Biodi e si y Da a Jou nal 13: e167521
doi: 10.3897/BDJ.13.e167521
Taxonomy & In en o ies
Fi s desc ip ion o he male and hemipenial
mo phology o Opis ho opis haihaensis Ziegle e
al., 2019 (Se pen es, Na icidae), wi h upda ed
diagnosis and dis ibu ion
Tie ui Zhang , Jinlong Ren , Maoliang Li , Yuhao Xu , Xinge Wang , Nikolay A. Poya ko ,
Tan Van Nguyen , Song Huang
‡ 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, China
§ CAS Key Labo a o y o Moun ain Ecological Res o a ion and Bio esou ce U iliza ion, Ecological Res o a ion and
Biodi e si y Conse a ion Key Labo a o y o Sichuan P o ince, Chengdu Ins i u e o Biology, Chinese Academy o
Sciences, Chengdu, China
| S a e Key Labo a o y o Pla eau Ecology and Ag icul u e, Xining, China
¶ Depa men o Ve eb a e Zoology, Lomonoso Moscow S a e Uni e si y, Moscow, Russia
# The School o Medicine & Pha macy, Duy Tan Uni e si y, Da Nang, Vie nam
¤ Cen e o En omology & Pa asi ology Resea ch, Duy Tan Uni e si y, Da Nang, Vie nam
« 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, W, China
Co esponding au ho : Tan Van Nguyen ( an.si as @gmail.com), Song Huang ([email protected])
Academic edi o : Chelmala S ini asulu
Recei ed: 02 Aug 2025 | Accep ed: 09 Oc 2025 | Published: 20 No 2025
Ci a ion: Zhang T, Ren J, Li M, Xu Y, Wang X, Poya ko NA, Nguyen TV, Huang S (2025) Fi s desc ip ion o
he male and hemipenial mo phology o Opis ho opis haihaensis Ziegle e al., 2019 (Se pen es, Na icidae),
wi h upda ed diagnosis and dis ibu ion. Biodi e si y Da a Jou nal 13: e167521.
h ps://doi.o g/10.3897/BDJ.13.e167521
Abs ac
Backg ound
The Haiha Moun ain S eam Keelback, Opis ho opis haihaensis Ziegle , Pham, Nguyen,
Nguyen, Wang, Wang, S ua & Le, 2019, was o iginally desc ibed based on a single
emale specimen om no he n Vie nam. To da e, only wo emale specimens ha e been
epo ed, ende ing O. haihaensis one o he leas s udied membe s o i s genus.
‡ § § | ‡ ¶
#,¤ «
© Zhang T 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.
New in o ma ion
Du ing ecen he pe ological su eys in Guangxi Zhuang Au onomous Region, sou he n
China, one male and one emale specimen o Opis ho opis we e collec ed om Qinzhou
Ci y. Phylogene ic analyses and de ailed mo phological compa isons iden i ied hese
indi iduals as O. haihaensis. Based on hese new specimens, we p o ide he i s
desc ip ion o he male o his species, including hemipenial mo phology, along wi h
e ised diagnos ic ea u es and new da a on i s na u al his o y and dis ibu ion.
Keywo ds
Na icidae, Opis ho opis, sexual dimo phism, sub opical mois b oadlea o es
In oduc ion
The genus Opis ho opis Gün he , 1872 (Na icinae) comp ises s eam-dwelling snakes
dis ibu ed om sou he n China and he Indochina Peninsula o he Ryukyu A chipelago,
Suma a, and he Philippines (Poya ko e al. 2023, Ue z e al. 2025). Cu en ly, 25
species a e ecognized in he genus, including nine desc ibed wi hin he pas wo
decades (Teynié e al. 2014, Wang e al. 2017, Wang e al. 2017, Ren e al. 2017, Ziegle
e al. 2019, Wang e al. 2020). Owing o hei aqua ic and c yp ic li es yles, many
Opis ho opis species emain poo ly known and a e o en ep esen ed solely by hei ype
se ies (S ua and Chuaynke n 2007, Teynié e al. 2014, Wang e al. 2020, Gao e al.
2024).
One such example is Opis ho opis haihaensis Ziegle , Pham, Nguyen, Nguyen, Wang,
Wang, S ua & Le, a membe o he O. maculosa complex oge he wi h O. maculosa
S ua & Chuaynke n and O. hung ai Wang, Lyu, Zeng, Lin, Yang, Nguyen, Le, Ziegle &
Wang. Species wi hin his complex sha e he ollowing combina ion o ea u es: a single
p e on al scale; do sal scale ows in 15–15–15; smoo h do sal scales; a da k b own o
blackish do sum on he body and ail, wi h each scale bea ing a pale yellow spo ; yellow
chin shields wi h da k mo ling; and yellow en al su aces wi h da k b own la e al
ma gins and sca e ed b own lecks (S ua and Chuaynke n 2007, Yang e al. 2011,
Ziegle e al. 2018, Wang e al. 2020).
The maculosa complex was i s eco ded in China by Yang e al. (2011), based on
specimens om Guangdong P o ince and Guangxi Zhuang Au onomous Region (ZAR).
Subsequen ly, Ziegle e al. (2019) desc ibed O. haihaensis as a dis inc species based
on a single emale om Tai Chi Village, Quang Son Commune, Hai Ha Dis ic , Quang
Ninh P o ince, no he n Vie nam. This specimen had p e iously been misiden i ied as O.
maculosa sensu S ua & Chuaynke n, which was o iginally desc ibed om Phu Wua
Wildli e Sanc ua y, Boong Kla Dis ic , Nong Khai P o ince, Thailand (Nguyen e al.
2018). La e , Wang e al. (2020) e-examined he Chinese eco ds assigned o O.
maculosa by Yang e al. (2011), desc ibing O. hung ai as a new species om Guangdong
2Zhang T e al
P o ince and Guangxi ZAR. They also con i med he p esence o O. haihaensis in China
o he i s ime, based on an adul emale specimen om Shiwandashan Na ional Na u e
Rese e, Fangchenggang Dis ic , Guangxi ZAR, and ecommended he emo al o O.
maculosa om he Chinese he pe o auna.
