Contribution to the taxonomy of the genus Lycodon H. Boie in Fitzinger, 1827 (Reptilia: Squamata: Colubridae) in China, with description of two new species and resurrection and elevation of Dinodon septentrionale chapaense Angel, Bourret, 1933

Abstract

While considerable progress has been made in the taxonomic studies of the genus Lycodon in Asia, questions remain to be clarified regarding the taxonomy of certain groups, particularly those containing species in China. Not only do many regions in China remain overlooked by herpetologists, resulting in the possibility of undiscovered new species, but the surveyed areas also have suspicious records of recognized congeners that require taxonomic confirmations. Combining both morphological and genetic data, we tackle these outstanding issues in the taxonomy of Lycodon in China. In particular, we discover two new species of Lycodon: one from the previously neglected hot-dry valley in the northern Hengduan Mountain Region close to Tibet, and another recluse and cryptic species from the L. fasciatus complex in the downtown park of a major city in southern Sichuan Province. Additionally, we clarify the distribution of L. septentrionalis in China and resurrect and elevate its junior synonym subspecies, Dinodon septentrionale chapaense, as a full, valid species, and we synonymize the recently described L. namdongensis to the resurrected L. chapaensis comb. nov .. Lycodon chapaensis comb. nov . thus represents a new national record of reptilian fauna of China. Lastly, based on literature review, we also correct some of the erroneous records of L. fasciatus and L. ruhstrati in China, point out remaining taxonomic issues of the genus for future research, and update the dichotomous key and distribution of the 20 species of Lycodon currently recorded from China.

Full text


ContributiontothetaxonomyofthegenusLycodonH.
BoieinFitzinger,1827(Reptilia:Squamata:
Colubridae)inChina,withdescriptionoftwonew
speciesandresurrectionandelevationofDinodon
septentrionale chapaenseAngel,Bourret,1933
KaiWang1,2,#,*,Zhong-BinYu1,3,#,GernotVogel4,JingChe1,*
1State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming,
Yunnan 650223, China
2Sam Noble Oklahoma Museum of Natural History, Department of Biology, University of Oklahoma, Norman, Oklahoma 73072, USA
3Kunming College of Life Science, University of the Chinese Academy of Sciences, Kunming, Yunnan 650204, China
4Society for Southeast Asian Herpetology, Heidelberg D-69115, Germany

ABSTRACT
While considerable progress has been made in the
taxonomic studies of the genus Lycodon in Asia,
questions remain to be clarified regarding the
taxonomy of certain groups, particularly those
containing species in China. Not only do many
regions in China remain overlooked by
herpetologists, resulting in the possibility of
undiscovered new species, but the surveyed areas
also have suspicious records of recognized
congeners that require taxonomic confirmations.
Combiningbothmorphologicalandgeneticdata,we
tackle these outstanding issues in the taxonomy of
LycodoninChina.Inparticular,wediscovertwonew
species of Lycodon: one from the previously
neglected hot-dry valley in the northern Hengduan
MountainRegioncloseto Tibet,andanotherrecluse
andcrypticspeciesfromtheL. fasciatuscomplexin
the downtown park of a major city in southern
Sichuan Province. Additionally, we clarify the
distribution of L. septentrionalis in China and
resurrectandelevateits juniorsynonymsubspecies,
Dinodon septentrionale chapaense, as a full, valid
species, and we synonymize the recently described
L. namdongensis to the resurrected L. chapaensis
comb. nov.. Lycodon chapaensiscomb. nov. thus
representsanewnationalrecordofreptilianfaunaof
China. Lastly, based on literature review, we also
correctsomeoftheerroneousrecordsofL. fasciatus


Received:29September2020;Accepted:17 December2020; Online:
18December2020
Foundation items:This research is supported by the Second Tibetan
Plateau Scientific Expedition and Research (STEP) program
(2019QZKK0501),Biodiversity Survey and Assessment Projectof the
Ministry of Ecology and Environment (2019HJ2096001006), China's
Biodiversity Observation Network (Sino-BON), Southeast Asia
Biodiversity Research Institute, CAS (Y4ZK111B01:
2017CASSEABRIQG002), and the Animal Branch of the Germplasm
BankofWildSpecies,CAS(LargeResearchInfrastructureFunding)to
J.C.andNSFGRFP2017216966toK.W.
#Authorscontributedequallytothiswork
*Corresponding authors, E-mail: kai.wang-2@ou.edu; chej@mail.kiz.
ac.cn
DOI:10.24272/j.issn.2095-8137.2020.286
Open Access
This is an open-access article distributed under the terms of the
Creative Commons Attribution Non-Commercial License (http://
creativecommons.org/licenses/by-nc/4.0/), which permits unrestricted
non-commercial use, distribution, and reproduction in any medium,
providedtheoriginalworkisproperlycited.
Copyright ©2021 Editorial Office of Zoological Research, Kunming
InstituteofZoology,ChineseAcademyofSciences


Received:29September2020;Accepted:17 December2020; Online:
18December2020
Foundation items:This research is supported by the Second Tibetan
Plateau Scientific Expedition and Research (STEP) program
(2019QZKK0501),Biodiversity Survey and Assessment Projectof the
Ministry of Ecology and Environment (2019HJ2096001006), China's
Biodiversity Observation Network (Sino-BON), Southeast Asia
Biodiversity Research Institute, CAS (Y4ZK111B01:
2017CASSEABRIQG002), and the Animal Branch of the Germplasm
BankofWildSpecies,CAS(LargeResearchInfrastructureFunding)to
J.C.andNSFGRFP2017216966toK.W.
#Authorscontributedequallytothiswork
*Corresponding authors, E-mail: kai.wang-2@ou.edu; chej@mail.kiz.
ac.cn
DOI:10.24272/j.issn.2095-8137.2020.286
Open Access
This is an open-access article distributed under the terms of the
Creative Commons Attribution Non-Commercial License (http://
creativecommons.org/licenses/by-nc/4.0/), which permits unrestricted
non-commercial use, distribution, and reproduction in any medium,
providedtheoriginalworkisproperlycited.
Copyright ©2021 Editorial Office of Zoological Research, Kunming
InstituteofZoology,ChineseAcademyofSciences
ZOOLOGICAL RESEARCH
62 SciencePress Zoological Research42(1):62−86,2021

and L. ruhstrati in China, point out remaining
taxonomic issues of the genus for future research,
and update the dichotomous key and distribution of
the 20 species of Lycodon currently recorded from
China.
Keywords: Erroneous records; Guangdong;
Hunan; Misidentification; Serpentes; Sichuan;
WolfSnake;Yunnan
INTRODUCTION
After major generic revisions (Guo et al., 2013; Siler et al.,
2013), Wolf Snakes of the genus Lycodon Fitzinger, 1826
represent one of the most diverse snake radiations in Asia,
including66 recognizedspecies todate(Ganesh etal.,2020;
Uetzetal.,2020).Majorityofthecurrentlyrecognizedtaxaare
knowninhabitingtropicaltosubtropicalforestsatamid-to-low
elevation (Luu et al., 2018; Vogel & Brachtel, 2008; Vogel et
al.,2009;Wangetal.,2020a),wherespeciesofthegenusare
known to feed heavily on reptiles, particularly on lizards
(Zhang&Wang,2014;Zhaoetal.,1998).
In China, 17 species have been recorded (Janssen et al.,
2020;Wangetal.,2020b),includingL. aulicus(Linnaeus,1758),
L. cathaya Wang, Qi, Lyu, Zeng, Wang, 2020, L. fasciatus
(Anderson, 1879), L. flavozonatus (Pope, 1928), L.
futsingensis (Pope, 1928), L. gongshan Vogel, Luo, 2011, L.
laoensisGünther,1864,L. liuchengchaoiZhang,Jiang,Vogel,
Rao, 2011, L. multizonatus (Zhao, Jiang, 1981), L.
meridionalis (Bourret, 1935), L. pictus Janssen, Pham, Ngo,
Le, Nguyen, Ziegler, 2019, L. ruhstrati (Fischer, 1886), L.
rosozonatus(Hu,Zhao,1972),L. rufozonatusCantor,1842,L.
septentrionalis(Günther,1875),L. subcinctusBoie,1827,and
L. synaptor Vogel, David, 2010. In the Hengduan Mountain
Region (HMR) in Southwest China alone, seven recognized
species have been recorded from Yunnan and Sichuan
Provinces,namelyL. fasciatus,L. gongshan,L. liuchengchaoi,
L. ruhstrati,L. multizonatus,L. septentrionalis,andL. synaptor
(Chenetal., 2018a, 2018b; Guo et al., 2007;Vogel&David,
2010; Vogel & Luo, 2011; Yang & Rao, 2008; Zhang et al.,
2011b; Zhao, 2004; Zhao & Yang, 1997). Of these seven
species found in the HMR, only L. multizonatus is from the
high-elevation regions in the northeast (Lei et al., 2014). As
mostpartsofthe HMR have not been surveyed in details for
herpetological diversity, and given previous studies already
suggestedthatthe northern parts oftheHMRactually harbor
asurprisingnumberofundocumentedreptiliandiversity(Peng
etal.,2014b;Wangetal.,2021),itislikelythatthediversityof
LycodoninthenorthernHMRisalsounderestimated.
In relation to the overlooked diversity, many recognized
specieshaveoutstanding taxonomicissues.Speciescurrently
recorded from HMR are known by having wide distribution
ranges that expand across distinct zoogeographic regions
(Zhao&Adler,1993),particularlyL. fasciatus,L. ruhstrati,and
L. septentrionalis(Zhao,2006;Zhaoetal.,1998). As studies
have suggested that cryptic diversity and misidentification of
recognizedcongenersexplain someoftheexistingsuspicious
records(Vogel&David,2010;Vogel&Luo,2011;Vogeletal.,
2009),thecurrentremainingrecords of these species across
ChinaandSoutheastAsiawarrantfurtherconfirmations.
Inthisstudy, we combined both morphological andgenetic
data to shed lights into the current taxonomy of Lycodon in
China. As results, we discover two new species of Lycodon:
one species from northern HMR that has never been
documented before, and another one from the previously
identified population of L. fasciatus in Panzhihua, Sichuan.
Additionally, we found that the previously identified  “ L.
septentrionalis” in Yunnan Province represent the same
lineage as the recently described species L. namdongensis
from northern Vietnam, and this lineage matches the
diagnosis of an existing synonym, Dinodon septentrionale
chapaense Angel, Bourret, 1933 (=Lycodon septentrionalis
chapaensis after generic revision), which we resurrect and
elevate to full species status. We provide an expanded
description of the poorly known L. chapaensiscomb. nov.
based on additional specimens from China. Furthermore, we
confirm that the questionable records of  “ L. fasciatus” from
Hunan and Guangdong represent misidentifications over L.
liuchengchaoi, and records of  “ L. ruhstrati” in Yunnan
represent clear misidentifications over L. chapaensiscomb.
nov. and L. gongshan. Lastly, we provide an updated
dichotomouskeyanddistribution tothe recognizedspeciesin
China and discuss some remaining taxonomic issues for
futurestudies.
MATERIALS AND METHODS
Taxonomic sampling
Atotalof13specimensandanon-voucheredgenetictissueof
the genus Lycodon were collected from Southwest China
between 2016 and 2020 (Figure 1; Table 1; Appendix I, II).
Liverormuscletissuesweretakenafter the specimens were
euthanized, and the voucher specimens were fixed in 10%
bufferedformalin in the field,transferredto 70% ethanol after
48h for permanent storage, and deposited at the Zoological
MuseumofKunmingInstituteofZoology,ChineseAcademyof
Sciences(KIZ).
Specimens of recognized congeners were examined in
museum collections, including Natural History Museum
(BMNH), California Academy of Sciences (CAS), Chengdu
InstituteofBiology,ChineseAcademyofSciences(CIB),Field
Museum of Natural History (FMNH), KIZ, and Henan
University(HENU)(AppendixII).Forspeciesthatwecouldnot
examineinperson,datawere obtained from literature (Angel
& Bourret, 1933; Boulenger, 1893; Janssen et al., 2019; Luu
et al., 2019; Peng et al., 2014a, 2015, 2017; Vogel et al.,
2009; Wang et al., 2020a; Zhang, 2019). Additional
abbreviations of voucher collections included herpetological
collectionofDr.Guo Peng at Yibin University (GP), Muséum
National d’Histore Naturelle (MNHN), and Vietnam National
University of Forestry (VNUF). Photos of the holotype of
Dinodon septentrionale chapaense were obtained from the
Zoological Research42(1):62−86,2021 63

