis, and G.
pauci ube cula us, oge he o ming a dis inc e olu ion-
a y assemblage. The close gene ic ela ionships among
zse.penso .ne
Xu, Y. e al.: A new Gekko species
2238
Table 5. Compa ison o mo phological cha ac e s among Gekko essela us sp. no ., G. engshanensis, G. kwangsiensis, G. liboensis,
and G. pauci ube cula us; di e ences a e shown in bold.
Species G. essela us sp. no . G. engshanensis G. kwangsiensis G. liboensis G. pauci ube cula us
N = 14 N = 6 N = 6 N = 2 N = 2
SVLmax
(mm)
65.3 79.9 69.7 79.7 85.9
NS 3 o 4 3 3 3 3
INS 0 o 1 0 0 o 1 0 0
SL 10 o 11 9–13 10–13 11 11
IL 9–12 10–13 11–13 9–11 9–10
IOS 22–26 22–27 29–31 32–35 37
PO 12–15 15–19 18–20 17–18 14–18
PM 2 ( a ely 1) 2 2 2 2
GP 3–6 3–6 4–6 4–6 4–6
DTR 12–15 9–11 9–11 9–10 4
GSDT 7–9 9–10 8–10 9–10 8
SMC 130–157 197–213 185–208 183–195 189–192
MBSR 98–106 149–161 143–156 131–140 136–142
VS 31–39 40–49 41–45 38–41 42–44
LF1 7–10 11–13 10–13 12–13 10–11
LF4 9–12 12–16 12–14 14–17 12–13
LT1 7–9 12–14 11–13 12–13 11
LT4 9–12 13–15 14–18 14–15 11–13
PP 6–8 9–12 9–10 9 12
PAT 1 ( a ely 2) 1 1 1 1
Web Absen P esen , weakly P esen , weakly P esen , weakly Pesen , weakly
Fo e
ube cles
Absen Absen Absen Absen Absen
Hind
ube cles
P esen Absen Absen Absen Absen
Tail
ube cles
P esen Absen P esen – Absen
Sou ces This s udy Huang e al. (2025) Yang e al. (2015) Huang e al. (2025) Wang e al. (2024)
hese axa sugges a po en ially sha ed e olu iona y o -
igin. Howe e , despi e i s phylogene ic a ini y wi h he
ka s -dwelling species, G. essela us sp. no . exhibi s
ma kedly di e en ecological and geog aphic cha ac e -
is ics. The new species inhabi s highly wea he ed ocky
slopes wi hin ho –d y alleys o no hwes e n Sichuan
P o ince, in con as o i s congene s, which a e con ined
o wa m limes one ka s o es s in sou he n China. These
con as ing habi a p e e ences sugges ha en i onmen-
al he e ogenei y has likely played a signi ican ole in
d i ing di e gence wi hin his clade.
Geog aphically, Gekko essela us sp. no . is widely
sepa a ed om i s closes ela i es. The nea es known
ock-dwelling species, G. liboensis, occu s nea ly 900 km
away. I is he e o e plausible ha addi ional undesc ibed
ock-dwelling species o Japonigekko exis in hese un-
de explo ed a eas. This obse a ion unde sco es he
need o mo e comp ehensi e and sys ema ic ieldwo k
in sou hwes e n and cen al China o imp o e ou unde -
s anding o species di e si y, dis ibu ion pa e ns, and
e olu iona y his o y wi hin he subgenus Japonigekko.
The disco e y o Gekko essela us sp. no . aises he
o al numbe o species in he subgenus Japonigekko o
39, wi h 26 eco ded om China, including se en spe-
cies cu en ly known om Sichuan P o ince: Gekko es-
sela us sp. no ., G. alpinus, G. chinensis ( eco d ques-
ionable), G. cib, G. japonicus, G. jinjiangensis, and G.
scab idus (Cai e al. 2018; Hou e al. 2021; Lyu e al.