Based on wo emale specimens, O. haihaensis was ini ially diagnosed by he ollowing
combina ion o cha ac e s: o al leng h 500–509 mm in adul emales; ela i ely long ail
( ail leng h/ o al leng h a io 0.22); in e nasal no in con ac wi h lo eal; p e on al no
con ac ing sup aocula ; on al con ac ing p eocula ; one p eocula and one o wo
pos ocula s; empo als 1+1; eigh sup alabials, wi h he 4 and 5 en e ing he o bi ; 22–
24 maxilla y ee h; an e io chin shields longe han pos e io ; 164–169 en als (plus wo
p e en als); 75–79 subcaudals; nasal cle di ec ed owa ds i s sup alabial; do sal
scales in 15–15–15 ows, smoo h; ail scales smoo h o indis inc ly keeled; chin shields
yellow wi h da k mo ling; and body and ail do sum da k, each scale wi h a pale yellow
spo (Ziegle e al. 2019, Wang e al. 2020). Despi e i s ecen desc ip ion, he lack o male
specimens has hinde ed a comp ehensi e unde s anding o he species’ diagnos ic
cha ac e s and dis ibu ional limi s.
Du ing a ecen su ey in Qinbei Dis ic , Qinzhou Ci y, Guangxi ZAR, China, wo
specimens (one male and one emale) e e able o Opis ho opis we e collec ed.
Mo phological compa isons and phylogene ic analyses con i med hei iden i y as O.
haihaensis. In his s udy, we p o ide he i s de ailed mo phological desc ip ion o he
male, including hemipenial mo phology, upda e he species’ known dis ibu ion, and
e ise i s diagnosis based on new ma e ial.
Ma e ials and me hods
Sampling
Two specimens o Opis ho opis we e collec ed om Qinbei Dis ic , Qinzhou Ci y,
Guangxi ZAR, China. These specimens we e humanely eu hanized wi h 0.7% icaine
me hanesul ona e (MS222) solu ion. F esh li e issue was ex ac ed and immedia ely
p ese ed in 95% e hanol. Specimens we e p ese ed in 75% e hanol o pe manen
s o age, and deposi ed in Anhui No mal Uni e si y Museum (ANU). Sampling p ocedu es
in ol ing li e snakes we e app o ed by he Animal E hics Commi ee o Anhui No mal
Uni e si y and complied wi h he Wild Animals P o ec ion Law o China.
Phylogene ic analyses
To al genomic DNA was ex ac ed om p ese ed li e issue wi h OMEGA Tissue DNA
Ki D3396 (Omega Bio-Tek, No c oss, GA, USA). A agmen o he mi ochond ial
cy och ome b (Cy b) gene was ampli ied using he p ime pai L14910 (5’-
GACCTGTGATMTGAAACCAYCGTTGT-3’) and H16064 (5’-CTTTGGTTTACAAGAA
CAATGCTTTA-3’) (Bu b ink e al. 2000). The double-s anded p oduc s we e sequenced
by Sangon Bio ech (Shanghai, China), and aw sequences we e assembled using
h h
Fi s desc ip ion o he male and hemipenial mo phology o Opis ho opis ... 3
SeqMan in he DNASTAR so wa e package (Bu land 2000). The sequences ob ained
om his s udy we e uploaded o DNA Da a Bank o Japan (DDBJ).
Following Gao e al. (2024), 34 sequences om 18 known Opis h opis species and h ee
ou -g oup species, Hebius johannis (Boulenge ), Na ix na ix (Linnaeus) and
Rhabdophis leona di (Wall) we e ob ained om GenBank and inco po a ed in o ou
da ase (Table 1). DNA sequences we e aligned by he Clus al W algo i hm wi h de aul
pa ame e s (Thompson e al. 1997) and immed wi h gaps pa ially dele ed in MEGA X
(Kuma e al. 2018). Bayesian in e ences (BI) was conduc ed in MRBAYES . 3.2.7a
(Ronquis e al. 2012) unde he GTR + I + G model on Phylosui e 1.2.3 (Zhang e al.
2020, Xiang e al. 2023). In he BI analysis, h ee independen uns we e conduc ed wi h
1 × 10 gene a ions and sampled e e y 1000 gene a ions wi h he i s 25% samples
we e disca ded as bu n-in. In he ML analysis, he boo s ap consensus ee was in e ed
om 1000 eplica es. Maximum likelihood (ML) was conduc ed unde he bes - i
subs i u ion model (GTR + I + G) in RaxmlGUI 1.3 (Sil es o and Michalak 2012).
Boo s ap p opo ions (BSP) we e in es iga ed wi h 1,000 boo s ap eplica es using he
as -boo s apping algo i hm. Unco ec ed pai wise gene ic dis ances (p-dis ance) o Cy
b gene among congene s we e calcula ed wi h MEGA X (Kuma e al. 2018).
Species Vouche No. NCBI/DDBJ acc.
No.
Locali y
Opis ho opis ande sonii SYS 001020 KY594732 China: M . Wu ong, Shenzhen, Guangdong
Opis ho opis ande sonii SYS 001423 KY594730 China: Tai Tam, Hong Kong
Opis ho opis cheni YBU 071040 GQ281779 China: Guangdong
Opis ho opis cheni SYS 001422 KY594741 China: Shimen ai NR, Yingde, Guangdong
Opis ho opis dao an ieni CIB 109024 PQ726905 Vie nam: K’Bang, Gia Lai
Opis ho opis dao an ieni VNMN 2019.01 PQ726906 Vie nam: K’Bang, Gia Lai
Opis ho opis du andi NCSM 80739 MK941137 Laos: Phongsaly
Opis ho opis guangxiensis GP 746 GQ281776 China: Guangxi
Opis ho opis haihaensis IEBR A.
2016.34
MK941139 Vie nam: Hai Ha, Quang Ninh
Opis ho opis haihaensis SYS 000537 MN890017 China: Shiwandashan NR, Guangxi
Opis ho opis haihaensis ANU 20240076 LC896244* China: Qinbei, Qinzhou, Guangxi
Opis ho opis haihaensis ANU 20240077 LC896245*China: Qinbei, Qinzhou, Guangxi
Opis ho opis hung ai SYS 000538 MN890018 China: M . Wuhuang, Guangxi
Opis ho opis hung ai SYS 000946 KY594748 China: Heishiding NR, Fengkai,
Guangdong
Opis ho opis jacobi IEBR 4329 MG545601 Vie nam: Tam Dao NP, Vinh Phuc
7
Table 1.
In o ma ion o samples and sequences used in molecula analysis. No es: Is = Island; NP = Na ional
Pa k; NR = Na u e Rese e; WS = Wildli e Sanc ua y; * = DDBJ Accession Numbe .