website of MNHN (https://science.mnhn.fr/institution/mnhn/
collection/ra/item/1 933.11?listIndex=25&listCount=253).
Morphological data
Withtheexceptionof total length, snout–vent length, and tail
length, which were taken using a string and a ruler to the
nearest 1mm, measurements were taken using a digital
caliper to the nearest 0.1mm. Morphometric and pholidosis
charactersand theirmeasurement/countingmethods followed
Wang et al. (2020a) and include: eye diameter (ED), head
length (HL), head width (HW), snout–vent length (SVL), TaL
(Tail length), total length (ToL); supralabial count (SL),
infralabial count (IL), chin shield count (CS), preocular count
(PrO), postocular count (PtO), loreal count (LoR), loreal
enteringorbit(LoR-E),temporalcount(TMP),preventralscale
count(PrV),ventralscalecount(VEN), subcaudalcount(SC),
dorsal scale rows at one head length posterior to the neck
(DSRH), dorsal scale rows at midbody (DSRM), dorsal scale
rowsat one headlengthanterior tothevent (DSRV), number
of maxillary teeth (MT), body scale texture (BST; smooth vs.
keeled), numbers of light bands on the dorsum (NDB; which
excludesthecollar-bandonhead),andnumbersoflightbands
on the tail (NTB). All paired head pholidosis characters were
given in the left/right order. Maxillary teeth formula are
recordedas A-B-C format, where fromleftto right each letter
represents the number of teeth in that specific tooth group
from anterior end to posterior end of maxillary bone, and  “ -”
indicates the presence of a gap. Hemipenis morphology was
describedbased on Dowling & Savage (1960), and the color
descriptionfollowedKöhler(2012)formaximumcomparability.
ForSL,scalecountwasgivenin“A-B-C”format,whereAis
the number of anterior SL that do not enter the orbit, B is
number of SL that enter the orbit, and C is the number of
remainingSLthatareposteriortoanddonotcontacttheorbit.
For IL, scale counts were given in  “A(B)” format, where A is
the total number of IL, and B is the number of IL that are in
contact with the anterior chin shield. For TMP, scale count
wasgiven in“A+B” format, whereAand Barethe numberof
anterior and posterior temporal scales, respectively. For
posterior temporal scale count, paraparietal scale was
included.
Additionally, the following morphological characters were
alsorecorded:verticaleye diameter(VED), measuredlinearly
between superior most and inferior most points of eye;
distance between head and first body cross-band (DHB),
measuredbetweentheposterior meetingpoint ofparietaland
theanterioredgeoffirstdorsalcross-bandalongthevertebral
line;position of first body cross-band (PBB), recorded as the
number of the anterior most ventral scale at which the first
body cross-band is located; paraparietal scale count (PPT),
definedasthenumberofenlargedscalesborderingthepartial

Figure 1 Distributions of focal members of the genus
Lycodon
in Southwest China and nearby countries for this paper
Differentshapesrepresentdifferentspecies,andnumbers indicatedifferentlocalitieswherethespecieshave beenrecorded(detailsseeAppendix
I).Number1alwaysindicatesthetypelocalityofthatgivenspecies.Bifurcatingwhitelinesindicatesympatricdistributionoftwospeciesatthesame
site. Lycodon serratussp. nov. (brown star): Geyading, Deqin County, Yunnan, China. Lycodon obvelatussp. nov.(red star): Panzhihua City,
Panzhihua, Sichuan, China. Lycodon chapaensiscomb. nov.(triangle): the orange triangle indicates the type locality of its junior synonym, L.
namdongensis at Quan Son, Thanh Hoa Province, Vietnam; gray triangle indicates remaining localities of the species. Lycodon fasciatus sensu
stricto(circle);L. gongshan(square);L. multizonatus(trapezoid);L. septentrionalissensustricto(pentagon);andL. synaptor(hexagon).

64www.zoores.ac.cn

Table 1 Samples and their Genbank accession Nos. in the present study

Genus Species VoucherNo. Locality GenBankaccessionNo.
Lycodon chapaensiscomb. nov. KIZ035013 Lushui,NujiangPrefecture,Yunnan,China MW353742
Lycodon chapaensiscomb. nov. KIZ038282 Fugong,NujiangPrefecture,Yunnan,China MW353744
Lycodon chapaensiscomb. nov. KIZ035113 Dulongjiang,NujiangPrefecture,Yunnan,China MW353743
Lycodon chapaensiscomb. nov. KIZ027593 Tengchong,Baoshan,Yunnan,China MW353741
Lycodon chapaensiscomb. nov. KIZ034331 Xichou,WenshanPrefecture,Yunnan,China MW353739
Lycodon chapaensiscomb. nov. KIZ006753 Mengzi,HonghePrefecture,Yunnan,China MW353737
Lycodon chapaensiscomb. nov. KIZ049166 Daweishan,HonghePrefecture,Yunnan,China MW353738
Lycodon chapaensiscomb. nov. KIZ047084 Jingdong,Puer,Yunnan,China MW353740
Lycodon gongshan KIZ035112 Dulongjiang,NujiangPrefecture,Yunnan,China MW353748
Lycodon gongshan KIZ035114 Dulongjiang,NujiangPrefecture,Yunnan,China MW353749
Lycodon gongshan / YunlongNatureReserve,Dali,Yunnan,China MW353747
Lycodon obvelatussp. nov. KIZ040146 Panzhihua,Sichuan,China MW353745
Lycodon septentrionalis CIB117521 Medog,NyinchiPrefecture,Tibet,China MW353736
Lycodon serratussp. nov. KIZ038335 Deqin,Yunnan,China MW353746
Boiga cynodon KU324614 NegrosOccidental,Philippines KC010340
Dasypeltis atra CAS201641 Kabale,Uganda AF471065
Lycodon albofuscus USMHC1457 – KX822584
Lycodon alcalai KU327848 MunicipalityofSabtang,Batanes,Philippines KC010345
Lycodon banksi VNUFR.2015.20 Khammouane,Laos MH669272
Lycodon bibonius KU304589 Cagayan,Philippines KC010351
Lycodon butleri LSUHC9137 Perak,Malaysia KJ607891
Lycodon butleri LSUHC8365 Perak,Malaysia KJ607892
Lycodon capucinus MVZ291704 – MK844523
Lycodon capucinus MVZ291703 – MK844522
Lycodon cathaya SYSr001542 LongshengCounty,Guangxi,China MT602075
Lycodon cavernicolus LSUHC10500 Perlis,Malaysia KJ607890
Lycodon cavernicolus LSUHC9985 Perlis,Malaysia KJ607889
Lycodon chapaensiscomb nov.
(septentrionalis)– Yunnan,China MK201305
Lycodon chrysoprateros KU307720 Cagayan,Philippines KC010360
Lycodon dumerilii PNM7751 – KC010363
Lycodon dumerilii KU305168 – KC010362
Lycodon effraenis LSUHC9670 – KC010376
Lycodon effraenis KU328526 Karome,NakhonSiThammarat,Thailand KC010364
Lycodon fasciatus CAS234875 Mindat,ChinState,Myanmar KC010365
Lycodon fasciatus CAS234957 Mindat,ChinState,Myanmar KC010366
Lycodon fasciatus SYSr001654 Xishuangbanna,Yunnan,China MK201559
Lycodon fasciatus KIZ014125 Xishuangbanna,Yunnan,China MK201557
Lycodon flavozonatus SYSr000640 Huangganshan,Jiangxi,China MK201413
Lycodon flavozonatus HS15101 Huangshan,Anhui,China MK201312
Lycodon gongshan GP3548 Lingcang,Yunnan,China KP901026
Lycodon gongshan GP3547 Lingcang,Yunnan,China KP901025
Lycodon gongshan GP3546 Lingcang,Yunnan,China KP901024
Lycodon gongshan GP3516 Lingcang,Yunnan,China KP901022
Lycodon jara CAS235387 Putao,Kachin,Myanmar KC010367
Lycodon laoensis KU328529 Karome,NakhonSiThammarat,Thailand KC010371
Lycodon laoensis FMNH258659 Salavan,Laos KC010368
Lycodon liuchengchaoi(fasciatus) GP2094 Nanling,Guangdong,China KC733201
Lycodon liuchengchaoi(fasciatus) GP2097 Chebaling,Guangdong,China KC733202
Lycodon liuchengchaoi JK201704 Ningshan,Shaanxi,China MK201563
Lycodon liuchengchaoi SYSr001654 Shennongjia,Hubei,China MK201580
Zoological Research42(1):62−86,2021 65

Continued
Genus Species VoucherNo. Locality GenBankaccessionNo.
Lycodon liuchengchaoi DL14315 – KF732928
Lycodon meridionalis VNUFR.2017.123 ThanhHoa,Vietnam MH669270
Lycodon meridionalis VNUFR.2012.4 BacKan,Vietnam MH669271
Lycodon multizonatus SySr002411 BaishuijiangNationalNatureReserve,Longnan,
Gansu,China MT625863
Lycodon multizonatus KIZ01623 Luding,Sichuan,China KF732926
Lycodon multizonatus(liuchengchaoi) HS11252 Sanjiazhai,Yunnan,China MK201303
Lycodon chapaensiscomb. nov.
(namdongensis)VNUFR.2017.23 ThanhHoa,Vietnam MK585007
Lycodon pictus ZFMK93747 CaoBang,Vietnam MN395830
Lycodon pictus ZFMK93746 CaoBang,Vietnam MN395829
Lycodon rufozonatus LSUMZ44977 – AF471063
Lycodon rufozonatus GP133 Tongjiang,Sichuan,China KC733194
Lycodon ruhstrati GP2243 Ruyuan,Guangdong,China KC733208
Lycodon ruhstrati GP2049 Chebaling,Guangdong,China KC733200
Lycodon ruhstrati GP991 Gongcheng,Guangxi,China KC733197
Lycodon ruhstrati GP285 Junlian,Sichuan,China KC733195
Lycodon ruhstrati SYSr001631 Huangping,Guangxi,China MK201538
Lycodon ruhstrati SYSr001555 Yangmeiao,Guangxi,China MK201521
Lycodon ruhstrati SYSr001309 Jiulianshan,Jiangxi,China MK201473
Lycodon ruhstrati SYSr001275 Jiangshi,Fujian,China MK201467
Lycodon ruhstrati HS12069 Chebaling,Guangdong,China MK201310
Lycodon ruhstrati DL12678 – KF732925
Lycodon sealei KU327571 Palawan,Philippines KC010384
Lycodon sealei KU309447 Palawan,Philippines KC010385
Lycodon semicarinatus – RyukyuArchipelago,Japan AB008539
Lycodon sidiki MZB.Ophi.5980 – KX822583
Lycodon stormi JAM7487 – KC010380
Lycodon striatus FBRC_DNA205 – MK089444
Lycodon subcinctus MVZ291678 Indonesia MK844529
Lycodon subcinctus MVZ291679 Indonesia MK844530
Lycodon subcinctus MVZ291680 Indonesia MK844531
Lycodon subcinctus MVZ291681 Indonesia MK844532
Lycodon subcinctus MVZ291682 Indonesia MK844533
Lycodon subcinctus MVZ291683 Indonesia MK844534
Lycodon subcinctus MVZ291684 Indonesia MK844535
Lycodon subcinctus MVZ291685 Indonesia MK844536
Lycodon subcinctus LSUHC5016 Pahang,WestMalaysia KC010382
Lycodon synaptor GP2188 Yunnan,China KC733204
Lycodon synaptor SYSr001775 Darongshan,Guangxi,China MK201582
Lycodon synaptor SYSr001800 Dawuling,Guangdong,China MK201581
Lycodon synaptor HS13002 Honghe,Yunnan,China MK201309
Lycodon synaptor HS12087 Kunming,Yunnan,China MK201308
Lycodon synaptor HS11006 Mengzi,Yunnan,China MK201304
Lycodon synaptor GP3515 Lingcang,Yunnan,China KP901021
Lycodon synaptor GP3288 Maandi,Yunnan,China KP901020
Lycodon synaptor GP3270 Daweishan,Yunnan,China KP901019
Lycodon synaptor GP3545 Yunnan,China KP901023
Lycodon zawi CAS239944 Kyaukpyu,RakhineState,Myanmar KC010386
Lycodon zawi CAS210323 Thabakesay,Saging,Myanmar AF471040
Newsequencesareindicatedinbold.Speciesnameinparenthesesindicatestheprevioussynonymormisidentifiedspeciesnameusedforthe
givensequence.“/”indicatesnon-voucherdata,“–”indicatesthatinformationisnotavailableorcouldnotbelocated.
66www.zoores.ac.cn

