Molecular Phylogeny of the Snake Genus Oligodon (Serpentes ... · ii Molecular Phylogeny of the Snake Genus Oligodon (Serpentes: Colubridae), with an Annotated Checklist and Key Marc

Molecular Phylogeny of the Snake Genus Oligodon (Serpentes: Colubridae), with an Annotated Checklist and

Key

by

Marc David Green

A thesis submitted in conformity with the requirements for the degree of Master of Science

Ecology and Evolutionary Biology University of Toronto

© Copyright by Marc David Green 2010

ii

Molecular Phylogeny of the Snake Genus Oligodon (Serpentes:

Colubridae), with an Annotated Checklist and Key

Marc David Green

Master of Science

Ecology and Evolutionary Biology

University of Toronto

2010

Abstract

The snake genus Oligodon, known for its egg-eating feeding behaviour, is a taxonomically and

systematically challenging group from South and Southeast Asia. This work provides the first

phylogenetic hypothesis for the genus, and includes a checklist and key to the species. I use

approximately 1900 base pairs of mitochondrial DNA sequence data to infer the relationships of

these snakes, and I examine congruence between the molecular phylogeny and hemipenial

characters. A hypothesis for the position of Oligodon within the Colubridae is also proposed. I

discuss the implications of the phylogeny for previous taxonomic groupings, and consider the

usefulness of the trees in analysis of behaviour and biogeography.

iii

Acknowledgments

I gave considerable thought to doing something clever (or pretentious) with these

acknowledgements. The best idea that was suggested, was doing it all in limerick form.

Considering the amount of content in this document on snake genitalia, that could have been

amusing. However, in the end, I decided to take the straightforward approach.

First, Bob, without whose constant, unwavering support over the years, I never would have

finished. To put it most succinctly: It has been a most stimulating journey.

I would also like to thank my defense committee, Profs. Doug Currie, Debbie McLennan and

Chris Darling. Beyond giving the thesis a critical and fair review, I look back and thank each of

you for contributing to my education over the years.

My time in connection with the Murphy Lab spanned a few comings and goings, a few people

stand out especially. Amy and Ross: Any quality research produced here was facilitated by these

two fine human beings, who taught me my early laboratory and museum skills, and provided

many quality scientific interactions. Johan and Andre: My cohort mates. We endured most of our

academic trials together, and I learned as much from you guys as I did from classes and books. I

could go on a bit more about Andre, but will stop with this thought: That was nasty.

My family, Mother, Father, Janet, Larry and Chris, thank you for your love and support.

Anything good about my character has been formed by you, and anything positive I leave behind

for posterity will be a result of hard work flowing from that disposition. Over the years you have

provided many great times, and much needed encouragement and approval.

Finally, Sarah, without whose constant, unwavering support over the years, I never would have

finished. Words fail. You deserve credit for the continued development of my skills and

personhood. All my love and thanks for your endurance, tireless help, amazing and amusing

mind, dedication and example. This work is dedicated to you.

There once was a boy, scion of the Greens | Decided to look at snakes and their genes | Many

years later | With experience greater | He made much to do of their penes!

iv

Table of Contents

Acknowledgments .....................................................................................................................iii

Table of Contents....................................................................................................................... iv

List of Tables............................................................................................................................. vi

List of Figures ..........................................................................................................................vii

List of Appendices...................................................................................................................viii

1 INTRODUCTION ................................................................................................................. 1

2 MATERIALS AND METHODS............................................................................................ 9

2.1 Specimens Examined ...................................................................................................... 9

2.2 DNA Extraction, Amplification and Sequencing ........................................................... 14

2.3 DNA Sequence Alignment and Phylogenetic Analysis .................................................. 20

3 RESULTS............................................................................................................................ 26

3.1 Sequence Variability in Oligodon.................................................................................. 26

3.2 Phylogeny ..................................................................................................................... 28

3.3 Hemipenial Characters .................................................................................................. 32

3.4 Position of Oligodon in the Colubridae.......................................................................... 34

4 DISCUSSION...................................................................................................................... 36

4.1 Molecular Variation ...................................................................................................... 36

4.2 Phylogeny and Previous Hypotheses of Relationships ................................................... 37

4.2.1 Philippine Taxa.................................................................................................. 37

4.2.2 Oligodon cyclurus-taeniatus Group ................................................................... 38

4.2.3 Oligodon cinereus ............................................................................................. 40

4.2.4 Oligodon torquatus-planiceps-cruentatus-theobaldi .......................................... 40

4.3 Hemipenes and Phylogeny ............................................................................................ 41

4.4 Colour, Confusion and Biogeography............................................................................ 43

v

4.5 Taxonomic Implications................................................................................................ 44

4.6 Oligodon: Position Within the Colubridae/Colubrinae................................................... 45

5 CONCLUDING REMARKS ............................................................................................... 47

References ................................................................................................................................ 49

Appendix A .............................................................................................................................. 84

Appendix B ............................................................................................................................ 151

vi

List of Tables

Table 2.1 A list of specimens chosen for DNA sequencing, their voucher numbers and

locality data.. ............................................................................................................................ 10

Table 2.2 Primer sequences used to PCR amplify and sequence fragments of the mtDNA

genome in this study.. ............................................................................................................... 16

Table 2.3 Colubroidea sequences from GenBank used to determine the relationship of

Oligodon to other colubrids....................................................................................................... 23

Table 3.1 A list of total sites, variable sites and uncorrected p-distances for the three mtDNA

genes in the data set.. ................................................................................................................ 27

vii

List of Figures

Figure 1.1 Photograph of the everted hemipenes of Oligodon chinensis (ROM34581) showing

the bifurcate (forked) condition................................................................................................... 5

Figure 1.2 Photograph of the everted hemipenes of Oligodon formosanus (ROM35634)

showing the bifurcate (forked) condition as well as large papillae.. ............................................. 6

Figure 1.3 Drawing of everted Oligodon albocinctus hemipenes showing non-bifurcate (non-

forked) condition, apical flounces and calyces. ........................................................................... 7

Figure 1.4 Drawing of everted Oligodon sublineatus hemipenes showing non-bifurcate (non-

forked) condition and spines.. ..................................................................................................... 8

Figure 2.1 Composite satellite photograph of Southeast Asia, showing relief and topographic

features, collection localities of specimens are plotted. .............................................................. 12

Figure 2.2 A representation of a linear double stranded section of the mitochondrial genome

including 12S rRNA, tRNAval

, 16Sr RNA and location of primers used in this study. ............... 17

Figure 3.1 Maximum parsimony phylogeny of Oligodon.. ................................................... 29

Figure 3.2 Bayesian inference phylogeny for Oligodon.. ...................................................... 30

Figure 3.3 Hemipenial character state mapping for Oligodon ............................................... 33

Figure 3.4 Parsimony analysis of 48 colubrids including Oligodon and outgroups using

sequences of 12SrRNA, 16SrRNA, c-Mos, Cytochrome b, NADH4, NADH1, NADH2, COI

and Protein loci......................................................................................................................... 35

viii

List of Appendices

Appendix A: Annotated checklist of extant species of Oligodon ............................................... 84

Appendix B: Key to the species of the genus Oligodon ........................................................... 151

1

1 INTRODUCTION

The genus Oligodon Fitzinger 1826, the kukri snakes, contains 70 species distributed

throughout much of Southeast Asia (Appendix A). The common name derives from a

distinctively shaped Nepalese knife, the kukri. The hind teeth of these snakes are broad and

strongly recurved, much like the shape of the kukri knife. These non-venomous snakes are

usually nocturnal and often brightly coloured. They feed primarily on the eggs of birds and

reptiles. The morphology of their teeth is very effective for cutting open eggs. Although some

work has been done on this aspect of their life history, little can be inferred about the evolution

of this unique feeding strategy without a phylogeny.

Several attempts have been made to study the phylogenetic relationships and taxonomy

of Oligodon (Campden–Main, 1969; David et al., 2008b; Leviton, 1963a; Pope, 1935; Smith,

1943; Tillack and Günther, 2009; Wall, 1923b; Wallach and Bauer, 1996) and these efforts have

met with mixed success. Smith (1943) made the most ambitious review examining 34 species

and commenting on their relationships. However, his work predates the formalization of

phylogenetic methods (Hennig, 1966). Very little progress has been made since then and

Oligodon has yet to receive a modern phylogenetic treatment using either morphology or

molecules. Leviton (1963a) studied six Philippine taxa and Zhao et al. (1998) added new data in

their examination of Chinese species. Neither study constructed a phylogenetic tree. Most works

on Oligodon have not been comparative and they have focused on small, politically defined

geographic regions. Thus, in the absence of a comparative framework, some authors have

perpetuated errors and confusion. Herein, I propose a phylogenetic hypothesis for Oligodon that

could be applied to future questions on behaviour, biogeography and conservation.

2

An important first step in my study of Oligodon was producing an updated checklist and

key to the entire genus. This key consolidates single species and regional treatments of Oligodon

and supports species identifications and inferences on biology of Oligodon in light of the

phylogenetic results. The checklist and key are presented in the appendices. The checklist

indicates where previous works have hypothesized relationships among Oligodon species. In

particular, Smith (1943) gave several groups (e.g. cyclurus-group, taeniatus-group, cyclurus-

taeniatus-group and cinereus-group) that will be referred to herein.

Oligodon’s preference for eggs has been noted many times (Broadley, 1979; Coleman et

al., 1993; Golf, 1980; Hu and Zhao, 1987; Minton and Anderson, 1963; Ota and Lin, 1994; Wall,

1921c). The mechanisms and behaviour of their egg-eating have been explored (Coleman et al.,

1993; Kiran, 1981; Kiran, 1982; Minton and Anderson, 1963; Ota and Lin, 1994). The snakes

enthusiastically ingest the contents of opened eggs presented to them, and will ingest small

reptile and bird eggs whole. An interesting and unusual aspect of this natural history is their

apparent adaptation to taking eggs too large to swallow. They use their uniquely shaped teeth and

jaw articulation to saw a hole in an egg-shell large enough to insert their head, and feed on the

contents. Stomach content studies reveal that Oligodon also eat small rodents, tadpoles, frogs and

insects (Batchelor, 1958; Cox, 1991; David and Vogel, 1996; Hu, 2001; Hwang et al., 1965;

Meggitt, 1931; Schulz, 1988; Shi and Zheng, 1985; Toriba, 1987; Wall, 1921c; Wall, 1923b).

Juveniles appear to be primarily insectivorous. It is not known whether they hunt or scavenge for

food.

The feeding habits of Oligodon have aroused interest in Chinese agriculture. At least one

influential Asian reference stated they are harmful to crop production (Hu and Zhao, 1987),

presumably because they eat the eggs of other beneficial insectivorous reptiles.

3

Some other behaviours of Oligodon may have an underlying phylogenetic basis. Most

species are docile (David and Vogel, 1996; Pope, 1935; Stuebing and Inger, 1999; Wall, 1921c).

However, at least O. fasciolatus has an aggressive disposition (Smith, 1943; Taylor, 1965). It

also displays an interesting defensive behaviour in which a tail-up posture is taken and the bright

red hemipenes are extruded repeatedly, contrasting noticeably with the white ventrum (Wüster

and Cox, 1992).

Feeding, behaviour and the apparent large number of species in Oligodon provoke

interest in its phylogenetic position within the Colubridae. Phylogeny has been shown to be

useful for discussing the evolution of egg eating in snakes (de Queiroz and Rodríguez-Robles,

2006) and phylogenetic relationships within Oligodon could facilitate inferences on the causes

for radiation within the clade. Phyllorhynchus uses a similar egg-contents extracting strategy for

feeding (Gardner and Mendelson III, 2003) and it and Oligodon have been proposed to share a

close phylogenetic relationship (Dowling, 1974; Dowling et al., 1996; Lawson et al., 2005).

However the position of these snakes within the larger Colubridae has not been resolved

(Dowling, 1974; Dowling et al., 1996; Kraus and Brown, 1998; Lawson et al., 2005; Lopez and

Maxson, 1996; Zhang et al., 1984). Although de Queiroz and Rodriguez-Robles discussed

Oligodon and Phyllorhynchus, they did not include them in their tree-based character-

optimization analysis.

The geographic distribution of Oligodon is complex. Several species are widespread and

many have overlapping ranges. Those ranges are all threatened due to the rapid pace of

development in Southeast Asia (Dang and Nhue, 1995; de Silva, 1998). Many ranges have

become fragmented and it is unclear what impact this has had on the genetic diversity of isolated

populations.

4

Although hemipenial defensive behaviours have not been extensively surveyed in

Oligodon, the hemipenis itself has been used to infer historical relationships. Cope (1895) drew

the hemipenis of O. ancorus for use in the species’ description suggesting that they might

indicate historical relationships. Wall (1923) first looked at hemipenes in Oligodon

comparatively. Pope (1935) compiled a list of hemipenial characters in an attempt to distinguish

Holarchus from Oligodon. However, he was unable to do so because O. bitorquatus, the type

species for the genus, was intermediate in condition. Smith (1943) compiled observations on the

hemipenes of Oligodon. Leviton (1963a) used hemipenial data to reconstruct the evolutionary

history of the Philippine taxa. The groupings used today within Oligodon are primarily based on

Smith’s work and they are thought to reflect historical descent. Based on hemipenial

morphology, Oligodon was considered to be a member of the Colubrinae (Zaher, 1999), now the

Colubridae (Zaher et al., 2009). Figures 1.1, 1.2, 1.3 and 1.4 show the diversity of Oligodon’s

hemipenial morphology and provide a foundation for subsequent hemipenial character coding.

Mitochondrial DNA (mtDNA) sequences can be used to reconstruct the phylogenetic

relationships of snakes (de Queiroz and Lawson, 1994; Kraus and Brown, 1998; Lawson et al.,

2005; Lopez and Maxson, 1996; Murphy et al., 2002; Rodríguez-Robles and de Jesús-Escobar,

1999; Slowinski and Lawson, 2002; Vidal et al., 2000). Herein, I present a molecular phylogeny

for Oligodon based on mtDNA data and discuss correlation of relationships and hemipenial

morphology. Because Oligodon is such a speciose group, and tissue samples are not available for

most species, my study suffers somewhat from an under sampling of taxa. Nevertheless, it forms

the foundation for future comparative work for examining the evolution of structures, behaviours

and biogeography in the group.

5

Figure 1.1 Photograph of one everted, bifurcate (forked) hemipenis of Oligodon

chinensis (ROM34581).

6

Figure 1.2 Photograph of the everted hemipenes of Oligodon formosanus

(ROM35634) showing the bifurcate (forked) condition and large papillae (arrows).

7

Figure 1.3 Drawing of everted hemipenes of Oligodon albocinctus showing the

non-bifurcate (non-forked) condition. Small papillae present in O. albocinctus are not visible on

this view. Some apical flounces and calyces are shown. Redrawn and modified from a

combination of Wall (1923; everted drawing), and Smith (1943; dissected view).

8

Figure 1.4 Drawing of everted hemipenes of Oligodon sublineatus showing

non-bifurcate (non-forked) condition. Spines are typical for many of the Indian Oligodon.

Redrawn and modified from Wall (1923).

9

2 MATERIALS AND METHODS

2.1 Specimens Examined

In total, 48 specimens of Oligodon were sequenced (Table 2.1), including three

unidentified specimens, two of which appeared very similar to Oligodon chinensis, but would

have represented a substantial and unusual range extension for that species. The sequenced

individuals represented 17 of 70 named species and the collecting localities were mapped (Figure

2.1). The sequence data for O. modestus were a synthesis of a partial sequence from ROM17490

for 12S rRNA, tRNAval

and the 16S rRNA data of Lopez and Maxson (1996).

Sequenced specimens of Oligodon were primarily from Vietnam and Myanmar.

Specimens were selected firstly to maximize species representation in the analysis. I also

attempted to evaluate inter- and intra-population genetic variation within species when possible

by including multiple specimens.

I selected Phyllorhynchus decurtatus, a putative close relative of Oligodon, as the initial

outgroup taxon and sequenced a specimen de novo for the equivalent mtDNA regions used in the

ingroup. Sequences in GenBank (http://www.ncbi.nlm.nih.gov/GenBank) for more distantly

related colubrids, including Dinodon semicarinatus and Pituophis lineaticollis, were also used as

additional outgroup taxa.

10

Table 2.1 A list of specimens chosen for DNA sequencing, their voucher numbers

and locality data. For some specimens in the ROM collection, denoted *, field numbers are

provided in lieu of a catalogue number. ROM=Royal Ontario Museum; CAS=California

Academy of Science; TNHM=Texas Natural History Museum; UMMZ=University of Michigan

Museum of Zoology; USNM=United States National Museum.

Species

Voucher

reference Locality

Dinodon semicarinatus GenBank AB008539

Oligodon arnensis ROM 19155 Assam, India

O. barroni ROM 32464 Krong Pa, Gai Lai, Vietnam

O. chinensis ROM 25757* Chi Linh, Hia Duong, Vietnam

O. chinensis ROM 34540 Chi Linh, Hia Duong, Vietnam

O. chinensis ROM 35626 Quang Thanh, Cao Bang, Vietnam

O. chinensis ROM 26940* Quang Thanh, Cao Bang, Vietnam

O. chinensis ROM 31032 Tam Dao, Vinh Phu, Vietnam

O. chinensis ROM 30824 Pac Ban, Tuyen, Vietnam

O. sp. ROM 30971 24km W of Con Cuong, Nghe An, Vietnam

O. sp. ROM 30970 24km W of Con Cuong, Nghe An, Vietnam

O. cinereus ROM 25755* Chi Linh, Hia Duong, Vietnam

O. cinereus ROM 32462 Chi Linh, Hia Duong, Vietnam

O. cinereus ROM 29552 Tam Dao, Vinh Phu, Vietnam

O. cinereus ROM 37092 Cat Tien, Dong Nai, Vietnam

O. cinereus CAS 210159 Alaungdaw Kathapa Ntl. Pk., Sagaing Div., Myanmar

O. cinereus CAS 215597 Alaungdaw Kathapa Ntl. Pk., Sagging Div., Myanmar

O. cinereus CAS 205028 Rakhine Yoma Mtn. Rng., Rakhine State, Myanmar

O. cinereus CAS 213379 Hlaw Ga Park, Yangon Div., Myanmar

O. cinereus CAS 215261 Kalaw Tnshp., Shan State, Myanmar

O. cinereus ROM 30969 24km W of Con Cuong, Nghe An, Vietnam

O. cruentatus CAS 213271 Hlaw Ga Park, Yangon Div., Myanmar

O. cruentatus CAS 213408 Hlaw Ga Park, Yangon Div., Myanmar

O. cyclurus CAS 204963 Mwe Hauk, Ayeyarwady Div., Myanmar

O. cyclurus CAS 215636 Alaungdaw Kathapa Ntl. Pk., Sagging Div., Myanmar

O. formosanus ROM 25827* Chi Linh, Hia Duong, Vietnam

O. formosanus ROM 35806 Chi Linh, Hia Duong, Vietnam

O. formosanus ROM 35629 Quang Thanh, Cao Bang, Vietnam

O. formosanus ROM 35630 Quang Thanh, Cao Bang, Vietnam

O. formosanus ROM 30939 Ba Be, Cao Bang, Vietnam

O. formosanus ROM 30823 Pac Ban, Tuyen, Vietnam

11

O. formosanus ROM 30826 Tam Dao, Vinh Phu, Vietnam

O. maculatus TNHC 59846 Barangay Baracatan, Mindinao Is., Philippines

O. modestus ROM 17490 Valencia, Negros Is., Philippines

O. modestus 16S mRNA from Lopez and Maxson 1995

O. ocellatus ROM 32261 Yok Don, Dak Lak, Vietnam

O. octolineatus UMMZ 201913 3Km E of Tutong, Tutong Dist., Brunei

O. planiceps CAS 213822 Shwe Set Taw Wldlf. Sanc., Magwe Div., Myanmar

O. splendidus CAS 204855 Kyauk Se Tnshp., Mandalay Div., Myanmar

O. splendidus USNM 520626 2Km WNW Chatthin, Chatthin Wldlf. Sanc., Burma

O. taeniatus USNM 520625 2Km WNW Chatthin, Chatthin Wldlf. Sanc., Burma

O. taeniatus ROM 32260 Yok Don, Dak Lak, Vietnam

O. taeniatus ROM 37091 Cat Tien, Dong Nai, Vietnam

O. theobaldi CAS 210710 Naung U Tnshp., Mandalay Div, Myanmar

O. theobaldi CAS 213896 Shwe Set Taw Wldlf. Sanc., Magwe Div., Myanmar

O. torquatus CAS 210693 Pakokku Tnshp., Magwe Div., Myanmar

O. torquatus CAS 210695 Pakokku Tnshp., Magwe Div., Myanmar

O. torquatus CAS 215976 Min Gone Taung Wldlf Sanc, Mandalay Div,.Myanmar

O. venustus ROM 19156 India

O. sp. ROM 27049* Quang Thanh, Cao Bang, Vietnam

Phyllorhynchus

decurtatus

ROM 14153 Ocotillo, Imperial County, California, United States

Pituophis

lineaticollis

GenBank AF512746

12

Figure 2.1 Composite satellite photograph of Southeast Asia, showing relief and topographic features CONT’D NEXT PAGE… 1

2

13

CONT’D FIGURE 2.1 and the collection localities of specimens of Oligodon from Vietnam and Myanmar. The localities from

Brunei, India and the Philippines are indicated by arrows in their approximate directions. CB=Ba Be & Quang Thanh, Cao Bang,

Vietnam; PB=Pac Ban, Tuyen, Vietnam; CL=Chi Linh, Hia Duong, Vietnam; TD=Tam Dao, Vinh Phu, Vietnam; CC=24km W of Con

Cuong, Nghe An, Vietnam; KP=Krong Pa, Gai Lai, Vietnam; YD=Yok Don, Dak Lak, Vietnam; CT=Cat Tien, Dong Nai, Vietnam;

VP=Valencia, Negros Is., Philippines; BP=Barangay Baracatan, Mindinao Is., Philippines; TB=3Km E of Tutong, Tutong Dist., Brunei;

IN=India; MH=Mwe Hauk, Ayeyarwady Div., Myanmar; HG=Hlaw Ga Park, Yangon Div., Myanmar; RY=Rakhine Yoma Mtn. Rng.,

Rakhine State, Myanmar; KS=Kyauk Se Tnshp., Mandalay Div., Myanmar; KT=Kalaw Tnshp., Shan State, Myanmar; SS=Shwe Set Taw

Wldlf. Sanc., Magwe Div., Myanmar; NU=Naung U Tnshp., Mandalay Div, Myanmar; MGT=Min Gone Taung Wldlf Sanc, Mandalay

Div,.Myanmar; PT=Pakokku Tnshp., Magwe Div., Myanmar; AK=Alaungdaw Kathapa Ntl. Pk., Sagging Div., Myanmar; CWS=2Km

WNW Chatthin, Chatthin Wldlf. Sanc., Myanmar.

13

14

Tissue samples included either skeletal muscle or liver tissue preserved in 95% ethanol at the

time of collection and subsequently stored in either ethanol or frozen at -80°C. Voucher

specimens are preserved in the collections of the California Academy of Sciences (CAS), the

Royal Ontario Museum (ROM), Texas Memorial Museum (TNHC), Museum of Zoology,

University of Michigan (UMMZ) and National Museum of Natural History (USNM).

2.2 DNA Extraction, Amplification and Sequencing

Three contiguous mtDNA genes were selected for phylogeny reconstruction, including

12S ribosomal RNA (rRNA), 16S rRNA and tRNA Valine (tRNAval

). Total DNA was obtained

from alcohol preserved and frozen tissue samples using a preparation of 500!L 0.1M STE, 50!L

20% SDS and 12.5!L 20mg/mL Proteinase K (GIBCO) added to 0.2g of well-macerated tissue.

The preparation was incubated at 37°C for 24–48h with several intervals of mixing.

DNA was purified from the cell lysate using a phenol-chloroform-isoamyl alcohol (PCI),

chloroform-isoamyl alcohol (CI) extraction. A volume of 450!L PCI was added to the digested

DNA, mixed well and centrifuged at 13000 x g for 5min. The supernatant was transferred to a

new tube and the PCI step was repeated 2–3 times. Subsequently, the DNA supernatant was

cleared of remaining phenol with a 450!L CI wash, centrifugation and final supernatant draw-

off. Final volume of DNA suspension product was 30–120!L. The presence of unsheared

mtDNA was tested by adding 10!L of product to a 1% agarose/TAE buffered electrophoretic gel

with ethidium bromide. Results were visualized on a ultra-violet light table and recorded with

Polaroid film or a digital camera. The DNA extraction procedure was repeated if the sample

failed to amplify with the polymerase chain reaction (PCR).

15

Specimens of O. arnensis, O. modestus and O. venustus were collected predating the

practice of separately preserving tissues for molecular investigation, and were fixed with at least

some formalin rendering them refractory to analysis with the above extraction protocol. Thus, a

extraction protocol modified after Chatigny (2000) was used. Small pieces (>0.1g) of loose or

free tissue were harvested from the specimen/container. These were transferred to 1.5mL tubes

and frozen using a dry-ice ethanol mixture and then ground to a fine powder using a pestle sized

for 1.5mL conical bottom centrifuge tubes. These powdered tissues were digested with the

STE/SDS/Proteinase K buffer above, but with the following changes: Proteinase K concentration

was doubled; DTE (Dithioerythritol) was added as 2.5 !L of 20mg/mL stock solution to reverse

protein cross-linkages; incubation temperature was increased to 55°C. The samples were

incubated for 24h, then an additional volume of Proteinase K and DTE was added and the

samples incubated for a further 48h. The extracted DNA was subjected to the PCI/CI clean-up

described above. This produced DNA from which short PCR amplicons could be made.

The primers 12S1L, located at approximately 410 base pairs (bp) from the beginning of

the 12S ribosomal RNA (rRNA) region, and 16S2H, located approximately 200 bp from the

terminus of the 16SrRNA region, were used for amplification of purified mtDNA. These primers

flank a region of approximately 1800 continuous bp. The general amplification and sequencing

strategy was to use these primers to obtain an initial, continuous PCR amplicon, and to use

internal primer sites to sequence overlapping regions. However, in many cases, and especially

with the formalin-fixed samples, shorter amplicons were produced and sequenced leaving some

internal sequence corresponding to the priming region unknown. The complete list of primers

used in this study and their relative positions are presented in Table 2.2 and Figure 2.2.

16

Table 2.2 Primer sequences used to PCR amplify and sequence fragments of the

mtDNA genome in this study. See also Figure 2.2.

Name Sequence 5' to 3'

12S1L CCA ACT GGG ATT AGA TAC CCC ACT AT

12S2LM ACA CAC CGC CCG TCA CCC T

16S3H GTA GCT CAC TTG ATT TCG GG

16S4H CCA GCT ATC ACC AAG TTC GGT AGG CTT TTC

16S1LM CCG ACT GTT GAC CAA AAA CAT

16S1H GGC TAT GTT TTT GGT AAA CAG

16S5H CTA CCT TTG CAC GGT TAG GAT ACC GCG GC

16S2H CCG GAT CCC CGG CCG GTC TGA ACT CAG ATC ACG

17

Figure 2.2 A representation of a linear double stranded section of the mitochondrial

genome including 12S rRNA, tRNAval

and 16Sr RNA. Arrows represent relative location of

primers used in this study.

18

Double stranded mtDNA was amplified in 25!L PCR reactions (Saiki, 1989). Reactions

were set up using 1!L of DNA (0.05 µg/!L), 1!L of each primer (10mM), 0.08!L of a dNTP

mix (containing 10mM of each of dATP, dCTP, dGTP and dTTP) (Perkin-Elmer Cetus Kit),

0.2!L of Amplitaq polymerase (Perkin-Elmer Cetus), 2.5!L of reaction buffer (containing 67

mM Tris, pH 8.8, 2 mM MgCl) (Sigma) and 19!L distilled deionized water. Thermocycling of

the reaction mix was performed on MJ Research PTL-200 and Perkin-Elmer GeneAmp 9700

machines. Cycling parameters were set as follows: 38 or 40 cycles; denature at 92°C for 30s;

anneal at 45°C for 45s; extension at 72°C for 45–90s, and a final extension cycle of 72°C for

5min. The annealing temperature was varied upward or downward to increase or decrease

primer-DNA specificity when using different primer combinations with different samples. The

extension time was increased when amplifying the longer fragments when using the 12s-1L

primer to either 16s-4H or 16s-2H. Some samples could not be amplified for the entire ~1800 bp

fragment; these were amplified in three overlapping segments of 12s-1L to 16s-4H, 12s-2LM to

16s-5H and 16s-1L to 16s-2H. Other less desirable amplicons (12S-2LM to 16S-1H) were also

used when sample DNA proved refractory to amplification owing to variation in the sequence at

the priming region. The combination of 12S-2LM and 16S-3H proved particularly effective for

the formalin fixed samples.

Double stranded DNA PCR product was subjected to electrophoresis in a 100mL 1%

agarose gel with 10!L ethidium bromide for 20–30min at 120V and 99 mA and visualized under

UV light. The amplified DNA fragments were cleaned of agarose and purified using either

GeneClean (Bio101/Can Scientific), a standard QiaQuick (Qiagen) protocol, or with a protocol

involving centrifuging the band through an aerosol resistant p20/200 pipette tip at 13000g for

5min. Cleaned DNA was suspended in distilled deionized water.