4Zhang T e al
Species Vouche No. NCBI/DDBJ acc.
No.
Locali y
Opis ho opis jacobi ZFMK 100818 MG545602 Vie nam: Tam Dao NP, Vinh Phuc
Opis ho opis kua unensis SYS 000998 KY594745 China: Qixiling NR, Yongxin, Jiangxi
Opis ho opis kua unensis SYS 001008 KY594746 China: Wulong, Shanghang, Fujian
Opis ho opis la e ialis SYS 001080 KY594744 China: M . Wu ong, Shenzhen, Guangdong
Opis ho opis la e ialis ZFMK 100806 MF477899 Vie nam: Tay Yen Tu NR, Bac Giang
Opis ho opis la ouchii GP 647 GQ281783 China: Fujian
Opis ho opis la ouchii SYS 000670 KY594742 China: Guadun, Wuyishan, Fujian
Opis ho opis laui SYS 001161 KY594738 China: Shangchuan Is, Taishan,
Guangdong
Opis ho opis laui SYS 001170 KY594739 China: Shangchuan Is, Taishan,
Guangdong
Opis ho opis maculosa FMNH 265798 MK941138 Thailand: Phu Wua WS, Nong Khai
Opis ho opis maxwelli SYS 000841 KY594736 China: Nan’ao Is, Guangdong
Opis ho opis maxwelli SYS 001053 KY594737 China: Huboliao NR, Nanjing, Fujian
Opis ho opis
shenzhenensis
SYS 001018 KY594727 China: M . Wu ong, Shenzhen, Guangdong
Opis ho opis
shenzhenensis
SYS 001021 KY594728 China: Sanzhou ian, Shenzhen,
Guangdong
Opis ho opis amdaoensis IEBR A.
2016.33
MF477901 Vie nam: Tam Dao NP, Vinh Phuc
Opis ho opis oquyi IEBR 4327 MG451046 Vie nam: Tay Yen Tu NR, Bac Giang
Opis ho opis oquyi VNMN 06315 MG451047 Vie nam: Tay Yen Tu NR, Bac Giang
Opis ho opis zhaoe mii CIB 109999 MG012800 China: Guzhang, Hunan
Opis ho opis zhaoe mii GZNU 320074 MK161471 China: Leishan, Guizhou
Hebius johannis GP 897 KJ685708 China: Yunnan
Na ix na ix MTD T 9269 HF680010 Denma k: Sønde Bo up, Zealand
Rhabdophis leona di SCUM 090009 KF800933 China: Panzhihua, Sichuan
Mo phological examina ion
Mo phological desc ip ions ollowed Ziegle e al. (2019), Wang e al. (2020) and Gao e
al. (2024). Body measu emen s and hei abb e ia ions a e as ollows: eye ho izon al
diame e (ED), head leng h (HL), maximum head wid h (HW), ail leng h (TaL), o al leng h
(TL), snou leng h (SnL), snou wid h (SnW), in e o bi al dis ance (IOD), dis ance be ween
he lowe ma gins o eye and o lip (SoL). All measu emen s, excep o ail leng h and
o al leng h, we e ob ained wi h a digi al slide-calipe o he nea es 0.01 mm.
Scala ion ea u es and hei abb e ia ions a e as ollows: do sal scale ow coun s (DSR),
sup alabial coun s (SL), in alabial coun s (IL), in e nasal coun s (IN), chin shield coun s
(CS), on al coun s (F), pa ie al coun s (P), p eocula coun s (P O), pos ocula coun s
Fi s desc ip ion o he male and hemipenial mo phology o Opis ho opis ... 5
(P O), en al coun s (VEN), subcaudal coun s (SC), p e on al in con ac wi h sup aocula
o no (P F-SpO), p eocula in con ac wi h on al (P O-F), lo eal en e ing o bi (L-o bi ),
in e nasals in con ac wi h lo eal (IN-L), sup alabials in con ac wi h o bi (SL-o bi ),
in alabials in con ac wi h an e io chin shields (IL-aCS), empo al coun s (TEM), cloacal
pla e en i e o di ided (CP), do sal scale su ace o ail (DST). The numbe o en al
scales was coun ed acco ding o Dowling (1951). Symme ic cha ac e s we e gi en as
le / igh and a e ages we e used in he analyses.
Gende o specimen was de e mined h ough dissec ion. The hemipenes o he male
specimen was e e ed om he le side o he hemipenial desc ip ion. The p epa a ion
and measu emen me hod o hemipenes ollowed Ren e al. (2022). The e e ed
hemipenes we e e-in la ed wi h colou ed pe oleum jelly and hen p ese ed in 75%
e hanol. Pho og aphs o he hemipenes we e aken using digi al came a a ached o a
ipod head and mul i ocal pho og aphs we e combined and mon aged using he Helicon
Focus (7.0.2 P o) so wa e. Hemipenial mo phology and e minology ollowed Dowling
and Sa age (1960), Zhang e al. (1984) and Ren e al. (2022).
Museum and collec ion abb e ia ions a e as ollows: ANU = Anhui No mal Uni e si y,
Wuhu, China; FMNH = Field Museum o Na u al His o y, Chicago, USA; IEBR = Ins i u e
o Ecology and Biological Resou ces, Vie nam Academy o Science and Technology,
Hanoi, Vie nam; KFBG = He pe ological Collec ion o Kadoo ie Fa m and Bo anic
Ga den, Hong Kong, China; SYS = Sun Ya -sen Uni e si y, Guangzhou, Guangdong,
China.