scales on each side, excluding the anterior temporal and
frontalscales; nuchal scale(NS),defined as thetotalnumber
ofsmall nuchal scalesborderingthe posterior endofparietal;
presence or absence of collar-band of occipital head (NCB),
werealsorecorded.
Genetic data
Thegenomic DNA wasextractedfrom liver ormuscletissues
with a standard three-step phenol-chloroform extraction
method (Sambrook et al., 1989). The fragment of the
mitochondrial cytochrome b (cyt b) gene was targeted using
published primers (Burbrink et al., 2000), and PCR and
sequencingprotocolsfollowedWangetal.(2020a).Datawere
filteredandtrimmedmanuallyusingGeneiousv.10.0,andthe
finalsequence foralignmentcontains 1117bp, andallnewly
generatedsequences were depositedinGenBank (accession
No.MW353736–353749;Table1).
In addition, available sequences of congeners were
downloaded from Genbank (Table 1). Boiga cynodon and
Dasypeltis atrawereselectedasoutgroupsfollowingprevious
phylogenetic studies (Lei et al., 2014; Siler et al., 2013).
SequenceswereeditedandalignedusingGeneiousv.10.0.
Both maximum likelihood analyses (ML) and Bayesian
inferences (BI) were conducted on the final cyt b alignment.
Partitioned Bayesian analyses were conducted using
MRBAYESv.3.2.7a(Ronquistetal.,2012)onCIPRES(Miller
et al., 2010). Sequence data was partitioned by three codon
positions, and the best model of nucleotide substitution was
selectedfor each partitionbythe Akaike InformationCriterion
(AIC), implemented in JMODELTEST2 v. 2.1.10 (Darriba et
al., 2012), which was GTR+Γ for all three partitions. Two
independent Markov chain Monte Carlo analyses were run,
eachwith four Metropolis-coupled chains. Bayesian analyses
were run for 90 million generations, with parameters and
topologiessampled every 1 000generations.Stationarity and
convergencewereassessedwithTRACERv.1.6.0(Rambaut
etal., 2013), andthefirst 20% ofsampleswere discarded as
burn-in.
Partitioned Maximum Likelihood analyses were performed
using RAxML-VI-HPC v. 8.2.10 (Stamatakis, 2014) using the
same partition strategy as for the Bayesian analyses. The
most complex model (GTR+Γ) was applied for all the
partitions, with 1000 replicate ML inferences run. Each
inferencewas initiated with arandomstarting tree, and nodal
supportwas assessedwith1000 bootstrappseudoreplicates.
Nodes having ML bootstrap values of 70 and above and BI
posteriorprobabilitiesof0.95andabovewereconsideredwell
supported. Pairwise uncorrected genetic distances were
calculatedusingPAUPv.4.0b10(Swofford,2002).
RESULTS
Molecular results
MLandBIyieldoverallsimilartopology,althoughsomenodes
havedifferent levelof supports(stronglysupported inonebut
not in the other) (Figure 2). Overall, with addition of most
available Indian and Southeast Asian taxa (i.e., L. alcalai, L.
chrysoprateros, L. dumerilii, L. jara, and L. zawi), our
phylogenyshows similar topology astorecent studies for the
well-supported nodes (Luu et al., 2019; Wang et al., 2020a)
(Figure 2). Although the genus Lycodon is still recovered as
monophyletic, the current dataset could not resolve higher
relationships among major clades (polytomy in BI and/or low
bootstrapsupport<60inML).
Thesamplesof“L. septentrionalis”fromChinaandholotype
of L. namdongensis together from a monophyletic clade
(Clade A, 0.93/93), with two distinct, genetically diverged
groups recovered within this clade: first group includes the
samplefromsouthernTibet,whichisclosetoandinthesame
zoogeographic region of the type locality (i.e., Khasi Hills in
East Himalaya) of L. septentrionalis; and the second group
(Clade B, 1.00/100) contains the specimens of “L.
septentrionalis” from western and southern Yunnan and the
holotype of L. namdongensis, with L. namdongensis nested
withinthe Yunnan“ L. septentrionalis”(Figure 2).TheYunnan
populations of  “ L. septentrionalis” show minimal divergence
from the holotype of L. namdongensis (uncorrected genetic
distance 0%–1.8%), but they have considerable divergences
from the Tibetan population of true L. septentrionalis
(5.7%–7.4%)(Table2).
Theputativenewspeciesfrom northern HMR is recovered
sister to L. multizonatus with strong supports (Clade D,
1.00/98),anditshowsaconsiderablegeneticdivergencefrom
L. multizonatus(3.6%–4.0%).The previouslyreported sample
of“L. liuchengchaoi” from Yunnan, China is nested within L.
multizonatus (1.00/99). Samples that are currently identified
as L. fasciatus are polyphyletic, consisting of three major
groups: the first well-supported group (1.00/100) contains
samples true L. fasciatus from Myanmar, southern Yunnan
andwesternYunnan,whichiswithinthecloseproximityofthe
type locality of the species, and this group forms a strongly
supportedcladewithL. butleri, L. gongshan, L. cavernicolus,
andL. sidiki(CladeC;1.00/100), althoughrelationshipswithin
Clade C remain unresolved (Figure 2); the second group
includessamplesof“L. fasciatus”fromGuangdong,whichare
nested within L. liuchengchaoi (Clade E, 1.00/100); and the
third group includes the sample of the putative new species
from Panzhihua, which forms a monophyletic group with L.
synaptor (Clade G, 1.00/100). These three groups are
geneticallydiverged:theGuangdong samplesof “L. fasciatus”
are nearly identical to L. liuchengchaoi (≤0.2%) and show
substantialgenetic divergence from the trueL. fasciatus from
Myanmar and Yunnan (9.4%–11.7%); the Panzhihua sample
is also substantially diverged from the true L. fasciatus
(13.3%–13.6%), and it is also substantially diverged from its
sisterspeciesL. synaptor(8.6%–9.5%;Table2).
Morphological results and taxonomic conclusion
All examined specimens of the currently identified “L.
septentrionalis”fromYunnanhaveoverlappingbodysizesand
tail ratios, same head pholidosis characters, similar dorsal
pholidosis characters, and same body coloration and
Zoological Research42(1):62−86,2021 67

ornamentation with respect to the holotype of L.
namdongensis and the holotype of Dinodon septentrionale
chapaense (Figures 3, 5; Table 3). On the other hand,
specimensof L. septentrionalis from southern Tibet, which is
close to its type locality, differs from the above Yunnan and
Vietnampopulations by havingmultiplerows of keeleddorsal
scales (vs. smooth or only posterior vertebral row feebly
keeled) and different number of maxillary teeth (8 vs. 11 or
12). Such morphological differentiation suggests that the
Yunnan population of  “ L. septentrionalis”, the holotype of L.
namdongensis, and the holotype of D. septentrionale
chapaenserepresentthesamelineage,whichisdifferentfrom
trueL. septentrionalis.
Thespecimenoftheputativenewspeciesfromthenorthern

Figure 2 Phylogenetic trees of the genus
Lycodon
inferred by Bayesian analyses (BI) based on 1 117 bp of mitochondrial gene cyt
b
Bothbootstrapsupports(BS)andBayesianposteriorprobabilities(BPP)areindicatedoneachofthecorrespondingnode.“–”indicatesadifferential
topologybetweenMLand BIresults.Supportvalues forstronglysupported(BS≥70,BPP≥0.95)intraspecificnodeswereomitted,exceptfewcases
wherethenodesarespecificallyreferredintext.

68www.zoores.ac.cn

HMR is morphologically most similar to L. multizonatus (i.e.,
coloration), but it shows morphological differentiations from
the latter and all remaining recognized species, including
differentheadshape, moreIL,moreDSRH,smoothDST,and
distinct ornamentation patterns (details see comparison
section in the taxonomic account below; Figures 4, 5;
Table4).
Thespecimenof  “ L. fasciatus”fromPanzhihua in southern
Sichuan Province is morphologically similar to the true L .
fasciatus,butit canbedifferentiatedfromthe trueL. fasciatus
readily by having a smaller body size, smooth dorsal scales,
fewer infralabials, and a distinct collar-band on neck in adult
(details see comparison section in the taxonomic account
below; Figures 4, 5; Table 4). Additionally, the Panzhihua
specimendiffersfromthetypeofDinodon yunnanensis,which
iscurrentlyconsidered asajuniorsynonymof L. fasciatusbut
was believed to be valid (Vogel & David, 2010; details see
comparisonsinthetaxonomicaccountbelow).
In conclusion, the Lycodon specimens from northern HMR
and from Panzhihua represent two distinct evolutionary
lineages that cannot be assigned to any recognized species.
Hence we describe them as two new species. Additionally,
populations of  “ L. septentrionalis” from Yunnan Province
representthesamelineageasL. namdongensisandDinodon
septentrionale chapaense from northern Vietnam, which are
distinctanddiverged from the true L. septentrionalisfromthe
East Himalaya both morphologically and genetically. We
resurrectD. septentrionale chapaense and elevateitasa full
species, L. chapaensiscomb. nov., and synonymize L.
namdongensis as its junior synonym. The distribution of L.
chapaensiscomb. nov. in Yunnan hence represents a new
nationalrecordofreptilianfaunaofChina.
Taxonomic account
Lycodon chapaensiscomb. nov. (Angel, Bourret, 1933)
(Figures3C–G,5F,G)
ProposedChinese common name:  沙坝白环蛇(Pinyin: Sha
BaBaiHuanShe)
ProposedEnglishcommonname:ChapaWolfSnake
Chresonyms:Dinodon septentrionale chapaense Angel &
Bourret,1933
Dinodon septentrionalis Smith, 1943 (in part); He& Zhou,
2000;Zhangetal.,2002
Dinodon septentrionale: Zhao & Yang, 1997; Zhao et al.,
1998;He&Zhou,2002;Zhao,2006;Yang&Rao,2008
Lycodon septentrionalis: Siler et al., 2013; Guo et al., 2013;
Caietal.,2015(inpart);Jiangetal.,2016;Wangetal.,2020b
Lycodoncf.septentrionalisYangetal.,2019
Lycodon namdongensisLuuetal.,2019
Holotype: MNHN-RA-1 933.001 1, adult female, from 20 km
SW of Lao-Kay (=Lao Cai), Tonkin, Vietnam. Collected by
BourretR.on01July1931.
Additional referred specimens:VNUFR. 2017.23(holotype
ofL. namdongensis) fromNamDong Nature Reserve,Thanh
Hoa Province, Vietnam; KIZ 06753, female from Mengzi,
Honghe Prefecture, Yunnan, China; KIZ 35113, male from
Dulongjiang, Gongshan Prefecture, Yunnan, China; KIZ
035594, male from Yongping, Dali, Yunnan, China; KIZ
027593,malefrom Tengchong,Baoshan,Yunnan,China;KIZ
038282, male from Fugong, Nujiang Prefecture, Yunnan,
China; and KIZ 035045, subadult female from Lushui,
GongshanPrefecture,Yunnan,China.
Diagnosis:Lycodon chapaensiscomb. nov. differs from
congeners by a combination of the following characters: (1)
bodysize large, ToL 691–1 114 mm;(2)taillength moderate,
TaL 17.1%–20.5% ToL; (3) dorsal scale rows 17-17-15,
mostlysmooth, excepttheposterior vertebral row,whichvery
feebly keeled; (4) VEN 200–225; (5) SC 74–84; (6) cloacal
plateentire;(7)lorealshort,notenteringorbit;(8)SL7or8,2-
3-3, 3-2-3, or 2-2-3; (9) IL 8–10, first 4 or 5 in contact with
anterior chin shield; (10) preocular single, in contact with
supraocular and prefrontal; (11) postocular 2; (12) temporal
2+2 or 2+3; (13) paraparietal much enlarged, single; (14)
maxillary teeth 11 or 12, forming four distinct groups
separated by three gaps (3-1-1-6 or 5-1-1-5), fourth and fifth
tooth largest, about 2.5 times larger than first; first gap twice
as wide as between the first two teeth; second gap largest,
Table 2 Uncorrected genetic distance (%) based on 1 117 bp fragment of cyt
b
among selected members of the genus
Lycodon

1 2 3 4 5 6 7 8 9 10 11
1L. serratussp. nov. −
2L. obvelatussp. nov. 11.9 −
3L. septentrionalis 14.3 14.2 −
4L. namdongensis 14.8 14.9 6.6 −
5 “L. septentrionalis” 13.7–15.2 14.3–15.0 5.7–7.4 0.1–1.7 0–1.8
6L. gongshan 10.1–10.6 11.3–12.3 12.5–14.6 12.5–14.5 11.3–14.7 0–2.2
7L. fasciatus 9.9–11.1 13.3–13.6 12.4–14.1 12.9–14.1 12.4–14.4 5.7–8.0 0–2.4
8L. pictus 8.3–8.4 12.2–12.3 13.5–13.7 13.0 12.5–13.3 10.5–11.4 10.7–12.0 0.6
9L. liuchengchaoi 5.2–5.4 11.1–11.5 12.6–14.2 13.2–14.5 12.9–14.6 8.1–10.5 9.4–11.7 7.9–8.4 0–0.2
10 L. multizonatus 3.6–4.0 11.3–12.0 12.5–14.0 13.1–14.3 12.8–14.6 8.2–11.2 9.4–11.7 8.2–8.9 5.4–6.4 0.5–2.2
11 L. synaptor 10.9–12.6 8.6–9.5 12.1–13.8 12.7–14.1 11.4–14.2 8.5–12.1 10.1–12.0 9.4–11.1 10.5–12.5 10.9–12.8 0–3.0
Thediagonalvaluesareintraspecificgeneticdistancesforthespecieswithmultipleavailablespecimens.
Zoological Research42(1):62−86,2021 69

aboutfourtimesas wide as between the first two teeth; third
gap in same width as in first gap; (15) hemipenis single, not
forked at tip, bulbous shaped, with medium sized spines on
distalendofstem,andspinoseand calyculate with spinulate
ridgeson bulb, apicalnude;(16) dorsal Jet Black(Color300)
or dark Indigo (Color 190) in life, with 23–37 white cross-
bandson dorsum, 11–16ontail; (17)cross-bandswithrather
clearly defined edges, not serrated or only slightly serrated,
single scale width dorsally, widen ventrolaterally; and (18)
ventral white, with black transverse bands or irregular
speckles.
Comparisons: Lycodon chapaensiscomb. nov. differs from
trueL. septentrionalisby havingsmoothor onlyfeeblykeeled
vertebral scale row on posterior body (vs. much more

Figure 3 Comparisons between true
Lycodon septentrionalis
(A, B);
L. namdongensis
(C), and Yunnan specimen of “
D. septentrionalis
”
(D–G)
A, B: Closeup and dorsolateral overview of a un-vouchered individual from Medog, Tibet, China; C: Holotype of L. namdongensis (VNUF R.
2017.23) from Quan Son, Thanh Hoa, Vietnam; D: Un-vouchered individual from Daweishan Nature Reserve, Honghe Prefecture, Yunnan; E:
Vouchered adult male from Yongping County, Dali, Yunnan, China (KIZ 035594); F: Vouchered adult male from Tengchong County, Baoshan,
Yunnan(KIZ027593);G:HemipenisofKIZ027593after preservation.Photos ofL. namdongensistaken fromLuu etal. (2019),remaining photos
byChaoWu,KaiWang,ShaobinHou,WeiliangXie,andZhongbinYu.