2021; Ma e al. 2024). Wi h he excep ion o he newly
desc ibed species, all o he abo e men ioned axa ex-
hibi a ying deg ees o adap abili y o human-modi ied
en i onmen s, such as buildings, b idges, o conc e e
embankmen s. In con as , ield obse a ions indica e
ha G. essela us sp. no . is almos ne e ound wi hin
human se lemen s. E en a si es adjacen o illages o
owns, indi iduals we e only obse ed on nea by unde-
eloped ocky slopes, sugges ing ha he species may be
mo e sensi i e o human dis u bance han i s congene s.
Gi en he cu en lack o da a on popula ion size, ends,
and dis ibu ion ange, we ecommend ha Gekko es-
sela us sp. no . be p elimina ily classi ied as Da a De-
icien (DD) unde he IUCN Red Lis c i e ia. Fu he
ield su eys in adjacen mon ane egions a e essen ial
o be e unde s and i s conse a ion s a us and o as-
sess whe he he species may quali y o lis ing unde a
h ea ened ca ego y in he u u e.
Zoosys . E ol. 101 (4) 2025, 2221–2242
zse.penso .ne
2239
Table 6. Mo phological cha ac e s o Gekko (Japonigekko); bold on s indica e di e ences om he new species; Y = “Yes” o p esen , N = “No” o absen ; di e ences a e shown in bold o
cla i y.
No. Species SVLmax SL IL IOS DTR SMC MBSR VS LT 1 LT4 Web Fo e ube cles Hind ube cles Tail ube cles PP
1G. essela us sp. no . 65.3 10 o 11 9–12 22–26 12–15 130–157 98–106 31–39 7–9 9–12 N N Y Y 6–8
2G. aa onbaue i 80 13–14 10–11 34–37 0 – 98–104 39–43 14–17 14–16 Y NN N 3–4
3G. adle i 75.3 10–15 9–13 27–36 7–11 168–190 123–144 35–44 11–14 11–15 YN Y Y 17–21
4G. alpinus 74.16 9–13 8–10 22–28 12–17 158–189 92–114 32–39 8–11 13–15 NYY Y 4–7
5G. au i e ucosus 69 9–11 9–11 26–27 16–20 – – – 6–8 6–8 N YY Y 8–11
6G. bonkowskii 69.2 12–14 10–11 49–50 0 154–169 117 37–40 11–13 15 Y NN N 6
7G. canhi 99.2 14 10–12 49–50 11–12 168–170 205–227 49–51 13–16 14–17 N N Y N5
8G. chinensis 72 10–14 9–13 35–48 10 156–167 118–140 37–39 8–10 9–12 YN Y Y 17–27
9G. cib 66.4 10–12 10–14 28–36 0 171–196 128–149 37–45 9–13 9–17 YNN N 7–9
10 G. engshanensis 79.9 9–13 10–13 22–27 9–11 197–213 149–161 40–49 12–14 13–15 Y NN N 9–12
11 G. guishanicus 64 – – – 0– – – 8–10 8–10 YNN N 6–8
12 G. hokouensis 70 10–14 8–11 30–33 12–18 153–174 119–130 36–43 8–11 15–18 N N NY 5–9
13 G. ichangensis 66.5 7–11 7–11 27–36 10–12 174–195 127–145 39–47 12–15 13–18 Y Y Y Y 5–8
14 G. japonicus 74 9–13 8–13 32–35 9–14 169–188 130–144 39–44 10–12 14–16 NYY Y 4–9
15 G. jinjiangensis 61.6 7–10 6–9 20–24 12–16 146–169 111–149 31–47 8–11 11–15 N YY Y 4–5
16 G. kaiyai 64.99 9–12 9–13 22–33 11–18 157–209 99–121 30–43 8–9 7–11 N YY Y 9–12
17 G. khunkhamensis 75.2 9–10 9–10 31–32 0 181–185 127–138 42–45 13–14 14–15 Y NN N 0