Taxon ea men
Opis ho opis haihaensis Ziegle , Pham, Nguyen, Nguyen, Wang, Wang,
S ua & Le, 2019
Ma e ials
a. scien i icName: Opis ho opis haihaensis; axonID: u n:lsid:biosci.ohio-
s a e.edu:osuc_names:275502; class: Rep ilia; o de : Squama a; amily: Na icidae;
genus: Opis ho opis; speci icEpi he : haihaensis; scien i icNameAu ho ship: Ziegle ,
Pham, Nguyen, Nguyen, Wang, Wang, S ua & Le, 2019; coun y: China; coun yCode:
CN; s a eP o ince: Guangxi; coun y: Qinbei; municipali y: Qinzhou; locali y: Tianmushan
Na ional Na u e Rese e, M . Xian ending; e ba imLocali y: 570 m; e ba imEle a ion:
1200 m; e ba imLa i ude: 22.006365°N; e ba imLongi ude: 108.206833°E;
e ba imCoo dina eSys em: WGS84; e en Da e: 25-Jun-24; e en Rema ks: collec ed by
Tie ui Zhang; indi idualCoun : 1; sex: male; li eS age: adul ; ca alogNumbe : ANU
20240076; language: en; collec ionCode: Rep ilia; basisO Reco d: P ese edSpecimen;
occu enceID: 37B587F0-02CC-5439-B859-E7923E8C247A
b. scien i icName: Opis ho opis haihaensis; axonID: u n:lsid:biosci.ohio-
s a e.edu:osuc_names:275502; class: Rep ilia; o de : Squama a; amily: Na icidae;
genus: Opis ho opis; speci icEpi he : haihaensis; scien i icNameAu ho ship: Ziegle ,
Pham, Nguyen, Nguyen, Wang, Wang, S ua & Le, 2019; coun y: China; coun yCode:
CN; s a eP o ince: Guangxi; coun y: Qinbei; municipali y: Qinzhou; locali y: Tianmushan
6Zhang T e al
Na ional Na u e Rese e, M . Xian ending; e ba imLocali y: 570 m; e ba imEle a ion:
1200 m; e ba imLa i ude: 22.006365°N; e ba imLongi ude: 108.206833°E;
e ba imCoo dina eSys em: WGS84; e en Da e: 25-Jun-24; e en Rema ks: collec ed by
Tie ui Zhang; indi idualCoun : 1; sex: emale; li eS age: adul ; ca alogNumbe : ANU
20240077; language: en; collec ionCode: Rep ilia; basisO Reco d: P ese edSpecimen;
occu enceID: AAD9F7D6-CE03-591A-AD77-98734F6D0E95
Desc ip ion o male specimen ANU 20240076
Measu emen s and scala ion. Body slende and cylind ical (TL 428.0 mm); head
sho and b oad, do sally dep essed, ba ely dis inc om neck (HL/HW 1.87);
snou
mode a e (SnL/SnW 0.74); eyes in medium size (ED/SoL 0.97); pupil ound;
in e o bi al dis ance la ge (IOD/HW 0.66); minu e g anula aspe i ies absen on head
scales; ail ela i ely long, ape ing pos e io ly (TaL 113.9 mm, TaL/TL 0.27); nos il
o al-shaped, loca ing in he uppe middle pa o nasal, di ec ed upwa ds.
Do sal scales in 15 ows h oughou he body; he ou e mos ows o do sal scales
sligh ly enla ged, while he es ones homogeneous in size, smoo h h oughou ;
e eb al scales no enla ged; p e en als 1, en als 168; cloacal pla e di ided;
subcaudals 87 pai ed, wi h single e minal igid ip, smoo h en i ely.
Ros al c escen -shaped, wide han high, isible om abo e; nasals la ge, wide
han high, di ided below nos il by a dis inc u ow; nasals in con ac wi h he i s wo
sup alabials, os al, in e nasal, p e on al and lo eal; in e nasals pai ed, apezoid,
longe han wide, cu ed ou wa d pos e io ly, in con ac wi h os al, nasal, and
p e on al; p e on al single, much b oade han long, an e io ly poin ed, in con ac
wi h in e nasals, nasals, lo eals, p eocula s, and on al; on al single, pen agonal o
nea equila e al iangle, almos equal in wid h and leng h, ape ing pos e io ly, equal
o i s dis ance om ip o snou ; pa ie als nea ly 1.8 imes as long as wide, pa ie al
su u es nea ly equal o on al in leng h; 1/1 sup aocula , dis inc ly longe han wide,
no in con ac wi h p e on al; 1/1 lo eal, pen agonal, wide han high, no en e ing
o bi , su ounded by second and hi d sup alabials, nasal, p e on al and p eocula ;
1/1 p eocula , la ge, hexagonal, highe han wide, eaching on al; subocula absen ;
2/2 pos ocula , uppe pai is almos wice he leng h and wid h o lowe pai ; 7/7
sup alabials, 1 –5 highe han wide, only 5 en e ing o bi on le la e al, 4 and 5
en e ing o bi on igh la e al (4-1-2/3-2-2), 7 la ges ; 1+1/1+1 empo als, an e io
one e y long and na ow, in b oad con ac wi h 5 –7 sup alabials and pa ie als,
pos e io one mo e s ongly de eloped; in alabials 8/8, i s pai in con ac behind
small men al, 1 –5 / 1 –5 in con ac wi h an e io chin shields, 5 in alabial
la ges ; an e io chin shields longe han pos e io ones, pos e io chin shields do no
con ac wi h each o he .
Den i ion. Maxilla y ee h 22, subequal, densely se , wi hou dias ema.
Colo a ion in li e (Figs 1, 2). Scales on do sum o head glossy black wi h sca e ed
yellow lecking; en al o head yellow wi h b ownish black mo ling on he ma gins o
each scale; uppe lip yellow wi h b ownish-black an e io and la e al ma gins on each
s h h h h
h
h h
s h s h h
Fi s desc ip ion o he male and hemipenial mo phology o Opis ho opis ... 7
sup alabial; do sum o body and ail glossy black wi h i idescence abo e, wi h single
b igh yellow spo on each scale, yellow spo s becoming la ge on sides o body;
en als yellow wi h b ownish black la e al ma gins; subcaudals yellow wi h b ownish
black an e io and la e al ma gins.
Colo a ion in p ese a ion. Colo a ion and pigmen a ion pa e ns o newly p ese ed
specimens (wi h 75% e hanol) signi ican ly aded, b igh yellow spo s on do sals and
sca e ed yellow lecking on do sum o head u n milky whi e, backg ound colo
o
en als ades o pale yellow. Gloss o scales emains dis inc .
Hemipenis (Fig. 3). Fully e e ed hemipenis is ela i ely long, measu ing 15.71 mm in
leng h and 2.52 mm in wid h; slende , single, cylind ical, sligh ly cla a e apically.
O gan en i ely spinose, wi h spines ela i ely uni o m in size and dis ibu ion; spines
sligh ly spa se in he p oximal egion; no basal nude a ea p esen . Basal hook
absen . Sulcus spe ma icus un o ked, cen ipe al, ex ending s aigh o he apex;
sulcus lips well de eloped, b oad, and smoo h along he en i e leng h. When
e ac ed, he hemipenis eaches he le el o he 31 subcaudal scale.