19

DNA sequencing was performed using either 33

P incorporation and autoradiograph film

or with dye-labeled nucleotides and fluorescent detection during electrophoresis. For radio-

sequencing, the Thermo Sequenase 33

P-labeled cycle sequencing kit (Amersham) was used on

Perkin-Elmer Cetus and Perkin-Elmer GeneAmp 9700 thermocyclers using the manufacturers

protocols. The sequencing reaction mix consisted of 8!L dH2O, 1!L of light or heavy strand

primer, 1.5!L of buffer, 1.5!L of sequenase, 3!L DNA and 6!L termination mix, divided into 4

(A, C, G, T) tubes. Subsequently, 0.37!L of the appropriate 33

P labeled ddNTP was added to

each ACGT tube and overlaid with a drop of paraffin oil. Upon completion of cycling, 3!L of

stop solution (Amersham) was added to each tube and samples were stored at -20°C until

electrophoresis (usually within 24hr).

Products of the isotope-sequencing reactions were resolved in a 40cm long 7M urea-5%

polyacrylamide-0.6% TBE buffer gel electrophoresed at 65V for 2hr or 6hr (short run and long

run). The gels were blotted onto filter paper, dried under heat and vacuum and exposed for 2–5

days on Kodak XAR autoradiograph film. Gels were developed for 5min and fixed for 2min with

Kodak developer and fixer.

Alternatively, some sequencing reactions were performed with the BigDye Terminator v

3.1 Cycle Sequencing Kit (Applied Biosystems) using: 1!L BigDye, 2!L 5x BigDye Terminator

Buffer, 2!L ddH20, 1!L of 10 mM primer solution, and 4!L PCR product. Following

sequencing reactions, samples were cleaned and precipitated with sodium acetate and ethanol.

These products were resolved on an ABI377 (Applied Biosystems) automated sequencer using

the manufacturer protocols.

20

2.3 DNA Sequence Alignment and Phylogenetic Analysis

Sequences were either read from X-ray films manually and entered into BioEdit 3.0

(Hall, 1997), or extracted from ABI pherogram files, and initially aligned using ClustalW 1.7

(Thompson et al., 1994) with a gap-opening penalty of 15 and extension penalty of 6.66. Aligned

sequences were imported into MacClade 4.0 (Maddison and Maddison, 1992) and final

alignment was achieved by eye. Nucleotide ratios and proportions including

transition/transversion ratios were calculated in MacClade. PAUP* 4.0 (Swofford, 2001) was

used to calculate pairwise divergences among samples using uncorrected p-distances.

The aligned sequences were analyzed with PAUP*4.0b10 (Swofford, 2001) using

maximum parsimony (MP) as the criterion for tree selection. Phylogenetically uninformative

sites were excluded, and heuristic searches were performed using random addition sequence with

10000 replicates while retaining minimal trees only, tree bisection-reconnection branch

swapping and collapse zero length branches. All multistate characters were evaluated as

nonadditive (unordered) and unweighted. Gaps were treated as “missing data”.

Bayesian inference (BI) was also used to infer phylogenetic relationships. MrModeltest

v.2.3 (Nylander, 2004) was used to obtain evolutionary models using the Akaike Information

Criterion (Akaike, 1974; Akaike, 1979). BI was performed using MrBayes v.3.1.2 (Huelsenbeck

and Ronquist, 2001). Two simultaneous runs of four chains were run for 3 X 106 generations,

sampling every 100 generations. Stationarity was assessed when likelihood scores reached a

stable equilibrium. A burn-in value of 7500 was implemented to discard topologies with low

likelihood scores prior to generating a 50% majority rule consensus tree. Node support within the

21

Bayesian consensus tree was assessed from the posterior distribution of topologies (Erixon et al.,

2003).

Nodal support was evaluated using non-parametric bootstrap proportions (BSP;

Felsenstein, 1985) and Bremer Decay Indices (DI; Bremer, 1988). BSP was based on 10000

replicates calculated in PAUP* and nodes receiving >50% support were noted on the most

parsimonious trees (MPTs). DIs were calculated by searching for trees in PAUP* using run

instructions produced by AutoDecay v4 (Eriksson, 1998) decay values are noted on the MPT on

nodes that also have >50% BSP.

Because hemipenial characters are potentially phylogenetically informative, they were

included as data in a total evidence approach. The data set was reduced to 17 ingroup taxa by

creating consensus DNA sequences for the nominal species. Variable base pair characters within

species were coded as multi-state characters. Three unordered, unpolarized hemipenial characters

were appended to the molecular data set. Hemipenial data for Phyllorhynchus were determined

from literature (Klauber, 1935; Savage and Cliff, 1954) to be as follows: hemipenis with spines,

no apical papillae, and partially forked (or “lobed”, a condition in some Oligodon, but not any

species for which sequence data were available). Zaher’s (1999) assignment of both these species

to the Colubrinae (Colubridae) is based on them bearing distal calyces and the single sulcus in

those Oligodon with undivided, unlobed (non-quadrapenes) organs.

Hemipenial characters were mapped on to phylogenetic tress using MacClade. Specimens

in Table 2.1 and literature records (Appendix A) were used to determine character states.

Hemipenial structure was described as three characters as follows: hemipenis with a forked

division (quadrapenis), unforked or partially forked; apical papillae present or absent; spines

present or absent.

22

Available sequences from GenBank were used to test the relationship of Oligodon with

Phyllorhynchus and to investigate where Oligodon might belong within the Colubridae. Table

2.3 shows 255 sequence fragments from eight genes plus known protein encoding loci shared

between Oligodon and the other colubrids. These sequences were assembled into a file creating

48 synthetic genera. Concatenation produced 6971 aligned nucleotide positions. An unweighted

MP analysis in PAUP* with 10000 random addition heuristic replicates and TBR branch

swapping was applied to the data set.

23

Table 2.3 Colubroidea sequences from GenBank used to determine the relationship of Oligodon to other colubrids. “x”

denotes a genes sequence that was not available for use in the sample, or synthetic representative. “+” denotes known protein loci data

from Dowling et al. (1996), their coding used. “1” indicates a sequence obtained from Lopez and Maxson, 1995. 12S/16S sequences for

Oligodon cinereus denoted “this study” are from specimen CAS205028. 12S/16S sequences for Pseudoxenodon bambusicola “this study”

are ROM30825. 12S/16S sequences for Lycodon fasciatus “this study” are ROM25654. Higher taxon names follow Zaher et al. (2009).

Genus Species 12S 16S c-MOS CytB NADH4 NADH1 NADH2 COI

Protein

loci

Colubridae incertae sedis

Grayia ornata AF158434 AF158503 AF544684 AF544663 x x x

smythii DQ112080 DQ112077

Calamariidae

Calamaria pavimentata x x AF471103 AF471081 x x x x +

Colubridae

Ahaetulla fronticincta x x AF471161 AF471072 x x x x +

Boiga cynodon AF139568 Z46525 x x x +

dendrophila AF471128 AF471089 U49303 x x x

Cemophora coccinea x 1(RH54131) AF471132 AF471091 AF138754 x x x +

Coluber constrictor U96794 L01770 AY486937 AY486913 AY487040 AY486963 AY487001 AY122649 +

Coluber dorri x AY188081 AY188001 AY188040 AY487042 AY486964 AY487003 x

Coluber zebrinus x AY188084 AY188004 AY188043 AY487058 AY486980 AY487019 x

Coronella girondica AY122835 AY643353 AF471113 AF471088 AY487066 AY486988 AY487027 AY122751

Crotaphopeltis tornieri x x AF471112 AF471093 x x AF428016 x

Dasypeltis atra x AF471136 AF471065 x x x x

scabra x 1(RH64432) x x x x +

Dinodon rufozonatum AF471163 AF471063

semicarinatus AB008539 AB008539 AB008539 AB008539 AB008539 AB008539 AB008539 +

Dipsadoboa unicolor x x AF471139 AF471062 x x AF428017 x

Dispholidus typus x AY188051 AY187973 AY188012 U49302 x x x

Elaphe quatuorlineata AY122798 AF215267 AY486955 AY486931 AY487067 AY486989 AY487028 AY122714 + 23

24

Eirenis modestus AY039143 AY376780 AY486957 AY486933 AY487072 AY486994 AY487033 AY039181

Gastropyxis smaragdina AF158435 AF158504 DQ112078 DQ112075 x x x x

Gonyosoma oxycephalum AY122679 Z46490 AF471105 AF471084 x x x AY122661 +

Hemerophis socotrae AY039132 AY188083 AY188003 AY188042 AY487055 AY486977 AY487016 AY039170

Hemorrhois nummifer AY039163 AY376771 AY376800 AY376742 AY487049 AY486971 AY4870010 AY039201

Hierophis jugularis AY039152 AY376769 AY486941 AY486917 AY487046 AY486968 AY487007 AY039190

Lycodon capucinus U49317 x x x

fasciatus This study This study x x x

laoensis Z46455 Z46485 x x x +

zawi AF471111 AF471040 x x x

Lytorhynchus diadema AY647229 AY188064 AY187986 AY188025 x x x x

Macroprotodon cucullatus AY643296 AY643337 AF471145 AF471087 AY487064 AY486986 AY487025 x

Masticophis flagellum AY122668 1(SM) AY486952 AY486928 AY487061 AY486983 AY487022 AY122650 +

Oligodon cinereus This study This study AF471101 AF471033 x x x +

octolineatus U49316 x x x

Oocatochus rufodorsatus AY122800 x AF435015 AF036016 x x x AY122716

Opheodrys aestivus x 1(RH61940) AF471147 AF471057 x x x x +

Oxybelis aeneus AF158416 AF158498 AF471148 AF471056 x x x x +

Pantherophis obsoletus AY122844 Z46493 AF471140 AF283644 AF138757 x x AY122760 +

Philothamnus heterodermus x 1(RH61045) AF471149 AF471055 x x x x +

Phyllorhynchus decurtatus AF544783 AF544812

AF471098 &

AF544728 AF471083 x x x x +

Platyceps rogersi AY039127 AY188082 AY188002 AY188041 AY487052 AY486974 AY487013 AY039165

Pseustes poecilonotus x 1(RH62677) x x x x x x +

Ptyas korros AF236680 AY486953 AY486929 AY487062 AY486984 AY487023 AY122652 +

mucosus 1(RH56036)

Rhinechis scalaris AY122802 x AY486956 AY486932 AY487068 AY486990 AY487029 AY122718 +

Salvadora mexicana x AY486958 AY486934 AY487075 AY486997 AY487036

grahamiae AY122847 1(RH56651) AY122662 +

Sonora semiannulata x x AF471164 AF471048 x x x x

Spalerosophis diadema AY039144 AF471155 AF471049 AY487059 AY486981 AY487020 AY039182 +

cliffordi 1(RH58139)

Spilotes pullatus x x AF471110 AF471041 x x x x

24

25

Tantilla melanocephala AF158424 AF158491 x x x x +

relicta AF471107 AF471045 x x x x

Telescopus fallax x AY188078 AF471108 AF471043 x x x x +

Thelotornis capensis x x AF471109 AF471042 x x x x

Toxicodryas pulverulenta x x AF471118 AF471047 x x x x

Natricidae

Natrix natrix AY122682 AF158530

AF471121 &

AF544697 AF471059 AY873736 AY873760 AY870640 AY122664

Thamnophis godmani AF471165 AF420135 AF420138 AF420136 AF420137 x

sirtalis AF402647 1(LM3071) x +

Pseudoxenodontidae

Pseudoxenodon bambusicola This study This study x x x x

karlschmidti AF471102 AF471080 x x x x

Dipsadidae

Alsophis cantherigerus AF544694 x x x

portoricensis AF158448 AF158517 AF471126 AF471085 U49309 x x x +

25

26

3 RESULTS

3.1 Sequence Variability in Oligodon

The aligned data set contained 1885 bp of homologous DNA sequence sites from 48

ingroup samples and three outgroup taxa. In nine cases, multiple individuals of the same species

and locality did not vary and, in these cases, only one sample was selected for phylogenetic

analysis. In the case of Oligodon cinereus CAS205028 from Rakhine Yoma, Myanmar and O.

cinereus CAS213379 from Hlaw Ga, Myanmar, no variation was discovered. Further, O.

theobaldi CAS213896 from Shwe Set Taw, Myanmar and O. theobaldi CAS210710 from Naung

U, Myanmar showed no sequence variation. However, these sequences were maintained in the

analysis to maximize geographic coverage.

DNA extracted from O. arnensis, O. modestus and O. venustus proved difficult to PCR

amplify. However, high quality sequences were obtained for approximately 500 bp (using 12S-

2LM and 16S-3H). About 400 additional bp were added to O. modestus from the literature.

The aligned data set includes a contiguous region of 507 bp from 12SrRNA, 64 bp of

tRNA valine (tRNAval) and 1314 bp of 16SrRNA for a total of 1875 bp. The number of variable

sites for each of the three gene segments is given in Table 3.1. The average and ranges of the

nucleotide frequencies across the ingroup taxa are as follows: A, 0.389 (0.380–0.400); C, 0.237

(0.224–0.246); G, 0.159 (0.157–0.172) and T, 0.214 (0.194–0.221). Overall, 713 (38%) sites

vary among ingroup and outgroup taxa, of which 513 are potentially phylogenetically

27

Table 3.1 A list of total sites, variable sites and uncorrected p-distances for the three mtDNA genes in the data set. Inter-generic measures range and mean of divergences between the ingroup and outgroup samples, intra-ingroup measures range and mean of p-distances among ingroup samples.

Gene

Total bp

variable sites (potentially parsimony informative), Ingroup

Range in pairwise divergence (average pairwise divergence) inter-generic intra-ingroup

12SrRNA 507 161 (116) 0.115 – 0.266 (0.166) 0.000 – 0.149 (0.083) tRNAval 64 16 (11) 0.000 – 0.198 (0.114) 0.000 – 0.186 (0.063) 16SrRNA 1314 441 (312) 0.110 – 0.188 (0.141) 0.000 – 0.149 (0.087) Total 1885 618 (439) 0.121 – 0.202 (0.147) 0.000 – 0.125 (0.084)

28

informative. Within the ingroup, 618 characters are variable, 439 of which are potentially

phylogenetically informative.

The overall transition:transversion ratio was 1.13:1. Uncorrected p-distances, the pair-

wise absolute number of bp differences, are summarized in Table 3.1, with average and range of

sequence divergence within the ingroup and an intergeneric comparison between outgroups and

the ingroup samples. Eight species in the analysis have representatives from multiple localities

included as terminal nodes. The p-distances for the average inter-locality variation for these

samples are as follows: O. chinensis 0.027; O. cinereus 0.036; O. cyclurus 0.040; O. formosanus

0.014; O. splendidus 0.011; O. taeniatus 0.029; O. torquatus 0.018.

3.2 Phylogeny

The unweighted maximum parsimony analysis using all DNA data resulted in one MPT

(2124 steps, CI = 0.488, RI = 0.710) with many nodes receiving strong support (Fig. 3.1). The

heuristic search found the tree island with the MPT in 99.4% of all replicates.

MrModeltest selected the GTR+I+G model of nucleotide evolution for these data. The BI

tree is shown in Figure 3.2 along with posterior probabilities. Maximum parsimony and BI

produced very similar topologies, differing only in the placement of O. modestus, O. maculatus

and O. octolineatus.

29

Figure 3.1 Maximum parsimony phylogeny of Oligodon. Values denoted above internodes are bootstrap proportions, numbers below internodes are decay index values.

30

Figure 3.2 Bayesian inference phylogeny for Oligodon. Values denoted behind the nodes are posterior probabilities.

31

Almost all species received high support (BS > 98%, DI > 10+) with

localities/populations showing monophyly. Exceptions to this were O. taeniatus and O.

chinensis. Both were monophyletic, but O. chinensis only received high support values when the

specimens from Con Cuong, Vietnam were included. Support for monophyly in O. taeniatus was

high (BS = 75%, DI = 4), even though the Chatthin (Myanmar) and Cat Tien, Yok Don

(Vietnamese) specimens represented extreme east-west sampling of the range and the localities

showed considerable genetic divergence with respect to each other.

A cyclurus-group (sensu Smith, 1943) is resolved within the MPT, albeit with moderate

support only (BS = 62%, DI = 4). It consists of O. cyclurus, O. ocellatus, O. formosanus, and O.

chinensis, and within this group a clade of three Vietnamese-Chinese species—O. ocellatus, O.

formosanus and O. chinensis—receives higher support. The BI analysis differs in not uniting O.

cyclurus with O. ocellatus+O. formosanus+O. chinensis.

A taeniatus-group (sensu Smith, 1943), consisting of O. taeniatus and O. barroni (BS =

99%, DI = 12), is resolved and this is sister to the cyclurus-group in the MPT. Philippine and

Sunda Is. O. octolineatus and O. modestus show a weakly supported relationship with the

cyclurus-group + taeniatus-group. The cyclurus-group + taeniatus-group is only recovered in the

BI tree if O. octolineatus is included.

Indian O. arnensis (a widespread species) and O. venustus (a restricted-range species)

show a well-supported association with the cyclurus-group, taeniatus-group, O. octolineatus and

O. modestus. The other major clade recovered in the tree consists mostly of mainland Asian taxa

including O. cinereus (a widespread and highly variable species), O. splendidus, O. maculatus

(Philippines), and ((O. torquatus, O. planiceps) (O. theobaldi, O. cruentatus)). This latter clade

32

contains three taxa with quite restricted distributions and O. theobaldi, which is less restricted in

distribution. The ranges of these species might overlap in Myanmar.

Oligodon, as a genus, received very strong support. It is the sister group of

Phyllorhynchus + Spilotes.

3.3 Hemipenial Characters

Reduction to 17 taxa allowed the branch-and-bound algorithm to be used for tree

building, which resolved a single MPT of 1420 steps, CI 0.598, RI 0.577 (Fig. 3.3). The topology

of the tree with the hemipenial characters was very similar to the MPT produced by the DNA

data alone (Fig. 3.1). The two trees differed only at weakly supported nodes. In one case, O.

splendidus moved from being the sister of O. cinereus to becoming the sister of O. maculatus (as

is found in the BI tree). In the second case, O. chinensis and O. formosanus obtained a sister

relationship to O. ocellatus. Tree instability was not due to the addition of new data, but rather to

the deletion of taxa with uncertain identity and those having sequences from combined

individuals. The branching pattern in Figure 3.3 was obtained in the molecular-only data set

when one taxon of uncertain identity (ROM27049) and one locality of O. formosanus (Quang

Thanh, Cao Bang, Vietnam) were deleted. Regardless, the hemipenial characters had the same

transformations on both trees.

Optimizing the three hemipenial characters on the phylogeny (Figure 3.3, A–C)

33

Figure 3.3 Hemipenial character state mapping for Oligodon. A, presence, absence, or partially divided/forked hemipenis. B, presence or absence of apical papillae on hemipenis (under ACCTRAN, equivocal internodes are papillate, under delayed transformation optimization they are non-papillate). C, presence or absence of spines on hemipenis. Several clades have been labeled to facilitate discussion. All characters optimized unordered.

34

shows that the bifurcate hemipenis arose once only. Apical papillae and spines appear to have

more instances of parallel, homoplastic state changes, but they are apomorphic and homologous

for some clades. Accelerated (ACCTRAN) and delayed (DELTRAN) transformation

optimizations display the same mapping for the hemipenial characters with one exception.

Mapping apical papillae character states produces two equivocal internodes within the ingroup.

Under ACCTRAN these become unequivocal for the presence of papillae, under DELTRAN

they are non-papillate.

3.4 Position of Oligodon in the Colubridae

Meta-analysis of colubrine sequences recovered a single MPT on a tree island found in

40% of the replicates (Figure 3.4, length 14314 steps). The tree had a clade consisting of

Oligodon + (Phyllorhynchus + Spilotes), the same as found in the MP analysis of Lawson et al.

(2005).

35

Figure 3.4 Maximum Parsimony analysis of 48 colubrids including Oligodon and outgroups using published sequences of 12SrRNA, 16SrRNA, nuclear gene c-Mos, Cytochrome b, NADH4, NADH1, NADH2, COI and protein loci (Acp-2, Ldh-2, Mdh-1 and Pgm from Dowling et al., 1996).

36

4 DISCUSSION

4.1 Molecular Variation

Slowly evolving 12S and 16S ribosomal RNA genes are appropriate for resolving older

divergences (~150 Ma) (Mindell and Honeycutt, 1990). Divergence in the rRNA sequences is

moderate for the 48 samples of Oligodon. Among 507 bp sequenced for 12S, 32% are variable

and 23% are potentially informative. Of 1314 sites in 16S, 34% are variable and 24% are

potentially informative. The ratio of A:C:G:T for Oligodon is similar to that of other snakes

(Kraus and Brown, 1998; Rodríguez-Robles and de Jesús-Escobar, 1999; Slowinski and Lawson,

2002) and typical of mtDNA. Analysis of p-distances for each gene reveals a similar range and

average of variation as described in other studies, such as on Bufo (Liu et al., 2000) and snakes

(Vidal et al., 2000). Taken together, this suggests that these characters are useful for

reconstructing history in Oligodon at the taxonomic level of species and possibly the haplo-

history between populations.

Evaluation of mtDNA genes as a single data partition is appropriate because mtDNA

genes do not freely segregate and recombine but instead are inherited as a single linkage group.

The data could be partitioned by transcriptional unit, but it is not necessarily true that these data

partitions are appropriate or “natural” (Eernisse and Kluge, 1993). There are no a priori grounds

to believe that knowledge of the transcriptional structure indicates that the whole sequence is

under the same evolutionary forces. The most appropriate scale for data partitioning is the

individual base pair.

37

The finding of no intra-locality variation within species for these genes is interesting, but

can be explained several ways. It may be that faster evolving genes are needed to assess genetic

variation at this level, that more samples need to be sequenced, or that populations may in fact be

quite isolated from each other. Nuclear gene sequences will be required to assess gene flow.

The p-distances within species of Oligodon with multiple populations were less than 50%

of the average p-distance among all populations and species. They ranged from being as low as

13% of the variation seen among the whole ingroup. The p-distances were generally correlated

with increasing geographic distance between localities (r2 = 0.664) indicating that isolation-by-

distance was a factor to consider.

4.2 Phylogeny and Previous Hypotheses of Relationships

Many relationships recovered in this phylogeny agree with prior proposals. Although

several species of Oligodon are absent from this study, the corroboration between genetic

characters and hemipenial morphology allows for sound speculation about the relationships of

several other taxa.

4.2.1 Philippine Taxa

Taylor (1922–25, 1965), Taylor and Elbel, (1958) and Leviton (1963a, 1963b) believed

that O. maculatus was closely related to Philippine O. ancorus, and then to O. purpurascens,

which was not included in this study. Oligodon maculatus is more closely related to O.

38

splendidus and O. cinereus. Oligodon purpurascens is a widespread species extending onto the

Malay Peninsula. Oligodon maculatus, O. splendidus and O. cinereus all have quite similar

hemipenes in their being papillate, non-spinose and proximally calyculate. The relationship of O.

purpurascens, which also has large hemipenial papillae, with O. cinereus or O. splendidus has

not been hypothesized. It is possible that the large papillae of O. purpurascens are homologous

with those in a clade that includes widespread and complex O. cinereus. However, another set of

relationships is also possible for O. purpurascens.

Like O. maculatus, Leviton (1963a) believed that O. modestus was related to O. ancorus

and O. purpurascens. However, this phylogeny refutes a close relationship between O.

maculatus and O. modestus. While they share large hemipenis papillae, this condition is not a

defining synapomorphy. Philippine diversity might be best explained by multiple invasions of

the islands by different clades. A close relationship between O. purpurascens and O. modestus

would allow O. maculatus to represent an invasion by an ancestor more closely allied with the

cinereus-group.

4.2.2 Oligodon cyclurus-taeniatus Group

Smith (1943) suggested that O. cyclurus and O. chinensis were most closely related, that

O. taeniatus and O. barroni were most closely related, and then these two groups were sisters.

He placed several other species in these groups as well. The larger grouping, which Smith

referred to as the cyclurus-taeniatus complex, is recovered in the MPT but not in the BI tree.

This study contains a fair sampling of the species Smith assigned to these groups.

Notwithstanding, there are several recent taxonomic changes in this group.

39

David et al. (2008b) described three new species, O. deuvei, O. pseudotaeniatus, and O.

moricei, and allied them with O. taeniatus. These new species suggest that genetic information

might be helpful for identifying cryptic species of Oligodon. This study included a specimen of

O. taeniatus from Myanmar (Chatthin Wldlf. Sanc., USNM520625). This animal was discussed

by David et al. (2008b) who reported that it was incorrectly identified by Zug (1998; catalogue

number incorrect in publication). I have not seen the voucher specimen, nor did David et al., who

reported it as lost, yet Zug’s description (Zug, pers. comm.) and the sequence data leave little

doubt that it is O. taeniatus. I did not examine USNM520624, which David et al. referred to as

“most likely” being O. theobaldi.

Several recent publications (David et al., 2004; David et al., 2008a; David et al., 2008b;

Pauwels et al., 2003; Pauwels et al., 2002; Stuart and Emmett, 2006; Stuart et al., 2006; Teynie et

al., 2004) draw attention to an opinion (Wagner, 1975) that O. fasciolatus is a species distinct

from O. cyclurus. Although this study does not include O. fasciolatus, that species occurs in the

center of what was once the nearly pan-Southeast Asian range of O. cyclurus. Future studies

should include populations of this taxon and additional populations of O. cyclurus to evaluate the

validity of this change. This study includes another taxon historically included in the variation of

O. cyclurus, i.e. O. ocellatus, as well as O. cyclurus sensu stricto. Oligodon ocellatus does not

form the sister group of O. cyclurus.

This study’s sampling of the cyclurus-taeniatus group contains two specimens of

uncertain identity (O. sp., from the vicinity of Con Cuong). Genetically these are quite similar to

O. chinensis, yet the locality is far outside that species’ known range. These snakes might

represent an undescribed species. A morphological examination of the specimens is underway.

40

4.2.3 Oligodon cinereus

This species has a considerable geographic range and is highly variable in colouration.

These samples are from the western and eastern areas of its range, but not the center. Predictably,

p-distance between these samples is moderately high (0.036) relative to other species in this

study. The samples used here include some of the colour varieties: ROM37092 from southern

Vietnam, Cat Tien National Park, represents the “palidocinctus” colour form, while the

specimens from Chi Linh, northern Vietnam, represent the moderately inornate form (Smith’s

type II). Other specimens are closest to Smith’s form III. Given that O. taeniatus is a species

complex (David et al., 2008b) and because the Myanmar and Vietnamese haplotypes are not

sister lineages, it is likely that O. cinereus is also complex of cryptic species. Additional

sequenced specimens will be required to solve this problem, especially specimens from the

central region of the range.

4.2.4 Oligodon torquatus-planiceps-cruentatus-theobaldi

Smith (1943) proposed a close relationship among O. torquatus, O. planiceps, O.

cruentatus and O. theobaldi, but was unsure of where to place O. torquatus. His grouping is

supported here and O. torquatus is resolved as the sister taxon of O. planiceps. The

autapomorphic loss of hemipenial spines in O. torquatus serves to diagnose that species. This

group differs from other spinose hemipenial Oligodon by the presence of apical papillae. Smith

described additional details of spine size and placement, which taken together indicated that my

character coding of spinose/not-spinose might have been overly simplistic.

41

One unidentified specimen (ROM27049, O. sp.) was associated with this clade, albeit

with low measures of support. Nikolai L. Orlov (Russian Academy of Sciences) is currently

examining the specimen. It could represent a major range extension of a known species, or

perhaps represents an undescribed species. Its hemipenes are unknown, but are hypothesized to

be spinose and have papillae.

4.3 Hemipenes and Phylogeny

Fifty of 70 recognized species have described hemipenes. Of the remaining species, most

are rare (from a highly restricted range, and/or not very abundant) or very rare (in some instances

known only from the types). Pope (1935) compiled hemipenial data for 12 Chinese species of

Oligodon. Smith (1943) described the condition in 33 species, including nine surveyed by Pope,

and divided Oligodon into seven groups based on hemipenial condition. Leviton (1963) added

data for four Philippine species and proposed recognizing three hemipenial groups of Oligodon.

Leviton (1953; 1960) noted the condition in O. annamensis. Zhao and Jiang (1981) described

and illustrated the organ of O. multizonatus. David et al. (2008a,b) noted the organ when recently

describing new species.

A survey of three museum catalogues (Field Museum of Natural History: Alan Reastar,

pers. comm.; CAS: Joseph B. Slowinski, pers. comm.; and ROM), indicates that a majority of

Oligodon are not sexed. This situation precludes group assignment of many species of Oligodon.

Unfortunately, this pivotal character is present in males only and its diagnostic features have

been neglected in authoritative but less academically oriented literature (Chan–Ard et al., 1999;

Cox et al., 1998; Manthey and Grossmann, 1997). Hemipenial morphology has also been

42

neglected or not examined comparatively in several important academic treatments (Campden–

Main, 1969; Campden–Main, 1970; Campden–Main, 1970 (1984); in den Bosch, 1994; Lazell et

al., 1999; Mahendra, 1984; Tillack and Günther, 2009; Yang and Inger, 1986; Zhao et al., 1998).

Using hemipenial microstructure, Smith (1943) divided the spinose hemipenial snakes

into four groups. I recover two spinose groups consistent with Smith’s system, and have no

refutation for the other two, although Smith reports that spines are more plastic then either

papillae or division of the organ into quadrapenes. Leviton (1963) proposed three groups as

follows: deeply forked with papillae present or absent; unforked, papillae usually present, spines

usually absent; and unforked, spines usually present, papillae usually absent. Only one of these

groups is recovered herein. Leviton groups all species with deeply forked hemipenis together,

but indicates that the individuals of both other groups could be spinose or have papillae. It is

possible that there are two distinct clades with spines. One clade could be usually (=most species

in the group, not most organs on individuals) without spines but with papillae and this would be

comprised of O. theobaldi, O. cruentatus, O. planiceps, O. torquatus, O. cinereus, and other

unsampled taxa. The other clade could correspond to species with spines but without papillae,

including O. arnensis, O. venustus, and others. However, these groupings will not capture all

species of Oligodon that also have unforked hemipenes.