18 G. kwangsiensis 69.7 10–12 11–13 29–31 9–11 185–208 143–156 41–45 11–13 13–18 Y NNY9–11
19 G. liboensis 79.7 11 9–11 32–35 9–10 183–195 131–140 38–41 12–13 14–15 Y NN–9
20 G. liui 56.67 11–13 11–12 37–42 7–8 140–148 103–118 34–39 10–11 12–13 YNNY20–23
21 G. melli 80.3 10–13 9–12 34–40 0 171–192 148–160 44–46 10–12 11–14 YNN N 9–11
22 G. nadenensis 77.1 12–14 10–12 28–30 0 175–185 123–140 38–40 13–15 14–16 Y NN N 6
23 G. palma us 79.7 11–15 9–13 27–36 4–12 160–191 116–147 36–47 10–13 10–16 YNNY23–30
24 G. pauci ube cula us 85.9 11 9–10 37 4 189–192 136–140 42–44 11 11–13 YNN N 12
25 G. p ep 75.7 14 11 37–38 12 144–152 122–126 38–40 11–12 12–13 YNNY24
26 G. scien iad en u a 73 12–14 9–13 41–51 0 118–140 139–143 38–48 12–15 14–17 Y NNY23–30
27 G. scab idus 64 9–11 9–11 30 17–21 – – – 6–9 7–9 N YY Y 10–15
28 G. sengchan ha ongi 77.3 8–10 6–7 28–32 0 175–184 120–135 35–43 11–14 13–17 YNN N 4–5
29 G. shiba ai 70.9 10–13 10–14 37–52 5–14 – 114–134 –11 o 12 9–16 N N NY0–3
30 G. similignum 58.9 12–14 11 46–48 11 –144–153 –11–13 12–14 YNNY17
31 G. subpalma us 65.8 8–12 7–12 28–37 0 144–190 129–156 39–46 9–12 11–14 YNN N 5–9
32 G. swinhonis 66 7–12 7–11 23–24 6–8 – – 40 6–9 6–9 N YY – 7–9
33 G. aibaiensis 69 9–10 8–10 28 – – – – 6–7 7–8 Y – – – 4–6
34 G. awaensis 71 15 13 –0– – – 10 12 N N N N 0
35 G. hakhekensis 79.2 12–14 10–11 22–26 0 165–174 110–116 32–40 11–13 14–15 Y NN N 1–5
36 G. uongi 95.9 13–15 11–13 45–48 0 160–172 131–143 35–36 11–13 15–17 N N N N 10–11
37 G. e eb alis 69.2 10–15 10–15 35–50 2–12 – 112–139 10–12 – 9–17 N N N N 0–1
38 G. wenxianensis 59 12 11 – 10 – – 42–44 6 9 N N Y – 6–8
39 G. yakuensis 72 12–13 9–13 – – – – – 10 15 N N NY 6–8
zse.penso .ne
Xu, Y. e al.: A new Gekko species
2240
Acknowledgmen s
We would like o exp ess ou g a i ude o M . Jiaxiang Wu
(China), Danyang Zhou (China), and Maozhou Xu (Chi-
na) o hei help wi h ieldwo k and pho og aphy. We also
hank M . Tie ui Zhang (AHU, China), Hanming Song
(SYS, China), and Hao ian Wang (HAINNU, China) o
assis ance wi h da a analysis and manusc ip polishing.
We a e g a e ul o 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 he
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], he China Biodi e si y
Obse a ion Ne wo ks (Sino BON–Amphibian & Rep ile),
and he Russian Science Founda ion (RSF g an Nº 22-14-
00037-P, suppo ing he wo k o D . Nikolay A. Poya ko ).
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un ouche ed adul male indi idual om Se’e gu, Heishui, Sichuan, China; B. Gekko essela us sp. no ., un ouche ed ju enile indi-
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zse.penso .ne
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