Desc ip ion o addi ional emale specimen ANU 20240077
Measu emen s and scala ion. Body slende and cylind ical (TL 485.0 mm); head
sho and b oad (HL/HW 1.76), do sally dep essed, ba ely dis inc om neck; snou
mode a e (SnL/SnW 0.60); eye small, (ED/SoL 0.81); pupil ound; in e o bi al dis ance
la ge (IOD/HW 0.60); minu e g anula aspe i ies absen on head scales; ail ela i ely
long, ape ing pos e io ly (TaL 94.9 mm, TaL/TL 0.20); nos il o al-shaped, loca ing in
he uppe middle pa o nasal, di ec ed upwa ds.
s
Figu e 1.
Li e pho og aphs o Opis ho opis haihaensis (specimen ANU 20240076, male) in si u om
Qinzhou, Guangxi, China. Pho og aphed by T.R. Zhang.
8Zhang T e al
Do sal scales in 15 ows h oughou he body; he ou e mos ows o do sal scales
sligh ly enla ged, while he es ones homogeneous in size, smoo h h oughou ;
e eb al scales no enla ged; p e en als 1, en als 165; cloacal pla e di ided;
subcaudals 71, pai ed, wi h single e minal igid ip, smoo h en i ely.
Ros al c escen -shaped, wide han high, isible om abo e; nasals la ge,
wide
han high, di ided below nos il by a dis inc u ow; nasals in con ac wi h he i s wo
sup alabials, os al, in e nasal, p e on al and lo eal; in e nasals coalesced in o one,
pen agonal, wide han long, conca ed inwa d pos e io ly, in con ac wi h os al,
nasal, and p e on al; p e on al single, much b oade han long, an e io ly poin ed, in
con ac wi h in e nasals, nasals, lo eals, p eocula s, and on al; on al single,
pen agonal o nea equila e al iangle, almos equal in wid h and leng h, ape ing
pos e io ly, equal o i s dis ance om ip o snou ; pa ie als nea ly 1.8 imes as long as
wide, pa ie al su u es nea ly equal o on al in leng h; 1/1 sup aocula , igh
sup aocula small and pa ially coalesced wi h uppe pos ocula , dis inc ly longe
han wide, no in con ac wi h p e on al; 1/1 lo eal, pen agonal, wide han high, no
en e ing o bi , su ounded by second and hi d sup alabials, nasal, p e on al and
Figu e 2.
Opis ho opis haihaensis p io o p ese a ion (specimen ANU 20240076, male): A do sal iew
o whole body; B en al iew o whole body; C do sal iew o head; D en al iew o head; E
igh la e al iew o head; F le la e al iew o head; G close-up o mid-body do sal scales.
Pho og aphs by T.R. Zhang.
Fi s desc ip ion o he male and hemipenial mo phology o Opis ho opis ... 9
7The 6 sup alabial is no elonga ed, app oxima ely equal o o sho e han
hal he leng h o he an e io empo al O. haihaensis
–The 6 sup alabial is dis inc ly elonga ed and nea ly equal in leng h o he
an e io empo al O. hung ai
8A la e al black s ipe sepa a ing pale en e om da k do sum; lo eal no
en e ing o bi O. la e alis
– No such la e al s ipe; lo eal en e ing o bi o no 9
9 Lo eal no en e ing o bi , 1.4–1.7× as long as deep; las sup alabial longes O. maxwelli
–Lo eal en e ing o bi o no , mo e han 1.7× as long as deep; las
sup alabial smalle han p eceding one 10
10 Lo eal in con ac wi h second sup alabial; ail leng h 15–20% o o al leng h;
do sum wi hou yellow longi udinal s ipes o c ossba s O. ande sonii
–Lo eal no con ac ing second sup alabial; ail leng h 20–23% o o al leng h;
do sum wi h yellow longi udinal s ipes o c ossba s 11
11 Body size small, o al leng h sho e han 419 mm; maxilla y ee h ≤ 25 O. la ouchii
–Body size mode a e o la ge, o al leng h longe han 419 mm; maxilla y
ee h ≥ 25 12
12
Maxilla y ee h 25–28; an e io empo als elonga ed, maximum an e io
empo al leng h / dep h a io 2.63–3.63; do sum da k oli e wi h yellow
c ossba s
O. cheni
–
Maxilla y ee h 28–30; an e io empo als sho , maximum an e io
empo al leng h / dep h a io 1.74–2.04; do sum wi h longi udinal yellow
s ipes
O. zhaoe mii
Analysis
In his s udy, 1 060 base pai s (bp) o Cy b sequences we e success ully ob ained o
alignmen , including wo newly gene a ed sequences o O. haihaensis. Bo h ML and BI
analyses yielded highly cong uen opologies (Fig. 6), consis en wi h indings om
p e ious s udies (Ren e al. 2017, Wang e al. 2017, Ziegle e al. 2019, Wang e al.
2020).
The genus Opis ho opis was eco e ed as a monophyle ic g oup wi hin he scope o he
cu en sampling (BSP = 83, BPP = 0.99). All O. haihaensis indi iduals clus e ed wi hin a
dis inc and well-suppo ed clade (BSP = 100, BPP = 1.00), nes ed wi hin he
Opis ho opis clade. In aspeci ic unco ec ed p-dis ances among he O. haihaensis
h
h
16 Zhang T e al
specimens we e 0.2% –4.5% (Suppl. ma e ial 1). The p-dis ances be ween O. haihaensis
and all analyzed Opis ho opis species a ied om 13.3%–13.4% ( o O. hung ai) o
18.0%–18.3% ( o O. dao an ieni O lo , Da e sky & Mu phy). The ollowing is a de ailed
desc ip ion o he addi ional specimens o O. haihaensis.
Discussion
Wang e al. (2020) conside ed he numbe o sup alabials o be a key diagnos ic
cha ac e dis inguishing Opis ho opis haihaensis om O. hung ai, wi h he o me
bea ing eigh sup alabials and he la e se en. Howe e , ou indings indica e ha
sup alabial coun s may a y depending on he posi ion o he o al issu e ela i e o he
pos e io ma gin o he se en h sup alabial, po en ially leading o misin e p e a ion. We
also e-checked he sup alabial coun s o O. haihaensis specimen SYS 000537 and
ound i ac ually has 7 sup alabials (8 in Yang e al. (2011), Wang e al. (2020)). To
minimize diagnos ic ambigui y caused by a iable sup alabial coun s, we p opose an
al e na i e diagnos ic cha ac e : in O. haihaensis, he 6 sup alabial is no elonga ed and
is app oxima ely equal o o sho e han hal he leng h o he an e io empo al, whe eas
in O. hung ai, he 6 sup alabial is dis inc ly elonga ed and nea ly equal in leng h o he
an e io empo al.
h
h
Figu e 6.
