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Snakes of the World
Snakes of the World
A Supplement

Jeff Boundy
First edition published 2021
by CRC Press
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and by CRC Press

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© 2021 Taylor & Francis Group, LLC

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 Dedication

To Ken Williams, 1934–2017

Ken is my academic brother, not only in a titular sense by both of us receiving our doctoral
degrees from Douglas Rossman, but via our passion for maintaining a complete compendium
of literature toward a summary of all the world’s snakes, living and extinct.
Contents
Introduction.................................................................................................................................................................................. 1

Novelties��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 3

Nomenclatural Matters............................................................................................................................................................... 5

Papers Covering Museum Collections and/or Types................................................................................................................ 7

Classification................................................................................................................................................................................ 9

Classification and List of Genera............................................................................................................................................. 13

Species Accounts by Genus....................................................................................................................................................... 17

Geographical References......................................................................................................................................................... 199

Literature Cited....................................................................................................................................................................... 203

Index.......................................................................................................................................................................................... 269

vii
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Introduction

“We found only a few species missing from the book.” had forwarded to me, and I proceeded to “catch up” with new
literature. As with the Wallach, Williams, and Boundy book,
–Uetz, Davison & Ellis, 2014, from their review
Chuck Crumly offered a publishing outlet, and we agreed on
of Wallach, Williams, & Boundy, 2014.
a cutoff date of end of 2019.
That quote from Uetz et al. may be the highest acclaim that Aside from the literature document, Ken had begun sec-
we could have wished for a work documenting (almost) 3783 tions on papers dealing with valid vs. invalid names, papers
snakes with 860,000+ words. The present work is meant to discussing museum collections, species accounts for species for
be used as a companion to that first taxonomic summary of which there were additions or taxonomic changes, and a sum-
all the world’s snakes, both living and fossil, Snakes of the mary of geographical literature. To these I added a summary of
World: A Catalogue of Living and Extinct Species, by Van papers that cover snake classification at the suprageneric level,
Wallach, Ken Williams, and Jeff Boundy. Williams and and a list of genera in each family or subfamily. Because the
Wallach conceived the world snake project during the early alphabetic arrangement by genus is continued from the origi-
1980s, based on shared interests in producing a taxonomic nal book, this latter list is offered as a cross-reference for indi-
summary of snakes that was geographically and temporally viduals wishing to locate each genus in particular families. Ken
complete. Boundy, independently working on a similar sum- was creating accounts for those species for which there were
mary, joined that project in the mid-2000s, and in 2012 a taxonomic changes and/or pertinent biological or distributional
decision was made to use the end of 2012 as a cutoff date data. I have added all recognized species to provide an updated
for new material. We finalized the manuscript during 2013, list of all snake species. In comparison, up to the end of 2012
Van already having secured CRC Press as publisher through there were 651 genera and 3783 species of snakes, and at the
Chuck Crumly. The book was published in April 2014, but end of 2019 there are 685 genera and 4182 species. Within those
Ken had already been caching literature since the first days of totals, the number of extinct (fossil) genera increased from 112
2013. From his seamless efforts, Ken moved forward to work- to 138, and species increased from 274 to 314.
ing on two documents: new species accounts of novel taxa Aside from a half-dozen papers, all of the publications
with new data for existing species, and a new literature cited. cited in the literature are in JB’s possession. That fact limits
Once the “WWB” book was published, he offered that we the geographical sources section of this work due to finan-
three begin work on an addendum. Van declined to partici- cial constraints of buying the many new books that have been
pate, and I began sending Ken PDFs and literature citations recently published.
for a rapidly growing pair of manuscripts that the two of us
planned to eventually publish.
ACKNOWLEDGEMENTS
Ken’s work remained on pace through initial diagnoses and
treatments for cancers during 2014–2016, but incarcerations I thank Ken’s widow, Viola (“Vi”) Williams, for her help in
in medical facilities during much of 2017 resulted in little sorrowful times. Van Wallach, Bill Lamar, and Alex Pyron
additional work to the manuscript pair, and Ken passed away provided helpful comments and hard-to-get literature, and I
at home in November of that year. Within a month, his widow, thank everyone who sent PDFs. Lastly, I thank Peter Uetz for
Viola, blessed me with access to his computer from which creating and maintaining his Reptile Database, with which I
I retrieved the last versions, compared them to the latest he was able to compare our respective compilations.