Figure 3.3 shows that taxa tend to undergo subsequent modification of papillae and

spines, especially secondary loss. However, neither character is freely plastic. Secondary loss or

reduction of these ornamentations may be a key that reinforces speciation.

Hemipenial characters appear to be robust and diagnostic (Leviton, 1960; Pope, 1935;

Smith, 1943; Wall, 1923b), providing relief from reliance on colour and scalation for species

identification and phylogenetic corroboration (Eernisse and Kluge, 1993; Kluge, 1997; Siddall

43

and Kluge, 1997). The heritability of colour and scalation characters has been questioned in

Oligodon in particular (Smith, 1943), and, in general, their use in phylogenetics often rests on

invalid evolutionary assumptions (Murphy and Doyle, 1998).

4.4 Colour, Confusion and Biogeography

Much confusion in Oligodon has occurred because of extensive intraspecific variation in

colour and patterning. Most species of Oligodon have a fairly invariant colour pattern, but a few

species, namely O. cyclurus, O. fasciolatus, O. cinereus and O. taeniolatus, are highly variable.

The venters are described for some species of Oligodon, but not others. In some species,

individuals may be checkered or not. The dorsal colour of Oligodon ranges from brown and

black to green and orange with highlights in black, white, red and yellow. Overall colour can

change with age and contrast in pattern can be lost. The overall dorsal pattern can be very plain

with loose, indistinct reticulation confined to scale margins, as in one form of both O. cyclurus

and O. formosanus. It can also be more complex with varying numbers of longitudinal stripes,

crossbars, or spots ranging in size and shape. Some species combine elements of all of the above.

Taken together, Oligodon cyclurus and O. fasciolatus have five distinct colour morphs, O.

taeniolatus has five, and O. cinereus has four. Geographic overlap and intergradations of the

colour morphs are poorly understood.

The highly variable species are also widespread. Oligodon cyclurus and O. fasciolatus

range from India eastward through Thailand (at approximately 11° 15' N), to Vietnam and

southwestern China (Yunnan). A southerly separation of about 500 km on the Thai peninsula

separates O. fasciolatus from O. purpurascens, which continues throughout Indonesia. The

44

distribution of O. cinereus closely matches cyclurus+fasciolatus. Oligodon taeniolatus ranges

throughout India to eastern Iran and northward to southern Turkmenistan.

Many of Smith’s (1943) seven hemipenial groupings are corroborated here. None are

refuted. If his assignments are mostly correct, then several clades will contain a morphologically

variable widespread species with a range that will encompass that of other clade members. This

morphologically variable species will also encompass much of the colour and pattern variation

found in other clade members. Thus, the sympatric species will have similar colours and

patterns.

Pleistocene ecological changes (Hall and Holloway, 1998) may have lead to the isolation

of populations of widespread species that subsequently accrued morphological changes. This is

consistent with the idea that many of the extant species of Oligodon appear to have a subset of

the variation of one of the widespread species. Oligodon can be found on many Pleistocene

landbridge islands. Glacial cycles certainly could have played a role in speciation throughout the

Philippines and Indonesia (Karns et al., 2000).

4.5 Taxonomic Implications

Kukri snakes were originally partitioned into two genera based on dentition. The genus

Simotes Duméril, Bibron and Duméril 1854 was established to include kukri snakes that had

palatine and pterygoid teeth (Günther, 1864), whereas Oligodon Fitzinger 1826 was reserved for

those lacking this dentition (Günther, 1864; Boulenger, 1894). Holarchus Cope, 1886 (Pope,

1935; Stejneger, 1907) was used later to include most of the same species originally assigned to

45

Simotes Duméril, Bibron and Duméril 1854, but this group was redefined (by Pope) based on the

absence of hemipenial spines and the highly correlated condition of having a divided anal scale.

As late as 1970, Deuve continued to recognize Holarchus and Leviton (1963) speculated that

there might be two clades of Oligodon, one that could carry the name Holarchus. Most of the

kukris surveyed herein were originally assigned to Simotes based on dentition. Oligodon

modestus is an important exception in lacking palatine and pterygoid teeth. Based on hemipenial

and anal scale conditions, two clades in this study could be assigned to Oligodon: clade A and

clade C, Fig. 3.3. However, these two lineages are not monophyletic, and the clade that would

carry the name Oligodon is uncertain because the type species of Oligodon, O. bitorquatus, is

not included. Regardless, this arrangement precludes a monophyletic Simotes and/or Holarchus.

The division of Oligodon drawing on any previous formulation would require several additional

genera to be erected.

Nomenclatorial stability should be a primary concern. A phylogenetically informed

taxonomy contains the most information within a hierarchical system (Brooks and McLennan,

1991; Brooks and McLennan, 2002; Farris, 1967; Wiley, 1980). Until a far more complete

sampling of species is possible, Oligodon should not be subdivided into multiple genera.

4.6 Oligodon: Position Within the Colubridae/Colubrinae

To determine whether Oligodon is an unusually speciose genus of snakes, it is necessary

to determine its sister group relationships, and not just an appropriate outgroup (Brooks and

McLennan, 1991). It is also important to establish the sister group of Oligodon because outgroup

46

choice can be critical in phylogenetic reconstruction (Milinkovitch and Lyons-Weiler, 1998;

Tarrío et al., 2000; Ware et al., 2008).

Relationships within the Colubridae have been discussed elsewhere (Dowling, 1974;

Dowling et al., 1996; Kraus and Brown, 1998; Lawson et al., 2005; Lopez and Maxson, 1996;

Zhang et al., 1984) and they will be the subject of future studies. Oligodon has been included in

several of these studies, yet no one has used all of the available data from molecular

investigations to hypothesize the position of Oligodon relative to other colubrines. Kelly et al.

(2003) used much of the available molecular data in a meta-analysis of snakes, but used too few

colubrines (and no Oligodon) to address this question. Most studies agree that Oligodon falls

within the Colubrinae. Two studies (Dowling et al., 1996; Lawson et al., 2005) suggest that

Oligodon is related to New World Phyllorhynchus, but they disagree on the placement of these

two genera within the Colubrinae. The close relationship between a relatively speciose Asian

Oligodon and a relatively depauperate North American genus (Phyllorhynchus) remains an

unintuitive result.

This analysis (Fig. 3.4) did not refute a close relationship between Oligodon and

Phyllorhynchus (+Spilotes). However, this clade did not share a close relationship with other

New World colubrines. It diverged early from the remaining Colubrinae. This finding is

consistent with proposal of the Oligodontini by Dowling (1974), Dowling and Duellman (1978)

and Dowling et al. (1996). The positioning of Oligodon herein is also consistent with that of

Zhang and Zhao (1984).

47

5 CONCLUDING REMARKS

Oligodon is a genus worthy of future investigation. The rarity of specimens in collections

and in the field across most, if not all of their distributions, will continue to be a key challenge to

future work. There are relatively few specimens of Oligodon in museum collections, and even

fewer have tissues separately preserved for molecular investigation. Political instability across

much of the range of these species in the twentieth century often forces researchers to take an

artificially geographically restricted approach to work. Notwithstanding these limitations, this

study produced several interesting results.

The hemipenial morphology of Oligodon is phylogenetically constrained and, therefore,

informative. The hemipenes are useful in species identification, and it is tantalizing to speculate

that the ornamented organ may play a role in speciation or maintenance of species. Speculation

on cladistic relationships by Smith (1943), based on hemipenial morphology, are validated

herein, even where they have been challenged by others. Thus, new species of Oligodon should

be based on a type series that includes males with described hemipenes.

Oligodon needs taxonomic stability. Division of the genus is not justified at this time.

Oligodon may have diverged early from the other colubrids accounting for its speciose nature.

Possible connections of Asia and North America, traversable by snakes, are discussed by

Burbrink and Lawson (2007). If Phyllorhynchus arrived in North America in the early Oligocene

with rat snakes, Oligodon may well have been the lone member of its clade in Asia for up to 20

Ma, undergoing repeated habit fragmentation and reconnection. As molecular clock methods

improve, this hypothesis will become increasingly testable. The membership of Spilotes in a

48

clade with Phyllorhynchus and Oligodon should be treated with great skepticism. Out of all of

the nucleotides that were aligned for the colubrid analysis, only a minimum subset is known for

Spilotes.

My phylogeny may be applied to future surveys of feeding and defensive behaviors. Egg-

eating and defensive displays in this genus may not be uniform. If they are not ubiquitous, then

the diversity can be mapped to this phylogeny thus inferring phylogenetically unique events.

Better defined distributions will facilitate biogeographic investigations of Oligodon and these

might reveal historical connections between areas and identify barriers to dispersal in Southeast

Asia. Range overlap between kukri snakes that are relatively distantly related might reveal

historical biogeographic patterns in sub-clades. Phylogeny and biogeography of Oligodon could

improve our understanding of speciation modes and they could be taken into consideration when

planning for conservation in Southeast Asia.

49

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Appendix A

Annotated checklist of the extant species of Oligodon

This checklist is designed to consolidate and update the disparate regional accounts on

the genus Oligodon, and describe each species with consistent terminology. Tillack and Günther

(2009) pointed out that reductions in the number of head scales or their fused states is not

uncommon in burrowing and fossorial snakes, like Oligodon. In Oligodon, fusion and reduction

seems especially common in supralabials, temporals and the loreal. This list follows Wall

(1923b) in describing a correlated loss of a supralabial and gain of a temporal as usually being

more accurately described as a supralabial excluded from the labial border. Inconsistency among

authors examining this character has added much spurious variation to the literature.

Notwithstanding, this list follows the current convention and departs from Wall in describing the

subocular as a different kind of scale, whereas Wall held it to be a supralabial. Inconsistency

among authors in the treatment of this character, as well as distinguishing creased parietals from

additional temporals, has increased the perception of variation in many species. Some species of

Oligodon have up to 2 preoculars plus a distinct subocular, others have one or two preoculars.

This list identifies second preocular, if present, but it can be categorized as a presubocular.

Many species exhibit some combination of head markings often called “typical” for the

genus. These include: an ocular bar passing from the supralabials under the eye, through the eye

and usually meeting its partner on the prefrontals; a temporal bar, from the corner of the mouth

or below, to the temporals and often continuing to the parietals; a nauchal chevron, varying in

shape, often from the neck on either side forward to an apex on the parietals or frontal, often

confluent at the rear with other body markings. Many works describe confluent temporal bars

and the ocular, or a “prefrontal bar,” as chevron-shaped. This list uses the term “chevron” for the

mark on the neck, even when the triangular space between the arms of the chevron are more or

less filled. A “subocular spot” is taken to be part of an incomplete ocular bar. Colour is described

in the form of dark markings on a light background to facilitate comparisons between species.

Except for a few cases where the darkest colours dominate, most authors follow this convention.

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Ventral colour in many species is confounding due to a lack of consistency in the literature.

Ventral colouration is inconsistently described from life or in preservative. Whenever possible,

this list gives the colour in life. The ventral colour of Oligodon can be whitish or yellowish to

pinkish and reddish with or without black or brown spotting. Contrary to some authors, ventral

colour does not appear to vary in life, at least within some species. Literature sources are also

inconsistent on the effect of preservative on dorsal colour. Consequently, presence or absence of

dorsal longitudinal stripes in a colour pattern should be treated very carefully. It seems that many

species are variable for faint longitudinal stripes that fade quickly in preservative.

Oligodon Fitzinger 1826.

DIAGNOSIS: Maxillary teeth 6–16, the posterior very enlarged and compressed; palatine teeth

well developed or vestigial; head short, not distinct from neck; round pupil. Rostral large and

when viewed from above, protruding. Cylindrical body, paired subcaudals. Usually 1 preocular.

First pair of infralabials usually in contact behind the mental. Anterior chin shields usually

longer then posterior. Dorsal scale rows at the neck and 2 head lengths behind the head are

usually equal to those at midbody, especially in smaller species, but there are many cases of an

increase or reduction after the occipit and neck. There is potential for confusion in some species

in which there is a scale row reduction near mid-body. Tillack and Günther (2009) have pointed

out that measuring midbody by total length as opposed to the middle ventral count location can

make a difference. Mid-ventral location be used to determine mid-body position.

Synonymy: Oligodon H. Boie in Fitzinger 1826: 25 (type Coluber bitorquatus) Simotes Duméril, Bibron and Duméril 1854: 624 (nec. Fischer 1817, Mammalia) Wall 1923b: 305 (argues for synonymy of Simotes with Oligodon) Tripeltis Cope 1886: 487 (type O. brevicauda Günther) Holarchus Cope 1886: 488 (type later designated as O. formosanus Günther by Pope 1935) Stejneger 1907: 353 (indicates Simotes preoccupied, designates Holarchus to stand instead) Pope 1935: 288 (argues for continued recognition of Holarchus based on hemipenes) Dicraulax Cope 1893: 480 (type S. trinotatus Günther)

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Oligodon affinis

Oligodon affinis Günther 1862: 58 (Anamallays, India: BMNH) DISTRIBUTION: India (Western Ghats south of the Goa Gap). COLOURATION: Dominant dorsal colour brown. Head markings black, with an ocular bar,

temporal bars and small chevron all present, but confluent on the frontal and parietals. Body with

indistinct darker reticulations and narrow (5–7 rows broad), often broken and indistinct,

crossbars. Crossbars often with lighter edging. Ventral colour (in alcohol) white with black

quadrangular spots, many confluent across ventrals.

SCALATION: Nasal divided. Two internasals. No loreal (most authors, but see Wall, 1923b:

“Loreal. Rarely present.”). Posterior nasal sometimes in contact with preocular. Two postoculars.

Temporals 1+2. Seven supralabials, third and fourth in contact with eye. Seven infralabials.

Dorsal scales in 17 rows at midbody. Ventrals 129–145. Anal divided. Caudals 23–37.

DENTITION: Seven maxillary teeth.

HEMIPENIS: Not forked. No papillae. Distally, with longitudinal folds and flounces and very

small spines. Proximally spinose.

COMMENT: Its IUCN status is “Lower Risk near threatened”. Key Literature: (Boulenger,

1894; Günther, 1864; Smith, 1943; Wall, 1923b). Figured by Smith (1943, Fig. 79). Max. length:

340mm (tail 50).

/RGB 209 46 53 Oligodon albocinctus

Coronella albocincta Cantor 1839: 50 (Cherrapungi, Assam, India: Sketch in Bodleian Library) Simotes albocinctus Boulenger 1890a: 312 [refers albocinctus to Simotes] Oligodon albocinctus Wall 1923b: 326 [new combination] Holarchus albocinctus Pope 1935: 289 [new combination] Oligodon albocinctus Smith 1943: 211 [rejection of Holarchus] Coronella puncticulatus Gray 1853: 389 (Khassia Hills, India: BMNH) Simotes punctulatus Günther 1864: 217 [refers puncticulatus to Simotes] Oligodon purpurascens vars. D, E Günther 1858: 25 [referred into punctulatus by Günther 1864] Simotes amabilis Günther 1868: 416 (Arrakan Hills, Myanmar: BMNH) Simotes albocinctus Boulenger 1890a: 312 [synonymizes punctulatus and amabilis] DISTRIBUTION: Eastern Nepal, Bangladesh, India (Assam; Sikkim; Arunachal Pradesh), Myanmar

(north of the Arrakan Hills), China (Xizang; Yunnan).

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COLOURATION: Dominant dorsal colour various shades of brown to red. Head markings

dark (in the reddest specimens they may be yellow), with an ocular bar, temporal bars and

chevron. The head markings may or may not be confluent on the frontal, the chevron blending at

the back with the body colour. Body with 17–27+4–8 distinct white-yellow black edged

crossbars (the amabilis form/Boulenger var. B, double this number, spaced twice as dense).

Occasionally faint longitudinal stripes are seen. Ventral colour whitish to pinkish with black

quadrangular spots

SCALATION: Nasal divided. Two internasals. Loreal present. One (rarely 2 on one side)

preoculars. Two postoculars (rarely 1 on one side). Temporals 1+2 (rarely 2 anterior or 3

posterior on one side). Seven supralabials, third and fourth in contact with the eye (rarely 8

(4,5)). Eight infralabials. Dorsal scales in 19 rows at midbody (very rarely 21). Ventrals 177–

210, laterally angulate. Anal undivided. Caudals 42–69.

DENTITION: Ten to 12 maxillary teeth.

HEMIPENIS: Not forked. No spines. Distally, with longitudinal folds, flounces and papillae.

Proximally calyculate.

COMMENT: Despite relatively many known collections, its IUCN status is “Data Deficient”.

Smith (1943), describes two sympatric colour forms which do not intergrade but his figure

caption (Fig. 67) transposes 67A and 67C. Figure 67C is form I, forma typica. One colour is

apparently O. juglandifer (see comments under that taxon). Key literature: (Boulenger, 1894;

Smith, 1943; Wall, 1923b; Zhao et al., 1998). Figured by Wall (1914a, Pl. XX), Smith (1943,

Fig. 67, body only) and Zhao et al. (1998, Fig. 50, head only). Max. length: 915mm (tail 100).

/ 209 241 53 Oligodon ancorus

Xenodon ancorus Girard 1858: 182 (Manila, Luzon, Philippines: USNM5521) Holarchus ancorus Cope 1895: 200 [new combination] Simotes purpurascens var. C Günther 1858: 25 [in part] Simotes phaenochalinus Cope 1860: 244 (Manila, Luzon, Philippines: ANSP3523) Holarchus phaenochalinus Griffin 1911: 259 [new combination] Holarchus ancorus Taylor 1918: 361 [corrects priority of ancorus over phaenochalinus] Simotes aphanospilus Cope 1860: 245 (Philippines: ANSP3521) Simotes ancoralis Jan 1863a: 233 (no loc.: NMW) Simotes phaenochalinus Boulenger 1894: 225[syns. ancoralis, aphanospilus, purpurascens] Simotes russellii Jan 1863b: 45 [in part]

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Holarchus burksi Taylor 1918: 365 (Sumagui, Mindoro, Philippines: CM2575) Simotes burksi Werner 1929: 134 [new combination] Oligodon ancorus Leviton 1963a: 463 [full evaluation, synonymizes burksi, russelli] Oligodon rhombifer Werner 1924: 36 (Padang, Indonesia: NHMW23436a,b) DISTRIBUTION: Philippines (Luzon and Mindoro islands). A highly detailed list of collection

localities and discussion of spurious reports is given by Leviton (1963a).

COLOURATION: Dominant dorsal colour brownish-lavender. Head markings dark, with an

ocular bar, temporal bars and thick chevron. Temporal bars may be confluent with each other

and the point of the chevron or chevron point may reach between separate temporal bars and

touch ocular bar. Body with 14–22 dark, black-edged spots. Traces of narrow reticulations or

irregular spots between the well defined ones. Often a yellowish vertebral streak present. Ventral

colour cream-yellow with occasional dull brown spots.

SCALATION: Nasal partially divided rarely entire or completely divided. Two internasals.

Loreal present. One, rarely 2 preoculars. Two, rarely 1 postoculars. Temporals usually 1+2 rarely

1+1, 2+2 or 1+3. Normally 7 supralabials rarely 6 or 8 on one side, normally third and fourth in

contact with eye, rarely fourth and fifth. Normally 8 infralabials, sometimes 6 or 7. Dorsal scales

in 17 rows at midbody. Ventrals 147–173. Anal undivided, rarely divided. Caudals 32–46.

DENTITION: Seven to 8, rarely 9 maxillary teeth.

HEMIPENIS: Not forked. Two large papillae. No spines. Distally, with fine longitudinal folds

merging into a proximal calyculate half.

COMMENT: Its IUCN status is “Near threatened”. Thoroughly reviewed by Leviton (1963a),

who believes it was derived from an isolated population of a widespread “purpurascens-like

ancestor”. Recently, the type material of O. rhombifer in NHMW (Tiedemann and Häupl, 1980;

Werner, 1924) was reviewed and placed in the synonymy of O. ancorus (Tillack, 2008).

Werner’s type locality was in error. Key Literature: (De Rooij, 1917; Leviton, 1963a; Taylor,

1922). Figured by Jan and Sordelli (1865, L.11, Pl. 4, Fig. 2) and Taylor (1918, Pl. 2). Max.

length: 666mm (tail 108).

/209 46 209 Oligodon annamensis Oligodon annamensis Leviton 1953: 423 (Blao, Haut Donai, Vietnam: USNM90408)

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DISTRIBUTION: Vietnam (Blao, Haut Donai). COLOURATION (in alcohol): Dominant dorsal colour brown, scales often darker edged and with

fine dark flecks. Head markings are black-edged white blotches. Instead of ocular and temporal

bars, there are whitish marks in front and behind the eye, meeting just above the eye, but not

confluent across the top with those from the other side. Thin whitish chevron marks extend from

the neck to the parietals, but may or may not be confluent with a spot there. Body with

approximately 10, more or less distinct, black-edged white crossbars. A white spot on the tip of

the tail. Ventral colour white with black quadrangular spots, some confluent across the ventrals.

SCALATION: Nasal undivided or partially divided. Two internasals. No loreal. One postocular.

Temporals 1+2. Six supralabials, third and fourth in contact with eye. Six infralabials. Dorsal

scales in 13 rows at midbody. Ventrals 159–170, laterally angulate. Anal undivided. Caudals 30–

44.

DENTITION: Eight maxillary teeth.

HEMIPENIS: Deeply forked. Thin papillae present, extending half the length of the fork. No

spines.

COMMENT: Known from just 2 specimens. Thought by Leviton (1960), on the basis of the

hemipenis, to be part of the “cyclurus-taeniatus” complex. Key Literature: (Campden–Main,

1970 (1984); Leviton, 1953; Leviton, 1960). Photographed in Leviton (1953, Fig. 1). Max.


/14 233 209 Oligodon annulifer

Simotes annulifer Boulenger 1893a: 524 (north Borneo, Malaysia: BMNH 1946.1.3.34) Holarchus annulifer Barbour 1912: 195 [new combination] Oligodon annulifer annulifer Haas 1950: 554 [designates three subspecies] Oligodon annulifer Tillack and Günther 2009 [returns taxon to monotypic state] DISTRIBUTION: Brunei Darussalam?, Malaysia (Sabah, Borneo). COLOURATION: Dominant dorsal colour reddish-brown to dark brown, head more

yellowish. Head markings black, with an ocular bar and temporal bars. Chevron present,

extending from the frontal back to the nape and can enclose the first spot on the back. Body with

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20–25 black enclosed, oval, yellow crossbands., some united in a vertebral line Ventral colour

(in alcohol) white-yellow, sometimes with black edges or spots.

SCALATION: Nasal divided. Two internasals. A small loreal present. Two postoculars.

Temporals 1+1–2. Seven supralabials, third and fourth in contact with eye. Seven infralabials.

Dorsal scales in 15 rows at midbody. Ventrals 151–162. Anal undivided. Caudals 44–50.


HEMIPENIS: Not known.

COMMENT: Known from four specimens and perhaps closely related to O. pulcherrimus

(Tillack and Günther, 2009). Tillack and Günther carefully reviewed the literature for this

species (Das, 1995; Das, 2000; David and Vogel, 1996; Despax, 1912; Leviton, 1963a; Lidth de

Jeude, 1922; Stuebing and Inger, 1999; Werner, 1924) and referred off-Bornean occurrences to

other species. Figured by Boulenger (1894, PL. VIII, Fig. 3). Holotype and additional specimen

photographed by Tillack and Günther (2009, Fig. 8).

/40 25 195 Oligodon arnensis

Coluber Arnensis Shaw 1802: 526 (Vizagapatum and “Arni,” India: Russell, 1796, Fig. 38) Coluber russelius Daudin 1803: 395 (Vizagapatum and Arni, India: Russell, 1796, Figs. 35, 38) Simotes Russelii Duméril, Bibron and Duméril 1854: 628 [refers russelii to Simotes] Coronella Russelii Schlegel 1837: 79 [also from Russell, 1796] Simotes russellii Günther 1858: 24 [synonymizes Coronella Russelii] Coluber monticolus Cantor 1839: 52 (Nepal: Sketch in the Bodleian Library) Oligodon purpurascens var. A Günther 1858: 25 [referred into albiventer by Günther 1864] Simotes albiventer Günther 1864: 213 (near Kandy, Ceylon: BMNH) Simotes arnensis Boulenger 1890a: 314 [arnensis priority over russelii. monticolus, albiventer] Oligodon arnensis Wall 1921c: 231 [new combination] Oligodon arnensis albiventer Deraniyagala 1936: 89 [advocates Ceylonese ssp.] DISTRIBUTION: Sri Lanka, India, eastern Pakistan, Nepal. COLOURATION: Dominant dorsal colour various browns, reddish to purplish. Head

markings black, often light-edged, with an ocular bar, united temporal bars and separate chevron.

Body with black crossbars, sometimes centrally and laterally indented, sometimes edged light.

Ventral colour usually immaculate white but sometimes a few dark spots.

The number of crossbars may correlate roughly with geography: Ceylon 13–18+3–6;

India South of 20ºN 18–30+4–16; North of 20º 7–20+7–20.

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SCALATION: Nasal divided. Two internasals. Loreal present in specimens north of latitude 20º,

absent south of that. Two postoculars. Temporals 1+2 (rarely 2+2). Seven supralabials, third and

fourth in contact with eye, the sixth frequently excluded from the lip. Seven infralabials. Dorsal

scales in 17 rows at midbody. Ventrals 155–203, laterally angulate. Anal divided. Caudals 39–

59.

DENTITION: Eight to 11 maxillary teeth.

HEMIPENIS: Not forked. Spinose throughout, spines. decreasing in size distally.

COMMENT: Its IUCN status is “Lower Risk least concern”. Its karyotype is 2n=46 (Bhatnagar,

1959; Das and Ota, 1988; Trinco and Smith, 1971) and its jaw morphology has been studied

(Kiran, 1981; Kiran, 1982). Key Literature: (Deraniyagala, 1936; Smith, 1943; Wall, 1921c;

Wall, 1923b). Figured by Russell (1796, Pls. 35 and 38), Smith (1943, Fig. 76), Wall (1914a, Pl.

XX) and others. Max. length: 640mm (tail 100).

/231 158 144 Oligodon barroni

Simotes barroni Smith 1916: 46 (Hup Bon, Sriracha, Thailand: BMNH1946.1.3.21/.22/.26) Holarchus barroni Bourret 1936: 242 [new combination] Oligodon barroni Smith 1943: 210 [synonymizes H. taeniatus caudaensis] Holarchus taeniatus caudaensis Bourret 1934c: 173 (Cauda near Nha-trang, Vietnam: MNHNP) DISTRIBUTION: Thailand, southern Vietnam, southern Laos, Cambodia. COLOURATION: Dominant dorsal colour light brown. Head markings light-edged dark-

brown, with an ocular bar, temporal bars and heart shaped chevron. The head markings may or

may not be confluent with a spot on the frontal. Body with 10–14 transverse dark spots confluent

with smaller lateral spots. Between the spots, 3 or more, less distinct, crossbars formed by

darkened scale edges. Some (younger?) specimens may also have faint longitudinal stripes

formed by white edging of scales. Ventral colour coral red with black quadrangular spots

confined to the outside of the ventrals.

SCALATION: Nasal divided. Two internasals. Loreal present. One preocular. Two postoculars.

Temporals 1+1/2/3. Seven or 8 supralabials, third and fourth or fourth and fifth in contact with

eye. Eight or 9 (rarely 7) infralabials. Dorsal scales in 17 rows at midbody. Ventrals 135–160,

laterally angulate. Anal undivided. Caudals 28–48.


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HEMIPENIS: Deeply forked. Large papillae. No spines. Coarse folds at the proximal end.

COMMENT: Recently reviewed (David et al., 2008b), it is a member of the “taeniatus-group”.

Key Literature: (Saint-Girons, 1972b; Smith, 1943; Taylor, 1965). Figured by Smith (1943, Fig.

66, body only), photographed by David et al. (2008, Figs. 25–28). Max. length: 401mm (tail 55).

/250 167 58 Oligodon bitorquatus Oligodon bitorquatus Boie 1827: 519 (Java: RMNH) Calamaria oligodon Schlegel 1837: 40 (Java: RMNH) Oligodon subquadratus Duméril, Bibron and Duméril 1854: 55 (Java: MNHNP) Oligodon bitorquatus Steindachner 1867: 60 [establishes priority of Boie’s name] Rabdosoma amboinense Bleeker 1860b: 42 (“Amboyna”: BMNH) Oligodon bitorquatus Boulenger 1894: 237 [synonymizes amboinense] DISTRIBUTION: Indonesia (southern Sumatra?; Java; Sumbawa). COLOURATION: Dominant dorsal colour purple to blackish. Head markings black, with an

ocular bar, thick, confluent temporal bars and thin collar shaped chevron. Between the temporal

bars and chevron there is a brighter yellow collar. Sometimes the area between the ocular and

temporal bars is brighter. Body with yellow and red dots, usually also with a vertebral series of

larger spots. Ventral colour red with black quadrangular spots.

SCALATION: Nasal divided. Two internasals. Loreal usually present. Two postoculars.

Temporals 1+2 or 2+2. Seven supralabials, third and fourth in contact with eye. Seven

infralabials. Dorsal scales in 17 rows at midbody. Ventrals 130–166. Anal undivided. Caudals

30–46.

DENTITION: Six to 8 maxillary teeth.

HEMIPENIS: Not forked. Two small papillae. Proximal third with a few small spines. Distal

two thirds with transverse folds.