70www.zoores.ac.cn

distinctivelykeeled onmedial3–5 rows),more maxillaryteeth
(11or 12vs.8), anddifferentmaxillary teethformula(3-1-1-6
or5-1-1-5, forthandfifth teethlargest,second gapwidest,as
fourtimesofdistance as in between first two teeth vs. 4-2-2,
last two teeth largest, two gaps about same length, as twice
asinbetweenfirsttwoteeth).
Additionally, L. chapaensiscomb. nov. further differs from
L. butleri, L. cavernicolus, L. davisonii, L. dumerilii, L.
fasciatus, L. gibsonae, L. gongshan, L. gracilis, L.
liuchengchaoi, L. multizonatus, L. nympha, L. orientalis, L.
philippinus, L. pictus, L. sealei, L. sidiki, L. subcinctus, L.
submaculatus,andL. tristrigatus byhaving lorealnotentering
orbit (vs. entering); from L. albofuscus, L. aulicus, L.
capucinus, L. flavicollis, L. flavomaculatus, L. hypsirhinoides,
L. jara,L. kundui, L. laoensis, L. mackinnoni, L. meridionalis,
L. stratus, L. tessellatus,L. tiwartii, and L. zawi by having a
singlecloacal plate (vs. divided); fromL. alcalai, L. banksi, L.
cathaya, L. bibonius, L. cardamomensis, L. carinatus, L.
chrysoprateros, L. davidi, L. effraenis,L. fausti, L. ferroni, L.
flavozonatus, L. futsingensis, L. gammiei, L. kundui, L.
muelleri, L. multifasciatus, L. rosozonatus,L. rufozonatus, L.
ruhstrati,L. semicarinatus,L. solivagus,L. stormi,L. synaptor,
L. travancoricus,L. zoosvictoriaebyhaving alargermaximum
body size (ToL >1 000 mm vs. <1 000 mm); and from L.
paucifasciatusbyhavinglowernumberofdorsalscalerowsat
midbody (17 vs. 19); and from L. ophiophagus by having a
shortertail(TaL17.1%–18.4%vs.20.1%–22.8%).
Description of L. chapaensis comb. nov. based on
holotype and referred materials:LargeLycodon,maximum
ToL 1 114 mm; tail moderate, TaL 17.1%–20.5% ToL; head
oval, rather wide, moderately distinct from neck; eye large,
ovalinshape,notlaterallycompressed.Rostrallarge,broader
than height, pentagonal, visible from above; nasal divided,
anteriorhalf borderingrostral,first supralabial,and internasal,
posterior half bordering first and second supralabials, loreal,
internasal, and prefrontal; internasal paired, roughly
rectangular,widerthanlongorsubequalto,muchsmallerthan
prefrontals;prefrontalpaired, hexagonal, bordering preocular,
supraocular,and frontal posteriorly; lorealrathershort, longer
than wide, bean-shaped or spear-shaped, separated from
orbitbypreocular andthirdsupralabial;preocularsingle,taller
than wide; supralabials 8 (rarely 7), third to fifth or third and
fourthenteringorbit;postocular2,superioronelarger;anterior
temporal 2, superior one longer and narrower, inferior one
shorter and wider; posterior temporal 2 or 3 (including
paraparietal); frontal pentagonal, spear-like tip pointing
posteriorly; supraocular elongated; parietal paired, inlaying
spear tip of frontal anteriorly, bordering supraocular and
superior postocular anteriorly, paraparietal and 1–4 nuchal
scales posteriorly; paraparietal single, much enlarged and
elongated. Infralabials 8–10, anterior most pair enclosing
mental and meeting medioposteriorly; 5 or 6 infralabials
borderingchinshields,first to fouth or fifth bordering anterior
chin shield, fifth or sixth bordering posterior chin shield,
respectively; anterior chin shield much longer, forming V-
shape, inlaying tip of first pair of infralabials anteriorly;
posteriorchinshieldsslightlysmaller,notseparatedfromeach
other by distinct mental groove. Dorsal body scales smooth,

Figure 4 The dorsolateral view (1) and ventral view (2) of the holotype of
Lycodon serratus
sp. nov. (KIZ 038335) (A) and
L. obvelatus
sp.
nov. in life (KIZ 040146) (B) (Photos by Wen-Jie Dong and Kai Wang)

Zoological Research42(1):62−86,2021 71

except vertebral row that only feebly keeled toward very
posteriorportioninsomeindividuals; dorsal scale rows 17 at
one head-length posterior to neck, 17 at midbody, 15 one-
head length anterior to vent. Preventral 1 or 2; ventrals
200–225, angulate; cloacal plate entire; subcaudal paired,
74–85 excluding tail tip. DHB 4.5%–7.9% SVL,PBB at
15th–22thventralscale.
Maxillary teeth 11 or 12, forming four distinct groups
separatedbythreegaps.Firstthreeorfiveteethinfirstgroup,
gradually enlarged; single tooth in second and third group,
respectively, both significantly enlarged (about 2.5 times of
second tooth); remaining six or four teeth in last group,
graduallydecreaseinsize, eventuallyabout thesamesizeas
firstor second tooth. Firstgaptwice as wide asregularwidth
betweenfirsttwoteeth; second gap largest, about four times
wider as regular; last gap about same as in first gap. Most
teethcurvedposteriorlytowardstip,exceptfirsttwoorthree.
Hemipenis morphology based of KIZ 027593: hemipenis

Figure 5 Comparisons of (1) dorsal overview, (2) ventral overview, (3) lateral head, (4) dorsal head, (5) ventral head, and (6) dorsum
close-up among
Lycodon serratus
sp. nov. (KIZ 038335; holotype) (A),
L. obvelatus
sp. nov. (KIZ 040146; holotype) (B),
L. multizonatus
(KIZ 01623; topotype) (C),
L. fasciatus
(KIZ 74II0262) (D),
L. gongshan
(KIZ 730 034; holotype) (E), “
L. septentrionalis
” (KIZ 035594; from
Dali, Yunnan, China) (F),
L. chapaensis
comb. nov. (MNHN-RA-1 933.001 1, holotype; from Chapa, Tonkin, Vietnam) (G), and
L.
septentrionalis
(CIB 117 521; from Medog, Tibet, China) (H) (Photos of the holotype of
L. chapaensis
comb. nov. are obtained from the
website of Muséum National d’Histore Naturelle, remaining photos by Zhong-Bin Yu and Jin-Long Ren)

72www.zoores.ac.cn

single, bulbous shape, with single sulcus spermaticus; rather
short, reaching only fifth caudal scale from cloaca when
everted,lengthunknownatretractedstate;proximal1/4length
with some shallow transverse flounces; middle 1/4 densely
covered with medium sized spines; distal bulbous structure
large, about 1/2 of total length, spinose toward basal end,
gradually transition to calyculate with spinulate ridges toward
2/3 of bulbous, and eventually back to flounced toward very
tip;apicalnude(Figure3).
Coloration:ThedorsalsurfacesoftheheadandbodyareJet
Black (Color 300) or sometimes dark Indigo (Color 190). A
single white collar-band is present on the occipital region of
juveniles,butnotinadults.Whitecross-bandsaresingle-scale
broaddorsally andwideninto triangularshape ventrolaterally.
A total of 23–37 cross-bands are present on the body and
11–16onthe tail.Theventralsurfaceofthehead andbodyis
white,withsomeMediumNeutralGray(Color298)patcheson
the anterior infralabials and the gular region. The ventral
surfaceofthe body is white toLightBuff(Color 2), with Dark
Neutral Gray (299) to Jet Black (Color 300) cross-bands,
transverse groups of speckles, or random speckles. Ventral
surfaceofthetailisnearlycompletelyDarkNeutralGray(299)
to Jet Black (Color 300), with white to Light Buff (Color 2)
cross-bands, transverse groups of speckles, or random
speckles.
Natural history:Lycodon chapaensiscomb. nov. inhabits
subtropical and tropical evergreen and sometime mixed
forests(i.e.,withplanted coniferous trees in Dali, Yunnan) at
mid to low elevation (from 616 m at Nam Fong Nature
Reserve,QuanSonDistrict,ThanhHoa Province,Vietnam,to
2 030 m at Dahaoping, Tengchong, Yunnan, China). The
speciesis nocturnal,whereall individualswerefound atnight
actively foraging when collected in China. Unlike other
congenersthat feed heavilyonreptiles, L. chapaensiscomb.
nov. have been reported to feed mostly on rodents, and
sometimes frogs (Yang & Rao, 2008; Zhao & Yang, 1997).
Yang & Rao (2008) stated the specimens from Yunnan are
oftenfound in areasnearagriculture fieldswhererodentsare
abundant, and individuals from Lushui in western Yunnan
wereobservedactivelyhuntingforrodentsinvillagehouses.
Table 3 Comparisons between holotype of
Dinodon septentrionale chapaense
, holotype of
Lycodon namdongensis
, true
L.
septentrionalis
from Tibet, and “
L. septentrionalis
” from Yunnan Province

Species Lycodon septentrionalis Dinodon septentrionale
chapaense L. namdongensis “L. septentrionalis”
VoucherNo. CIB117521,CIBM20150607 MNHN-RA-1933.0011(holotype) VNUFR.2017.23(holotype) KIZ006753,027593,035594,
038282,034331
Samplesize 2(1) 1 1 5
ToL 1187 1051+ 723 691–1114
SVL 945–955 890 575 564–915
TaL 242 161(incomplete) 148 127–199
TaL/ToL 20.4% – 20.5% 17.1%–18.4%(average17.6%)
DSRH 17 17 17 17
DSRM 17 17 17 17
DSRV 15 15 15 15
SL 8(2-3-3) 8(2-3-3) 8(2-3-3) 7or8(2-3-3,3-2-3,or2-2-3)
IL 9(4) 9(5) 10(5) 8–10(first4or5incontactwith
anteriorchinshield)
PrO 1 1 1 1
PtO 2 2 2 2
LoR 1 1 1 1
LoR-E No No No No
TMP 2+3 2+3 2+3 2+2or2+3
VEN 207–212 224 218 200–225
SC 78 56+ 85 74–84
CloacalPlate Entire Entire Entire Entire
MT 8 – 12 11or12
NCB Absence Absence Absence Absence
NDB 33–35 28 23 23–37
NTB 19 11 14 11–14
BST Keeled Smooth Smooth Smoothoronlyposterior
vertebralrowfeeblykeeled
Abbreviationsareexplainedinmethods.“–”indicatesdatanotavailable.ThenumberinparenthesesfortheSamplesizerowindicatedifferential
samplesize fortotal length andtail lengthdueto theincomplete tailofone ofthe specimen (CIBM20150607). Exceptthe holotype ofL.
namdongensis,allremainingspecimenswereexaminedinpersonbyauthors.
Zoological Research42(1):62−86,2021 73

Currently L. chapaensiscomb. nov. has been recorded
fromwestern(GaoligongMountainRange,includingBaoshan,
Dali, Dehong Prefecture, and Nujiang Prefecture), south
central (Puer and Lincang Prefectures), and southeastern
(Honghe and Wenshan Prefectures) Yunnan Province in
China and Lao Cai and Thanh Hoa Provinces in northern
Vietnam (Luu et al., 2019). Based on the reported
questionable records of  “ L. septentrionalis” in eastern
Myanmar, northern Laos, and northern Thailand, it is likely
thatthespeciesisalsofoundinthesecountriesaswell(see
Discussionbelow).
Lycodon serratussp. nov.(Figures4A,5A;Table2)
ZooBankLSID:355B3EDA-546E-417B-9E16-7BC92789DE81
ProposedChinese common name:  “ 锯纹白环蛇” (Pinyin:Ju
WenBaiHuanShe)
ProposedEnglishcommonname:Serrate-bandedWolfSnake
Holotype:KIZ038335,adultmale,collectedbyZhong-BinYu
and Wen-Jie Dong on 25 July 2020 from the Jinsha River
Valley near Geyading Village, Deqin County, northwest
Yunnan Province, China (N28.7720º, E99.1128º, WGS84,
elevation2200ma.s.l.).
Etymology: The Latin species name serratus means
“serrated”, which describes the diagnostic narrow, serrated
blackcross-bandofthenewspecies.
Diagnosis:Lycodon serratussp. nov.canbediagnosedfrom
recognized congeners by a combination of the following
morphological characters: (1) body size moderate, slender,
ToL628 mm, SVL480mm;(2) tail long,TAL23.6% ToL; (3)
head flat, distinct from neck, snout narrow; (4) eye large,
vertical ellipse in shape; (5) dorsal body scales smooth, 19
rowsatone-head-lengthbehindtheneck,17rowsatmidbody,
and15 rowsatone-head-length beforevent;(6) ventralscale
Table 4 Comparison of key morphological characters between the holotypes of
Lycodon serratus
sp. nov.,
L. obvelatus
sp. nov., and
morphologically similar congeners that are also found in the Hengduan Mountain Region (i.e.,
L. gongshan
,
L. fasciatus
,
L. multizonatus
,
and
L. liuchengchaoi
)