ML phylogene ic ee es ima ed om Cy bsequences depic ing phylogene ic ela ionships o
Opis ho opis (numbe s abo e b anches a e BSP/BPP).
Fi s desc ip ion o he male and hemipenial mo phology o Opis ho opis ... 17
Among he 25 cu en ly ecognized species o Opis ho opis, hemipenial mo phology has
been desc ibed in only 13 species (see Table 5) and se e al o hese desc ip ions emain
b ie o lack su icien de ail (Pope 1935, Zhang e al. 1984, S ua and Chuaynke n 2007,
Yang e al. 2011, Ziegle e al. 2018, Ren 2019, Gao e al. 2024). Based on he mos
ecen summa ies by Gao e al. (2024), he gene al hemipenial mo phology o
Opis ho opis is cha ac e ized by he ollowing ai s: ela i ely sho and s ou hemipenis,
shallowly bilobed in a “Y” shape, noncalycula e and noncapi a e (excep in O. alcalai and
O. maculosa); uni o m co e age wi h small spines o spinules; a single enla ged basal
hook on he p oximal egion o he uncus (excep in O. maculosa); and a single,
undi ided, cen ipe al sulcus spe ma icus ex ending o he base o dis al po ion o he
inne igh lobe (excep in O. dao an ieni). Wi hin his amewo k, Opis ho opis
haihaensis and O. maculosa exhibi unique hemipenial mo phology in possessing a
single, cylind ical hemipenis en i ely spinose and lacking a basal hook dis inguishing
hem om o he congene s. P esence o absence o he basal hook and mo phology o
sulcus se e as signi ican axonomic diagnos ic c i e ia, as hey could di ec ly in luence
ep oduc i e beha io and e lec adap i e e olu ion in ep oduc i e s a egies. The
hemipenis o Opis ho opis hung ai, he species mos closely ela ed o O. haihaensis,
emains undesc ibed. Yang e al. (2011) b ie ly no ed ha he hemipenis o specimen
SYS 0538 is cylind ical and ex ends o he le el o he 12 subcaudal and p o ided an
image o KFBG 2002.01. Howe e , he s uc u e shown does no appea o be ully
e e ed, ende ing any mo phological in e ence inconclusi e. De ailed documen a ion o
hemipenes in Opis ho opis hung ai and o he poo ly cha ac e ized o undocumen ed
species is essen ial o e alua ing pa e ns o hemipenial a ia ion and hei po en ial
phylogene ic signi icance.
Species Shape O namen a ion Enla ged
basal
hook
Sulcus
spe ma icus
Rela i e
leng h o
e ac ed
hemipenis
Sou ce
O. alcalai Single Uni o mly spinose
h oughou
1 Undi ided,
ex ends o
apical ip
11 B own and
Le i on (1961)
O.
ande sonii
? Bese wi h spines
p oximally and papilla-
like p ocesses dis ally
1 ? ? Pope (1935)
O.
dao an ieni
Bilobed,
lobes well-
de eloped
Spinose h oughou ,
spines app oxima ely
homogeneous in size,
p oximal ou h spa se ,
wi hou basal naked
a ea
1 Bi u ca e,
cen ipe al,
eaching apex
o each lobe
17 Gao e al.
(2024)
h
h
h
Table 5.
Documen ed hemipenial mo phology o Opis ho opis species. No es: Rela i e leng h o e ac ed
hemipenis was de ined by he subcaudal i can each a he dis almos ; “?” indica es missing da a.
18 Zhang T e al
Species Shape O namen a ion Enla ged
basal
hook
Sulcus
spe ma icus
Rela i e
leng h o
e ac ed
hemipenis
Sou ce
O.
guangxiensis
Bilobed,
sho and
hin, sligh ly
s ou e a
he p oximal
hal
Spinose h oughou ,
p oximal hi d spa se .
1 Undi ided,
eaching apex
o he igh
lobe
?Ren (2019),
Gao e al.
(2024)
O.
haihaensis
Single,
cylind ical,
sligh ly
cla a e
apically
Spinose h oughou ,
spines app oxima ely
homogeneous in size,
p oximal spa se
sligh ly, wi hou basal
naked a ea.
Absen Undi ided,
cen ipe al,
eaching apex
o hemipenis
31 This s udy
O. hung ai Cylind ical ? ? ? 12 Yang e al.
(2011)
O. jacobi Bilobed,
lobes sho ,
inwa ds
cu ed
Co e ed wi h small
spines, cu ed
backwa ds. The
e minal a ea o he
hemipenis be ween he
lobes wi hou spines.
Uppe uncus o
hemipenes wi h ing o
enla ged spines
1 Undi ided,
s e ching o
he sligh ly
smalle lobe
only.
?Ziegle e al.
(2018)
O.
kua unensis
Sligh ly
di ided a
he ip
Spinous, spines a e se
in longi udinal ows and
connec ed by leshy
idges, ecu ed
p oximally and
g adually dec ease in
dis al.
1 Undi ided 10 Pope (1935)
O. la e alis ? Bese wi h spines
p oximally and papilla-
like p ocesses dis ally
2 ? ? Pope (1935)
O. la ouchii Bilobed,
sligh ly
o ked a
he ip
Bese wi h hook-
shaped spines
p oximally and papilla-
like p ocesses dis ally
1 Undi ided,
eaching apex
o he igh
lobe
6 –8 Pope (1935),
Zhang e al.
(1984), Ren
(2019)
O. laui Bilobed,
sligh ly
elonga ed
Bese wi h hook-
shaped spines
p oximally and papilla-
like p ocesses dis ally
1 ? ? Ren (2019)
O. maculosa Single,
cylind ical
Basal a ea naked,
emainde o hemipenis
uni o mly o namen ed
wi h spines
Absen Undi ided,
cen ipe al,
eaching apex
o hemipenis
12 S ua and
Chuaynke n
(2007)
s
h
h
h h
h
Fi s desc ip ion o he male and hemipenial mo phology o Opis ho opis ... 19
Species Shape O namen a ion Enla ged
basal
hook
Sulcus
spe ma icus
Rela i e
leng h o
e ac ed
hemipenis
Sou ce
O. oquyi Bilobed,
lobes sho ,
inwa ds
cu ed
Co e ed wi h small
spines, cu ed
backwa ds. The
e minal a ea o he
hemipenes be ween
he lobes wi hou
spines. Uppe uncus
o hemipenes
wi h ing o enla ged
spines.