COMMENT: A widespread Indonesian species, this is the type for Oligodon yet one of the least

published upon species of the genus. Rejected for Sumatra by David and Vogel (1996) who

agree with de Haas (1950), Mertens (1930) and Barbour (1912) that Bleeker mixed localities.

Contrary to this view, Edeling (1864a) records it from Sumatra. Key Literature: (Barbour, 1912;

Boulenger, 1894; De Rooij, 1917; Schlegel, 1837). Figured by Russell (1810, Pl. 39), De Rooij

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(1917, Fig. 54), Manthey and Grossmann (1997, Fig. 277, photo) and Hoesel (1959, Figs. 14,

15). Max. length: 370mm (tail 67).

/247 222 19 Oligodon booliati Oligodon booliati Leong and Grismer 2004: 12 (Pulau Tioman, Malaysia: ZRC.2.5153) DISTRIBUTION: Pulau Tioman, Pahang, Peninsular Malaysia. COLOURATION: Dominant dorsal colour deep maroon red. Head markings without

distinctive patterns. Body with 19–22 indistinct, dark brown transverse bars starting from nape.

Ventral salmon pink without dark spots.

SCALATION: Nasal divided. Internasals present. Loreal present, absent or fused. One

postocular. Temporals 1+2 (from Leong and Grismer’s Figure 3). Six to 7 supralabials, second

and third or third and fourth in contact with eye. Seven infralabials. Dorsal scales in 17 rows at

midbody. Ventrals 143–153. Anal single. Caudals 54–60.

DENTITION: Not described.


COMMENT: Known from 3 specimens, 2 females and a subadult male, from Pulau Tioman

only. Colour photographs provided in the original description. Max. length: 510mm (tail

121mm). Although the description does not state O. booliati has a divided nasal scale, Figure 3,

Table 1 and the use of the term “postnasal” in connection with Oligodon suggest it does.

Variability of scalation in O. signatus entirely encompasses that of O. booliati (Tillack and

Günther, 2009). Thus, colour pattern only potentially diagnoses O. booliati. Consequently, the

taxon is likely a colour pattern class, and not a species. Key Literature: (Leong and Grismer,

2004).

/57 222 184 Oligodon brevicauda Oligodon brevicauda Günther 1862: 58 (Anamallays, India: BMNH) Tripeltis brevicauda Cope 1886: 487 [new genus proposed] DISTRIBUTION: India (Western Ghats south of the Goa Gap).

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COLOURATION: Dominant dorsal colour brown. Head markings dark, with an ocular bar,

temporal bars and chevron. A spot on the frontal, may connect chevron and ocular bar. Body

with a light vertebral stripe, bordered by two dark stripes. These dark stripes marked with paired

spots in the anterior, forming short bars posteriorly. Also with lighter lateral stripes and a fine

black line on scale row 3. Ventral colour red with black quadrangular spots many confluent

across the ventrals.

SCALATION: Nasal divided. No internasals. No loreal. Two postoculars. Temporals 1+2,

sometimes 2+2 if there is a detached segment of parietal. Seven supralabials, third and fourth in

contact with eye. Seven infralabials. Dorsal scales in 15 rows at midbody. Ventrals 158–173.

Anal divided. Caudals 25–29.

DENTITION: Seven or 8 maxillary teeth.

HEMIPENIS: Not known; most (all?) of the known specimens are females.

COMMENT: Poorly collected within its restricted range. Its IUCN status is “Lower Risk near

threatened”. Key Literature: (Beddome, 1863; Smith, 1943; Wall, 1923b). Figured by Smith

(1943, Fig. 80, pattern only) and Günther (1864, Pl. XIX, Fig. A). Max. length: 500mm (tail 55).

/57 222 184 Oligodon catenata Calamaria catenata Blyth 1854: 287 (Assam, India: type lost) Oligodon catenatus Smith 1940: 481 [refers Calamaria catenata to Oligodon] Oligodon herberti Boulenger 1905: 235 (Mogok, Myanmar: BMNH) Oligodon herberti var. eberhardti Pellegrin 1910: 30 (TamDao, TongKing, Vietnam: MNHNP) Macrocalamus wongii Fan 1931: 101 (Lohsiang, Kwangsi, China) Oligodon eberhardti Pope 1935: 300 [elevates eberhardti synonymizes M. wongii] Oligodon catenata Smith 1943: 232 [synonymizes herberti, h. eberhardti; latter persists in lit.] Oligodon catenata Zhao et al. 1998: 195 [affirms synonymy of Smith] DISTRIBUTION: Eastern India, northern Myanmar, northern Vietnam, Laos, southern China, and

perhaps Cambodia.

COLOURATION: Dominant dorsal colour purplish-grey to brown. Head markings dark on a

lighter background, with an ocular bar, thick temporal bars and chevron. A spot on the frontal

which may or may not be connected to the chevron and the ocular bar. At the back the chevron is

confluent with the stripes. Body with two dark lateral lines and a lighter vertebral stripe bordered

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by black. If the black border interrupts the light stripe it is catenata/eberhardti if not is herberti.

Ventral colour red with black quadrangular spots on the edges of alternating ventrals.

SCALATION: Nasal undivided. No internasals. No loreal. One postocular (Zhao et al. (1998)

state 2, but this may be a typographical error in the text and a transposition of ocular and

temporal columns in the table). Temporals 1+2. Six supralabials, third and fourth in contact with

eye. Six infralabials. Dorsal scales in 13 rows at midbody. Ventrals 165–212. Anal divided.

Caudals 29–43.


HEMIPENIS: Not forked. No papillae. Distally, with spine edged longitudinal folds and a

proximally spinose area.

COMMENT: As currently constituted, this species does not respect the Song Hong faunal

break. Generally, eberhardti represents the eastern form, herberti the western form, and an

overlap occurs only in northern Myanmar. Saint-Girons (1972b) reports that it does not occur in

Cambodia as originally reported by Werner (1924), without locality. Deuve (1970) used herberti,

and Hu et al. (1980) and Zhao and Adler (1993) used eberhardti without discussion. Toriba

(1994) reiterates Smith’s case for synonymy and Zhao et al. (1998) follow Smith. Key Literature:

(Bourret, 1934a; Bourret, 1935d; Bourret, 1936; Bourret, 1937a; Bourret, 1939a; Gong et al.,

2007; Hu et al., 1980; Smith, 1943; Zhao et al., 1998). Figured by Smith (1943, Fig. 81, pattern

only) and Hu et al. (1980, Pl. 54). Max. length: 565mm (tail 75).

/178 79 125 Oligodon chinensis Simotes chinensis Günther 1888: 169 (mountains north of “Kiu Kiang”, Jiangxi, China: BMNH) Holarchus chinensis Schmidt 1927a: 537 [new combination] Oligodon chinensis Smith 1943: 206 [rejects Holarchus] Simotes longicauda Boulenger 1903: 351 (Man-son Mts. Tong-king, Vietnam: BMNH) Holarchus chinensis Pope 1935: 291 [synonymizes longicauda] Holarchus violaceus longicauda Bourret 1935c: 265 [synonymized by Smith, 1943] DISTRIBUTION: Northern Vietnam, southern China (Kiangsu; Anhwei; Jiangxi; Chekiang; Fujian;

Guangdong; Guangxi; Yunnan; Hainan).

COLOURATION: Dominant dorsal colour light grey-brown to reddish. Head markings dark,

with an ocular bar and chevron, no temporal bars. Body with 10–15+3–4 black crossbands,

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sometimes a faint vertebral line. Ventral colour grayish white sometimes with irregular black

spots. Tail pink.

SCALATION: Nasal divided. Two internasals. Loreal present, sometimes 2. One sometimes 2

preoculars, very rarely a subocular. Two postoculars, rarely 1. Temporals usually 1+2 or 2+2,

rarely 3 anterior. Usually 8 supralabials, fourth and fifth in contact with eye, sometimes 7, rarely

9 or 6. Seven to 9 infralabials, usually 7. Dorsal scales in 17 rows at midbody. Ventrals 170–199,


DENTITION: Nine to 10 maxillary teeth.

HEMIPENIS: Deeply forked. No papillae, but a conspicuous diagonal ridge from the fork to the

tip with a free proximal end that may be homologous with an intermediate condition. No spines.

Calyculate at the tip, proximally with oblique flounces.

COMMENT: Thoroughly reviewed by Pope (1935). Smith (1943) considered it a close relative

of O. cyclurus. Key Literature: (Pope, 1935; Smith, 1943; Wu et al., 1985; Zhao et al., 1998).

Figured by Pope (1935, Pl. XI, Figs. F–I). Max. length: 729mm (tail 104).

/0 115 177 Oligodon cinereus Simotes cinereus Günther 1864: 215 (Cambodia: BMNH) Oligodon cinereus Smith 1940: 481 Oligodon cinereus Smith 1943: 215 [discards violaceus, referring Cantor’s name to cyclurus] Simotes swinhonis Günther 1864: 215 (“Amoy”, Fujian, China: BMNH) Simotes multifasciatus Jan and Sordelli 1865: L. 12, Pl. IV, Fig. 2 Simotes semifasciatus Anderson 1871a: 16 (Naga Hills, Assam, India: ZSI) Holarchus dolleyanus Cope 1895: 423 (Hainan Island, China) [synonymized by Pope 1935: 297] Holarchus lightfooti Rendahl 1937: 24 (Tauyggyi, Myanmar: NHRM3023) Simotes violaceus Boulenger 1890: 312[nonCantor, syn swinhonis, multifasciatus, semifasciatus] Simotes violaceus fasciatus Wall and Evans 1901b: 615 Holarchus violaceus Cochran 1930: 29 Oligodon violaceus Wall 1923b: 318 Holarchus violaceus swinhonis Mell 1931: 212 Simotes violaceus pallidocinctus Bourret 1934d: 18 (Saigon, Vietnam: MNHNP) Holarchus violaceus tamdaoensis Bourret 1935c: 265(TamDao, TonKing, Vietnam: MNHNP) Holarchus violaceus multifasciatus Bourret 1936: 237 Holarchus violaceus poilani Bourret 1939b: 40(DongTamVe, Vietnam)[tentative, Smith 1943] Holarchus violaceus plurimaculatus Bourret 1941: 40 (Chaîne, Quang-tri) Oligodon cinereus swinhonis Taylor 1965: 759 [recognizes subspecies] Oligodon cinereus multifasciatus Taylor 1965: 760 [recognizes subspecies] Oligodon cinereus tamdaoensis Hu et al. 1973: 160 [recognizes subspecies]

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[the name swinhonis persists as a specific in the Chinese literature well past 1980] DISTRIBUTION: India (Assam; Arunachal Pradesh), Bangladesh, Cambodia, Laos, Vietnam,

Myanmar, Thailand, northwest Malaysia?, southern China (westward to Yunnan, eastward to

Hong Kong and Hainan).

COLOURATION: Dominant dorsal colour grayish to reddish-brown or pinkish. Head

uniform brown without markings in most cinereus and swinhonis. In tamdaoensis, a distinct

chevron, sometimes also an ocular bar and temporal bars. In pallidocinctus, only a chevron in the

young. In multifasciatus, a distinct ocular bar, a narrow chevron with detached spot on frontal

and thin temporal bars with detached parietal spots. In all forms the ventral colour is white to

yellowish-white with spotting as follows. The pure cinereus colour form is inornate dorsally,

with little to no ventral spotting. The body of swinhonis has black edged scales forming

reticulations or wavy crossbars, little ventral spotting. The multifasciatus body is as swinhonis

but with distinct ventral spots. The tamdaoensis form has distinct crossbars with interspersed

reticulations and black quadrangular ventral spotting. The pallidocinctus form is most distinctive

with 27–34+3–4 light black-edged crossbars and unspotted ventrals.

SCALATION: Nasal divided (Saint-Girons records Cambodian examples as entire or semi-

divided). Two internasals. Loreal present. Two, sometimes 1 preoculars, the second a subocular.

Two postoculars, rarely 1. Temporals 1+2 or 2+2. Seven or 8 supralabials, third and fourth or

fourth and fifth in contact with eye. Eight, rarely 7 or 9 infralabials. Dorsal scales in 17 rows at

midbody. Ventrals 151–197, laterally angulate. Anal undivided. Caudals 26–57.


HEMIPENIS: Not forked. Two large papillae of unequal length. No spines. Distally, with

longitudinal folds merging into a proximally calyculate area.

COMMENT: An abundant and widespread species on mainland South and Southeast Asia.

Stuebing and Inger (1999) reject it as a Bornean species. Smith (1943) identified 4 intergrading

colour forms, all of which were widespread. Currently, 4 subspecies are recognized plus the

colour form “pallidocinctus”, and these partially conform to different colour forms, given

uncertain extents of range overlap. Taylor’s Thai subspecies multifasciatus occurs in the middle

of the species range. The South Chinese specimens appear to be mostly swinhonis, whose

western border is unclear. Perhaps it intergrades with cinereus in Myanmar and it may respect

98

the Song Hong boundary. The pallidocinctus colour form occurs in southern Vietnam, and in

Cambodia (type loc.) cinereus is squeezed in between this and multifasciatus. Northern Vietnam

is home to tamdaoensis. Taxonomically, this is a very confusing taxon in need of further

examination using samples from multiple populations. Its IUCN status is “Endangered” in India.

Key Literature: (Boulenger, 1893b; Bourret, 1936; Bourret, 1942; Deuve, 1970; Saint Girons,

1972b; Smith, 1943; Taylor, 1965; Zhao et al., 1998). Figured by Smith (1943, Fig. 70, pattern

only) and Taylor (1965, Figs. 34–35, photos). Max. length: 760mm (tail 75).

/255 225 56 Oligodon cruentatus Simotes cruentatus Günther 1868: 417 (Pegu, Myanmar: BMNH) Oligodon cruentatus Wall 1923b: 317 [new combination] Holarchus cruentatus Bourret 1935a: 20 [new combination] Oligodon cruentatus Smith 1943: 221 [rejects Holarchus] DISTRIBUTION: Southern Myanmar. COLOURATION: Like O. theobaldi. Dominant dorsal colour grey-brown. Head markings

dark in the young, often lost in the adults. An ocular bar, temporal bars and chevron. The

temporal bars may or may not be confluent on the frontal. Body with many darkened scales

edges forming reticulations. A pale vertebral line bordered by thicker dark lines and a dark

lateral line. Ventral colour yellowish with black quadrangular spots, tail crimson with a spot at

the base and often one at the tip. Ventral colour (in alcohol) with black quadrangular spots.

SCALATION: Nasal divided. Two internasals. Loreal present, rarely absent. Rarely 2 preoculars,

usually 1. Two postoculars. Temporals 1+2. Eight supralabials, fourth and fifth in contact with

eye. Eight infralabials. Dorsal scales in 17 rows at midbody. Ventrals 148–173, laterally

angulate. Anal divided. Caudals 27–40.

DENTITION: Fourteen to 16 maxillary teeth.

HEMIPENIS: Not forked. Two large papillae. Proximal two thirds spinose, the spines increasing

in size basally.

COMMENT: Very similar to O. theobaldi but significant hemipenial differences seem to occur.

Ventral scale counts have minimal overlap. The easiest character to distinguish them might be

the presence of absence of tail spots. Key Literature: (Boulenger, 1894; Smith, 1943; Wall,

1923b). Never illustrated. Max. length: 365mm (tail 45).

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/236 0 95 Oligodon cyclurus Coronella cyclura Cantor 1839: 50 (probably India: Sketch in the Bodleian Library) Simotes cyclurus Boulenger 1890a: 311 [refers C. cyclura to Simotes] Holarchus cyclurus Smith 1920a: 96 [new combination] Oligodon purpurascens Wall 1923a: 631 [non Schlegel, briefly thought conspecific] Oligodon cyclurus Smith 1943: 202 [cyclurus reinstated] Oligodon cyclurus Campden-Main 1970: 763 [Neotype: Bangkok, Thailand: USNM72067] Coronella violacea Cantor 1839: 50 (Rangpur, Bengal, India: Sketch in the Bodleian Library) Oligodon cyclurus Smith 1943: 202 [synonymizes Cantor’s violacea] Simotes bicatenatus Günther 1864: 217 (no loc.: BMNH) Simotes cyclurus Boulenger 1890a: 311[synonymize bicatenatus, fasciolatus, cochinchinensis] Simotes albocinctus var. dorsolateralis Wall 1910c: 898 (Jalpaiguri Dist., India: no type) Oligodon cyclurus Smith 1943: 202 [refers variety to cyclurus and synonymizes] Oligodon dorsolateralis Welch 1988: 83 [lists as a species, later authors follow] Oligodon kheriensis Acharji and Ray 1936: 519 (North Kheri Div., India: ZSI) Oligodon cyclurus Smith 1943: 202 [synonymizes kheriensis] DISTRIBUTION: India (Assam), Bangladesh, northwestern Myanmar. COLOURATION: Dominant dorsal colour brownish-grey, may be reddish/pinkish-brown in

life. Head markings dark, usually quite distinct, with an ocular bar, temporal bars and elongate

chevron. Temporal bars rarely confluent with chevron. Body with more or less distinct darker

reticulations or thin crossbars, these crossbars usually not as strong or blotch like as fasciolatus

(see comments). The cross-marks can be very thin and irregular, the black only on scale edges,

or somewhat more regular wavy crossbars. Sometimes longitudinal stripes are present, which

may change with maturity in some individuals. Ventral colour yellowish white with or without

black quadrangular spots.

SCALATION: Nasal divided. Two internasals. Loreal present. Two preoculars, 1 a subocular,

rarely absent. Two postoculars. Temporals 1/2+2. Eight, sometimes 7, supralabials, fourth and

fifth, or third and fourth in contact with eye. Nine infralabials. Dorsal scales in 19 rows at

midbody (see comments). Ventrals 167–197, laterally angulate. Anal undivided. Caudals 35–63.

DENTITION: Nine to 10 maxillary teeth. Palatine: 6–8 teeth, no anterior edentulous space.

HEMIPENIS: Deeply forked. No papillae. No spines. Distally, with transverse flounces,

calyculate at the tip.

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COMMENT: See also comments under O. fasciolatus. Oligodon cyclurus is a member of the

“cyclurus-group”, which also includes O. chinensis, O. formosanus, O. fasciolatus, O ocellatus

and O. saintgironsi. The group is defined primarily on hemipenial condition. Among species in

the group, O. chinensis and O. formosanus have long been separated by morphology and range.

The Red River (Song Hong) serves as a faunal break between them and the rest of the group

(Pope, 1935; Yang et al., 1980). Within the group, morphology varies with geography. Taylor

(1965) reports 21 mid-body scale rows in the center of the group’s range and this appears to be

true in Yunnan, China (Zhao et al. 1998). The blotched colour pattern also tends to be more

common in the center of the range. In the southeast (southern Vietnam) and northwest (Myanmar

and adjoining India), 19 scale rows dominate. Collection records show some parapatry among

the species, but it is not clear to what extent this occurs. Notwithstanding, the Northwest Indian

19 scale row group contains O. cyclurus by original description, the southeast 19 scale row group

is O. ocellatus, and the central 21 row group takes the next available name, O. fasciolatus. Thus,

Campden-Main’s neotype from Bangkok, which has 21 scale rows typical of O. fasciolatus,

cannot stand for O. cyclurus. This position is resurgent in the literature (David et al., 2004; David

et al., 2008a; David et al., 2008b; Pauwels et al., 2003; Pauwels et al., 2002; Stuart and Emmett,

2006; Stuart et al., 2006; Teynie et al., 2004) based on the unpublished thesis of Wagner (1975).

The western limit of O. cyclurus is probably the Ganges, but Anderson (1871b) reports a

specimen from Calcutta. Oligodon kheriensis is resurrected as a valid species by Das (1996) but

without justification. Das (1996) also lists O. dorsolateralis as a species, as do Welch (1988) and

Taylor (1965). However, Taylor (1965) uses O. dorsolateralis for a Thai specimen of O.

fasciolatus, and the Das’ Indian specimens are referable to O. cyclurus (David et al., 2004;

Smith, 1943; Wagner, 1975). Stoliczka (1873) lists S. bicatenatus as occurring in Sumatra and

Panang, and it is not clear what species these specimens represent. Its IUCN status is

“Endangered”. Key Literature: (Boulenger, 1894; Smith, 1943; Wall, 1923b). Figured by Smith

(1943, Fig. 63, A and B only).

/102 135 56 Oligodon deuvei Oligodon deuvei David, Vogel and Rooijen 2008b: 28 (“Arboretum de Trang Bom”, Dong Nai, Vietnam: MNHN1974.1366)

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DISTRIBUTION: southern Vietnam (Dong Nai; Ho Chi Minh District), Laos (Vientiane), Cambodia

(Pursat).

COLOURATION: Dominant dorsal colour in life, grayish-brown to reddish-brown, scales

narrowly edged dark. Head markings most like O. taeniatus, with an ocular bar, temporal bars

and narrow chevron. The chevron is usually detached from its point on the frontal. The chevron

arms do not approach the ventrals. Body with a distinct, wide, light (orange to rusty-red in life)

vertebral line flanked by faint brown lines or small brown spots. Sometimes a faint dorsolateral

series of spots. Ventral colour yellowish or grey (in life: cream anteriorally progressing to red

posteriorly), with dark quadrangular spots on the margins. The tail usually lacks the dorsal spots

that characterize O. mouhoti, except in the holotype were very small spots in the same position

are found.

SCALATION: Nasal divided. Two internasals. Loreal present. Two preoculars. Two postoculars.

Temporals 1(rarely 2)+2. Seven, rarely 8 supralabials, third and fourth in contact with eye. Eight

or 9 infralabials. Dorsal scales in 17 rows at midbody. Ventrals 142–163, laterally angulate. Anal

undivided. Caudals 31–47.

DENTITION: Twelve to 15 maxillary teeth.

HEMIPENIS: As O. taeniatus.

COMMENT: A member of the “taeniatus-group” (David et al., 2008b). Figured by David et al.

(2008b, Figs. 19–24, photos). Photographed in life by Grismer et al. (2008) as O. sp., Fig. 19.

Max. length: 530mm (tail 70).

Oligodon dorsalis Elaps dorsalis Gray and Hardwicke 1835: Pl. 85 (Chittagong, India: BMNH) Oligodon dorsalis Günther 1858: 22 [refers E. dorsalis to Oligodon] DISTRIBUTION: India (Assam), Bhutan, Bangladesh, Myanmar. COLOURATION: Dominant dorsal colour brown, darker laterally, some specimens dark

brown. Head markings very indistinct or absent, with hints of an ocular bar, and chevron,

confluent on the frontal. Body with a light vertebral stripe, edged with black or black dots.

Another fine, dark lateral line on scale rows 2 and 3. Ventral colour white with equal proportion

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black quadrangular spots many confluent across the ventrals. Tail crimson to orange with a bar at

the base and another one or two at the tip.

SCALATION: Nasal undivided. Two internasals. Loreal present. One postocular, very rarely 2

on one side. Temporals 1+2. Seven supralabials, third and fourth in contact with eye. Seven

infralabials. Dorsal scales in 15 rows at midbody. Ventrals 160–188. Anal divided. Caudals 27–

51.


HEMIPENIS: One third forked. No papillae. A few large basal spines. Distally, with oblique

flounces.

COMMENT: Its IUCN status is “Vulnerable”. Said to be rare (Cox, 1991). Key Literature:

(Anderson, 1871b; Das and Palden, 2000; Smith, 1943; Wall, 1910b; Wall, 1923b). Figured by

Wall (1908c) and Cox et al. (1998, photo). Max. length: 415mm (tail 80).

/163 145 184 /166 124 82 Oligodon erythrogaster Oligodon erythrogaster Boulenger 1907: 216 (Nagarkot, Nepal: ZSI15850(16108?)) DISTRIBUTION: India (eastern Himalayas), Nepal. COLOURATION: Dominant dorsal colour purplish-grey to brown. Head markings black,

with an ocular bar, and thick temporal bars. Chevron present, confluent with body stripes, may or

may not be detached from spot on frontal. Body with a light vertebral stripe, and on either side

adjoining it: a thick dark stripe and two thin dark stripes separated by light stripes (very similar

to O. hamptoni). Ventral colour coral red, with black quadrangular spots many confluent.

SCALATION: Nasal undivided. Two internasals. No loreal. Two postoculars. Temporals 1+2.

Seven supralabials, third and fourth in contact with eye. The sixth supralabial not known to reach

the lip (Boulenger, 1907, counted the labials 6 due to this). Seven infralabials. Dorsal scales in

17 rows at midbody. Ventrals 163–186. Anal divided. Caudals 42–59.

DENTITION: Seven to 8 maxillary teeth.

HEMIPENIS: Not forked. No papillae. No spines. Proximally with thick smooth longitudinal

folds, distally with flounces becoming finer at the tip.

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COMMENT: Known from very few specimens. The type is in the ZSI collection (Murthy et al.,

1993). Its IUCN status is “Endangered”. Wall (1910e, 1923b) identifies Mr. Hodgson’s S.

octolineatus sketches (Günther, 1861b; Günther, 1864) as this species. Key Literature:

(Boulenger, 1907; Smith, 1943; Wall, 1910e; Wall, 1913b; Wall, 1923b). Figured by Wall

(1910e). Max. length: 450mm (tail 75).

/153 134 117 Oligodon erythrorhachis Oligodon erythrorhachis Wall 1910d: 923 (Namsang, Jaipur district, Assam, India: BMNH) DISTRIBUTION: India (Naga Hills, Arunachal Pradesh). COLOURATION: Dominant dorsal colour dark grey (light grey with black flecks). Head

markings black, with an ocular bar, temporal bars and narrow chevron, the chevron extends to

the prefrontals and is confluent with the ocular bar. Body with a bright red vertebral stripe

covering the vertebral row and 1 and a half scales on each side; also many black, light-edged

crossbars. Ventral colour whitish, with a pink central stripe at the back, with black quadrangular

spots mostly at the edges of the ventral scales.

SCALATION: Nasal undivided. Two internasals. No loreal. Two postoculars. Temporals 1+2.

Seven supralabials, third and fourth in contact with eye. Eight infralabials. Dorsal scales in 15

rows at midbody. Ventrals 154. Anal divided. Caudals 46.

DENTITION: Seven or 8 maxillary teeth (from Smith, 1943, Table p200; notably absent from

Wall 1910d and 1923b).


COMMENT: Known only from the type (Murthy et al., 1993). Its IUCN status is “Data

Deficient”. Key Literature: (Smith, 1943; Wall, 1910d; Wall, 1923b). Figured in Wall (1910d).


/239 71 77 Oligodon everetti

Oligodon everetti Boulenger 1893a: 524 (Mt. Kina Balu, Sabah, Borneo, Malaysia: BMNH) DISTRIBUTION: Malaysia (Borneo; Sabah), Indonesia (Kalimantan).

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COLOURATION: Dominant dorsal colour grey-brown. Head markings dark brown to black,

with an ocular bar, temporal bars confluent with a spot on the frontal and looking much like a

chevron. In the usual chevron position the body stripes meet and point forward to the backs of

the parietals. Body with 3 darker stripes. The vertebral stripe is 3 scales wide and encloses

yellow-white and coral marks shaped like thin bars to small rhomboids. The side stripes have

small, sometimes absent white spots. Ventral colour coral red, no spots.

SCALATION: Nasal divided. Two internasals. Loreal present. Two postoculars. Temporals 1+2.

Seven supralabials, third and fourth in contact with eye. Seven infralabials. Dorsal scales in 15

rows at midbody. Ventrals 132–154. Anal undivided. Caudals 46–72.



COMMENT: Also see comments for O. vertebralis and O. annulifer. Poorly collected, but

thought to exist throughout Borneo (Stuebing and Inger, 1999). The range in scale counts is

based on variation in the type, six specimens in FMNH and one in SMF. At least one specimen

also exists in SSM and probably another in SMK. Key Literature: (Boulenger, 1893a; De Rooij,

1917; Stejneger, 1922; Stuebing, 1991; Stuebing, 1994; Stuebing and Inger, 1999). Figured by

Boulenger (1894, Pl. XI, Fig. 1), photographed by Stuebing and Inger (1999, p105). Max. length:

500mm.

/250 98 95 Oligodon fasciolatus Simotes fasciolatus Günther 1864: 218 (Petchabun, Thailand: BMNH) Simotes cochinchinensis Günther 1864: 219 (Laos Mts.: BMNH) Simotes cyclurus Boulenger 1890a: 311[synonymizes fasciolatus, cochinchinensis into cyclurus] Oligodon purpurascens Wall 1923b: 328 [non Schlegel, briefly thought conspecific] Oligodon purpurascens maculatus Wall 1923b: 329 [part] Holarchus purpurascens Cochran 1930: 27 [new combination] Holarchus purpurascens cyclurus Bourret 1934b: 2 Simotes smithi Werner 1925: 58 (“Don-Rek”, Thailand: NMW) Simotes purpurascens smithi Smith 1928: 497 [puts smithi in the synonymy of cyclurus] Holarchus smithi Bourret 1936: 233 [continues to recognize smithi] Oligodon dorsolateralis Taylor 1965: 763 (Chiang Mai, Thailand: ?77) [non Wall?] Oligodon cyclurus smithi Taylor 1965: 765 [distinguishes fasciolatus material from cyclurus] Oligodon cyclurus superfluens Taylor 1965: 768 (Doi Stuhep, Thailand: EHT77) DISTRIBUTION: southeastern Myanmar, Thailand , Laos, Cambodia, Vietnam, China (Yunnan).