Species
Lycodon
serratus
sp. nov.
L. obvelatus
sp. nov. L. multizonatusL. gongshan L. fasciatus L. liuchengchaoi
Sex M M M M F M F M F
Samplesize 1(holotype) 1(holotype) 4 2 2 1 3 2(1) 5(3)
ToL 628 551 440–520 928–963 753–1 003 696 418–613 747 389–615
SVL 480 447 350–428 691–740 589–798 553 341–493 595–676 309–481
TaL 148 104 90–96 223–237 164–207 143 71–120 152 80–138
TaL/ToL 23.6% 18.9% 17.7%–20.5% 23.1%–23.2% 21.8%(n=1) 20.5% 17.0%–19.6% 20.3% 20.6%–24.8%
DSRH 19 17 17 17 17 17 17 17 17
DSRM 17 17 17 17 17 17 17 17 17
DSRV 15 15 15 15 15 15 15 15 15
SL 8or9(2-3-3
or2-4-3) 8(2-3-3) 8(2-3-3) 8(2-3-3) 8(2-3-3) 8(2-3-3) 8(2-3-3) 7(2-3-2)or8
(2-3-3) 8(2-3-3)
IL 10(5) 8(4or5) 8(4) 9(4) 9(4) 9(5) 9(4)or9(5) 8(4) 8(4)
PrO 1 1 1 1 1 1 1 1 1
PtO 2 2 2 2 2 2 2 2 2
LoR 1 1 1 1 1 1 1 1 1
LoR-E Yes Yes Yes Yes Yes Yes Yes/No(*) Yes Yes
TMP 2+2 2+2or2+3 2+3 2+2or2+3 2+2 2+2 2+2or2+3 1+2or2+2 1+2,2+2,or
2+3
VEN 198 199 191–195 210–212 209–215 198 205–211 202–204 200–228
SC 84 76 63–75 94–96 92(n=1) 84 66–95 68–69+ 75–81
Cloacalplate Divided Entire Divided Entire Entire Entire Entire Divided Divided
MT 12(6-1-1-4or
6-1-2-3) 11(7-1-1-2) 11(nodistint
gap)(n=2)
10(7-1-2)or
11(7-2-2)
11(5-3-3)
(n=1) 12(8-2-2) 12(8-2-2) 8 –
CBinadults Presence Presence Presence Absence Absence Absence Absence Presence Presence
NDB 66 31 55–62 37or38 32–36 34 31–37 40–45 33–45
NTB 26 13 11–19 15or16 13(n=1) 16 11–17 10–15 11–13
BST Smooth Smooth Keeled Keeled Keeled Keeled Keeled Keeled Keeled
Abbreviationscanbefoundinmethods.“–”indicatesnotavailableduetoincompletetail.“*”indicatesonlylorealsofasinglespecimen(KIZ75I473)
donotenter orbit.The numberinparenthesesforthe SampleSizerowindicatesdifferential samplesize fortotallengthandtail lengthforL.
liuchengchaoi.DataforthefemaleofL. gongshanandforallL. liuchengchaoiwereobtainedfromliterature(L. gongshan:Vogel&Luo,2011;L.
liuchengchaoi:Pengetal.,2014,2017,2018;Zhangetal.,2011b;Zhangetal.,2019).M:Male;F:Female.
74www.zoores.ac.cn

count 198; (7) subcaudal scale count 84; (8) cloacal plate
divided;(9)supralabials8or9,2-3-3or2-4-3;(10)infralabials
10(5); (11) preocular single, postocilar 2; (12) loreal scale
entering orbit; (13) enlarged paraparietal 2, bordered by 7
additional nuchal scales other than posterior upper temporal
andparietal; (14)maxillary teeth12in fourgroups(6-1-1-4or
6-1-2-3),sixth and seventhmuchlarger, first andsecondgap
about same size, twice as distance between first two teeth;
(15)dorsal surface dirty Tawny Olive (Color 17) with narrow,
stronglyserratedJetBlack(Color300)cross-bands,two-scale
broad middorsally at anterior 1/7 of body, single-scale broad
for the remaining ones; (16) single collar-band on neck, 66
cross-bandsondorsum,26ontail;(17)firstdorsalcross-band
at 10th ventral scale; (18) anterior 1/3 of ventral surface
uniform white, remaining 2/3 of ventral body and whole tail
speckledwithDarkNeutralGray(Color299).
Comparisons:Lycodon serratussp. nov. is morphologically
mostsimilarandcloselyrelatedtoL. multizonatus,whereboth
species have divided cloacal plate, large eyes, and similar
number of black bands across the body. However, the new
species can be differentiated from L. multizonatus by having
moreDSRH(19vs.17),moreIL(9or10 vs.8), moreIL-aCS
(5vs.4),aflatterheadthatisdistinctfromtheneck(vs.robust
and indistinct), a narrower snout (vs. wide), much narrower
black bands on the middle to posterior body (mostly single
scale broad, rarely two vs. ≥3 scales), more black bands on
the tail (26 vs. 11–19), and a distinct ventral ornamentation
patterns(irregularspecklesvs. regularlypaired blackspotsor
completeblackbands)(Figure5).
Forremainingspecies that are foundintheclose proximity
inthe HengduanMountainRegion(L. fasciatus, L. gongshan,
L. liuchengchaoi, L. ruhstrati, L. chapaensiscomb. nov.,and
L. synaptor), L. serratussp. nov. differs from all by having
more DSRH (19 vs. 17), smooth dorsal scales (vs. feebly or
distinctively keeled medially), a narrow snout (vs. robust and
wide), larger and laterally compressed eyes (ED 15.2% HL,
VED 17.5% HL vs. not laterally compressed, <12%), more
cross-bandsonthebodyandtail(66onbody,26ontailvs.L.
fasciatus19–37 onbody, 7–21ontail; L. gongshan 32–40on
body, 13–15 on tail; L. liuchengchaoi 40–45 on body, 10–15
on tail; L. ruhstrati 33–46 on body, 14–28 on tail; L.
chapaensiscomb. nov. 23–37 on body, 11–16 on tail; L.
synaptor 30 or 31 on body, 9 on tail), different-shape and
width of the bands (narrow (mostly single-scale broad) and
stronglyserratedvs.broader(mostlytwo-tothree-scalebroad
and less serrated) and less serrated), and a distinct body
coloration(TawnyOlive (Color 17) with Jet Black (Color 300)
bandsvs.JetBlack(Color300)withwhiteoryellowishbands).
Additionally, L. serratussp. nov. differs from L. fasciatus,L.
gongshan,andL. ruhstratibyhavingdividedcloacalplate(vs.
entire), presence of neck collar-band in adult (vs. absence),
and a distinct ventral ornamentation pattern (randomly
speckled vs. regular transverse bands (L. fasciatus, L.
gongshan) or mostly uniform white (L. ruhstrati and L.
chapaensiscomb. nov.)); from all but L. fasciatus and L.
chapaensiscomb. nov.byhavingmoreIL-aCS(5vs.4);from
all but L. chapaensiscomb. nov. by having non-overlapping
SC (84 vs. L. gongshan 92–96; L. liuchengchaoi 68–77; L.
ruhstrati97–114; L. synaptor68 or69);from L. liuchengchaoi
byhaving fewer VEN (198 vs. 202–206); from L. chapaensis
comb. nov. and L. synaptor by having loreal entering orbit
(vs.separated)andadividedcloacalplate(vs.entire).Forthe
junior synonym of L. fasciatus that is currently available, L.
serratus differs from Dinodon yunnanensis Werner, 1922 by
moreDSRH (19vs.17), smoothorfeebly keeleddorsalbody
scale rows (strongly keeled), a divided cloacal plate (vs.
entire),andmorecross-bandsonthedorsum(66vs.23).
Forremainingthreespecies thathave geneticdataandare
in the same clade (i.e., L. butleri, L. pictus, and L.
cavernicolus),L. serratussp. nov.differsfromallbyhavinga
distinctbodycoloration(TawnyOlive(Color17)withJetBlack
(Color 300) bands vs. Jet Black (Color 300) with white or
yellowishbands). Additionally,thenew speciesdiffers fromL.
pictusby moreDSRH(19 vs. 17),adivided cloacalplate(vs.
entire), more cross-bands on the body (66 vs. 28 or 29) and
tail (26 vs. 13), much narrower cross-bands (mostly single
scale-broad, rarely two vs. 2–4 scale broad), and by the
presence of collar band in adult (vs. absence); from L.
cavernicolus by having smooth dorsal scales (vs. keeled),
fewerSL(8 vs. 9 or 10), and more NDB (66 vs.36–45);and
from L. butleri by having smooth dorsal scales (vs. keeled)
andadividedcloacalplate(vs.entire).
For all the remaining 55 species of the genus, L. serratus
sp. nov.differsfrom all by having a distinct dorsal coloration
(Tawny Olive (Color 17) with Jet Black (Color 300), strongly
serratedbandsvs.blackorbrownishwithwhite,yellow,orred
cross-bands that are less serrated or smooth, or with no
complete cross-bands but reticulated ornamentations).
Additionally, L. serratussp. nov. differs from all except 18
species(i.e.,L. cardamomensis,L. carinatus,L. flavozonatus,
L. futsingensis, L. hypsirhinoides, L. jara, L. laoensis, L.
mackinnoni,L. meridionalis,L. nympha,L. orientalis,L. sealei,
L. septentrionalis,L. sidiki,L. striatus,L. tessellatus,L. tiwarii,
andL. zawi)byhavingadividedcloacalplate(vs.entire).For
the excluded 17 species, the new species differs from L.
cardamomensis,L. carinatus,L. flavozonatus,L. meridionalis,
L. nympha, L. sealei, and L. sidiki by having smooth dorsal
scales(vs.keeled); from L. futsingensis, L. hypsirhinoides, L.
jara,L. laoensis,L. mackinnoni,L. striatus,L. tessellatus,and
L. zawibyhavingmoreDSRH(19vs.17);fromL. orientalisby
the presence of preocular scale (vs. absence); and from L.
tiwariibyhavingfewerventralscales(198vs.218–237).
Description of holotype: KIZ 038335, Adult male, medium
sized Lycodon, SVL 480mm, TaL 148mm. Body slender; tail
long, TaL 23.6% of ToL; head elongated, flat, snout narrow,
HW9.7mm,HL 12.2mm,distinctfromneck;eyelarge,slightly
compressedlaterally,ED2.4mm,VED2.8mm,ED19.9%HL,
VED 23.0% HL; pupil vertically oriented. Rostral pentagonal,
broader than height, visible from above; nasal laterally
elongated, divided, anterior one bordering rostral, first
supralabial, and internasal, posterior one bordering first and
second supralabials, loreal, internasal, and prefrontal;
Zoological Research42(1):62−86,2021 75