1 Undi ided,
s e ching o
he sligh ly
smalle lobe
only.
?Ziegle e al.
(2018)
O. zhaoe mii Bilobed,
igh lobe
sho e han
he le lobe
Co e ed wi h small
spines. The e minal
a ea o he hemipenis
be ween he lobes and
basal a ea naked.
Spines a ound basal
hook mode a ely
enla ged, spines on he
emainde o hemipenis
app oxima ely
homogeneous in size
1 Undi ided,
eaching apex
o he igh
lobe
?Gao e al.
(2024)
Opis ho opis haihaensis is cu en ly known om only ou specimens collec ed a h ee
locali ies in no he n Vie nam and sou he n China. I inhabi s clea , slow- lowing
moun ain s eams wi hin e e g een b oad-lea ed o es s habi a s ha a e inc easingly
h ea ened by de o es a ion and o he an h opogenic dis u bances. Gi en i s na ow
known dis ibu ion and ex emely limi ed numbe o eco ded specimens, we ecommend
a p o isional IUCN Red Lis s a us o Nea Th ea ened (NT). This classi ica ion is
p ecau iona y, pending u he ield su eys and popula ion assessmen s, and e lec s he
u gen need o mo e comp ehensi e ecological and conse a ion da a.
Acknowledgemen s
We hank Ying-Yong Wang and Shuo Qi om Sun Ya -sen Uni e si y o hei assis ance
in examining he Opis ho opis haihaensis specimen SYS 000537. We hank Jun-Jie
Huang om Chengdu Ins i u e o Biology o pho og aphing he hemipenis. We also hank
Ngoc Quynh Nguyen and Duc T ong Nguyen (SIFASV, Vie nam) o hei assis ance in
p epa ing he igu es and map. We a e since ely g a e ul o he anonymous e iewe o
hei aluable commen s and sugges ions, which helped imp o e he manusc ip . This
esea ch was suppo ed by Science and Technology P ojec s o Xizang Au onomous
Region, China (XZ202301ZY0036G), he Na ional Na u al Science Founda ion o China
(NSFC 31471968), he Ru o d Founda ion (G an No. 45888-2-P; da a analysis), he
Russian Science Founda ion (RSF G an No. 22-14-00037-P: da a analysis) and
Na ional Animal Collec ion Resou ce Cen e , Sun Ya -sen Uni e si y (The Museum o
Biology), Guangzhou, China.
20 Zhang T e al
Re e ences
• B own WC, Le i on AE (1961) Disco e y o he snake genus Opis ho opis in he
Philippine Islands: wi h desc ip ion o a new species. Occasional Pape s o he Na u al
His o y Museum o S an o d Uni e si y.
• Bu b ink FT, Lawson R, Slowinski JB (2000) Mi ochond ial DNA phylogeog aphy o he
poly ypic No h Ame ican a snake (Elaphe obsole a): a c i ique o he subspecies
concep . E olu ion 54: 2107‑2118. h ps://doi.o g/
10.1554/00143820(2000)054[2107:MDPOTP]2.0.CO;2
• Bu land T (2000) DNASTAR's Lase gene Sequence Analysis So wa e. Bioin o ma ics
Me hods and P o ocols 13: 71‑91. h ps://doi.o g/10.1385/1-59259-192-2:71
• Dowling H, Sa age JM (1960) A guide o he snake hemipenis: a su ey o basic s uc u e
and sys ema ic cha ac e is ics. Zoologica: Scien i ic con ibu ions o he New Yo k
Zoological Socie y 45 (2): 17‑28. h ps://doi.o g/10.5962/p.203350
• Gao ZY, Zhang Y, Nguyen VDH, Jiang K, Nguyen TT, Li JT, Ren JL (2024) Redisco e y
and e ision o he diagnos ic cha ac e s o Opis ho opis dao an ieni O lo , Da e sky,
and Mu phy, 1998 (Squama a: Na icidae) om Sou he n Vie nam, wi h no es on i s
hemipenial mo phology and de ensi e beha io . Asian He pe ological Resea ch 16 (1):
49‑65. h ps://doi.o g/10.3724/ah .2095-0357.2024.0048
• Kuma S, S eche G, Li M, Knyaz C, Tamu a K (2018) MEGA X: molecula e olu iona y
gene ics analysis ac oss compu ing pla o ms. Molecula Biology and E olu ion 35:
1547‑1549. h ps://doi.o g/10.1093/molbe /msy096
• Nguyen TV, Nguyen TQ, Pham CT, Ziegle T (2018) Fi s coun y eco d o Opis ho opis
maculosa S ua e Chuaynke n, 2007 om Vie nam. Russian Jou nal o He pe ology 25:
74‑78. h ps://doi.o g/10.30906/1026-2296-2019-25-1-74-78
• Pope CH (1935) The ep iles o China. Tu les, c ocodilians, snakes, liza ds. X. Ame ican
Museum o Na u al His o y., New Yo k. h ps://doi.o g/10.5962/bhl. i le.12104
• Poya ko NA, Nguyen TV, Popo ES, Geissle P, Pawangkhanan P, Neang T,
Suwannapoom C, Ananje a NB, O lo NL (2023) Recen p og ess in axonomic s udies,
biogeog aphic analysis, and e ised checklis o ep iles in Indochina. Russian Jou nal o
He pe ology 30 (5): 255‑476. h ps://doi.o g/10.30906/1026-2296-2023-30-5-255-476
• Ren JL, Wang K, Jiang K, Guo P, Li JT (2017) A new species o he Sou heas Asian
genus Opis ho opis (Se pen es: Colub idae: Na icinae) om wes e n Hunan, China.
Zoological Resea ch 38 (5): 251‑263. h ps://doi.o g/10.24272/j.issn.2095-8137.2017.068
• Ren JL (2019) Sys ema ics and biogeog aphy o Opis ho opis (Se pen es: Colub idae).