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COLOURATION: Dominant dorsal colour brownish-grey, sometimes reddish or yellowish-

brown. Head markings dark, usually quite distinct, with an ocular bar, temporal bars and

elongate chevron. Temporal bars rarely confluent with chevron. Body with darker reticulations

or crossbars, in much of its range with blotches of widely varying regularity and thickness. Wavy

crossbars sometimes found in a series of 3 between wider transverse blotches. Transverse spots

often rhomboidal with paired spots laterally. The larger spots in a series of 9–18+2–4. The spots

can be indented medially any amount to the point of being a pair of spots. Rarely, longitudinal

stripes are present. Ventral colour yellowish white with or without black quadrangular spots.

SCALATION: Nasal divided. Two internasals. Loreal present. Two preoculars, one a subocular,

rarely absent. Two postoculars. Temporals 1/2+2. Eight, sometimes 7, supralabials, fourth and

fifth, or third and fourth in contact with eye. Nine infralabials. Dorsal scales in 21 rows at

midbody, very rarely 23. Ventrals 147–210, laterally angulate. Anal undivided. Caudals 33–61.


HEMIPENIS: Like O. cyclurus.

COMMENT: Duméril et al. (1854), as well as Tirant (1885a, 1885b), described Simotes

trinotatus from China. Günther (1864) noted the Chinese locality, synonymized Duméril’s snake

into O. purpurascens, and described O. fasciolatus from Thailand. Currently, variation in the

colour of O. fasciolatus (Deuve, 1963a; Deuve, 1963b) encompasses that of S. trinotatus

allowing for the later to be the oldest available name. See also comments under O. cyclurus.

Boulenger (1890a) synonymized O. fasciolatus into O. cyclurus, a decision maintained by Smith

(1943). However, recent work (David et al., 2004; David et al., 2008a; David et al., 2008b;

Pauwels et al., 2003; Pauwels et al., 2002; Stuart and Emmett, 2006; Stuart et al., 2006; Teynie et

al., 2004) has followed Wagner's (1975) unpublished thesis in recognizing O. fasciolatus. The

use of the epithet smithi to describe blotched specimens from the center of the range that have

high (21/23) mid-body scale row occurs often in literature. Taylor (1965) resurrected O.

dorsolateralis. However, whereas Wall’s 1910, Indian O. dorsolateralis were referable to O.

cyclurus, Taylor’s Thai example was referable to O. fasciolatus in having 21 mid-body scale

rows.

Specimens from Pulo Condore (=Con Son) (Saint Girons, 1972a; Saint Girons, 1972b; Smith,

1920a; Tirant, 1885a) have a mixture of scale row formulas and colouring like either

106

smithi/fasciolatus or cyclurus. Their characters appear to be incompatible with O. ocellatus,

although the range of this species lies between this remote landbridge island and the rest of the

accepted range of O. cyclurus/fasciolatus. Their tail ratio is close to 20%. They might prove to

be one or more subspecies of validity equal to O. ocellatus.

Some defensive behaviour in O. fasciolatus has been recorded (Wüster and Cox, 1992). Key

Literature: (Campden–Main, 1970; Saint Girons, 1972b; Smith, 1943; Taylor, 1965; Taylor and

Elbel, 1958; Zhao et al., 1998). Figured by Günther (1864, Pl. XX, Figs. B), Zhao and Adler

(1993, Pl. 36, Figs. C–D) and Taylor (1965, Figs. 36, 37, photos; 1958, Fig. 25, photo).

Photographed often, but presently labeled O. cyclurus. Max. length: 807+mm.

Oligodon forbesi Simotes forbesi Boulenger 1883: 387 (Timor Laut, Indonesia: BMNH) Oligodon forbesi Brongersma 1933: 28 [new combination]

DISTRIBUTION: Indonesia (Tanimbar Islands; Damar; Babar Islands). COLOURATION: Dominant dorsal colour grayish-brown, scales darker edged. Head

markings darker, no ocular bar. A somewhat trident shaped mark in place of the chevron and

temporal bars, extending on to the frontal, but confined to the head scales and not wrapping

around the nape. Body with a pair of fine dark brown lines either side of the vertebral row,

separated by 3 scale rows. Sometimes these fine lines are as broad as 1- and 2- half rows. Ventral

colour yellowish with brown spots at the edges, sometimes forming lines.

SCALATION: Nasal divided. Two internasals. Loreal present. Two postoculars. Temporals 1+2

(See comments). Seven supralabials, third and forth in contact with eye. Eight infralabials.

Dorsal scales in 17 rows at midbody. Ventrals 150–170, slightly laterally angulate. Anal




COMMENT: Known from only a few specimens. Key Literature: (Boulenger, 1883;

Brongersma, 1933; De Rooij, 1917). Figured by Boulenger (1883, Pl. XLII), and it may contain

an error in showing 6 supralabials, while both he and De Rooij count 7 in the types. The

underside view in the plate seems to show labials in position 6 excluded from the edge of the lip.

107

This is probably the accurate figure as Brongersma describes a specimen with this condition.

Therefore, Brongersma’s description should be given as labials 7 and temporals 1+2, not 6 and

2+2. De Rooij’s note of temporals 1+2 or 2+2 should probably only be 1+2 only. Max. length:

305mm (tail 58).

/166 106 117 Oligodon formosanus Simotes swinhonis Swinhoe 1863: 225 [non Günther, irregular citation of Rept. Brit Ind. Günth.] Simotes formosanus Günther 1872b: 20 (“Takao”, Formosa: BMNH) Simotes formosanus Stejneger 1898: 221 [swinhonis Swinhoe nomen nudum] Holarchus formosanus Stejneger 1907: 354 [new combination] Oligodon formosanus Smith 1943: 208 [rejects Holarchus] Simotes bicatenatus Müller 1878: 595 (Fumum, Kwangtung: ZMB27681) [non Günther] Simotes formosanus Schenkel 1901: 161 [synonymizes bicatenatus] Simotes hainanensis Boettger 1894: 133 (Hainan, China: SMF8222a) Simotes formosanus Boettger 1898: 73 [synonymizes hainanensis] Holarchus formosanus hainanensis Barbour 1909: 70 (Tingan and Wuchih Mts., Hainan) Holarchus nesiotis Barbour 1908: 318 (“Tingan”, Hainan, China: MCZ7107)

Simotes nesiotis Mell 1922: 121 [new combination]

Holarchus formosanus Pope 1935: 293 [synonymizes nesiotis] Simotes vaillanti Stanley 1914: 27 [non Sauvage]

Simotes cyclurus Stanley 1914: 27 [non Cantor] Holarchus formosanus Pope 1935: 293 [refutes Stanley’s identifications] Holarchus purpurascens bicatenatus Mell 1931: 211 [Hainan] Holarchus formosanus violaceoides Mell 1930: 323 (Yaoshan, “Kwangsi”, China) Holarchus formosanus brunnea Mell 1930: 323 (Yaoshan, “Kwangsi”, China) Holarchus formosanus Pope 1935: 293 [discusses hainanensis, does not recognize any ssp.] DISTRIBUTION: Northern Vietnam, Taiwan, southern China (Chekiang; Jiangxi; Fujian;

Guangdong; Nan Ao Island; Hainan; Guangxi; Hong Kong).

COLOURATION: Dominant dorsal colour light brown. Head markings dark-edged brown or

dark-edged white, with an ocular bar, temporal bars and chevron. The point of the chevron may

be detached on the frontal, top ends of the temporal bars may be detached on the parietals. Body

with dark scale edges forming many more or less distinct crossbars. Also, often with a red or

yellow vertebral line flanked by brown lines and brown lateral lines on scale row 4. Ventral

colour yellowish or brown with a few black marks and a white ventro-lateral line.

SCALATION: Nasal divided. Two internasals. Loreal present. Two, very rarely 1 preoculars.

Two, very rarely 1 postoculars. Temporals 1+2, rarely 2+2. Eight supralabials, fourth and fifth in

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contact with eye, very rarely 7 (3, 4). Eight, rarely 9 infralabials. Dorsal scales in 19, rarely 17

rows at midbody. Ventrals 154–189, laterally angulate. Anal undivided. Caudals 38–59.


HEMIPENIS: Deeply forked, uneven parts. Short calyculate papillae. No spines. Proximally,

with transverse folds, merging into a distally calyculate region.

COMMENT: This species apparently treats the Song Hong (Red River) in Yunnan and Vietnam

as its range boundary (Yang et al., 1980). It is easy to confuse O. formosanus for O. taeniatus

when the lines are strong and the reticulations weak. Stomach contents of a few Oligodon have

been published, mostly from O. formosanus. In addition to reptile eggs, lizards and insects have

been found (Coleman et al., 1993; Golf, 1980; Hu, 2001; Hwang et al., 1965; Meggitt, 1931;

Minton and Anderson, 1963; Ota and Lin, 1994; Shi and Zheng, 1985; Toriba, 1987). Some

putative defensive behaviours of O. formosanus have been recorded (Mori et al., 1992). Its

karyotype is 2n=36 (Nakamura, 1935; Trinco and Smith, 1971). Key Literature: (Maki, 1931;

Pope, 1935; Smith, 1943; Stejneger, 1907; Zhao et al., 1998). Figured by Pope (1935, Pl. XI, Fig.

D, E) Zhao and Adler (1993, Pl. 36E) and Maki (1931, Figs. 86, 87). Max. length: 942mm (tail

182).

/129 129 129 Oligodon hamptoni Oligodon hamptoni Boulenger 1918: 9 (Mogok, Myanmar: BMNH) DISTRIBUTION: Northern Myanmar. COLOURATION: Head markings dark brown, with a thick ocular bar, temporal bars and

chevron. Chevron detached from a spot on the frontal and confluent with the dark body stripes.

Body with a yellow, vertebral stripe covering the vertebral row and a half for on each side. The

yellow stripe bordered by black-edged dark stripes of equal size. Two more dark stripes on

bluish-grey sides. The most latero-ventral stripe interrupted. Ventral colour red with alternating


SCALATION: Nasal undivided. No internasals. Loreal present or absent. Two postoculars.

Temporals 1+1. Five supralabials, second and third in contact with eye. Six infralabials. Dorsal

scales in 15 rows at midbody. Ventrals 160–175, laterally angulate. Anal divided. Caudals 30–

32.

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HEMIPENIS: Not forked. No papillae. Distal half with spine edged flounces, proximal half with

thick spines.

COMMENT: Perhaps known from as few as two specimens. Uetz (1996) caused confusion by

adding Ablabes hamptoni BOULENGER 1900; Liopeltis hamptoni WALL 1924a: 865;

Opheodrys hamptoni SMITH 1943: 180; Cyclophiops hamptoni TORIBA 1989 to the synonymy

of O. hamptoni in his online database. At one point he commented “Current name and synonymy

unclear (see SMITH 1943).” However, Smith does not mention any lack of clarity in either

Oligodon or Opheodrys. Boulenger (1900, 1918) shows the only point of similarity between

these snakes is the collector and the locality. This species appears to have 2 postoculars even

though Boulenger (1918) gives it as 1. The figure in the description shows 2 postoculars and

Wall (1925b) states that the second specimen has 2. Key Literature: (Boulenger, 1918; Smith,

1943; Wall, 1925a; Wall, 1925b). Figured by Boulenger (1918). Max. length: 590mm (tail 75).

/0 101 145 Oligodon inornatus Simotes inornatus Boulenger 1914: 68 (Nong Kai Ploi, southeast Thailand: BMNH) Holarchus inornatus Bourret 1936: 242 [new combination] Oligodon cinereus Smith 1943: 215 [synonymizes inornatus] Oligodon inornatus Taylor 1965: 762 [resurrects inornatus] DISTRIBUTION: Eastern Thailand, Cambodia. COLOURATION: Dominant dorsal colour pale brown (to salmon red?). Head uniform

brown. Body uniform (or with very faint, narrow grey crossbars?). Ventral colour yellowish-

white. (See also comments)


Eight supralabials, fourth and fifth in contact with eye. Eight infralabials. Dorsal scales in 15

rows at midbody. Ventrals 171(–174?) , laterally angulate. Anal undivided. Caudals (36–?)42.

DENTITION: Not known with certainty, probably as O. cinereus.

HEMIPENIS: Not known with certainty, but given Smith’s treatment, probably as O. cinereus.

COMMENT: Only one specimen known when described and a second individual was referred

in footnote by Smith (1943). The species was synonymized into O. cinereus by Smith creating

110

variation in the midbody scale rows of that species. Taylor (1965) resurrected the species without

adding any specimens and believed the second specimen belonged somewhere else. Saint-Girons

(1972b) accepted this judgment. Another specimen appears in Smith and Kloss (1915), with a

somewhat different colour pattern, and Stuart and Emmett (2006) report a recent collection with

a quite divergent pattern (typical Oligodon head markings; ventrals with some quadrangular

black spots; dorsally with a vertebral paired-series of black-outlined red spots). Key Literature:

(Boulenger, 1914; Saint Girons, 1972b; Smith, 1943; Taylor, 1965). Photographed by Cox et al.

(1998). Max. length: 580mm (tail 90).

/149 92 166 Oligodon jintakunei Oligodon jintakunei Pauwels, Wallach, David and Chanhome 2002: 8 (Krabi, Thailand: QSMI 385) DISTRIBUTION: southern Thailand (Krabi province). COLOURATION: Dominant dorsal colour dark brown. Head markings dark on a light

background, with an ocular bar. Some spots on the frontal and parietals, but holotype lacking

clear temporal bars or chevron. Body with 11+3 narrow whitish rings/crossbands wrapping to the

ventrals. Ventral colour uniform whitish.

SCALATION: Nasal divided. No internasals (fused to prefrontals). Loreal present. One

preocular. One postocular. Temporals 1+1. Seven supralabials, third and fourth in contact with

eye. Seven infralabials. Dorsal scales in 15 rows at midbody. Ventrals 189, laterally angulate.

Anal divided. Caudals 46.

DENTITION: Six maxillary teeth.


COMMENT: Known only from the holotype. Pauwels et al. (2002) indicated the taxon might

bear pleisiotypic non-fossorial characters for Oligodon. They further speculated it might be most

closely allied with O. lacroixi, O. hamptoni and O. brevicauda. Key Literature: (Pauwels et al.,

2002). Figured by Pauwels et al. (2002, Figs. 1-5, photos). Max. length: 448mm (tail 78).

Oligodon joynsoni

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Simotes longicauda joynsoni Smith 1917: 276 (Maa Yome val., MuangNgow, Thailand: BMNH) Holarchus longicauda Smith 1920b: 208 [synonymized] Oligodon joynsoni Smith 1943: 218 [resurrected from O. chinensis] DISTRIBUTION: Northwest Thailand, Laos, Vietnam. COLOURATION: (in alcohol) Dominant dorsal colour purplish-brown. Head markings

typical, dark, with an ocular bar, temporal bars and chevron. Body with about 50 alternating dark

reticulations and crossbands. Ventral colour (in alcohol) whitish-yellow (red in life? According

to Smith only) with or without many black quadrangular spots.

SCALATION: Nasal divided. Two internasals. Loreal present. A subocular present or absent.

Two postoculars. Temporals 1+2 or 2+2. Eight supralabials (second and third fused on one side

of one specimen), fourth and fifth in contact with eye. Eight infralabials. Dorsal scales in 17

rows at midbody. Ventrals 187–195, slightly laterally angulate. Anal undivided. Caudals 40–50.

DENTITION: Eleven to 12 maxillary teeth.

HEMIPENIS: Not forked. Like O. cinereus. Large papillae. No spines. Distally shallow

longitudinal folds merging into a proximally calyculate area.

COMMENT: Smith (1943) knew this species from just four specimens. Taylor (1965) had no

new specimens, although now five individuals are identified as O. joynsoni in USNM. Originally

referred by Smith (1917) to S. longicauda as a subspecies, which was later synonymized with O.

chinensis, Smith (1943) resurrected this species. The colour of O. chinensis and O. joynsoni is

not reliable for distinguishing the two species. Taylor (1965) was suspicious of the species-level

status accorded to O. joynsoni but conceded to Smith’s (1943) judgment, which was based on

distinctive hemipenial morphology. Specimens have now been photographed (Cox et al., 1998),

and the species is now reported from northern Thailand, Laos and Vietnam. These specimens

have not had their hemipenes examined (even if there were males) and, thus, these records

should be treated with skepticism as they could easily be chinensis, although, that would mean a

significant range extension for that species. Further, O. cinereus is difficult to distinguish from

O. joynsoni. Hemipenis morphology caused Smith (1943) to ally O. joynsoni with O. cinereus,

the colour in some forms of cinereus also being very similar to O. joynsoni. Smith (1943)

separated the two species based on a non-overlap in subcaudal count. The subcaudal counts are

now known to overlap as do the ranges. Key Literature: (Smith, 1917; Smith, 1920b; Smith,

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1943; Taylor, 1965). Figured by Smith (1943, Fig. 72, pattern only). Max. length: 760mm (tail

105).

/0 112 164 Oligodon juglandifer Simotes albocinctus var. C Boulenger 1894: 221 [in part?] Simotes albocinctus var. juglandifer Wall 1909a: 349(Tindharia, Darjeeling, India: BNHM210) Simotes juglandifer Wall 1911a: 1163 [elevates variety] Oligodon juglandifer Wall 1923b: 327 [new combination] Oligodon albocinctus var. II Smith 1943: 211 [in part?] Oligodon albocinctus Mahendra 1984: 200 [synonymizes juglandifer without comment] DISTRIBUTION: Northeast India (Darjeeling area). COLOURATION: Dominant dorsal colour brown to black-brown. Many scales edges black

and buff to form variegation. Head markings black, with an ocular bar, temporal bars and

chevron. Chevron extends to frontal, may or may not be confluent with a spot there. Body with

22–28 +6–8 black spots, indented or bisected along the vertebral line. In younger specimens

there are additional small lateral spots adjoining these. In some specimens there smaller spots

alternate between larger ones. In a few specimens there are 4 dark longitudinal stripes. Ventral

colour white, buff or brown with black quadrangular spots on alternating sides of the ventrals.


Seven supralabials, third and fourth in contact with eye, sometimes fifth and sixth fused on one

side, sometimes labial 6 excluded from lip on one side. Eight infralabials. Dorsal scales in 19

rows at midbody. Ventrals 162–208, laterally angulate. Anal undivided. Caudals 53–68.

DENTITION: Ten to 12 maxillary teeth. Palatine: an anterior edentulous space 1–2 teeth in size;

6–8 teeth; a posterior edentulous space 2–3 teeth in size. (For O. albocinctus there should be no

anterior space, 8–10 teeth and a posterior space only 1–2 teeth in size. O. cyclurus palatine as O.

albocinctus, but with 6–8 teeth).

HEMIPENIS: Deeply forked. No papillae. No spines.

COMMENT: Known from very few specimens, but probably due to confusion with other

species. Only two specimens in ZSI (Murthy et al., 1993). Type lost, according to Smith (1943)

but according to Wall (1923b) the type is in BNHM, not BMNH. Wall (Wall, 1914a; Wall,

1923b) believed that all of Boulenger’s (1894) albocinctus form C belonged to O. juglandifer.

113

Smith (1943) placed O. juglandifer in a closer relationship to O. cyclurus based on hemipenis.

Strangely, Smith’s (1943) O. albocinctus section seems to also include Boulenger’s colour C as

colour form II, but with the caveat it may be yet another species. Perhaps this was an oversight,

the O. albocinctus part of Smith’s rather large manuscript may have been prepared before the

section on O. juglandifer. Tentatively, all O. albocinctus var. C sensu Boulenger should be

assigned to O. juglandifer. Mahendra (1984) has, without discussion, referred O. juglandifer to

the synonymy of O. albocinctus. As Mahendra’s Oligodon section is primarily a recapitulation of

Smith (1943) but with much omitted, this change would require some discussion to refute

Smith’s hemipenial characters. Its IUCN status is “Endangered”. Identification of O. juglandifer

is problematic because some colour forms of O. cyclurus, (O. albocinctus?) and O. cinereus look

like it. Oligodon cinereus should have 17 scale rows. Distinguishing O. juglandifer from O.

cyclurus and the less common colour form of O. albocinctus, assuming SmithII/BoulengerC is

not O. juglandifer, is more difficult without resorting to hemipenial characters. Wall (1923b) has

indicated the teeth and edentulous spaces can diagnose the species. Key literature: (Smith, 1943;

Wall, 1909a; Wall, 1911a; Wall, 1923b). Figured by Wall (1911a, Fig. 1). Max. length: 853mm.

/212 49 39 Oligodon kunmingensis Oligodon kunmingensis Kou and Wu 1993: 379 (Kunming, Yunnan, China: YU906452) DISTRIBUTION: China (Yunnan). COLOURATION: Dominant dorsal colour light yellow-brown or darker, some scales with a

dark edge. Head markings absent, top of head brown, sides above the labials grey. Body with

irregular, delicate longitudinal stripes. Ventral colour light yellow with black flecks.

SCALATION: Nasal divided. Two internasals. No loreal . Two postoculars, single on one side of

one specimen. One temporal, almost separated from the postoculars by the parietals. Five

supralabials, third in contact with eye. Six infralabials, first pair not in contact behind mental.

Dorsal scales in 15 rows at midbody. Ventrals 108–121. Anal undivided. Caudals 23–24.

DENTITION: Sixteen to 18 maxillary teeth.

HEMIPENIS: Not forked. No papillae. Spinose throughout.

COMMENT: Known from three specimens only. Believed to be related to O. ornatus. It has an

unusually low ventral scale count and unusually high maxillary tooth count for a species of

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Oligodon. It possesses a somewhat unusual colour and chin shield arrangement. Key Literature:

(Kou and Wu, 1993; Zhao et al., 1998). Figured by Kou and Wu (1993, Figs. 1–4). Max. length:

282mm (tail 36).

/249 133 103 Oligodon lacroixi Oligodon lacroixi Angel and Bourret 1933: 138 (Chapa [=Sa Pa], “Tong-king”, Vietnam: MNHNP) DISTRIBUTION: Northern Vietnam, China (Yunnan). COLOURATION: Dominant dorsal colour dark purplish-brown. Head markings (often

described inversely) brown on a lighter background, with a thick ocular bar, temporal bars and

chevron. Also a spot on the snout. Body with a vertebral series of 11–19+2–4 black-edged

orangish spots. Spots round or transversally oval, covering one scale and the edges of the

adjoining scales. Also four indistinct blackish longitudinal stripes. Ventral colour coral red with


SCALATION: Nasal undivided. No internasals. No loreal. Two postoculars. Temporals 1+2.

Five supralabials, second and third in contact with eye. Six infralabials. Dorsal scales in 15 rows

at midbody. Ventrals 162–178. Anal divided. Caudals 25–33.



COMMENT: Key Literature: (Angel and Bourret, 1933; Bourret, 1935b; He and Yang, 1979;

Smith, 1943; Zhao et al., 1998). Figured by Bourret (1936, p254, head only). Max. length:

700mm (tail 80).

/41 41 103 Oligodon lungshenensis Oligodon lungshenensis Zheng and Huang (in Huang et al.) 1978: 92 (Longsheng, Guangxi) Oligodon guizhouensis Li (in Wu et al.) 1979: 104 (Yongle, Leishan, Guizhou, China) Oligodon lungshenensis Zhao and Adler 1993: 244 [resolves priority] DISTRIBUTION: China (Guizhou).

115

COLOURATION: Much like O. ornatus. Dominant dorsal colour blackish-brown. Head

markings black, with an ocular bar, temporal bars and chevron. The chevron extends only to the

mid-parietals, the temporal bars are confluent on the frontal. Body with 4 darker longitudinal

stripes and 9–12 brown, wave-shaped crossbars with black margins. Ventral colour orange-red

with black quadrangular spots on one or both edges of most scales.

SCALATION: Nasal undivided. No internasals. No loreal. Two postoculars. Temporals 2+2,

rarely 2+1. Six supralabials, third and fourth in contact with eye. Six infralabials. Dorsal scales

in 15 rows at midbody. Ventrals 163–179, laterally angulate. Anal divided. Caudals 31–38.



COMMENT: This snake is very similar in scalation and basic colour pattern to O. lacroixi, a

montaine endemic. Zhao and Adler (1993) and Zhao et al. (1998) explain priority issues for the

name of this snake. Key Literature: (Huang et al., 1978; Li, 1985; Li, 1989; Wu et al., 1985; Wu

et al., 1979; Zhao and Adler, 1993; Zhao et al., 1998). Figured by Wu et al. (1985, Fig. 79). Max.


/238 69 145 Oligodon macrurus Simotes violaceus macrurus Angel 1927: 497 (Pointe Lagan, southern Vietnam: MNHNP) Holarchus violaceus macrurus Bourret 1936: 238 [new combination] Oligodon macrurus Smith 1943: 207 [elevation to full species] DISTRIBUTION: Southern Vietnam. COLOURATION: Dominant dorsal colour light brown. Head markings dark, with a spot

below the eye, and another behind the mouth, like truncated ocular and temporal bars. A chevron

from the neck to the parietals. Body with indistinct darker reticulations. Ventral colour whitish.

SCALATION: Nasal divided. Two internasals. A subocular, present or absent. Loreal present or

absent. Two postoculars. Temporals 1+2. Seven or 8 supralabials, third and fourth or fourth and

fifth in contact with eye. Eight infralabials. Dorsal scales in 17 rows at midbody. Ventrals 143–


DENTITION: Thirteen maxillary teeth.

HEMIPENIS: Deeply forked, no papillae, no spines. Like cyclurus.

116

COMMENT: Known only from two South Vietnamese specimens (Campden-Main, 1970b).

Elevated to full species by Smith (1943) and believed by him, on the basis of hemipenes, to be

related to O. cyclurus. Key Literature: (Angel, 1927; Bourret, 1936; Smith, 1943). Never

illustrated. Max. length: 447mm (tail 147).

/238 69 76 Oligodon maculatus Holarchus maculatus Taylor 1918: 364 (Bunawan, Agusan, Mindanao, Philippines: CM2571) Simotes maculatus Werner 1929: 140 [new combination] Oligodon maculatus Leviton 1963a: 469 [new combination] DISTRIBUTION: Philippines (Mindanao). COLOURATION: Dominant dorsal colour pale lavender. Head markings dark, with a broad

ocular bar, temporal bars and chevron. Chevron, temporal and ocular bars may all be separate or

confluent on the frontal. Body with 20–24 white-edged, dark crossbars, 6–8 scales wide in the

middle narrowing to 1–3 scales laterally. Alternate lighter crossbands 3–6 scales wide. Ventral

colour yellow with black quadrangular spots on the edges of alternating scales.

SCALATION: Nasal entire or perhaps sometimes partially divided. Two internasals. Loreal

present or absent. Two, sometimes 1 or 3 preoculars. Two postoculars. Temporals 1+2, 2+3, 1+3

or 2+2. Seven supralabials, fourth only in contact with eye. Seven infralabials. Dorsal scales in

17 rows at midbody. Ventrals 156–164. Anal undivided. Caudals 52–55.

DENTITION: Nine maxillary teeth.

HEMIPENIS: Not forked. Large spongy papillae extending half the length of the organ. No

spines. Proximal half of the organ calyculate.

COMMENT: Its IUCN status is “Least concern”. Only five specimens of this rare snake are

known in museums (Smith, 1993), although additional cursory records exist (Alcala, 1986). Key

Literature: (Leviton, 1963a; Smith, 1993; Taylor, 1918; Taylor, 1922–25 (1966)). Figured by

Taylor (1918, Pl. 1). Max. length: 299mm (tail 59).

/85 166 119 Oligodon mcdougalli Oligodon mcdougalli Wall 1905a: 251 (“Sandarang”, Myanmar: BNHM963)

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DISTRIBUTION: Myanmar (Sandoway). COLOURATION: Similar to O. catenata. Dominant dorsal colour dusky-black. Head

blacker, with a lighter rostral and collar bars. Body with a red-brown vertebral stripe on the

vertebral row and half row to each side. The stripe is edged with linear black spots. Also with a

lateral black line. Ventral colour black with fawn mottling, a lighter chin and centrally reddish

tail. A black spot at the base of the tail and at the tip.

SCALATION: Nasal undivided. Two internasals. No loreal. One postocular. Temporals 1+2.



DENTITION: Six maxillary teeth.

HEMIPENIS: Not forked. Probably no papillae. Distal end unknown. Proximally with spine

edged calyculate.

COMMENT: Known only from the type. Dowling and Jenner (1989) report that it is most

related to O. catenata and O. dorsalis. Key Literature: (Dowling and Jenner, 1989; Smith, 1943;

Wall, 1905a). Figured by Wall (1905a, p.251). Max. length: 349mm (tail 48).

/0 0 0 Oligodon melaneus Oligodon melaneus Wall 1909a: 349 (Tindharia, Darjeeling, India: BMNH) Oligodon taeniolatus melaneus Mahendra 1984: 203 [synonymizes melaneus without comment] DISTRIBUTION: India (Darjeeling area). COLOURATION: Dominant dorsal colour blackish-brown on top, merging to grey on the

sides, scales speckled. Head markings absent, lips whitish. Ventral colour blue-grey, female also

speckled with black, chin whitish.

SCALATION: Nasal partially divided. Two internasals. Very small loreal. Two postoculars.

Temporals 1+2. Seven supralabials, third and fourth in contact with eye (fourth and fifth on one

side of one specimen). Seven infralabials. Dorsal scales in 15 rows at midbody. Ventrals 152–

160. Anal divided (in the single male, only partially so). Caudals 39–40.


HEMIPENIS: Not forked. No papillae. Spinose throughout, the spines. of a uniform size.

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COMMENT: Its IUCN status is “Data Deficient”. Known from just two specimens in ZSI

(Murthy et al., 1993). Mahendra (1984) has, without any discussion, referred O. melaneus to a

subspecies of O. taeniolatus. This change would require some discussion to refute Smith’s

hemipenial characters. Key Literature: (Smith, 1943; Wall, 1909a; Wall, 1923b). Figured by

Wall (1909a, p350). Max. length: 330mm (tail 55).