internasal pentagonal; prefrontal paired, hexagonal, larger
than internasal; loreal elongated tear shape, entering orbit,
bordering posterior nasal, prefrontal, second and third
supralabials,andpreocular; preocularsingle;supralabials8/9,
thirdtofifthenteringorbitonleft,thirdtosixthenteringorbiton
right; postocular 2; temporal 2+2 (including paraparietal),
inferioroneoffirstpairmuchlarger;frontalpentagonal,spear-
like tip pointing posteriorly; supraocular elongated; parietal
paired in V-shape, inlaying spear-tip of frontal anteriorly,
bordering supraocular and superior postocular anteriorly,
paraparietal, and 7 nuchal scales posteriorly; paraparietal
single, enlarged. Infralabials 10/10, anterior most pair
enclosing mental and meeting medialposteriorly; anterior 5
infralabials bordering anterior chin shield on both sides, fifth
andsixthborderingposteriorchinshieldonbothsides;2pairs
ofchinshield,anteriorpairmuchelongated,meetingmedially,
forming V-shape and inlaying meeting tip of first pair of
infralabialsanteriorly;posteriorchinshieldmuchnarrowerand
shorter, separated from each other by rather wide section of
mental groove. Dorsal body scales smooth, 19 rows one-
head-length behind neck, 17 rows midbody, 15 rows one-
head-length before vent. Single preventral; ventral 198,
angular;cloacal platedivided;subcaudal paired,84 excluding
tailtip.DHB10.4mm,2.1%SVL;PBBat5thventralscale.
Maxillary teeth 12 (fifth lost), forming 4 distinct groups
separatedbythreegapsonbothsides.Sixteeth(firsttosixth)
infirstgroup:first four gradually increase in size, followed by
much enlarged sixth; single (seventh) tooth in second group,
alsomuchenlarged,samesizeassixth; single (eighth) tooth
inthirdgroup,samesizeasfourth;lastfour(ninthtotwelfth)in
lastgroup,ninth and tenth same sizeasfourth,eleventhand
twelfth same as second. Three gaps present between sixth
and seventh teeth (about 1.8 times regular width), seventh
and eighth (twice regular width), eighth and ninth (twice
regularwidth).
The hemipenis only partially everted, single; very proximal
endfree of spines; remainingpartfilledwith small to medium
sizedspines.
Coloration: In life, the background coloration of the dorsal
and lateral surfaces of the head and body is Tawny Olive
(Color 17). The dorsal surface of the head is speckled with
DarkNeutralGray(Color299),particularlyonthe frontal and
parietalscales.Asingle Jet Black (Color 300) collar-band on
neck, 66 cross-bands of the same color are present on the
dorsum, and another 26 cross-bands are present on the tail.
Cross-bands on the anterior 1/7 of SVL are broader,
expending across two dorsal scales in width, and the
remaining bands are rather narrow, expanding only a single
dorsal scale in width. All bands are strongly serrated. The
immediateborderingmarginsofeachblackbandarePaleBuff
(Color1).Ventralsurfaceof thehead iswhite.Theimmediate
bordering regions between infralabials and between
infralabials and chin shields are speckled with Dark Neutral
Gray (Color 299). The ventral surface of the body and tail is
white: the anterior 1/7 of the SVL is uniform white with no
patterns, and the remaining section of the ventral body and
the tail is speckled with Dark Neutral Gray (Color 300), with
the tail more heavily speckled. Coloration and ornamentation
patternsremainmostly thesameaftershort-termpreservation
(onemonth).
Distribution, natural history and conservation:Besides L.
multizonatus,L. serratussp. nov.istheonlyknownspeciesof
thegenusthatinhabitshot-dryvalleyhabitatsathighelevation
inthe northern HMR. Currently L. serratussp. nov. is known
from the type locality in Yunnan Province only, however, an
individual of the same species was photographed but not
captured in Derong County of Sichuan Province, which is
about21kmlinearlysoutheastfromthetypelocality(personal
communication with Mr. Di-Hao Wu). The habitats consist of
open rock outcrops and low bushes, and the annual
precipitation is very low (Figure 6A). While the distribution
range of the species remains unknown, habitat destructions
from road constructions were observed at and near the type
locality of the new species (Wang et al., 2021). We
recommend Data Deficient (DD) for the IUCN status of the
newspecies,andwe call for population studies to assess its
conservationstatusinthenearfuture.
The new species is sympatric with Diploderma sp., Gekko
scabridus,andScincella monticola(Wangetal.,2021;Yang&
Rao, 2008), and the holotype of L. serratussp. nov. was
found at night searching for food on a bush, where several
individualsof Diploderma sp.weresleeping on.Asthegenus
Lycodon is known to feed predominantly on lizards (Zhao et
al., 1998), it is likely that these sympatric lizard species
constitutemainpreysof thenew species.Otherherpetofauna
that are sympatric with the new species include Elaphe
carinata, E. taeniura, Protobothrops xiangchengensis,
Amolops jingshaensis,Bufo gargarzans,andScutigersp..
Lycodon obvelatussp. nov.(Figures4B,5B;Table4)
ZoobankLSID:D15F4F07-FADF-43D5-94B8-EA746655727B
Chresonyms:Lycodon fasciatus:Dengetal.,1991;Wuetal.,
1997;Zhaoetal.,1998;Zhao,2002,2004,2006
ProposedChinese common name:  隐士白环蛇(Pinyin: Yin
ShiBaiHuanShe)
ProposedEnglishcommonname:RecluseWolfSnake
Holotype:KIZ040146,adultmale,collectedbyKaiWangand
Ben-FuMiaofrom Panzhihua City Park, Panzhihua,Sichuan,
China (N26.5751º, E101.7174º, WGS84, elevation 1 243 m
a.s.l.)on19April,2018.
Etymology: The Latin species name, obvelatus, means
“hidden” or  “concealed”, which not only describes the
taxonomic confusions of the cryptic new species over L.
fasciatus, but it also highlights the fact that new species can
behiddeneveninmajorurbanareas.
Diagnosis:Lycodon obvelatussp. nov. can be diagnosed
fromcongenersby a combination of the following characters:
(1) body size small, ToL 551 mm; (2) tail moderate, TaL
18.9% ToL; (3) dorsal scale rows 17-17-15, all smooth; (4)
VEN 199; (6) SC 76; (7) cloacal plate entire; (8) loreal long
and narrow, entering orbit; (9) SL 8, 2-3-3; (10) IL 8(4 or 5);
76www.zoores.ac.cn

(11) preocular single, in contact with supraocular and
prefrontal; (12) postocular 2; (13) temporal 2+2 or 2+3; (14)
paraparietal enlarged, single; (15) frontal bordering 4 nuchal
scales; (16) maxillary teeth 11 in four groups (7-1-1-2),
seventh largest, first gap widest, four times wide as distance
between first two teeth; (17) hemipenis single clavate, nip at
distal end, spinose except very proximal end; spines larger
toward proximal end; (18) distinct collar band present on
occipitalhead,SalmonColor(Color251);(18)dorsalJetBlack
(Color 300) in life, with 31 Salmon Color (Color 251) cross-
bands on dorsum, 13 on tail; (19) cross-bands with serrated
edges, 2- or 3-scale broad medially, widen slightly toward
ventrolateral side; (20) first dorsal cross-band at 4th ventral
scale, DHB 14.1 mm; and (21) ventral pale Salmon Color
(Color251), withmoreor lessregularblack transversebands
andsomeirregularspeckles.
Comparisons: The new species is morphologically most
similar and was confused as L. fasciatus, but it can be
differentiated from the latter by having smooth dorsal scales
(vs. keeled), fewer infralabials (8 vs. 9 in most individuals),
andthepresenceofdistinct collarband onheadinadults(vs.
absence) (Figure 5). Lycodon obvelatussp. nov. further
differsfromDinodon yunnanensis,whichisstillconsideredthe
junior synonym of L. fasciatus but believed to be valid, by
havingsmooth dorsal scales (vs. keeled), more ventral scale
(199 vs. 193), more dorsal cross-band on body (32 vs. 23),
andfewersupralabials(8vs.9).
ForspeciesthatarealsosimilartoL. fasciatus,L. obvelatus
sp. nov. differs from L. gongshan by having smooth dorsal
scales (vs. keeled), fewer subcaudals (76 vs. 92–96), and a
smallerbodysize(ToL551mmvs.maximum963mm);fromL.
liuchengchaoibyhavingsmoothdorsalscales(vs.keeled),an
entirecloacalplate(vs.divided),andfewerdorsalcross-bands
(31vs.≥40); from L. pictus byhavingfewer ventrals (199 vs.
212–218), presence of collar-band in adults (vs. absence),
and a distinct coloration (dorsal Jet Black (Color 300), with
Salmon Color (Color 251) cross-bands vs. dorsal Brick Red
(Color 36) to Warm Sepia (Color 40), with dirty white cross
bands); and from L. synaptor by having loreal entering orbit
(vs.separatedfromorbitby preocular), smooth dorsal scales
(vs.keeled),andwiderdorsal cross-band(2-or3-scalebroad
dorsallyvs.singlescalebroad).
Lycodon obvelatussp. nov. differs from L. serratussp.
nov.byhavingfewerinfralabials(8vs.10),fewerASR(17vs.
19),farfewerdorsalcross-bands(31ondorsum,13ontailvs.
66 on dorsum, 26 on tail), a distinct coloration (dorsal Jet
Black(Color 300)withSalmon Color (Color251)cross-bands
vs. dorsal dirty Tawny Olive (Color 17) with Jet Black (Color
300)cross-bands),andwidercross-bands(expending2-or3-
scalewidedorsallyvs.mostlysingle-scalebroad)(Figure3).
Forremainingspecies,L. obvelatussp. nov.differsfromall
members of the L. ruhstrati species group (L. cathaya, L.
chapaensiscomb. nov., L. futsingensis,L. multifasciatus,L.
ophiophagus, L. paucifasciatus, L. ruhstrati, and L.
septentrionalis) and L. alcalai, L. banksi, L. bibonius, L.
cardamomensis, L. carinatus, L. chrysoprateros, L. davidi,L.
ferroni,L. flavozonatus,L. gammiei, L. kundui, L. muelleri, L.
rufozonatus, L. rosozonatus, L. solivagus, L. stormi, L.
travancoricus, and L. zoosvictoriae by having loreal entering
orbit (vs. separated); from L. effraenis by the presence of
loreal scale (vs. absence); from L. subannulatus by having
more DSRH (17 vs. 15) and DSRM (17 vs. 15); from L.
albofuscus, L. aulicus, L. capucinus, L. flavicollis, L.
flavomaculatus, L. hypsirhinoides, L. jara, L. laoensis, L.
mackinnoni, L. meridionalis, L. multizonatus, L. nympha, L.
orientalis, L. sealei, L. sidiki, L. striatus, L. subcinctus, L.
tessellatus,L. tiwarii,andL. zawibyhavinganentirecloacal
plate (vs. divided); from L. anamallensis by fewer temporals
(2+2or 2+3vs. 3+4);andfromL. philippinus bymore MT(11
vs.8)andfewerventralscales(199vs.216–225).
Description of holotype: KIZ 040146, adult male, medium
sizedLycodon,ToL551 mm,SVL 447mm.Bodyslender,tail
moderate,TaL18.9%ToL;headmoderate, flat,snoutnarrow,
HL 11.3 mm, HW 9.4 mm, distinct from neck; eye large, not
laterally compressed, ED 2.1 mm, 18.6% HL; pupil vertically
oriented. Rostral pentagonal, broader than height, slightly
visible from above; nasal divided, anterior half rectangular,
small, bordering rostral, first supralabial, and internasal,
posterior half hexagonal, much larger, bordering first and
second supralabials, loreal, internasal, and prefrontal;
prefrontal paired, hexagonal, much larger than internasal,
separated from orbit by preocular; loreal much elongated,
entering orbit, bordering posterior nasal, prefrontal, second
and third supralabials, and preocular; preocular single;
supralabials 8/8, third to fifth entering orbit; postocular 2;
temporals 2+3/2+2, inferior one of first pair much larger;
frontal pentagonal, spear-like tip pointing posteriorly;
supraocular elongated; parietal paired in V-shape, relatively
wide, inlaying spear-tip of frontal anteriorly, bordering
supraocular and superior postocular anteriorly, paraparietal,
andfoursmaller nuchalscalesposteriorly;paraparietalsingle,
enlarged.Infralabials8/8, anterior most pair enclosingmental
andmeetingmedioposteriorly; anterior5infralabialsbordering
anteriorchinshieldon left,4 onright;fifthandsixthbordering
posteriorchinshieldsonleft,fourthandfifthonright;twopairs
ofchinshield,anteriorpairwider,meetingmedially,formingV-
shape and inlaying meeting tip of first pair of infralabials
anteriorly; posterior chin shields much narrower and
elongated, separated from each other by rather wide section
of mental groove. Dorsal body scales smooth, 17 rows one-
head-length behind neck, 17 rows midbody, 15 rows one-
head-length before vent. Single preventral; ventral 199,
angular; cloacal plate entire; subcaudal paired, 76 excluding
tailtip.DHB14.1mm,3.1%SVL,PBBat4thventralscale.
Maxillary teeth 11, forming 4 distinct groups separated by
three gaps. Seven teeth in first group: first five gradually
increaseinsize,followedbymuchenlargedsixthandseventh;
smallereighthtoothinsecondgroup,samesizeasthird;ninth
toothin third group, same size as eighth; last two (tenthand
eleventh)inlastgroup,samesizeasfifth.Threegapspresent,
Zoological Research42(1):62−86,2021 77