Uni e si y o Chinese Academy o Sciences, Chengdu.
• Ren JL, Jiang K, Huang JJ, Da id P, Li JT (2022) Taxonomic e iew o he genus
He pe o eas (Se pen es: Na icidae), wi h he desc ip ion o a new species om Tibe ,
China. Di e si y 14 (2). h ps://doi.o g/10.3390/d14020079
• Ronquis F, Teslenko M, Van De Ma k P, Ay es DL, Da ling A, Höhna S, La ge B, Liu L,
Sucha d MA, Huelsenbeck JP (2012) M Bayes 3.2: E icien Bayesian phylogene ic
in e ence and model choice ac oss a la ge model space. Sys ema ic Biology 61 (3):
539‑542. h ps://doi.o g/10.1093/sysbio/sys029
• Sil es o D, Michalak I (2012) RaxmlGUI: a g aphical on -end o RAxML. O ganisms
Di e si y and E olu ion 12 (4): 335‑337. h ps://doi.o g/10.1007/s13127-011-0056-0
Fi s desc ip ion o he male and hemipenial mo phology o Opis ho opis ... 21
• S ua BL, Chuaynke n Y (2007) A new Opis ho opis (Se pen es: Colub idae: Na icinae)
om No heas e n Thailand. Cu en He pe ology 26 (1): 35‑40. h ps://doi.o g/
10.3105/1345-5834(2007)26[35:anoscn]2.0.co;2
• Teynié A, Lo ie A, DAVID P, Nguyen TQ, Vogel G (2014) A new species o he genus
Opis ho opis Gün he , 1872 om no he n Laos (Squama a: Na icidae). Zoo axa 3774
(2). h ps://doi.o g/10.11646/zoo axa.3774.2.4
• Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG (1997) The
CLUSTAL_X windows in e ace: lexible s a egies o mul iple sequence alignmen aided
by quali y analysis ools. Nucleic Acids Resea ch 25: 4876‑4882. h ps://doi.o g/10.1093/
na /25.24.4876
• Ue z P, F eed P, Aguila R, Hošek J (2025) The Rep ile Da abase. h p://www. ep ile-
da abase.o g.
• Wang J, Lyu ZT, Zeng ZC, Liu ZY, Wang YY (2017) Re-desc ip ion o Opis ho opis laui
Yang, Sung and Chan, 2013 (Squama a: Na icidae). Asian He pe ological Resea ch 8
(1): 70‑74. h ps://doi.o g/10.16373/j.cnki.ah .160032
• Wang J, Lyu ZT, Zeng ZC, Lin CY, Yang JH, Nguyen TQ, Le M, Ziegle T, Wang YY
(2020) Re-examina ion o he Chinese eco d o Opis ho opis maculosa (Squama a,
Na icidae), esul ing in he i s na ional eco d o O. haihaensis and desc ip ion o a new
species. ZooKeys 913: 141‑159. h ps://doi.o g/10.3897/zookeys.913.48622
• Wang YY, Guo Q, Liu ZY, Lyu ZT, Wang J, Luo L, Sun YJ, Zhang YW (2017) Re isions o
wo poo ly known species o Opis ho opis Gün he , 1872 (Squama a: Colub idae:
Na icinae) wi h desc ip ion o a new species om China. Zoo axa 4247 (4): 391‑412.
h ps://doi.o g/10.11646/zoo axa.4247.4.3
• Xiang CY, Gao F, Jako lić I, Lei HP, Hu Y, Zhang H, Zou H, Wang GT, Zhang D (2023)
Using PhyloSui e o molecula phylogeny and ee-based analyses. iMe a 2 (1): e87.
h ps://doi.o g/10.1002/im 2.87
• Yang JH, Wang YY, Zhang B, Lau MWN, Chou WH (2011) Re ision o he diagnos ic
cha ac e s o Opis ho opis maculosa S ua and Chuaynke n, 2007 wi h no es on i s
dis ibu ion and a ia ion, and a key o he genus Opis ho opis (Squama a: Na icidae).
Zoo axa 2785: 61‑68. h ps://doi.o g/10.11646/zoo axa.2785.1.4
• Zhang D, Gao FL, Jako lić I, Zou H, Zhang J, Li WX, Wang GT (2020) PhyloSui e: An
in eg a ed and scalable desk op pla o m o s eamlined molecula sequence da a
managemen and e olu iona y phylogene ics s udies. Molecula Ecology Resou ces 20
(1): 348‑355. h ps://doi.o g/10.1111/1755-0998.13096.
• Zhang FJ, Hu SQ, Zhao EM (1984) Compa a i e s udies and phylogene ic discussions on
hemipenial mo phology o he Chinese Colub inae (Colub idae). Ac a He pe ol Sinica 3:
23‑44. [In Chinese].
• Zhao EM (2006) Snakes o China. I. Anhui Science and Technology Publishing House
Publishing, He ei, 372 pp. [In Chinese].
• Ziegle T, Da id P, Ziegle NT, Pham TC, Nguyen QT, Le DM (2018) Mo phological and
molecula e iew o Jacob’s Moun ain S eam Keelback Opis ho opis jacobi Angel &
Bou e , 1933 (Squama a: Na icidae) wi h desc ip ion o a sibling species om no he n
Vie nam. Zoo axa 4374 (4). h ps://doi.o g/10.11646/zoo axa.4374.4.2
• Ziegle T, Pham CT, Nguyen TV, Nguyen TQ, Wang J, Wang YY, S ua BL, Le MD (2019)
A new species o Opis ho opis om no he n Vie nam p e iously misiden i ied as he
Yellow-spo ed Moun ain S eam Keelback O. maculosa S ua & Chuaynke n, 2007
22 Zhang T e al
(Squama a: Na icidae). Zoo axa 4613 (3): 579‑586. h ps://doi.o g/10.11646/zoo axa.
4613.3.9
Supplemen a y ma e ial
Suppl. ma e ial 1: Unco ec ed pai wise sequence di e gence
Au ho s: Tie ui Zhang, Jinlong Ren, Maoliang Li, Yuhao Xu, Xinge Wang, Nikolay A. Poya ko ,
Tan Van Nguyen, Song Huang
Da a ype: Phylogene ic
B ie desc ip ion: Unco ec ed pai wise sequence di e gence (%) among Cy b m DNA gene
sequences o Opis ho opis species.
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Fi s desc ip ion o he male and hemipenial mo phology o Opis ho opis ... 23