/0 0 57 Oligodon melanozonatus Oligodon erythrorhachis Annandale 1912: 48 [non Wall] Oligodon melanozonatus Wall 1922: 29 (Upper Rotung Valley, Abor Hills, India: ZSI16799) Oligodon melanozonotus [sic] Smith 1943: 199 Oligodon melazonotus Smith 1943: 213 Oligodon melanozonatus Zhao and Adler 1993: 245 DISTRIBUTION: India (Arunachal Pradesh), China (Xizang). COLOURATION: Dominant dorsal colour brown-red to crimson in life. Head markings dark,

with an ocular bar, temporal bars and chevron. The temporal bars are confluent in a chevron

shape on the frontal, the chevron is confined to the neck. Body with 20+4 irregular black edged

crossbars, centrally white like O. albocinctus in the younger specimen, faded to grey in the

larger. Perhaps a vertebral line in life for juveniles. Ventral colour white with black quadrangular

spots, many confluent across the ventrals.

SCALATION: Nasal undivided (in diagram, questionably divided in Wall, 1923b). Two

internasals. No loreal. Two postoculars. Temporals 1+2. Six (7 on one side of one specimen)

supralabials, third and fourth in contact with eye. Six infralabials. Dorsal scales in 17 rows at

midbody. Ventrals 171–173. Anal divided. Caudals 42–45.



COMMENT: Its IUCN status is “Data Deficient”. Known from only two female specimens in

ZSI (Murthy et al., 1993). Wall (1923) decided that in all cases of a 6 supralabial count, the fifth

labial looks creased or like it is two fused together. Key Literature: (Annandale, 1912; Smith,

1943; Wall, 1922; Wall, 1923b). Figured by Wall (1922, Fig. 1, head only). Max. length: 520mm

(tail 85).

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/114 0 57 Oligodon meyerinkii Simotes meyerinkii Steindachner 1891: 294 (Sulu Islands, Philippines: NMW25828) Simotes octolineatus var. C Boulenger 1892: 74 [synonymizes O. meyerinkii] Holarchus meyerinkii Barbour 1912: 118 [suggests reinstatement of species] Holarchus meyerlinkii [sic] Taylor 1918: 360 [reinstatement] Oligodon meyerinkii Leviton 1963a: 471 [new combination] DISTRIBUTION: Philippines (Bongao; Jolo; Papahang; Sibutu; Tawi-Tawi), Malaysia (Borneo;

Sabah)

COLOURATION: Striped, width of stripes varying. Head markings darker, but can be

indistinct or incomplete, with an ocular bar and temporal bars. No chevron. The temporal bars

often confluent. A stripe on the frontal, may contact ocular bar if present, extends back and

forms the beginning of the dark median body stripes. Body with a salmon vertebral stripe,

bordered by wide darker stripes, and a varying number of alternating light and dark lateral

stripes, like O. octolineatus. Ventral colour rosy-pink, finely peppered with dark brown, tail with

occasional brown spots.

SCALATION: Nasal divided. Two internasals. Loreal present. Two postoculars. Temporals 2+2,

first anterior often not touching oculars. Six supralabials, third and fourth in contact with eye.

Seven infralabials. Dorsal scales in 17 rows at midbody. Ventrals 154–169, laterally angulate.

Anal undivided. Caudals 38–57.

DENTITION: Nine maxillary teeth.

HEMIPENIS: Not forked. No papillae. No spines. Distal third with two longitudinal folds,

proximally calyculate.

COMMENT: Its IUCN status is “Endangered”. Boulenger (1892) believed that this species was

a variety of O. octolineatus. Both Taylor (1922) and Leviton (1963a) recognized its validity. The

ranges of O. octolineatus and O. meyerinkii overlap only in Sabah, Borneo. Hemipenes are the

most reliable way of diagnosing the species in Sabah, although some colour differences also

exist. Key Literature: (Gaulke, 1993; Gaulke, 1994; Leviton, 1963a; Steindachner, 1891).

Photographed by Gaulke (1993, Figs. 1–3), figured by Taylor (1922, Pl. 17, Figs. 6–7). Max

length: 379mm(tail 71).

/228 202 31

120

Oligodon modestus Oligodon modestus Günther 1864: 210 (Philippines: BMNH) Oligodon modestum Leviton 1963a: 473 DISTRIBUTION: Philippines (Luzon; Mindanao; Negros; Tablas; Panay). COLOURATION: Dominant dorsal colour dark brown. Head markings dark on yellow-

brown background, with an ocular bar, temporal bars and chevron. Temporal bars may be

confluent on the frontal, chevron may extend to frontal. Body with a salmon to red vertebral

stripe, most distinct on the tail. Dark top blends to lighter brown peppered yellow laterally.

Indistinct black reticulations. Ventral colour (in alcohol) yellowish with alternating black

quadrangular spots on the edges of the ventrals.

SCALATION: Nasal usually divided. Two internasals. Usually no loreal. One postocular.

Temporals 1+1/2/3. Six supralabials, third only in contact with eye. Six or 7 infralabials. Dorsal

scales in 15 rows at midbody. Ventrals 158–176. Anal undivided. Caudals 27–44.


HEMIPENIS: Not forked. No papillae. No spines. Distal third with narrow longitudinal folds,

proximal two thirds with transverse folds.

COMMENT: Its IUCN status is “Vulnerable”. Well reviewed by Leviton (1963a), who thought

it was closely related to O. purpurascens and O. ancorus. It also bears a resemblance to O.

waandersi. It has been collected very rarely outside of Negros and Panay islands. Specimens can

be found in PNM, SMF, CAS-SU, MCZ, MVZ, USNM and FMNH (Alcala, 1986; Brown and

Alcala, 1970; Ferner et al., 2001; Gaulke, 2001; Sison et al., 1995; Taylor, 1917). Key Literature:

(Leviton, 1963a). Figured by Boulenger (1894, Pl. 10, Fig. 3). Max length: 350mm(tail 55).

/182 146 81 Oligodon moricei Oligodon moricei David, Vogel and Rooijen 2008b: 40 (“Nha-Trang”, Khanh Hoa, Vietnam: MNHN1919.0137) DISTRIBUTION: Vietnam (Khanh Hoa). COLOURATION: Dominant dorsal colour dark grey-brown, many dark edged scales forming

irregular reticulations. Head markings dark, with an ocular bar, temporal bars, a spot on the

121

frontal detached from the heart shaped narrow chevron. A broad, lighter brown vertebral line,

bordered by distinct dark paravertebral stripes. Ventral colour yellowish with quadrangular black

spots. Ventral checks are united in the middle on the posterior half.

SCALATION: Nasal divided. Two internasals. Loreal present. One preocular. Two postoculars.

Temporals1+2. Eight supralabials, fourth and fifth in contact with eye. Nine infralabials. Dorsal

scales in 17 rows at midbody. Ventrals 175, laterally angulate. Anal undivided. Caudals 41.

DENTITION: Twelve maxillary teeth.

HEMIPENIS: Unknown.

COMMENT: Tentatively a member of the “taeniatus-group” (David et al., 2008b). Known from

the holotype only. Figured by David el al. (2008b, Figs. 29–34, photos). Max. length: 443mm

(tail 59).

Oligodon mouhoti Simotes taeniatus Günther 1864: 216 [re-description of taeniatus but using new specimen] Simotes quadrilineatus Müller 1887: 263 [part, non Jan] Holarchus taeniatus Barbour 1909: 70 Simotes taeniatus var. mouhoti Boulenger 1914: 70 (Cambodia: type uncertain? [see below]) Simotes toeniatus Bourret 1927: 234 Holarchus taeniatus mouhoti Cochran 1930: 29 Holarchus taeniatus taeniatus Bourret 1934c: 173 Oligodon taeniatus Smith 1943: 208 [Smith and others until 1969 held 17sc. rows as taeniatus] Oligodon mouhoti Campden-Main 1969: 296 Oligodon mouhoti David, Vogel and Rooijen 2008: 23 [offer holotype BMNH RR1946.1.3.32] DISTRIBUTION: Cambodia, central and eastern Thailand, southern Vietnam?. COLOURATION: Dominant dorsal colour grey to brown. Head markings dark with an very

thick ocular bar, frontal spot, temporal/parietal dots or bars. The heart shaped chevron is usually

detached from the spot on the frontal, often confluent with part of the temporal bar giving the

effect of a “W” chevron. The W shape either confluent or disconnected from the dots on the

parietals. The chevron/temporal band contacting the ventrals. Body with a light vertebral line

flanked by brown lines (sometimes faded). A fine black lateral line between scale rows 3 and 4.

Ventral colour reddish with black quadrangular spots usually confined to the edges of the

ventrals. Two dorsal black spots on the tail, base and tip.

122


Eight supralabials, fourth and fifth in contact with eye. Nine, rarely 10 infralabials. Dorsal scales

in 17 rows at midbody. Ventrals 143–169, laterally angulate. Anal undivided. Caudals 29–47.


HEMIPENIS: Deeply forked. Large papillae. No spines. Calyculate proximal to the fork.

COMMENT: Recently reviewed (David et al., 2008b), the species is a member of the

“taeniatus-group.” See also comments under O. taeniatus. Like O. taeniatus, specimens

BMNH1946.1.3.30–32 were implied but not designated as neotypes by Campden-Main. David et

al. (2008b) identified one specimen as being figured by Günther. Key Literature: (Campden–

Main, 1969; Saint Girons, 1972b; Smith, 1943; Taylor, 1965). Figured by Günther (1864, Pl. 20,

Fig. A), Cox et al. (1998, p60, photo) and David et al. (2008, Figs. 15–18, photos). Maximum

length: somewhat smaller then O. taeniatus.

/102 102 102 Oligodon multizonatum Oligodon multizonatum Zhao and Jiang 1981: 54 (Luding, Sichuan, China: CIB800289) Oligodon multizonatus Zhao and Adler 1993: 245 DISTRIBUTION: China (west Sichuan). COLOURATION: Dominant dorsal colour dull orange. Head markings black in juveniles,

fading in adults, consist of 3 irregular, more or less confluent patches around the eye, frontal and

parietals. On the neck a somewhat chevron shaped dark transverse blotch. Body with 54–47

black transverse stripes 1–3 scales wide, almost crossbar anteriorally, posteriorly increasingly

broken. On the tail 14–19 black crossbars. Ventral colour whitish with black quadrangular spots

at the edges, alternating 2 ventrals spotted, 1–3 not spotted.

SCALATION: Nasal divided. Two internasals. Loreal very long and touching the eye. One

preocular placed high may be present or absent. Two postoculars. Temporals 2+3, sometimes

only 1 anterior or 2 posterior temporals on one side. Eight supralabials, third, fourth and fifth in

contact with eye, sometimes fourth and fifth fused on one side. Eight infralabials, sometimes 7

on one side. Dorsal scales in 17 rows at midbody. Ventrals 190–195, laterally angulate. Anal

divided. Caudals 68–75.


123

HEMIPENIS: Not forked. No papillae. Proximally with a few small spines, distally, with long

furry spines.

COMMENT: Only four specimens recorded. Key Literature: (Jiang et al., 1983; Zhao et al.,

1998; Zhao and Jiang, 1981). Figured by Zhao and Jiang (1981, Fig. 1, head, 2 hemipenis; Pl. 1,

body).

/102 102 250 Oligodon nikhili Oligodon nikhili Whitaker & Dattatri 1982: 630 (Tiger shola, Tamilnadu, India: MSPT) DISTRIBUTION: Southern India (Palini Hills). COLOURATION: Dominant dorsal colour light brown, each scale with a black fleck. Head

markings dark brown, with an ocular bar, temporal bars and chevron. Chevron confined to the

top of the neck, confluent with the temporal bars on the frontal, confluent with the stripes at the

back. Body with two 1.5 scale wide dark stripes either side of the vertebral line, and two less

distinct dark stripes on scale row 3. Ventral colour whitish with black quadrangular spots at the

edges of the ventrals.

SCALATION: Nasal divided. Two internasals. No loreal. Two postoculars. Temporals 1+2.

Seven supralabials, third and fourth in contact with eye. Eight infralabials. Dorsal scales in 15


DENTITION: Not known.

HEMIPENIS: Not forked. No papillae. Distal half, with three longitudinal folds and flounces

edged with small spines. A middle quarter with equal size spines, and a smooth proximal quarter.

COMMENT: Known only from the holotype (Murthy et al., 1993). In their original discussion,

the authors did not regard it as being closely related to O. taeniolatus. The hemipenes are quite

distinct and the colour is very similar to one pattern in O. taeniolatus. Its IUCN status is

“Critically Endangered”. Key literature: (Whitaker and Dattatri, 1982), including a photograph.


/255 195 53 Oligodon ningshaanensis

124

Oligodon ningshaanensis Yuan 1983: 65 (Huoditang, Shaanxi, China: NWSUAF810218) DISTRIBUTION: China (south Shaanxi). COLOURATION: Somewhat like O. catenata. Dominant dorsal colour brownish-green.

Head markings very indistinct. Neck with three black-brown longitudinal stripes, vertebral short,

lateral two extending to tail. Two additional thin, dark ventro-lateral stripes. Ventral colour

whitish.

SCALATION: Nasal undivided. Two internasals. No loreal. Two postoculars. Temporals 1+2,

the lower posterior in a line with the supralabials. Six supralabials, third and fourth in contact

with eye. Six infralabials. Dorsal scales in 13 rows at midbody. Ventrals 163–170. Anal divided.

Caudals 64–73.

DENTITION: Five to 6 maxillary teeth.

HEMIPENIS: Forked only to a depth of 1/4. Some spines. on the apical end. No papillae.

COMMENT: Only three specimens known. Thought to be related to O. catenata. Key

Literature: (Yuan, 1983; Zhao et al., 1998). Figured by Yuan (1983, Fig. 1). Max. length:

655mm (tail 42).

/98 138 53 Oligodon ocellatus Simotes brevicauda Steindachner1867:61(Cochinchina: NHMW16530)[nec S. brevicauda Günth] Simotes ocellatus Morice 1875: 57 (Tay-ninh, Vietnam: MNHL1269–1272 (1569–1572?)) Holarchus purpurascens cyclurus Bourret 1934d: 19 [in part] Oligodon cyclurus Smith 1943: 202 [in part] Oligodon analepticos Campden-Main 1970: 763 [nomen novum]

Oligodon ocellatus Saint-Girons 1972a: 5 [corrects analepticos]

DISTRIBUTION: Cambodia, Vietnam. COLOURATION: Like O. cyclurus smithi. Dominant dorsal colour brown. Head markings

dark brown, with an ocular bar, temporal bars and chevron. Body with about 12+2 dark brown

blotches or crossbands, sometimes indented centrally and laterally like O. venustus. In between

each crossband, a series of three reticulations. Ventral colour (in alcohol, posterior ventrals may

be pink in life) immaculate white, no quadrangular spots, in several specimens.

SCALATION: Nasal divided. Two internasals. Loreal present. Two preoculars, one might be

considered a presubocular. Two postoculars. Temporals 2+2 or 1+2. Eight supralabials, fourth

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and fifth in contact with eye. Nine infralabials. Dorsal scales in 19 rows at midbody. Ventrals

154–180, laterally angulate. Anal undivided. Caudals 26–44.

DENTITION: Ten maxillary teeth.

HEMIPENIS: Like O. cyclurus.

COMMENT: See additional comments under O. cyclurus and O. fasciolatus. Probably many

specimens, but most will be identified as O. cyclurus. The Song Ba River drainage may form a

natural range boundary in northern Vietnam. Two reviews (Campden–Main, 1970; Saint Girons,

1972a) resurrected this species out of Smith’s (1943) synonymy but neither examined the

hemipenis. Saint-Girons (1972) rejected Campden-Main’s (1970) suggestion of allopatry. A few

records show specimens with 21 scale rows in the range of O. ocellatus. One important field

guide (Campden–Main, 1970 (1984)) appears to give descriptions of O. cyclurus extracted from

other sources, while actually discussing the range of O. ocellatus. Bourret (1934d) reports on

three specimens that conform to O. ocellatus. Key Literature: (Campden–Main, 1970; Morice,

1875; Saint Girons, 1972a; Saint Girons, 1972b; Steindachner, 1867). Figured by Steindachner

(1867, Pl. III, Fig. 13) and Smith in part (1943, “form III cyclurus”, Fig. 63d). Max. length:

663mm (largest certain ocellatus). The tail of Ô. ocellatus is supposed to be shorter then that of

O. cyclurus (Saint-Girons, 1972a,b), only 10–12% of total length in males, 9–10% females

versus closer to ~20% males and ~14% females in O. cyclurus.

/179 76 33 Oligodon octolineatus Elaps octo-lineatus Schneider 1801: 299 (no loc.) Coluber octolineatus Shaw 1802: 540 (no loc.: “described from specimens” in BMNH) Coronella octolineata Boie 1827: 539 [refers E. octolineatus to Coronella] Simotes octolineatus Duméril, Bibron and Duméril 1854: 634 [refer E. octolineatus to Simotes] Simotes octolineatus Günther 1858: 24 [synonymizes Coluber, Coronella octolineatus] Simotes octolineatus var. A and B Boulenger 1885: 389[colour vars., rejected by Barbour, 1912] Holarchus octolineatus Barbour 1912: 118 [new combination] Oligodon octolineatus Smith 1930: 55 [new combination] Oligodon octolineatus Mertens 1969: 340 [type locality “probably Java”] Oligodon octolineatus Tillack and Günther 2009 (Java: ZMB 2099) [neotype] DISTRIBUTION: Brunei Darussalam, Malaysia (Malaya and East Malaysia), Singapore Islands,

Indonesia (Nias; Sumatra; Riau Archipelago; Bangka; Billiton; Borneo; Java; Sulawesi), and

Philippines?

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COLOURATION: Alternating pale-brownish-orangish and dark brown stripes, thickness

varies, if black dominant it is Boulenger var. B, if yellowish then var. A. Head markings dark,

with an ocular bar and temporal bars. Chevron reduced to a longitudinal bar on the frontal

splitting posteriorly into the first dark stripes. Body with a reddish vertebral stripe and alternating

dark and light stripes. Ventral colour pinkish, with or without black quadrangular spots on the

edges of the ventrals, spots can be confluent into ventral lines.

SCALATION: Nasal divided. Two internasals. Loreal present. Two postoculars. Temporals

usually 2+2, lower anterior usually not in contact with oculars. Six, rarely 5 or 7 supralabials,

third and fourth in contact with eye. Seven or 8 infralabials. Dorsal scales in 17 rows at midbody.

Ventrals 155–197, slightly laterally angulate. Anal undivided. Caudals 42–63.


HEMIPENIS: Not forked. Two large papillae. No spines. Distal third with two longitudinal

folds, proximally calyculate.

COMMENT: Tillack and Günther (2009), working at ZMB with the type material, have

recently reviewed the species and designated a neotype. Part of a group of presumably related

species, including O. meyerinkii, O. woodmasoni, O. unicolor and O. forbesi. Geographically, O.

octolineatus inhabits the larger Indonesian islands, while others species are isolated on smaller

peripheral islands. This has lead authors to use the terms “derived from an octolineatus-like

ancestor” when referring to the other species. The group’s occurrence in the Philippines is

probably represented by O. meyerinkii. Key Literature: (David and Vogel, 1996; De Rooij, 1917;

Mertens, 1969; Stuebing and Inger, 1999; Tillack and Günther, 2009). Figured by Russell (1810,

Pl. 38), David and Vogel (1996, Fig. 4, head), Mertens (1969, Figs. 1, 2, stripes) Stuebing and

Inger (1999, p106, photo) and Tillack and Günther (2009, Fig 1, photos). Max length: 580mm

(tail 100). /152 193 33 Oligodon ornatus Oligodon ornatus Van Denburgh 1909: 53 (“Shinchiku”, Formosa: CAS18005) Holarchus torquatus konishii Oshima 1910: 196 (“Urai”, Shinko, Formosa: NMNS84) Oligodon ornatus Maki 1931: 134 [synonymizes torquatus konishii] Simotes musyi Roux 1919: 61 (“Fokien”, China: NHMB) Holarchus musyi Pope 1929: 452 [new combination]

127

Holarchus ornatus musyi Mell 1931: 211 [synonymizes musyi] Holarchus ornatus musyi Pope 1935: 302 [ssp. only separate by ventral count] Oligodon ornatus Zhao et al. 1986: 157 [ventral count now overlaps] Oligodon ornatus Zhao and Adler 1993: 245 [do not recognize ssp.] DISTRIBUTION: Taiwan, China (Sichuan; Hunan east to Fujian and north to Anhui). COLOURATION: Dominant dorsal colour brownish-grey. Head markings dark, with an

ocular bar, confluent temporal bars and narrow chevron. Chevron and temporal bars sometimes

confluent at the posterior outside corner of the parietals. Body with 7–9+2–3 dark, indented

crossbars. Also with either faint longitudinal lines or small paired spots between the crossbands.

Ventral colour yellowish with black quadrangular spots on the edges of the ventrals. Sometimes

a median red line on the ventrals.

SCALATION: Nasal partially divided or undivided. Two internasals. No loreal. One, rarely 2

preoculars. Two, rarely 1 postoculars. Temporals 1+2 (many authors 2+2, counting the sixth

labial). Seven supralabials, third and fourth in contact with eye, sixth excluded from the lip.

Seven infralabials. Dorsal scales in 15 rows at midbody. Ventrals 156–182. Anal divided.

Caudals 27–44.


HEMIPENIS: Not forked. No papillae. Spinose throughout, spines uniformly large in the

proximal two thirds, decreasing in size in the distal third.

COMMENT: Few specimens are known. Key Literature: (Gong and He, 2008; Maki, 1931;

Pope, 1935; Zhao et al., 1998). Figured by Roux (1919, Figs. 1–2) and Hu et al. (1980, Pl. 56).

Max length: 775mm (tail 115).

/152 84 130 Oligodon perkinsi Holarchus perkinsi Taylor 1925: 108 (Culion Is., Philippines: MCZ25725) Oligodon perkinsi Leviton 1963a: 476 [new combination] DISTRIBUTION: Philippines (Culion Islands). COLOURATION (in alcohol): Dominant dorsal colour grayish-lavender. Head uniform slate,

none of the usual markings. Body with numerous dark, transverse, irregularly shaped and broken

crossbars. Also a vertebral, broken line of dull red, from the parietals to the end of the tail. The

128

red line is bordered by irregular black lines which also form a border for some of the adjoining

spots. Ventral colour whitish with black quadrangular spots on alternating ventrals, either on the

left or right side.

SCALATION: Nasal divided or partially divided. Two internasals. Loreal present. One or 2

postoculars. Temporals 1+2+2 or 1+3. Seven supralabials, third and fourth in contact with eye.

Seven or 8 infralabials, unusual in not having the first pair in contact behind the mental. Dorsal

scales in 17 rows at midbody. Ventrals 183–188. Anal single or divided. Caudals 34–45.



COMMENT: Its IUCN status is “Near threatened”. Known from two female specimens, the

species is diagnosed by several points of scalation. Tentatively thought by Taylor (1922–5) to be

related to O. signatus. Omitted from a Philippine checklist (Brown and Alcala, 1970). Key

Literature: (Leviton, 1963a; Taylor, 1922–25 (1966)). Never illustrated. Max. length: 672mm

(tail 90).

/142 102 98 Oligodon petronellae Oligodon ornatus Roux 1914: 28 [nec Oligodon ornatus VanDenburgh] (Sumatra: MHNG767.2) Oligodon petronellae Roux in Rooij 1917: 131 [nomen novum at request of Roux] Oligodon rouxi Ruthven 1921: 20 [nomen novum] Oligodon petronellae Werner 1929: 136 [corrects rouxi] Oligodon annulifer var. confluens Werner 1924: 37 (Padang, Sumatra: NHMW16408) DISTRIBUTION: Indonesia (Sumatra). COLOURATION: Quite similar to O. pulcherrimus and O. annulifer. Dominant dorsal colour

grey-brown. Head markings dark brown, with an ocular bar, temporal bars and chevron. Chevron

may be disconnected from a spot on the frontal or connected to the ocular bar, and at the rear

confluent with the markings enclosing the rings on the body. Body with 20–25+3–5 dark

bordered lighter rings, the border continuous between the anterior spots, spots more rhomboid

posteriorly. Some white spots bordering the rings. Ventral colour brick-red with dark brown

quadrangular spots on the edges of every other ventral. Tail red in life.

SCALATION: Nasal partially divided, rarely entire. Two internasals. Loreal present. Two

postoculars. Temporals 1+2 or 2+2. Seven supralabials, third and fourth in contact with eye.

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Eight, rarely 7 infralabials. Dorsal scales in 15 rows at midbody. Ventrals 144–163. Anal




COMMENT: Formerly known only from the holotype, Tillack and Günther (2009) recently

reviewed the species, redescribed the holotype, and referred additional specimens to the species.

Figured by Tillack and Günther (2009, Fig. 2, photos). Max. length: 480mm. /

69 196 220 Oligodon planiceps Simotes planiceps Boulenger 1888: 597 (Minhlà, Myanmar: MSNG) Oligodon planiceps Wall 1923b: 307 [new combination] Holarchus planiceps Pope 1935: 289 [new combination] Oligodon planiceps Smith 1943: 221 [rejection of Holarchus] DISTRIBUTION: Southern Myanmar. COLOURATION: Dominant dorsal colour pale brown. Head markings darker in the young,

often faded in mature specimens. An ocular bar or subocular spot present. Temporal bars present

or faded, a transverse bar behind the parietals in place of the chevron. Body with indistinct black

reticulations on the scale edges. Ventral colour yellow (Wall, Smith; pinkish in Boulenger) with

pairs of black quadrangular spots on the outer edges of some ventrals.

SCALATION: Nasal undivided. Two internasals, the rostral completely separating the

internasals. Loreal present. Two postoculars. Temporals 1+1. Five supralabials, third in contact

with eye (in one specimen only 4 supralabials, the fourth being the size of a fourth and fifth. Six

infralabials. Dorsal scales in 13 rows at midbody. Ventrals 132–145, laterally angulate. Anal


DENTITION: Ten maxillary teeth.

HEMIPENIS: Not forked. Both spinose and papillate.

COMMENT: Very rare in the literature, Smith (1943) knew of only four specimens. At least

two more now occur in the CAS. The hemipenis is known from AMNH27454 (Pope, 1935). Key

Literature: (Boulenger, 1888; Smith, 1943; Wall, 1923a). Figured by Boulenger (1888, Pl. V,

Fig. 2). Max. length: 230mm (tail 22).

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/236 196 220 Oligodon praefrontalis Oligodon praefrontalis Werner 1913: 25 (Sabang, “Pulo Weh”, Indonesia: ZMH lost?) DISTRIBUTION: Indonesia (Pulau Weh). COLOURATION: In general like O. pulcherrimus. Dominant dorsal colour grey-brown.

Body with a yellow-brown stripe covering the vertebral row and half the next row, much as if O.

pulcherrimus’ spots were connected. Ventral colour yellowish with alternating black and white

ventral edges.

SCALATION: Nasal undivided. No internasals. No loreal. One postocular. Temporals 1+2.





COMMENT: Known only from the type, which was not located by Tillack and Günther (2009).

Thought by Werner to be very closely related to its Sumatran neighbor O. pulcherrimus. Key

Literature: (De Rooij, 1917; Werner, 1913). Never illustrated. Max. length: 241mm (tail 31).

/237 27 50 Oligodon propinquus Oligodon propinquus Jan 1862: 38 (“Java”: ZMHr04432) DISTRIBUTION: Indonesia (Java?). COLOURATION: Dominant dorsal colour black (?), in illustration sides and rear half dark,

but not black). Head uniformly dark, labials lighter in illustration. Body with yellowish dots and

a series of yellowish spots on the vertebral row. A light vertebral stripe on the tail. Ventral colour

(in alcohol?) whitish.

SCALATION: Nasal undivided. Two internasals. Loreal present. Two postoculars. Temporals

1+2. Seven supralabials, third and fourth in contact with eye. Seven infralabials. Dorsal scales in

15 rows at midbody. Ventrals 140. Anal undivided. Caudals 27.


131


COMMENT: Known only from the holotype, which lacks a specific locality. It may be

conspecific with O. waandersi (see also comments under O. waandersi). In colour they share a

light vertebral stripe on the tail, never noted in descriptions of O. propinquus but clearly shown

in illustration. A black latero-ventral line in the rear of O. propinquus is also never mentioned in

the literature. The anal scale is entire in O. propinquus but almost always divided in O.

waandersi. Oligodon propinquus has an undivided nasal but it is always partially divided in O.

waandersi. The single specimen of O. propinquus might represent a large O. waandersi. Its lack

of a light collar marking is consistent with larger O. waandersi, although large O. waandersi

usually have much more faded spots than O. propinquus. Key Literature: (Boulenger, 1894; De

Rooij, 1917; Jan, 1862). Figured by Jan and Sordelli (1876, L. 48, Pl. I, Fig. 1). Max. length:

285mm (tail 40).

/0 106 170 Oligodon pseudotaeniatus Oligodon pseudotaeniatus David, Vogel and Rooijen 2008b: 18 (“Khorat”, Thailand: BMNH1938.8.7.37) DISTRIBUTION: Thailand (Nakhon Ratchasima, Bangkok, Saraburi). COLOURATION: (In preservative) Dominant dorsal colour grey to brown, scales narrowly

edged with black. Head markings most like O. taeniatus in the figures except the holotype seems

to have confluent chevron and temporal bars. Head with an ocular bar, temporal bars and narrow

chevron. The chevron is very pointed to the frontal and may be detached from its point there.