namely between seventh and eighth teeth (largest, about 4
timesregular teeth width), eighthandninth (1.5 times regular
width),ninthandtenth(twiceregularwidth).
Hemipenisonlypartiallyeverted,singleclavate, nipatdistal
end, spinose except very proximal end; spines enlarged
towardproximalend;veryproximalendfreeofspines.
Coloration:Inlife,thedorsal andlateral surfacesofthehead
are Jet Black (Color 300), except the anterior portion of the
head: the internasal, prefrontal and anterior frontal are
speckledwithPaleNeutralGray(Color296);andtheposterior
half of nasal, loreal, and first four supralabials are nearly
uniform Pale Neutral Gray (Color 296). A distinct collar band
onoccipitalregionofthehead,dirtySalmonColor(Color251).
Dorsal surface of the body is Jet Black (Color 300). Salmon
Color(Color251)cross-bandsare present on the dorsal and
lateral surfaces of body and tail. Cross-bands have jagged
edges,andtheyaretwo-tothree-scalebroaddorsallyandare
further widen ventrolaterlly. A total of 31 cross-bands are
presentonthebody,and13areonthetail.Startingatthefifth
cross-bandsfromthe head,mostSalmonColored(Color251)
cross-bands of the body has a transverse row of black
speckles running through the middle, some of which even
forms a narrow and almost complete black transverse streak
(i.e., in number 20 and 21 cross-bands from the head). The
ventral surfaces of the head, body, and tail are pale Light
Flesh Color (Color 250) to white. Anterior five infralabials,
mental, and anterior portion of the anterior chin shields are
Medium Neutral Gray (Color 298). Dark Neutral Gray (Color
299)to Jet Black (Color300)cross-bands,transverse groups
of patches, or irregular speckles are present on the ventral
body, with the anterior nine cross-bands clearly defined. A
totaloftwelveJetBlack(Color300)cross-bands are present
onventraltail.
The ornamentations remain the same after two-year of
preservation, but coloration fades away. Specifically, the
SalmonColor(Color251)ofdorsalcross-bandsbecomespale
LightFleshColor(Color250),andthe ventral color becomes
almostLightBuff(Color2).
Distribution, natural history, and conservation:AlthoughL.
obvelatussp. nov.iscurrentlyonlyknownfromthePanzhihua
CityPark,it is possible that the new species is also found in
the nearby regions in Panzhihua and in the adjacent north-
centralYunnanProvince(i.e.,inYongrenCounty).Thehabitat
consists of both natural and horticultural plants of both
deciduousandevergreenspecies,androadsandothertourist
infrastructures fragmented the habitats (Figure 6B). The
holotype was found actively hunting for geckos on a stone
parapet at night. Other reptiles that are sympatric in the city
park include Naja kaouthia, Ptyas nigromarginata, Elaphe
taeniura, Achalinus sp., Pareas sp., Indotyphlops braminus,
Diploderma dymondi, Gekko sp.,Hemidactylus bowringii, and
Sphenomorphus indicus; and amphibian includes Kaloula
verrucosa, Polypedates sp., Odorrana grahami, and
Duttaphrynus melanostictus.
Although the type locality is at the center of a major city
(about10.8millionpeople), theoasis inthecityparkprovides
habitats for a surprisingly diverse group of reptiles and
amphibians. The natural habitats around the Panzhihua City
have been deforested in the mid 1900s, and the self-
recovering process of the fragile valley ecosystem is
particularlyslow.TheCityParkofPanzhihuapreservedfewof
theremainingnaturalmontaneevergreen forests in the area,
which provide important habitats for local wildlife. The
discovery of the new species highlights the conservation
importance of the remaining habitats in the city park.
Unfortunately,the current maintenancepracticeof the parkis
not ecofriendly, with rapid developments for tourist
infrastructure, replacements of native plants with exotic
horticultural plants, and the wide usage of pesticides. We
recommendtheparkmodifyitscurrentpracticesandconserve
theremainingnaturalhabitatsforthenativewildlife.
DISCUSSION
Additional cryptic diversity in the northern HMR
Thediscoveryofourtwonewspeciessupportsthenotionthat
the reptilian diversity in the northern HMR is still
underestimated. As the suitable habitats of reptiles (i.e., hot-
dry valleys) in the HMR are isolated and fragmented by
continuous mountain ranges over 4 000 m of elevation,
populations in different river valleys are allopatric to each
other,despitetheshortlineardistanceamongthem(Figure1).
Therefore, it is likely that nearby valleys along the upper
Mekong, Salween, and Yalong Rivers also harbor additional
undiscovereddiversity ofthegenus Lycodon.Further surveys
are needed to better inventory of the reptilian diversity and
assesstheirconservationstatusesinthenorthernHMR.
Problematic records of
Lycodon
species in China and SE
Asia
Lycodon fasciatus
and
L. liuchengchaoi
:For the
recognized species of the genus Lycodon in China, great
confusions exist in published literature regarding the
taxonomic identification and the resulting distribution range,
particularlyforL. fasciatus(Vogel&David,2010;Vogel&Luo,
2011).Much similar to other groupsofreptiles from the HMR
that represent species complexes (i.e., Gloydius strauchi,
Diploderma flaviceps; Shi et al., 2018; Wang et al., 2019a,
2021),L. fasciatuswasandstillisconsideredasawidespread
taxon, despite increasing evidence suggesting the existence
ofcrypticdiversity(Vogel & David, 2010; Vogel & Luo, 2011;
Zhangetal., 2011b). As the results of taxonomic confusions,
misidentifications and erroneous records of species are
prevalentinliterature.
Kanget al. (2009)reportedL. fasciatusasanew recordof
snake in Hunan Province based on specimens from
HupingshanNature Reserve. Later Bai et al. (2018) reported
L. liuchengchaoi from the very same nature reserve. Closer
examinationofthecorrespondingdescriptionsrevealsthatthe
referredspecimensbyKangetal.(2009)andBaietal.(2018)
both possess a divided cloacal plate, which matches the
diagnosisofL. liuchengchaoibutnotL. fasciatus(Zhangetal.,
78www.zoores.ac.cn

2011b).In additionto thepresenceof ayellow collar-bandon
the neck in figures of both Kang et al. (2009) and Bai et al.
(2018),which againcontradictto thediagnosisof L. fasciatus
but align with L. liuchengchaoi, it is clear that the previous
record of L. fasciatus from Hunan Province by Kang et al.
(2009)representamisidentificationofL. liuchengchaoi.
Li et al. (2012) first recorded L. fasciatus from Guangdong
Province, and the authors stated that the tail length of
Guangdongspecimens is 24.8%–25.8% of the total length in
sub-adults,whicharemuchlongerthanthetrueL. fasciatus(≤
22.5%;Vogel & Luo, 2011; Table 4). Later Guo et al.(2013)
providedthegeneticdataofL. fasciatusfromGuangdong,but
atthetime thereisnogeneticdatafromtopotypicL. fasciatus
to compare against. Recently, Peng et al. (2018) reported L.
liuchengchaoi from Guangdong based on morphological and
molecularevidence ofcyt bgene,and theauthorsstatedthat
the cyt b data of their specimens of L. liuchengchaoi from
GuangdongisnearlyidenticaltothepublishedsequenceofL.
liuchengchaoi on GenBank and share the same haplotype
withpreviouslypublisheddataofL. fasciatusfromGuoetal.
(2013). As results, Peng et al. (2018) confirms the previous
record of L. fasciatus in Guangdong represents
misidentification of L. liuchengchaoi. However, Peng et al.
(2018)didnotsubmittheirnew datatoGenBank,nordidthey
conductphylogeneticanalysesofthementionedsamples.
Ourphylogeneticstudyofavailablesequencessupportsthe
conclusion by Peng et al. (2018), where the Guangdong
samples of  “ L. fasciatus” from Guo et al. (2013) are nested
within available data of L. liuchengchaoi; and with the newly
available topotypic samples of L. fasciatus, the Guangdong
samplesare confirmed to be paraphyletic with respect to the
true L. fasciatus from Yunnan (Figure 2). In addition to our
revision of  “ L. fasciatus” in Panzhihua, it is clear that the
currentrecordsof“L. fasciatus”outsideofYunnanProvincein
China (i.e., in Anhui, Gansu, Guizhou, Hubei, Shaanxi, and
Zhejiang;Zhao et al., 1998) aredistantfromthe range of the
true L. fasciatus, and these questionable records likely
representeithermisidentificationsofrecognizedcongeners,or
additional cryptic diversity that warrant further investigations.
Future studies should focus on confirming the taxonomic
statusesof the questionable records of  “ L. fasciatus”outside
of Yunnan Province in China. Currently, L. fasciatus sensu
stricto has been confirmed in Yunnan Province of China and
Myanmar(Vogel&Luo,2011;presentstudy).
Regarding L. liuchengchaoi, Li et al. (2020) reported a
sample of  “ L. liuchengchaoi” from  “Sanjiazhai” in Yunnan
Province, which would expand the distribution range of the
speciesfurthersouthwestwardandrepresentsanewrecordof
herpetofauna of Yunnan Province. However, our phylogeny
showsthatthereferred sample of  “ L. liuchengchaoi” by Li et
al. (2020) is phylogeneticlly distinct from the true L.
liuchengchaoi,and it actually representsamisidentification of
L. multizonatus instead (Figure 2). With this correction of
taxonomy, this record in Yunnan still represents a range
extensionof the L. multizonatusand a new recordofYunnan
Province. However, Li et al. (2020) did not provide complete
information regarding the county or prefecture of the locality
name  “Sanjiazhai”. As multiple localities in Yunnan Province
are under this very same name, the distribution of L.
multizonatus in Yunnan remains unknown. Future studies
shouldverifytherecord.
Fortheremainingconfirmedrecord ofL. liuchengchaoi,itis
importantto notethatthereareconsiderable discrepanciesof
key morphological characters between the type series of the
species and the later reported records in China, particularly
regarding the number of dorsal cross-band and the state of
cloacalplate(Pengetal.,2018;Zhangetal.,2011b).Future
population-level studies are needed to better understand the
morphologicalvariationanddiagnosisofL. liuchengchaoi.
Lycodon ruhstrati
in Yunnan Province:Guo et al. (2007)
firstreportedL. ruhstratiasthenewrecordofreptilianfaunaof
YunnanProvincefromtheGaoligongMountainsinfarwestern

Figure 6 The habitats at the type locality of
Lycodon serratus
sp.
nov. near Geyading Village, Deqin County, Yunnan Province,
China (A) and
L. obvelatus
sp. nov. in Panzhihua City Park,
Panzhihua, Sichuan, China (B) (Photos by Zhong-Bin Yu and Ben-
Fu Miao)

Zoological Research42(1):62−86,2021 79

Yunnan.However,examinationof the description and photos
byGuo et al.(2007)reveals thatallthree referred specimens
by Guo et al. (2007) do not agree with the diagnosis of L.
ruhstrati: the first specimen (HNU 200 505 001) has a much
shortertail (TaL/ToL 18.7%),fewerventral scales (VEN 203),
fewer subcaudal scales (SC 68), and distinctively banded
ventralsurface ofthe bodythroughout(vs. intrueL. ruhstrati,
TaL/ToL 20.8%–24.8%, VEN 214–233, SC 90–116, and
ventral body either uniformly colored or speckled without
distinct cross-bands; Vogel et al., 2009); and the remaining
twospecimens(HNU 200 505 002and200 609 001)bothhave
lorealsenteringorbits(vs.intrueL. ruhstratinotenteringorbit;
Vogel et al., 2009; Zhao et al., 1998). Furthermore, the later
twospecimens have muchlongertails (TaL/ToL21.5–25.5%)
and more subcaudals (92–94) than the first specimen.
Therefore,even basedonthereportedmorphological databy
Guo et al. (2007) alone, it is clear that the three referred
specimens are neither true L. ruhstrati, nor do they even
represent the same taxa: HNU 200 505 001 is similar to L.
chapaensis,whileHNU200 505 001and 200 609 001matches
diagnosisofL. gongshan.
A year after Guo et al. (2007), Yang & Rao (2008) also
recorded L. ruhstrati from Yunnan. This time the record is
based on a different vouchered specimen, which has no
detailed locality information (KIZ 8 300 012,  “from Yunnan”;
Yang & Rao, 2008). Unfortunately, we could not locate the
referred specimen at KIZ (possibly lost), but upon review of
the description by Yang & Rao (2008), we found that the
specimen does not agree with the diagnosis of the true L.
ruhstrati,including havingfewerSC (81 vs.90–116),different
dorsal scale texture (feebly keeled vs. distinctively keeled),
and by the presence of white collar-band on neck (vs.
absenceinadults).Therefore,basedonthecurrentpublished
data, all reported voucher specimens of  “ L. ruhstrati” from
Yunnan do not agree with the diagnosis of true L. ruhstrati,
and there is no evidence confirming the presence of L.
ruhstratiinYunnanProvinceasoftodate.
Lycodon gongshan
in Yunnan and Sichuan:Lycodon
gongshan was described based on morphological characters
only, and the type series was collected from far western
Yunnan Province in the Dulongjiang Valley and adjacent
Nujiang valley (Vogel & Luo, 2011). Later Guo et al. (2015)
recorded the species from Lincang, Southwestern Yunnan
Provinceandprovidedgenetic dataof thespecies.Ournewly
collected topotypic materials from Dulongjiang confirm the
taxonomic identification by Guo et al. (2015) (Figure 2;
Table2).Furthermore,ourphylogeneticanalysesconfirmthat
our non-vouchered genetic sample from Yunlong Nature
Reserve in Yunlong County, Dali is also L. gongshan, which
expanditsrangeeastwards(Figure1).
Although our results expand the range of L. gongshan
furthereastwards,the speciesisstillendemictoYunnanonly,
and the existing records of the species in Sichuan Province
require further confirmation. Chen et al. (2018a) recorded L.
gongshanbasedontwospecimensfromHongbaoVillageand
Daheishan National Forest in Panzhihua, Sichuan. However,
the images that Chen et al. (2018a) provided show obvious
difference from the type series of L. gongshan in terms of
ornamentation pattern, and the recorded numbers of dorsal
cross-bands do not match with the bands of the actual
specimen in the photographs. Based on morphological data
alone, we cannot assign these two specimens to our new
species L. obvelatus from Panzhihua City (i.e., Hongbao
individualshavekeeled dorsal scales, where dorsal scalesof
L. obvelatusissmooth). It is likely that there are two species
of Lycodon in Panzhihua, similar to the genus Diploderma
(i.e.,D. dymondiisfound inPanzhihua City,where D. swildis
found in Hongbao Village; Wang et al., 2019b), but whether
theHongbao populationrepresentsmorphological variationof
L. gongshan or a distinct new species would require future
confirmation.
Remaining records of “
L.
septentrionalis
” in SE Asia and
validity of
L. ophiophagus
:Lycodon septentrionalis has
been recognized to have a wide distribution range, from the
Himalaya (i.e., India (Boulenger, 1893; Smith, 1943) and
Bhutan(Tshewang & Letro,2018))across Myanmar(Dowling
& Jenner, 1988) and Yunnan of China (Zhao et al., 1998;
Zhao, 2006) to Southeast Asia (including Vietnam (Smith,
1943;Van Sang et al., 2009),Laos(Deuveet al., 1961), and
Thailand(David et al.,2004)).Similar to theabove-discussed
congenersthatalsohavewidedistributionranges,thecurrent
recordsofL. septentrionalis likelycontain misidentificationsof
different lineages, particularly in Southeast Asia. With our
resurrectionof L. chapaensis, itleavestheremaining records
of L. septentrionalis in Laos, Vietnam, and Thailand
questionable. The taxonomic position of the Southeast Asian
populations of  “ L. septentrionalis” should be reconsidered in
futurestudies.
Additionally, our morphological comparison shows
overwhelmingly similar morphology between L. chapaensis
andL. ophiophagus.Theonlydifferencesare the relative tail
length(17.1%–20.5%inL. chapaensisvs.20.1%–22.8%inL.
ophiophagus) and number of subcaudal scales (74–85 vs.
87–90).However, given the small sample size (n=2) andthe
lackofmoleculardataofL. ophiophagus,wecannotconclude
onits taxonomic validity.Futureintegrative taxonomicstudies
are needed to confirm the validity of L. ophiophagus with
respecttoL. chapaensis.
Records of
L. aulicus
and
L. capucinus
in China:Owning
the nearly indistinguishable morphology and the lack of
geneticmaterialsfromtopotypicindividuals,taxonomistshave
notreachedagreementsregardingthevalidityofL. capucinus:
whetheritisjuniorsynonym of L. aulicus, valid but only as a
subspecies, or valid as a full species (O’Shea et al., 2018;
Ota,2000; Sileretal., 2013;Wostlet al.,2017).Although the
overalldistributions of L. capucinusandL. aulicus havebeen
relativelyconsistentinliterature(L. aulicusisfromSouthAsia,
whereL. capucinus isfromSoutheast Asia, andbothspecies
arehypothesizedto besympatricinMyanmar;David&Vogel,
1996; Lanza, 1999; Smith, 1943), the distribution of both
80www.zoores.ac.cn

species near the hypothesized contacting region remain
unclear,particularlyinChina(O’Sheaetal.,2018).
While considering L. capucinus as a subspecies of L.
aulicus,both L. a. capucinusand thenominatesubspeciesL.
a. aulicus have been recordedfromHong Kong (Pope,1935;
Romer, 1979). Most of the later authors did not consider the
subspecies or species status of L. capucinus, and only L.
aulicus have been recorded from China, with its distribution
ranging from southwestern Yunnan, Fujian, to Guangdong
Provinces (Wang et al., 2020b; Zhao & Adler, 1993; Zhao et
al.,1998;Zhao,2006). In contrary, Zhang et al. (2011b) only
record L. capucinus from China, without discussing the past
record of L. a. aulicusfrom Hong Kong (Pope, 1935) or the
possible distribution of L. aulicus from the Myanmar border
regionsinSouthwestYunnan.
Images of live individuals of the L. aulicus-capucinus
complex from China-Myanmar border in Yunnan and from
Hong Kong show nearly identical ornamentation patterns
(Figure 7), which matches the current diagnosis of L.
capucinus (O’Shea et al., 2018). Unfortunately, vouchered
genetic materials of the L. aulicus-capucinus complex from
China and from the type localities of the two corresponding
names are still unavailable to date. Given the L. aulicus-
capucinus complex is known by its profound variability in
ornamentation patterns (O’Shea et al., 2018), we cannot
determine the taxonomic identity of the Chinese populations
with confidence. Based on the current diagnosis of both
species, we here consider the Chinese populations as L.
capucinus, and we propose to maintain its Chinese common
nameas  白环蛇. Later taxonomic studies are needed to
further verify the validity of L. capucinus and confirm the
identityoftherelatedChinesepopulations.
Updated key and distribution of the genus
Lycodon
in
China:To facilitate future taxonomic studies of the genus
Lycodon in China, we provide an updated dichotomous key
andthe distributionsof the20recognized speciesofLycodon
species in China. The distribution data are based of Zhao et
al. (1998) and are further modified with new findings in this
present study and additional literatures published after 1998
(Appendix III).  “?” indicates possible but not yet confirmed
records based on photographic evidence or published
sequenceswithvaguelocalityandnomorphologicaldata;and
“!” indicate possible erroneous records that warrant future
confirmations.  “Dorsal background coloration” is defined as
thesamecolorationofthedorsalsurfaceofthehead.

Key to the species of
Lycodon
in China:
1a) Dorsal background coloration yellowish brown, dark
brown, or reddish brown; dorsal cross-bands Jet Black
(Color 300), relatively narrow and serrated, not widen
towards ventrolateral sides; cloacal plate divided; loreal
entering orbit....................................................................2
1b) Dorsal background coloration blackish, with white, gray,
yellowish, pinkish, or reddish dorsal cross-bands, usually
widen toward ventrolateral sides; or dorsal brownish with
no cross-bands but reticulated patterns; cloacal plate
dividedorentire;lorealentering orbitor not......................3
2a)Headdistinctfromneck;eyeslaterallycompressed;dorsal
scale rows 19 at one-head length behind neck; first five
infralabialsincontactwithanteriorchinshield;blackcross-
bands on the anterior dorsum strongly serrated, mostly
single scale broad, rarely two..........................L. serratus
(Yunnan;Sichuan?)
2b)Head indistinct from neck; eyesnotlaterally compressed;
dorsalscalerows17atone-headlengthbehindneck;first
four infralabials in contact with anterior chin shield; black
cross-bandson the anterior dorsum less serrated, mostly
2- or 3-scale broad....................................L. multizonatus
(Gansu,Sichuan,Yunnan?)
3a) Dorsal scale rows 19 at one-head length behind neck...
...........................................................................4
3b) Dorsal scale rows 17 at one-head length behind neck...
..............................................................................5
4a) Dorsal scale rows 19 at mid-body; dorsal cross-bands
wide, 28–35 on dorsum, 8–13 on tail..............................
......................................................................L. rosozonatus
(Hainan)
4b)Dorsalscalerowsmostly17atmid-body,rarely19;dorsal
cross-bands narrow, 51–87 on dorsum, 12–30 on tail
......................................................................L. rufozonatus

Figure 7 Photos of live
Lycodon aulicus-capucinus
complex from China
A:FromHongKong,China;B:FromYingjiangCounty,Yunnan,China.PhotosbyJin-LongRenandFrancoLeungKaWah.

Zoological Research42(1):62−86,2021 81

(Anhui,Beijing,Chongqing,Fujian,Gansu,Guangdong,
Guangxi,Guizhou,Henan,Hebei,Hubei,Hunan,Heilongjiang,
Jilin,Jiangsu,Jiangxi,Liaoning,Sichuan,Shandong,
Shanghai,Shanxi,Shaanxi,Taiwan,Tianjin,
Yunnan,Zhejiang)
5a)Cloacalplateentire.............................................................6
5b) Cloacal plate divided.....................................................16
6a) Dorsal body scale smooth................................................7
6b) Dorsal body scale feebly keeled or strongly keeled......
...............................................................................10
7a) Ventrals 212–218; subcaudals 90 or 91; infralabials 9 or
10; loreal entering orbit; maxillary teeth 13 or 14......
................................................................................L. pictus
(Guangxi)
7b) Ventrals≤210; subcaudals<90; infralabials ≥8; loreal
entering orbit or not....................................................8
8a)Infralabials8;lorealalways entering orbit; maxillary teeth
11; dorsal cross-bands Salmon color (Color 251) in
life.....................................................................L. obvelatus
(Sichuan;Yunnan?)
8b) Infralabials 9 or more; loreal mostly not entering orbit;
dorsal cross-bands Pale Rose Pink (Color 243), with
dense but faint gray speckles........................................9
9a) Dorsal cross-bands connecting with each other laterally,
separating ground black coloration into ellipse patches;
maxillary teeth 10.............................................L. cathaya
(Guangxi)
9b) Dorsal cross-bands separated from each other; maxillary
teeth 12–15..................................................L. futsingensis
(Fujian,Guangdong,Guangxi,Jiangxi,Zhejiang,
HongKong)
10a)Lorealenteringorbit;dorsalcross-bandswide(usually3-
scale wide) with strongly jagged edges.............................
........................................................................................11
10b) Loreal not entering orbit; dorsal cross-bands narrow
(usually1- to2-scale wide)withsmotheredges(except L.
ruhstrati)..................................................................12
11a) Tail long, TaL 23.1%–23.2% ToL in males, 21.5% in
female...............................................................L. gongshan
(Sichuan?,Yunnan)
11b)Tail short, TaL 19.8%–22.5% inmales,19.0%–21.9% in
females...............................................................L. fasciatus
(Yunnan,Anhui!,Gansu!,Guizhou!,Hubei!,Shaanxi!,
Zhejiang!)
12a) Dorsal cross-bands bright Sulphur Yellow (Color 80) in
life, 50–96 on dorsum.................................L. flavozonatus
(Anhui,Chongqing,Fujian,Guangdong,Guangxi!,Guizhou,
Hainan,Hunan,Jiangxi,Sichuan,Zhejiang)
12b) Dorsal cross-bands white or gray, <50 on dorsum.....
..........................................................................13
13a) Dorsal cross-bands dirty white (speckled with Drak
NeutralGray(Color 300)) or Cinnamon Drab (Color 259),
increasinglymore dirty posteriorly, 19–46ondorsum,3–4
dorsal-scale wide for most parts; intercepted black
segmentsrather short.........................................L. ruhstrati
(Anhui,Beijing,Chongqing,Fujian,Gansu,Guangdong,
Guangxi,Guizhou,Hainan,Henan,HongKong,Hubei,
Hunan,Jiangsu,Jiangxi,Sichuan,Shaanxi,Taiwan,
Tianjin,Zhejiang)
13b)Dorsalcross-bandsclearwhite,25–31ondorsum,1or2
scale wide on most parts; intercepting black segments
long.........................................................................14
14a)Body size small, ToL 463–487mm; subcaudal 68 or 69;
maxillary teeth 10, forming three groups, group one and
three each with two significantly enlarged teeth..............
...........................................................................L. synaptor
(Yunnan)
14b) Body size large, ToL >1 000 mm; subcaudal 74–85.....
...................................................................................15
15a) Medial 5–7 rows of dorsal scale keeled; maxillary teeth
8................................................................L. septentrionalis
(Tibet)
15b) Dorsal body scale completely smooth or only very
posterior portion of vertebral row feebly keeled; maxillary
teeth 11 or 12...............................................L. chapaensis
(Yunnan)
16a) Dorsal body scales smooth and glassy........................
........................................................................................17
16b) Medial rows of dorsal body scales strongly or feebly
keeled......................................................................18
17a) Frontal in contact with preocular; reticulated patterns
absentonbody...................................................L. laoensis
(Yunnan)
17b) Frontal not in contact with preocular; light reticulated
patterns present on lateral and sometimes dorsal body...
..............................................L. aulicus-capucinus complex
(Fujian,Guangdong,HongKong,Yunnan)
18a) Loreal not entering orbit; dorsal cross-bands Sulphur
Yellow (Color 80) to Olive Sulphur Yellow (Color 90);
distinctspecklesandreticulatedpatternspresentondorsal
andlateralhead;..........................................L. meridionalis
(Guangxi,Yunnan)
18b) Loreal entering orbit; speckles and reticulated patterns
absentondorsalandlateralhead....................................19
19a)Preocularabsent;prefrontal entering orbit; dorsal cross-
bands white....................................................L. subcinctus
(Fujian,Guangdong,Guangxi,Hainan)
19b) Preocular present; prefrontal not entering orbit; dorsal
cross-band creamy Dark Spectrum Yellow (Color 78) to
creamy Light Chrome Orange (Color 76).........................
...................................................................L. liuchengchaoi
(Beijing?,Guangdong,Henan,Hunan,Sichuan,Shanxi,
Shaanxi,Zhejiang)
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