The chevron arms do not approach the ventrals. Body with a light narrow vertebral line flanked

by brown lines fading posteriorly. A faint but wide darker dorsolateral line on scale rows three

and four. Ventral colour yellowish, with dark spots on the margins. These ventral spots fainter,

and rounder anteriorally, darker and more quadrangular posteriorly, terminating abruptly part

way down the tail.

SCALATION: Nasal divided. Two internasals. Loreal present. Two preoculars. Two postoculars.

Temporals 1+2. Eight supralabials, fourth and fifth in contact with eye. Nine infralabials. Dorsal

scales in 17 rows at midbody. Ventrals 137–156, laterally angulate. Anal undivided. Caudals 34–

46.

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DENTITION: Fifteen maxillary teeth.

HEMIPENIS: As in O. taeniatus.

COMMENT: A member of the “taeniatus-group” (David et al., 2008b). Known from four

specimens only that have been stored in preservative for a considerable length of time. Figured

by David el al. (2008b, Figs. 10–14, photos). Max. length: 320mm (tail 66).

Oligodon pulcherrimus Oligodon pulcherrimus Werner 1909: 227 (“Pudang”, Sumatra, Indonesia : ZMH lost?) Oligodon pulcherrimus Tillack and Günther 2009 (Balinka, Sumatra, Indonesia: ZMB70756) Oligodon annulifer var. bipartita Despax 1912: 204 (Indes Néerlandaises: MNHN1912.0049) Oligodon durheimi Baumann 1913: 269 (Battak Gebirge, Indonesia: NHM1017619) DISTRIBUTION: Indonesia (Sumatra). COLOURATION: Dominant dorsal colour brown. Head markings dark brown, with an ocular

bar, temporal bars and chevron. Chevron extending well onto the frontal and in the back

confluent with the spot borders. Additional dark spots on the rostral, below the nostril and

between labials, on their margins. Body with 28–38 light brown elongate-oval or hexagonal

spots enclosed by a more or less connected black border. Anterior spots may be confluent.

Laterally bluish grey flecked with black and white. Ventral colour yellow with approximately

alternating ventrals showing black spots extending from the margin to the center.

SCALATION: Nasal divided. Two internasals, or fused to the prefrontals. Loreal present. Two

postoculars. Temporals 1+2 or 1+1. Seven, rarely 6 supralabials, third and forth in contact with

eye, rarely only the third. Seven, rarely 6 infralabials. Dorsal scales in 17 rows at midbody.

Ventrals 152–176. Anal divided. Caudals 38–53.



COMMENT: Formerly known only from the holotype, Tillack and Günther (2009) have

recently reviewed the species, designated a neotype, and referred additional specimens

(Baumann, 1913; Despax, 1912; Lönnberg and Rendahl, 1925) to the taxon. Thought by Werner

to be related to O. praefrontalis. Tillack and Günther ally it with O. praefrontalis and O.

annulifer. This species has 17 scale rows at mid-body, if mid-body is determined by ventral

133

count. There are 15 scale rows at mid-body if total length is used (Tillack and Günther, 2009).

Max. length: 376mm. Figured by Werner (1909, Fig. 4) and by Tillack and Günther (2009, Fig.

3, photos).

/106 106 103 Oligodon purpurascens Xenodon purpurascens Schlegel 1837: 90 (Mt. Parang, Java: RMNH) Simotes purpurascens Bleeker 1857: 472/3 [Simotes purpurascens=Xenodon purpurascens] Simotes purpurascens var. C Günther 1858: 25 [the only var. of Günther that is purpurascens] Simotes purpurascens var. albocinctus Jan and Sordelli 1865: L.12, Pl.V [example: odd colour] Simotes purpurascens var. B Werner 1900: 489 Holarchus purpurascens Barbour 1912: 195 [new combination] Oligodon purpurascens Wall 1923b: 328 [at this point cyclurus is united with purpurascens] Oligodon purpurascens maculatus Wall 1923b: 329 [part] Oligodon purpurascens purpurascens Haas 1950: 557 Simotes tri-notatus Duméril, Bibron and Duméril 1854: 631 (“China”?: MNHNP) Simotes purpurascens Jan 1863b: 46 [synonymizes tri-notatus]

Simotes trinotatus Günther 1864: 219 [synonymizes X. purpurascens of Cantor] Dicraulax trinotatus Cope 1893: 480 [new combination] Dicraulax purpurascens trinotatus Cope 1895: 200 Simotes purpurascens var. trinotata Boettger 1898: 73 Simotes albocinctus Duméril and Bibron 1854: 633 (Indes orientales: MNHNP)[nec Cantor 1839] Simotes purpurascens var. C Günther 1858: 25 [synonymizes albocinctus] Calamaria brachyorrhos Motley and Dillwin 1855: 49 (no loc.) Simotes purpurascens Günther 1861b: 216 [synonymizes C. brachyorrhos] Simotes labuanensis Günther 1864: 217 (Laboan im Reich Deli: BMNH) Simotes purpurascens var. labuanensis Boettger 1898: 73 Simotes obscurus Theobald 1868: 48 (no loc.: ZSI) Simotes crassus Theobald 1868: 48 (no loc.: ZSI) Simotes cyclurus Boulenger 1890a: 311 [synonymizes obscurus and crassus into cyclurus] Oligodon purpurascens Smith 1943: 206 [obscurus, crassus from cyclurus to purpurascens] Simotes catenifer Stoliczka 1873: 121 (Penang and Jahore: ZSI) Simotes dennysi Blanford 1881: 218 (Singapore: RM) Simotes affinis Fischer 1885b: 4 (Nias, Indonesia: MTKD1299/1300) [nec Günther 1862] Simotes purpurascens Boulenger 1890b: 34 [syns. labuanensis, catenifer, dennysi, affinis] Simotes cyclurus Boulenger 1912: 149 [moves catinifer to cyclurus] Oligodon purpurascens Smith 1930: 53 [moves catinifer back] Oligodon purpurascens kangeanicus Mertens 1959: 13 (Kanisangka: SMF55144) DISTRIBUTION: Indonesia (Nias; Sumatra; Borneo; Java; Kalimantan; Karimata Archipelago;

Mentawai Archipelago; Riau Archipelago; We; Kangean Is.), Malaysia (Malaya and East

Malaysia), Singapore, South Thailand.

134

COLOURATION: Dominant dorsal colour purple to brown. Head markings dark, with an

ocular bar, temporal bars (often faded) and chevron. Body with approximately 10–18 wavy

crossbars, sometimes very faded or absent. The crossbars usually quite thick, either light-edged

dark or thinner dark-edged light. Most individuals have about five faint, dark reticulations

between bands. Ventral colour yellowish or pinkish with black quadrangular spots covering half

or all of alternating ventrals.

A few specimens are said to present a colouration just like O. albocinctus.

SCALATION: Nasal divided. Two internasals. Loreal present. One or 2 preoculars, 1 or 2

suboculars. Two or 3 postoculars. Temporals 2+3 or 2+2. Usually eight supralabials, fourth and

fifth in contact with eye, sometimes the seventh excluded from lip, sometimes fourth divided into

second subocular. Nine infralabials. Dorsal scales in 19–21 rows at midbody. Ventrals 150–210,


DENTITION: Nine to 10 maxillary teeth. Palatine: nine teeth, an anterior edentulous space 1–2

teeth in size.

HEMIPENIS: Not forked. Large papillae. No spines.

COMMENT: This species shows almost as much colour pattern variation as O. cyclurus and O.

fasciolatus. Oligodon purpurascens and O. cyclurus were once thought to be conspecific. A

faunal break of some 500Km south of lat. 11º15'N on the Malay Peninsula divides these species

and their hemipenes differ (Smith, 1943). Key Literature: (Boulenger, 1890b; Boulenger, 1894;

Boulenger, 1912; David and Vogel, 1996; De Rooij, 1917; Smith, 1930; Stuebing and Inger,

1999; Taylor, 1965; Tillack and Günther, 2009). Figured by De Rooij (1917, Fig. 52), Chan-Ard

et al. (1999, p175, spots well photographed) and Stuebing and Inger (1999, p106, photo). Max.

length: 950mm.

/125 92 146/0 30 0/109 128 92 Oligodon saintgironsi Oligodon saintgironsi David, Vogel and Pauwels 2008a: (“Arboretum de Trang Bom”, Vietnam: MNHN 1974.1264) DISTRIBUTION: Southern Vietnam (Dong Nai province). COLOURATION: Like O. ocellatus or the “smithi” pattern of O. cyclurus. Dominant dorsal

colour brown. Head markings dark brown, with an ocular bar, temporal bars and chevron. Body

135

with 10–13+2–4 large dark brown blotches or crossbands. In between the crossbands, a series

reticulations.

SCALATION: Nasal divided. Two internasals. Loreal present. Two preoculars, one might be

considered a presubocular. Two postoculars. Temporals 2+2. Eight supralabials, fourth and fifth

in contact with eye. Nine infralabials. Dorsal scales in 17–18 rows at midbody. Ventrals 166–



HEMIPENIS: Very long, reaching subcaudal 28–29. Deeply forked. No papillae. No spines.

COMMENT: Known from very few specimens, possibly more in collections as O. cyclurus

with 17 scale rows. See also comments in O. cyclurus. As currently constituted, 17(–18) scale

row “cyclurus”, have only been detected in a restricted southern Vietnam range, and are referable

to this taxon. May be distinguished from O. ocellatus (overlapping range) by having a longer tail

and fewer mid-body scale rows, and from O. chinensis (overlapping scale row counts) by

disjunct range, hemipenial length and a higher scale row count on the anterior third of the body

(19 as opposed to 17 in O. chinensis). Key Literature: (David et al., 2008a).

Oligodon signatus Oligodon purpurascens var. F Günther 1858: 26 [referred into signatus by Günther 1864] Simotes signatus Günther 1864: 215 (Singapore Island: BMNH 1946.1.3.20) Holarchus signatus Barbour 1912: 195 [new combination] Oligodon signatus Smith 1930: 55 [new combination] Simotes subcarinatus Günther 1872a: 595 (Sarawak, Malaysia :BMNH 1946.1.4.24) Holarchus subcarinatus Barbour 1912: 195 [new combination] Oligodon subcarinatus de Haas 1950: 558 [new combination] Simotes annulifer var. annulata Lidth de Jeude 1922: 245 (Padang, Indonesia: RMNH 4690) DISTRIBUTION: Singapore Island, Indonesia (Sumatra; Borneo; Java?), Malaysia (Malaya). COLOURATION: Dominant dorsal colour dark brown to red-brown. Head markings dark on

a yellowish background, with an ocular bar, temporal bars and chevron. Body with about 12–

17+3–5 dark-edged tan-yellow (in alcohol, red in life) oval spots or bands, the most anterior with

a chevron shape, the point towards the head and arms at the back extending laterally. Ventral

colour yellowish with brown quadrangular spots on the edges of the ventrals.

136

SCALATION: Nasal divided. Two internasals. Loreal present, sometimes fused to the prefrontal.

One postocular. Temporals 1+2 (rarely 1 + 3). Seven supralabials, third and fourth in contact

with eye. Seven or 8 infralabials. Dorsal scales in 17–15 rows at midbody. Ventrals 141–160,

may be weekly keeled. Anal undivided. Caudals 43–59.



COMMENT: A recent review (Tillack and Günther, 2009) synonymized O. subcarinatus and O.

annulifer annulata into O. signatus and summarized much of the locality and checklist records

for this species (Bartlett, 1895; Batchelor, 1958; David and Vogel, 1996; Mocquard, 1890;

Robinson and Kloss, 1920; Robinson and Kloss, 1923; Shelford, 1901; Stuebing, 1991;

Stuebing, 1994; Sworder, 1922; Werner, 1893; Werner, 1896). Rare on the peninsula and

Singapore (Hendrickson, 1966; Tweedie, 1953; Voris, 1977). Individuals with 15 as opposed to

17 scale rows are typically from Borneo. A few recently collected specimens differing somewhat

in colour (Leong and Grismer, 2004) are probably close relatives of O. signatus, if not

conspecific. Key Literature: (David and Vogel, 1996; De Rooij, 1917; Stuebing and Inger, 1999;

Tillack and Günther, 2009). Photographed by Tillack and Günther (2009, Figs. 5 and 6). Figured

by Günther (1864, Pl. 20, Figs. F, F'). Max. length: 525mm (tail 130).

/210 190 190 Oligodon splendidus Simotes splendidus Günther 1875: 231 (“Wynad”?: BMNH) Oligodon splendidus Wall 1923b: 331 [new combination] DISTRIBUTION: Central Myanmar. COLOURATION: Dominant dorsal colour light brown, each scale with a dark center. Head

markings dark, with an ocular bar, temporal bars and thick chevron. The ocular and temporal

bars usually indistinct. Body with 14–17+3–5 very large dark spots. The spots are indented in

front and behind, are edged with black and outlined by buff. Ventral colour whitish to yellowish

with black quadrangular spots on alternating sides of alternating ventrals.

SCALATION: Nasal divided. Four prefrontals and sometimes even 4 internasals, for a total of

eight (Wall and Evans, 1901a). Loreal present. Sometimes a postocular. Two postoculars, rarely

3. Temporals 2+3. Eight supralabials, fourth and fifth in contact with eye. Eight infralabials.

137

Dorsal scales in 21 rows at midbody. Ventrals 169–193, laterally angulate. Anal undivided.

Caudals 35–47.


HEMIPENIS: As O. cinereus. Not forked. Two large papillae of unequal length. No spines.

Distally, with longitudinal folds merging into a proximally calyculate area.

COMMENT: Possible error in type locality. Apparently common within its restricted range.

Key Literature: (Evans, 1905; Smith, 1943; Wall, 1923b; Wall, 1926; Wall and Evans, 1901a).

Figured by Smith (1943, Fig. 69, pattern only) and Günther (1875, Pl. XXXIII). Max. length:

730mm (tail 100).

/36 105 179 Oligodon sublineatus Oligodon sublineatum Duméril, Bibron and Duméril 1854: 57 (Ceylon: MNHNP) Oligodon sublineatus Günther 1858: 21 DISTRIBUTION: Sri Lanka, Nicobar Islands? (Annandale, 1905; discredited by Wall, 1921c). COLOURATION: Dominant dorsal colour brown, many scale edges black and white. Head

markings often faint, obscure with age. An ocular bar present, no temporal bars, the chevron

somewhat forward and W shaped. The chevron sometimes a spot on the frontal and two spots in

the area of the temporal bars. Body with 12–15+3–4 paired or alternating dark spots or crossbars.

Ventral colour yellowish with three longitudinal series of dark spots, centrally and laterally. The

lateral series often confluent, the median sometimes absent.

SCALATION: Nasal divided. Two internasals. Loreal present, very rarely absent on one side.

Two postoculars. Temporals 1+2. Seven supralabials, third and fourth in contact with eye. Seven

infralabials. Dorsal scales in 15 rows at midbody. Ventrals 130–161. Anal divided. Caudals 23–

39.


HEMIPENIS: Forked at the tip. No papillae. Uniform size spines throughout.

COMMENT: One of the most common of all Ceylonese snakes (Abercromby, 1910;

Abercromby, 1911). Neither included in the fauna of Nicobar by Das (1996; 1999), nor listed in

tables of erroneous reports. Key Literature: (De Silva, 1980; Smith, 1943; Wall, 1911b; Wall,

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1921a; Wall, 1921c; Wall, 1923b; Wall, 1924b). Figured by Jan and Sordelli (1876, L. 48, Pl. I,

Fig. 2.) and Taylor (1950, Pl. XVII, Fig. 1). Max. length: 350mm (tail 40).

/0 122 183 Oligodon taeniatus Simotes taeniatus Günther 1861a: 189 (Cambodia: type lost? [But see below]) Oligodon taeniatus David, Vogel and Rooijen 2008: 10 [give BMNH1946.1.3.27 as holotype] Simotes quadrilineatus Jan and Sordelli 1865: L. 12, Pl. 4, Fig. 3 S. quadrilineatus David, Vogel and Rooijen 2008: 13 [give MNHN1991.1819 as neotype] Simotes sexlineatus Morice 1875: 57 [Bourret 1936: 229, “probably H. taeniatus”] Holarchus taeniatus taeniatus Cochran 1930: 28 Oligodon toeniatus toeniatus Bourret 1934d: 18 Oligodon taeniatus taeniatus Bourret 1934e: 79 Simotes taeniatus Boulenger 1914: 69 [selects 19 scale rows for taeniatus] Oligodon quadrilineatus Smith 1943: 210 [given Günther 1964, chose next name for 19 scales] Oligodon taeniatus Campden-Main 1969: 295 Incertae sedis

See Campden-Main (1969), also Oligodon taeniatus Yang et al. 1980: 262 [discusses Chinese taeniatus as east of Red River, perhaps O. formosanus if scale rows = 19] Oligodon taeniatus Zhao et al. 1998: 208 [gives the scale rows 17-17-15 for very few specimens] DISTRIBUTION: Thailand , Laos, Cambodia, Vietnam, Myanmar?, China? (Yunnan). COLOURATION: Dominant dorsal colour grey to brown. Head markings dark with an ocular

bar, short temporal bars and narrow chevron. The chevron is very pointed to the frontal and may

be detached from its point there. The chevron arms do not approach the ventrals. Body with a

light vertebral line flanked by brown lines (sometimes faded). A fine black lateral line between

scale rows 3 and 4. Ventral colour whitish or red, tail red, with black quadrangular spots usually

confined to the edges of the ventrals. Lacking black spots on the tail.

SCALATION: Nasal divided. Two internasals. Loreal almost always present. One or 2

preoculars. Two postoculars. Temporals 1+2. Eight supralabials, fourth and fifth in contact with

eye. Nine, very rarely 10 infralabials. Dorsal scales in 19 rows at midbody. Ventrals 141–169,



HEMIPENIS: Deeply forked. Large papillae. No spines. Calyculate proximal to the fork.

139

COMMENT: Recently reviewed (David et al., 2008b), O. taeniatus gives its name to the

“taeniatus-group”, which is defined by hemipenial morphology. The group consists of O.

taeniatus, O. mouhoti, O. barroni, O. deuvei, O. pseudotaeniatus and O. moricei. The taxonomic

history of O. taeniatus and O. mouhoti are particularly entangled. It is not clear that either of

these species occurs any where near the Song Hong (Red River) much less east of it. The western

extent of its range is also unclear. Long thought to end in Thailand, and contrary to David et al

(2008), O. taeniatus does seem to occur in Myanmar (Dowling and Jenner, 1988; Zug et al.,

1998). It is confirmed in Laos (Teynie and David, 2007). Campden-Main (1969) and the recent

review discuss which specimens and publications refer to which name. Campden-Main implied

BMNH1946.1.3.24, 27 could serve as neotypes, but he did not officially designate them. David

et al. (2008) explicitly identified the holotype from the BMNH collection. Key Literature:

(Bourret, 1937b; Bourret, 1943; Campden–Main, 1969; Saint Girons, 1972b; Smith, 1943;

Taylor, 1965). Figured by Jan and Sordelli (1965, L. 12, Pl. 4, Fig. 3) and Cox et al.(1998, p61,

photo). Max. length: 447mm (tail 80).

/241 155 140 Oligodon taeniolatus Coluber taeniolatus Daudin 1803: 428 (Russell, 1796, Pl. XIX, nomen oblitum by Bauer (2003)) Coronella taeniolata Jerdon 1853: 528 (“common at Madras”) Xenodon dubium Jerdon 1853: 528 (north Canara: type lost) Oligodon subgriseum Duméril, Bibron and Duméril 1854: 59 (Pondicherry: MNHNP) Oligodon taeniolatus Wall 1921c: 239 [establishes priority of Jerdon’s name] Oligodon spilonotus Günther 1864: 207 (Madras and Malibar: BMNH) Oligodon fasciatus Günther 1864: 208 (Deccan: BMNH) Oligodon subgriseus Boulenger 1890a: 321 [synonymizes dubium, spilonotus and fasciatus] Oligodon elliotti Günther 1864: 207 (Madras: BMNH) Oligodon taeniolatus Smith 1943: 223 [synonymizes elliotti] Oligodon subgriseus alternans Bethancourt-Ferreira 1897: 224 (Goa: Lisbon) Oligodon taeniolatus var. typicus Wall 1921c: 239 Oligodon taeniolatus var. ceylonicus Wall 1921c: 240 Oligodon taeniolatus var. fasciatus Wall 1921c: 241 Oligodon taeniolatus var. spilonotus Wall 1921c: 241 Oligodon taeniolatus fasciatus Taylor 1950: 550 [recognizes subspecies] Oligodon taeniolatus ceylonicus Taylor 1950: 550 [recognizes subspecies] Contia transcaspica Nikolsky 1903: 11 (Kopet-Dag: ZISP9869) Oligodon taeniolatus Chernov 1935: 351 [synonymizes C. transcaspica] DISTRIBUTION: India (Maharashtra), Pakistan, Sri Lanka, southern Turkmenistan (Kopet-Dag),

eastern Iran, Afghanistan.

140

COLOURATION: Dominant dorsal colour light brown to buff. Head markings dark, with an

ocular bar, temporal bars and chevron. The head markings show a full range of confluent

permutations. The temporal bars are often confluent on the frontal, the chevron is often

connected to the temporal bars and the ocular bar. Sometimes the head is almost completely

dark.

taeniolatus: Body with narrow black crossbars on the scale edges. Also 4 longitudinal brown

stripes, 2 adjoining a light vertebral line, 2 on scale rows 2 and 3. Ventral colour yellowish with

or without black spots.

ceylonicus: As above, the stripes more or less present, the crossbars more or less obscure. This

may vary year to year.

fasciatus: Crossbars enlarged into sometimes asymmetrical median spots with paired lateral

spots, sometimes white edged. No longitudinal stripes

spilonotus: Crossbars enlarged to 13–22 medially indented spots. Faint stripes usually visible.

alternans: Large dark, black then white edged spots, paired or alternating either side of the

vertebral row.

SCALATION: Nasal divided. Two internasals. Loreal almost always present. One, very rarely 2

preoculars. Two postoculars, rarely 3. Temporals 1+2 (very rarely 2+2 with a segment of

parietal). Seven supralabials, third and fourth in contact with eye. Seven infralabials. Dorsal

scales in 15 rows at midbody (Taylor gives 19-17-15 for ceylonicus De Silva, aware of Taylor,

gives 15 midbody, so it would seem Taylor gave two reductions before midbody; Constable

(1949), gives 13 in a single specimen). Ventrals 152–218, slightly laterally angulate. Anal



HEMIPENIS: Two fifths forked at the tip. No papillae. Spinose from the base to the fork, spines

decreasing is size distally. The tip has four longitudinal folds.

COMMENT: Wall’s opinion on the ratio of stripe to crossbar intensity of the different forms

and localities changed from 1921 to 1923. Taylor’s (1950) opinion more closely matched that of

Wall’s earlier views, while Deraniyagala (1955) was nearer the later. Colour pattern may vary

with time. There is a variety of opinions as to what, if any, patterns should be accorded

subspecific rank. None of these opinions contains a coherent argument to refute Smith’s

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assertion that all the colour forms are continuous and there is considerable range overlap.

Mahendra (1984) referred O. melaneus and O. templetoni to subspecific status without

accounting for their distinctive hemipenial morphology. Its IUCN status is “Lower Risk near

threatened”. Key Literature: (De Silva, 1980; Dotsenko, 1984; Sharma, 1982; Smith, 1943;

Taylor, 1950; Wall, 1923b). Figured by Smith (1943, Fig. 76) and Jan and Sordelli (1876, L. 48,

Pl. I, Fig. 3). Max. length: 590mm (tail 63).

/241 122 5 Oligodon templetoni Coluber Calamarius Linnaeus 1758: 216 (“America”: NHRM) Oligodon templetonii Günther 1862: 57 (Ceylon: BMNH) Oligodon calamarius Andersson 1899: 8 [calamarius should have priority over templetoni] Oligodon calamarius Smith 1943: 228 [first to apply Anderson’s judgment] Calamaria Brongersma et al. 1966[revoke calamarius for Oligodon, templetoni next available] [O. calamarius continues in wide use] Oligodon taeniolatus calamarius Mahendra 1984: 203 [synonymizes templetoni, no comments] DISTRIBUTION: Sri Lanka. COLOURATION: Dominant dorsal colour brown. Head markings very indistinct. An ocular

bar, usually only strong below the eye. Short temporal bars and chevron, more or less confluent

on the frontal if present. Body with a more or less distinct vertebral streak. Also 18–24+5 dark,

very thin crossbars, which can be across the back or incomplete, on half the back, alternating

with the other side. Ventral colour yellowish or pinkish with equal proportion of black

quadrangular spots, paired or centrally distributed.

SCALATION: Nasal divided. Two internasals. Loreal present, sometimes in contact with eye.

Two or 3 postoculars. Temporals 1+2. Seven supralabials, third and fourth in contact with eye,

sometimes sixth excluded from lip. Seven infralabials. Dorsal scales in 15 rows at midbody.



HEMIPENIS: Not forked. No papillae. Uniformly dense spines throughout.

COMMENT: Said to be rare in its range. Confusion about the type species of genus Calamaria

(Andersson, 1899) has lead to a complex petition that would see the long standing name

templetoni stand for Oligodon calamarius. Brongersma et al. (1966) summarize the arguments.

Mahendra (1984) has, without any discussion, referred templetoni (calamarius) to a subspecies

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of taeniolatus. This change requires some discussion to refute Smith’s hemipenial characters.

The hemipenes differ only in a small amount of forking, and it is not clear if this is only known

from continental examples. Key Literature: (De Silva, 1969; Smith, 1943; Wall, 1921c; Wall,

1923a). Figured by Smith (1943, Fig. 78, pattern only) and Günther (1864, Pl. XIX, Fig. C).


/119 168 109 Oligodon theobaldi Simotes theobaldi Günther 1868: 417 (Pegu, Myanmar: BMNH) Oligodon theobaldi Wall 1923b: 322 [new combination] Simotes beddomii Boulenger 1890a: 229 (“Wynad”?, India: BMNH) Simotes theobaldi Wall 1914c: 170 [synonymizes beddomii; most certainly not from Wynad] DISTRIBUTION: Central Myanmar, India (Assam). COLOURATION: Dominant dorsal colour light brown. Head markings dark, with an ocular

bar, temporal bars and chevron. Temporal bars are confluent, chevron sometimes very thick

posteriorly. Body with many darkened scales edges forming reticulations. A pale vertebral line

bordered by thicker dark lines and a dark lateral line on scale rows 2–4. Ventral colour yellowish

or brownish, with or without black quadrangular spots on the edges of the ventrals.


Eight supralabials, fourth and fifth in contact with eye. Eight infralabials. Dorsal scales in 17

rows at midbody. Ventrals 163–180. Anal divided. Caudals 27–49.

DENTITION: Fifteen to 16 maxillary teeth.

HEMIPENIS: Not forked. Two large papillae. Proximal half with small uniform spines.

COMMENT: Constable (1949) challenged Wall’s rejection of South India as part of the range

of this species. His specimen originally bore the designation O. beddomii. Considering that many

species are endemic to the Western Ghat ranges, O. beddomii might be a distinct species, if from

this area. Key Literature: (Smith, 1943; Wall, 1914c; Wall, 1923b). Figured by Smith (1943, Fig.

74, body only) and Boulenger (1894, Pl. IX, Fig. 2). Max. length: 390mm (tail 47).

/128 62 38 Oligodon torquatus Simotes torquatus Boulenger 1888: 597 (Bhamo, Myanmar: MSNG)

143

Oligodon torquatus Wall 1923b: 309 [new combination] DISTRIBUTION: Myanmar (Irrawaddy Valley). COLOURATION: Dominant dorsal colour grey brown. Head markings faintly darker.

Usually described as having a black spot below they eye (where part of the ocular bar would be)

and a band on the neck. In one specimen seen by me, with the ocular spot continuing above the

eye, but not confluent across the top, temporal bars and the mark on the neck somewhat more a

chevron blotch, stopping at the back of the parietals, but continuing to the frontal in the

interparietal suture. Body with black and white reticulations confined to the scale edges and 4

darker stripes, 2 vertebral, 2 lateral. Ventral colour white with a few black quadrangular spots on

the posterior ventrals.

SCALATION: Nasal undivided. Two internasals. Loreal present. Two postoculars. Temporals


15 rows at midbody. Ventrals 144–169, slightly laterally angulate. Anal divided. Caudals 25–34.

DENTITION: Fifteen to 16 maxillary teeth.

HEMIPENIS: Not forked. No spines. Proximally, with longitudinal folds and distally with

spongy papillae.

COMMENT: Relatively rarely referred to in the literature, but apparently quite common within

its restricted range (Wall, 1925b). Key Literature: (Boulenger, 1888; Smith, 1943; Wall, 1925b).

Figured by Boulenger (1888, Pl. V, Fig. 1). Max. length: 292mm (tail 37).

/0 65 0 Oligodon travancoricus Oligodon travancoricum Beddome 1877: 685 (Tinnevelly Hills, India: BMNH) Oligodon travancoricus Boulenger 1890a: 318 DISTRIBUTION: India (Western Ghats south of Palghat). COLOURATION: Dominant dorsal colour grey-brown. Head markings darker, with a broad

ocular bar, temporal bars and chevron extending to the parietals. The head markings are often

confluent on the frontal. Body with 25–33 light-edged dark crossbands, 5 or 6 on the tail. These

crossbands are often indented along the vertebral line and are much like a thin O. venustus

marking. Ventral colour yellow or whitish with equal proportion black quadrangular spots.

144

SCALATION: Nasal divided. Two internasals. No loreal. Two postoculars. Temporals 1+2 or

2+3. Seven supralabials, third and fourth in contact with eye, the sixth often excluded from the

labial border (and mistaken for an anterior temporal?). Eight infralabials. Dorsal scales in 17

rows at midbody. Ventrals 145–155. Anal divided. Caudals 34–37.


HEMIPENIS: Not forked. No papillae. Distally with transverse flounces edged with small

spines. merging into a 1/3 densely spinose region at the base.

COMMENT: Very few specimens known. Although Wall synonymized this species with O.

venustus in 1914(b), he changed his opinion in 1923. Hemipenial characters distinguish the two

species. Its IUCN status is “Endangered”. Key Literature: (Beddome, 1877; Ferguson, 1895;

Smith, 1943; Wall, 1914b). Figured by Boulenger (1894, Pl. X, Fig. 2). Max. length: 465mm

(tail 65).

/117 68 154 Oligodon trilineatus Simotes trilineatus Duméril. Bibron & Duméril 1854: 636 (“grandes Indes”: MNHN 3541) Oligodon trilineatus Fischer 1885b: 7 [refers trilineatus to Oligodon] DISTRIBUTION: Indonesia (Nias; Sumatra; Mentawai). COLOURATION: Body, above and below, dark-reddish brown, head yellowish brown. Head

marked with dark temporal bars. Body with a yellow (red?) vertebral stripe and white latero-

ventral lines. Some specimens also with an additional fine yellow line on each side.

SCALATION: Nasal divided. Two internasals. Loreal present. One postocular. Temporals 1+2 or

2+1. Seven supralabials, third and fourth in contact with eye. Seven or 8 infralabials. Dorsal

scales in 17 rows at midbody. Ventrals 145–162. Anal undivided. Caudals 39–62.



COMMENT: Very few specimens are known. Tillack and Günther (2009) report an ocular bar

in colouration. Key Literature: (David and Vogel, 1996; De Rooij, 1917; Duméril et al., 1854;

Tillack and Günther, 2009). Figured by Jan and Sordelli ( 1865, L. 12, Pl. IV, Fig. 1), holotype

photographed by Tillack and Günther (2009, Fig. 7). Max. length: 507mm (tail 80).

145

/138 84 68 Oligodon unicolor Simotes unicolor Kopstein 1926: 107 (Jamdena, Tanimbars, Indonesia: RMNH) Oligodon unicolor Haas 1950: 558 [new combination] DISTRIBUTION: Indonesia (Tanimbar Islands). COLOURATION: Like O. forbesi varying slightly in head markings: with an ocular band.

Without any faint body stripes.


Six supralabials, third and fourth in contact with eye. On one side the third and fourth

supralabials partially fused. Dorsal scales in 17 rows at midbody. Ventrals 162, slightly laterally

angulate. Anal undivided. Caudals 41.



COMMENT: Known from the holotype only. Kopstein (1926) believed the species to be closely

related to O. forbesi. Both O. forbesi and O. unicolor are very similar to O. octolineatus. The

ranges of O. octolineatus and O. unicolor are not known to overlap, although O. forbesi and O.

unicolor appear to be sympatric. Oligodon forbesi and O. unicolor appear to have the same

scalation, perhaps differing in the number of supralabials/temporals, but see comments for O.

forbesi. Key Literature: (Kopstein, 1926). Never illustrated. Max. length: 300mm (tail 45).

/242 99 53 Oligodon venustus Xenodon venustum Jerdon 1853: 528 (North Canara dist., India) Simotes venustus Günther 1864: 213 [refers venustum to Simotes] Oligodon venustus Boulenger 1890a: 317 [refers venustus to Oligodon] Simotes bi-notatus Duméril, Bibron and Duméril 1854: 630 (Malibar dist., India: MNHNP) Oligodon venustus Boulenger 1894: 235 [synonymizes binotatus] Incertae sedis S. purpurascens Beddome 1862: 12 [Madras specimens from the collector of O. venustus and O. travancoricus, it could not be purpurascens if Indian, but it could be one of these.] DISTRIBUTION: India (Western Ghats south of the Goa Gap).

146

COLOURATION: Dominant dorsal colour grey-brown. Head markings dark (faded in older

examples), with an ocular bar, temporal bars and thick chevron. Temporal bars usually confluent

on the frontal, and chevron usually confluent with them. Body with 23–31+6–7 dark spots,

varying in shape from transversally oval to rhomboidal and deeply medially indented. Main spots

often paired with confluent lateral spots. Ventral colour yellowish to whitish with black

quadrangular spots in equal proportion. Fewer spots on the tail.

SCALATION: Nasal semi-divided. Two internasals. Loreal usually present, but frequently absent

(contra Smith). Two postoculars. Temporals 1+2. Seven, rarely 6 (very rarely 8), supralabials,

third and fourth in contact with eye. Seven infralabials. Dorsal scales in 17 rows at midbody.



HEMIPENIS: Not forked. No papillae. Proximal one third with coarse dense spines. Distally,

with transverse folds.

COMMENT: Its IUCN status is “Lower Risk near threatened”. One of several Oligodon in the

western Ghats with a very restricted range (Patel and Reddy, 1995; Vyas, 1998). Key Literature:

(Boulenger, 1894; Smith, 1943; Wall, 1919; Wall, 1923b). Figured by Smith (1943, Fig. 75,

body only) and Jan and Sordelli (1865, L. 11, Pl. VI, Fig. 3, as S. binotatus). Max. length:

490mm (tail 65).

/38 35 38 Oligodon vertebralis Simotes vertebralis Günther 1865: 91 (Bandjermasin, Borneo, Indonesia: BMNH) Oligodon vertebralis Boulenger 1894: 245 [refers vertebralis to Oligodon] Oligodon notospilus Günther 1873: 169 (Mindanao, Philippines: BMNH) Oligodon vertebralis notospilus Leviton 1963a: 477 [refers notospilus to subspecific status] Oligodon schadenbergi Boettger 1895: 4 (Busuanga, Philippines: MTKD) Oligodon iwahigensis Griffin 1909: 598 (Iwahig, Philippines: type lost) Oligodon vertebralis Thompson 1913: 216 [Philippine example, v. notospilus, as vertebralis] Oligodon iwahigensis Slevin and Leviton 1956: 548 [neotype CAS 62173] Oligodon vertebralis notospilus Leviton 1963a: 477 [synonymizes schadenbergi, iwahigensis] DISTRIBUTION: O. v. vertebralis: Indonesia (Borneo).

O. v. notospilus: Philippines (Calamian Islands; Palawan; Busuanga; Balabac; Mindanao).

147

COLOURATION: (in alcohol) Dominant dorsal colour dark grey to purplish-brown. Head

markings dark, with an ocular bar, confluent temporal bars and chevron. Body sometimes with

approximately 9+2 black-edged yellow or white dorsal blotches. Also additional small black-

edged light spots laterally. Ventral colour (in alcohol) immaculate yellow to cream, possibly red

in life.

SCALATION: Nasal divided. Two internasals. Loreal present. One or 2 postoculars. Temporals


15 rows at midbody. Ventrals 136–154. Anal single or divided. Caudals 32–54.


HEMIPENIS: Not forked. Two long papillae. No spines. Proximally smooth.

COMMENT: Leviton (1963a) extensively discussed the taxonomic history of O. vertebralis

notospilus. The ranges of O. v. vertebralis and O. v. notospilus appear to be disjunct. Stuebing

and Inger (1999) stated that based on colour and scalation O. v vertebralis and O. everetti may be

conspecific. Oligodon v. vertebralis is known from very few specimens. The name O. vertebralis

would have priority over O. everetti. Key Literature: (De Rooij, 1917; Leviton, 1963a; Stuebing

and Inger, 1999). Figured by Boulenger (1894, Pl. XI, Fig. 2) and Taylor (1922, Pls. 7, Fig. 2, 17

Figs. 3–5, 18 Fig. 1). Max. length: 345mm (tail 75).

/0 119 193 Oligodon waandersi Rabidon cruciatum Bleeker 1860c: 82 (domain of Boni, Indonesia: BMNH1946.1.3.12)

Rabidon Waandersi Bleeker 1860c: 83 (Boni, north Celebes, Indonesia: BMNH1946.1.2.52) Oligodon waandersi Günther 1865: 91 [refers waandersi to Oligodon synonymizes cruciatum] Oligodon taeniurus Müller 1897: 826 (Lilang and Pinogo, Indonesia: BMNH1946.1.3.15) Oligodon waandersi De Rooij 1917: 133 [synonymizes taeniurus] DISTRIBUTION: Indonesia (Sulawesi; Buton; Sula Islands). COLOURATION: Changes considerably from juvenile to adult. Dominant dorsal colour

reddish-brown, the head of juveniles often lighter. Head markings darker, most obvious in the

juveniles. Traces of an ocular bar below the eyes is present. Temporal bars are present, often

described as delineating a light coloured collar which remains in all but the largest specimens. A

dark blotch is often found along the edges of the suture between the parietals and the frontal.

Body with a reddish vertebral stripe on the tail. The juveniles also have a vertebral series of

148

widely spaced, small, black-edged orange spots; these alternate with a lateral series of paired

spots. The spots are sometimes more yellow or white and fade with age. Ventral colour (some

disagreement) yellowish with some dark spots (caudals may be pinkish) a black latero-ventral

line in juveniles.

SCALATION: Nasal partially divided (incorrectly entire in Boulenger (1894, but figured

correctly) and De Rooij (1917)). Two internasals. Loreal present or absent. One or 2 postoculars.

Temporals 1+2. Five to 7 supralabials, third and fourth in contact with eye. Six to 8 infralabials.

Dorsal scales in 15 rows at midbody. Ventrals 134–168. Anal divided (rarely entire). Caudals

18–29.



COMMENT: A detailed review of this snake was provided by in den Bosch (1994). The amount

of variation displayed by this species is remarkable, especially considering the small sample size.

Some of the examples bear a close resemblance to some O. vertebralis and O. everetti. Although

not well collected, in den Bosch’s suggestion that it is common on Sulawesi seems supported by

the number of locality records (Barbour, 1912; Boettger, 1898; De Rooij, 1917; Haas, 1950; in

den Bosch, 1985). Online, Uetz

(http://www.jcvi.org/reptiles/species.php?genus=Oligodon&species=waandersi; 24 February

2010), on Wagner’s authority places O. propinquus in the synonymy of O. waandersi. Key

Literature: (Bleeker, 1860c; in den Bosch, 1994; Kopstein, 1927; Kopstein, 1935; Smith, 1927).

Figured by Boulenger (1894, Pl. XI, Fig. 3) and in den Bosch (1994, Figs. 1–4). Max. length:

335mm (tail 36).

/244 65 73 Oligodon woodmasoni Simotes wood-masoni Sclater 1891: 235 (Andamans and Nicobars, India: ZSI8459) Oligodon woodmasoni Wall 1923b: 325 [new combination] DISTRIBUTION: India (Andaman and Nicobar Islands). COLOURATION: Very like O. octolineatus. Dominant dorsal colour brick-red. Head

markings darker, with an ocular bar, temporal bars and Y shaped chevron extending onto the

149

frontal. Body with seven yellowish (redish in life) stripes separated by darker stripes. Stripes

more pronounced in the juveniles. Ventral colour dusky red.

SCALATION: Nasal divided. Two internasals. Small loreal present on all but one side of one

specimen. Two postoculars. Temporals 1+2. Six supralabials, third and fourth in contact with

eye, or third excluded by a subocular. Seven infralabials. Dorsal scales in 17 rows at midbody.

Ventrals 180–190, laterally angulate. Anal undivided. Caudals 46–57.


HEMIPENIS: Not forked. No spines. At the base, with transverse folds, becoming calyculate

toward the tip. Distally with large papillae.

COMMENT: Previously known from just three specimens (Annandale, 1905; Das, 1996),

additional specimens were collected recently (Vijayakumar and David, 2006). Smith (1943)

stated it was related to O. octolineatus. In contrast, Boulenger (1894) firmly believed it to be

related to O. trilineatus. Its IUCN status is “Data Deficient”. Wall (1923b) gave specimen

numbers, one of which was probably a type and another probably the third specimen. Key

Literature: (Sclater, 1891; Smith, 1943; Wall, 1923b). Figured by Sclater (1891, Pl. VI, Fig. 2,

head), Smith (1943, Fig. 73, body), and photographed in life by Vijayakumar and David (2006,

Fig. 6). Max. length: 620mm (tail 120).

/244 133 141 Incertae sedis Oligodon dorsale Berthold 1859: 179 [nec Gray] Oligodon bertholdi Werner 1929: 138 [nomen novum for O. dorsatus, a single specimen in ZIUG, thought by Günther and Boulenger to be another Genus, Werner assigned a new name

without comment. Nomen oblitum]

Material introduced as, or once referred to Oligodon.

Some collections of Maculophis bella (until recently Elaphe) have been published as Oligodon

bella and O. bellus (Burbrink and Lawson, 2007; Ding and Zheng, 1974; Jiang et al., 2006;

Schulz, 1996; Schulz et al., 2000).

Holarchus roulei Angel and Bourret 1933: 135 (Chapa [=Sa Pa], northern Vietnam)

Presently in synonymy with Euprepiophis mandarinus (Smith, 1943; Utiger et al., 2002).

150

Simotes australis Krefft 1864: 180 (Port Curtis, Australia)

Presently in synonymy with Simoselaps australis (Cogger, 2000).

Simotes coccineus Duméril, Bibron and Duméril 1854: 637

Presently in synonymy with Cemophora coccinea (Williams, 1985).

Simotes Conradi Peters 1874: 376 (ZMB8186)

Simotes Herzi Boettger 1886c: 519 (Kiang-Su, China: SMF18383)

Both presently in synonymy with Elaphe rufodorsata (Stejneger, 1907).

Oligodon evansi Wall 1913a: 514 (Taounggyi, Myanmar)

Presently in synonymy with Plagiopholis nuchalis (Smith, 1943).

Oligodon melanocephalus septentrionalis Werner 1905: 411

Oligodon melanocephalus satunini — Chernov 1937: 349

Once referred to Oligodon, but are presently referable to Rhynchocalamus (Darevsky, 1970;

Reed and Marx, 1959).

Oligodon sauteri Steindachner 1913: 2

Presently in synonymy with Sinomicrurus sauteri (Ota et al., 1999; Slowinski et al., 2001).

Simotes semicinctus Peters 1862: 637 (probably Borneo: ZMB4553)

Initially revalidated as Oligodon by Wallach and Bauer (1996), presently in synonymy with

Coronella austriaca (Tillack et al., 2008).

Oligodon subpunctatus Duméril and Bibron 1854: 58 ("Malabar, Inde”, India: MNHN 3240)

Presently in synonymy with Sibynophis subpunctatus (Captain et al., 2004).

Simotes Vaillanti Sauvage 1876: 275

Presently in synonymy with Oreocryptophis prophyraceus (Bourret, 1935a; Utiger et al., 2005;

Werner, 1903; Zhao and Adler, 1993).

151

Appendix B

Key to the species of the genus Oligodon.

Every effort has been made to include range, colour and scale characters to distinguish each

species. In some cases, specimens of unknown origin may be difficult to identify.

Midbody dorsal scale rows(MDSR) are used as diagnostic characters. Wherever clarity permits,

larger than known ventral scale ranges are used. Only four species are known to have variation in

the condition of the anal scale and this is considered in the key. Regardless, creases in the anal

scale are known and thus the condition should be interpreted carefully.

1A. Internasals+prefrontals equal to or greater then 6; with large dark spots .………O. splendidus

1B. Internasals+prefrontals less then 6 ……………………………………………………………2

2A. Lacking internasals, 1 pair prefrontals ……………………………………………………….3

2B. 1 pair internasals, 1 pair prefrontals …………………………………………………………10

3A. Sunda Is. distribution; a vertebral stripe or series of connected spots ..………………………4

3B. Asian continental distribution ………………………………………………………………5

4A. 15 MDSR; Pulau We Is. N. of Sumatra; light vertebral stripe; 1 postocular …O. praefrontalis

4B. 17 MDSR; Sumatra; connected series of light vertebral spots; 2 postoculars…O. pulcherrimus

152

5A. 13 scale rows; light vertebral line, may be interrupted; no crossbars; 6 labials……O. catenata

5B. 15 scale rows …………………………………………………………………………………6

6A. No transverse markings, light and dark stripes; usually 5 supralabials …………O. hamptoni

6B. Transverse markings or paired dorsal spots present …………………………………………7

7A. Distinct stripes with inset paired dorsal spots or short bars; usually 7 labials …O. brevicauda

7B. With distinct transverse spots or crossbars; may have faded stripes …………………………8

8A. Wave shaped crossbars; 6 supralabials; usually 2 anterior temporals ………O. lungshenensis

8B. Usually 1 anterior temporal; light spots/crossbands ……………………………………………9

9A. 7 supralabials; ventrals >180, sc > 40; 11 whitish rings ..…………………………O. jintakunei

9B. 5 supralabials; ventrals < 180, sc < 40; light black-edged spots……..………………O. lacroixi

10A. Anal divided ………………………………………………………………………………11

10B. Anal entire …………………………………………………………………………………38

11A. 13 scale rows ………………………………………………………………………………12

11B. 15 or 17 scale rows …………………………………………………………………………15

12A. West of the Ganges; dark crossbars, sometimes stripes; ventrals >152 ………O. taeniolatus

12B. East of the Ganges delta ……………………………………………………………………13

13A. Shaanxi, China; ventrals 150–180; longitudinal stripes …………………O. ningshaanensis

13B. Southern Myanmar …………………………………………………………………………14

153

14A. Ventrals less then 160; 5 supralabials.; indistinct reticulations; 10 max. teeth.…O. planiceps

14B. Ventrals greater then 170; 7 supralabials.; stripes, some spots; 6 max. teeth.…O. mcdougalli

15A. 15 scale rows ………………………………………………………………………………16

15B. 17 scale rows ………………………………………………………………………………27

16A. Sunda Is. + Philippines distribution; light spots……………………………………………17

16B. Asian continental inc. Sri Lanka distribution ………………………………………………18

17A. Borneo–Sulu Arch.–Mindanao; caudals >30; small spots, larger vertebrally …O. vertebralis

17B. Sulawesi and south; caudals <30; small light spots and a red stripe on tail ……O. waandersi

18A. West of the Ganges and Sri Lanka …………………………………………………………19


19A. Sri Lanka ...…………………………………………………………………………………20

19B. Continental India ...…………………………………………………………………………22

20A. Each ventral scale with 3 brown spots 1 median, 2 lateral, forming lines .……O. sublineatus

20B. Lacking distinct ventral longitudinal lines formed by the ventral spots ...…………………21

21A. Ventrals <155; crossbars, many half width alternate sides; no hemipenis div…O. templetoni

21B. Ventrals >152; variable crossbars, may have larger spots; 2/5div. hemipenis…O. taeniolatus

154

22A. Four distinct brown stripes; ventrals <150?; hemipenis only part spinose, no div …O. nikhili

22B. Often stripes faded, with some crossbars/spots; hemipenis 2/5div., spinose .…O. taeniolatus

23A. East of the Red River (Song Hong) China; crossbars; no loreal …………………O. ornatus

23B. Myanmar, India, Thailand .…………………………………………………………………24

24A. Divided nasal; indistinct reticulations and indistinct longitudinal stripes ………O. torquatus

24B. Nasal entire or partially divided ……………………………………………………………25

25A. Partially divided nasal; dark above with faint black spots, dark gray below ……O. melaneus

25B. Nasal entire …………………………………………………………………………………26

26A. No loreal; a light (red) vertebral stripe, ~29+7 narrow dark crossbars ……O. erythrorhachis

26B. Loreal present; a light vertebral stripe, dark-edged or bordered by dark spots ...…O. dorsalis

27A. Sunda Is. or Philippines distribution ………………………………………………………28

27B. Asian continental inc. Sri Lanka distribution ………………………………………………30

28A. Philippines distribution; light vertebral line, crossbars or reticulations……………………29

28B. Sumatra endemic; 28–38 dark-bordered lighter oval spots …………..……O. pulcherrimus

29A. All the usual head markings; 14–22 crossbars, also reticulations; ventrals <175…O. ancorus

29B. No head marks; ventral >175; broken crossbars; 1st infralabials not in contact .…O. perkinsi

30A. West of the Ganges and Sri Lanka …………………………………………………………31


155

31A. Ventrals >155, angulate; caudals >39; 8–11 max. teeth; few/no ventral spots ..…O. arnensis

31B. Western Ghats; ventrals <165, not angulate; caudals <41; abundant ventral black spots.…32

32A. Head marks confluent, more dark then ground colour; 31–41 narrow crossbars……O. affinis

32B. Less then 33 crossbands or transversally arranged spots ..…………………………………33

33A. Body marks are indented bars/connected spots; hemipenes 1/3 spinose ...………O. venustus

33B. Semi-divided nasal; marks more narrow, bar-like; spinose hemipenes..……O. travancoricus

34A. China (Sichuan); divided nasal; 54–47+14–19 transverse spots/crossbars …O. multizonatum

34B. Xizang region, Myanmar, Nepal or India .…………………………………………………35

35A. 6 supralabials; entire nasal; ~20 whitish black-edged or black crossbars …O. melanozonatus

35B. Greater then 6 supralabials; distinct or faint stripes, very thin or no crossbars ……………36

36A. 7 supralabials; entire nasal; distinct light and dark stripes .…………………O. erythrogaster

36B. 8 supralabials; divided nasal; stripes, may be faint, faint reticulations/thin crossbars .……37

37A. Angulate ventrals; black spots at the base & tail tip; hemipenis 2/3 spinose ..…O. cruentatus

37B. Ventrals not angulate; hemipenes 1/2 spinose, spines uniform in size .…………O. theobaldi

38A. 13 scale rows; black-edged white crossbars, white tipped tail ..………………O. annamensis

38B. 15–21 scale rows ...…………………………………………………………………………39

156

39A. 21 or 23 scale rows ...………………………………………………………………………40

39B. 15–19 scale rows ...…………………………………………………………………………42

40A. Malay Pen. south of the isthmus of Kra and in the Sunda Is. Archipelago …O. purpurascens

40B. Distribution on mainland Asia north of Kra ………………………………………………41

41A. Brown to red with black-edged light crossbars; hemipenis unforked …………O. albocinctus

41B. Brown shades; dark reticulations/cross-blotches; hemipenis deeply forked ..…O. fasciolatus

42A. 19 scale rows ………………………………………………………………………………43

42B. 15 or 17/18 scale rows ..……………………………………………………………………49

43A. South of the isthmus of Kra and in the Sunda Is. Archipelago ..……………O. purpurascens

43B. Distribution on mainland Asia north of Kra..………………………………………………44

44A. Northeast of Song Hong; dark reticulations, more or less distinct stripes ……O. formosanus

44B. Southwest of Song Hong river distribution ...………………………………………………45

45A. Four dark longitudinal stripes, no crossmarks; 14–16 maxillary teeth .…………O. taeniatus

45B. Baring crossbars or reticulations; 12 or fewer maxillary teeth .……………………………46

46A. Brown to red with black-edged light crossbars; hemipenis unforked..………O. albocinctus

46B. Brown, darker reticulations, mesially bisected or rhomboidal spots; forked hemipenis ..…47

47A. 7 supralabials, subcaudals >53, an anterior palatine edentulous space ………O. juglandifer?

47B. 7 or 8 supralabials, subcaudals <59, no anterior palatine edentulous space .………………48

157

48A. Southern Vietnam and adjacent Cambodia and Laos, see checklist comments …O. ocellatus

48B. Eastern India and adjacent Myanmar ..……………………………………………O. cyclurus

49A. 15 scale rows .………………………………………………………………………………50

49B. 17/18 scale rows ……………………………………………………………………………60

50A. Mainland Asia distribution, 5 or 8 supralabials ……………………………………………51

50B. Sunda Is. distribution, 6 or 7 supralabials .…………………………………………………52

51A. Grey-brown, unmarked or faintly marked as cinereus; maxillary teeth 10–12 …O. inornatus

51B. Yellow-brown to brown, faint longitudinal lines; 16–18 maxillary teeth ..…O. kunmingensis

52A. 6 supralabials .………………………………………………………………………………53

52B. 7 (or 5) supralabials, third and fourth in contact with eye …………………………………54

53A. Supralabial 3 touches eye; Negros, Philippines; brown, a red vertebral stripe ….O. modestus

53B. Third and fourth supralabials touch eye; Indonesia, Sulawesi and vicinity ....…O. waandersi

54A. Subcaudals >30 (including 19+ of O. petronellae) ...………………………………………55

54B. Subcaudals <30; a light vertebral stripe on the tail ...………………………………………59

55A. Discrete yellow, white or red dots or small spots dorsally ..………………………………56

55B. Dorsally with large lighter rings or crossbands ……………………………………………57

158

56A. Dark brownish with yellow or white spots, enlarged along vertebral line .……O. vertebralis

56B. Yellow, white or red dots imbedded in three dark stripes, dots may be fine bars …O. everetti

57A. Dorsally with crossbands, anterior bands chevron shaped …….…………………O. signatus

57B. Vertebral series of black-bordered light rings, oval or rhombic, may be connected ..……58

58A. Dorsal spots more rhombic then oval, Sumatra endemic ………………………O. petronellae

58B. Dorsal spots more oval then rhombic, Borneo endemic …………………………O. annulifer

59A. Nasal partially divided …………………………………………………………O. waandersi

59B. Nasal entire; see checklist comments .…………………………………………O. propinquus

60A. Andaman Is. endemic distribution; alternating yellow and black stripes ……O. woodmasoni

60B. Any other distribution ...……………………………………………………………………61

61A. Malay Pen. south of Kra, Sunda Is., Philippines distribution ...……………………………62

61B. Mainland Asia distribution …………………………………………………………………72

62A. 6 supralabials .………………………………………………………………………………63

62B. 7 supralabials .………………………………………………………………………………65

63A. Grayish-brown with dark edged scales, No body stripes ...………………………O. unicolor

63B. Alternating pale-brown-orange and dark longitudinal stripes...……………………………64

64A. Philippines (and part of North Borneo); hemipenis without papillae …….....…O. meyerinkii

64B. Not Philippines; hemipenis with papillae .……………………………………O. octolineatus

159

65A. Philippines distribution ………………………………………………………….…………66

65B. Singapore, Malaysia, Sunda Islands ..……...………………………………………………68

66A. Ventrals >180; dark-edged red vertebral line, many dark spots/weak crossbars …O. perkinsi

66B. Ventrals <179; fewer then 30 well defined dark cross-blotches or saddles ..………………67

67A. Subcaudals >49; 2 preoculars; dark blotches large and extend to ventrals .……O. maculatus

67B. Subcaudals <48; normally 1 preocular; dark marks narrow, saddle shaped………O. ancorus

68A. No transverse marks ..………………………………………………………………………69

68B. Some faint or strong bands …………………………………………………………………71

69A. 1 postocular; Subcaudals >52; dark, red vertebral and fine light lateral lines .…O. trilineatus

69B. 2 postoculars; Subcaudals <51; lines if present fine and dark ..……………………………70

70A. Dark with yellow an red spots, enlarged along a vertebral series .……………O. bitorquatus

70B. Brown with a pair of fine dark vertebral lines ..……………………………………O. forbesi

71A. Body deep maroon red dorsally with 19–22 indistinct bands …………..………..O. booliati

71B. Body with tan-yellow oval spots …....…………………………………………….O. signatus

72A. Northeast of the Song Hong ………………………………………………………………73

72B. Southwest of Song Hong river..……………………………………………………………76

160

73A. No head markings, more or less distinct crossbars or reticulations …O. cinereus (swinhonis)

73B. A distinct head pattern, reticulations or crossbands...………………………………………74

74A. Hemipenis single; alternating very defined/indistinct crossbars .....O. cinereus(tamdaoensis)

74B. Hemipenis forked; dark reticulations or 3 reticulations alternating with bands ...…………75

75A. Widely-spaced transverse spots alternating with indistinct bands; 17 MDSR ..…O. chinensis

75B. Reticulations, usually longitudinal stripes; 19 MDSR (rarely17)..……………O. formosanus

76A. Subcaudals >71; light brown with indistinct dark reticulations ...………………O. macrurus

76B. Subcaudals <70 .……………………………………………………………………………77

77A. Predominantly striped………………………………………………………………………78

77B. Crossbars or reticulations or blotches………………………………………………………81

78A. Predominantly striped with black dorsal spots at the base and tip of the tail ……O. mouhoti

78B. No dorsal tail spots (or very small)…………………………………………………………79

79A. Vertebral and black paravertebral stripes broad, conspicuous; ventrally dark ……O. moricei

79B. Paravertebral stripes narrow; ventrally with more defined quadrangular spots ……………80

80A. Broad vertebral stripe; faint paravertebral stripes; 1 preocular/no presubocular……O. deuvei

80B. Narrow, faint vertebral stripe; dark paravertebral stripes; 2 preoculars …O. pseudotaeniatus

81A. Head lacking a distinct pattern, uniform brown or with dark reticulations ………O. cinereus

81B. Head with at least a chevron, or with a complete pattern …………………………………82

161

82A. Crossbars thin but very distinct, these bars dark or light with a dark outline ……O. cinereus

82B. Transverse marks dark, band edges irregular, reticulated, often a light vertebral line .……83

83A. Nasal entire; dorsal pattern with large blotches; no hemipenis papillae ..……O. saintgironsi

83B. Divided nasal; dorsally with small spots or thin bands and reticulations; large papillae …84

84A. Ventrals <170; dorsally, alternating reticulations and small transverse spots ……O. barroni

84B. Ventrals >171; dark brown, strong dark reticulations thickened to wavy bands …O. joynsoni

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