1
Novelties

Our objective is not to introduce new names or combinations

within this work, but in the following cases there is either suf-
ficient evidence to relocate a taxon to another genus, or a need,
based on the genera and their content that we recognize, to do
so. The following represent new combinations or changes to
existing nomenclature:

Bothrops lojanus to Bothrocophias (Viperidae)

Lytorhynchus levitoni to Rhynchocalamus (Colubridae)
Rhinophis grandis vs. R. saffragamus (Uropeltidae)
Sinonatrix yapingi to Trimerodytes (Natricidae)
Trimeresurus gunaleni to Parias (Viperidae)
Trimeresurus yingjiangensis to Popeia (Viperidae)
Sinovipera sichuanensis to Viridovipera (Viperidae)

3
Nomenclatural Matters

CHARINIDAE (2013) proposed a method of screening and standardizing

nomenclatural works, targeting AJH, and H. Kaiser et al.
Quintero-A. & Shear (2016) requested the International Code (2013a) argued further for a standardized procedure for pub-
of Zoological Nomenclature to suppress the family group lishing taxonomic novelties.
name Charinidae Gray due to its homonymy with Charinidae Hoser published several rebuttals (e.g., 2015a, b) to criti-
Quintero-A. (for Charinus, an arthropod). Savage & Crother cisms, but the arguments have ceased for several years pend-
(2017) recommended modification of the family group name ing decision of the ICZN. Case 3601 remains open as of
to Charinaidae, and Pyron (2017) requested to conserve the February 2020. Rather than produce a long list of names pro-
family-group name Charinidae Gray by modifying the arthro- posed by Hoser since 2012, we here refer to the Australasian
pod version to Charinusidae. This Case, 3688, remains open. Journal of Herpetology by issue number, and provide taxa
covered in each:
CALAMARIIDAE, PAREATIDAE,
16: Acrantophis, Anilios, Anomochilus, Atropoides, Boiga,
XENODERMATIDAE
Candoia, Chrysopelea, Cylindrophis, Eryx, Furina,
Savage (2015) provides the rationale for the correct spelling of Hydrophiinae, Malayopython, Pseudonaja
the family names for the type genera Calamaria, Pareas, and 17: Calliophis, Colubroidea families, Dendrelaphis,
Xenodermus, as Calamariidae, Pareidae, and Xenodermidae. Gerrhopilus, Liophidium, Liopholidophis, Suta,
Tropidophiidae, Ungaliophiinae, Uropeltidae, Viperidae
tribes
RAYMOND HOSER PUBLICATIONS
19: Bitis, Bothrops, Causus, Crotalinae
In Wallach et al. (2014), we considered names published by 20: Atractaspis, Atractus, Boulengerina, Chamaelycus,
Raymond Hoser in his journal Australasian Journal of Biology Dendrelaphis, Drysdalia, Gongylosoma, Hapsidophrys,
to be illegitimate (nomina illegitima), and we listed them as Hemachatus, Liopeltis, Lycophidion, Mopanveldophis
such under each taxon to which they pertained. Our rationale, 21: Dasypeltis
that their naming violated the spirit of the International Code 22: Acrochordus
of Zoological Nomenclature, would be confirmed or nullified 23: Acanthophis, Aparallactus, Aspidomorphus, Cacophis,
if ruled upon by the ICZN. Hoser (2013) challenged that his Chrysopelea, Cryptophis, Lycodon, Malayopython,
journal, and therefore his names, is legitimate, and submitted Oligodon, Suta, Xerotyphlops
a request for a ruling on the matter to the ICZN (Case 3601). 30: Demansia, Liasis, Vipera
His specific case was the priority of his name Spracklandus 31: Acanthophis, Achalinus, Candoia, Denisonia,
(2009) over Afronaja Wallach, Wüster & Broadley, which was Loveridgelaps, Salomonelaps, Tropidechis, Xenodermus
published six months later. 33: Aplopeltura, Boiga, Crotalus, Emydocephalus,
Rhodin (2015), with 69 co-signees, argued that Hoser’s Hoplocephalus, Malayopython, Montivipera, Oligodon,
megalomania was initiating a dual taxonomy of names that, Typhlopidae
whether properly published or not, would be ignored by most 35: Boiga, Telescopus
researchers. They requested that the issues of Australasian 36: Bungarus, Calloselasma, Morelia
Journal of Herpetology published to that point be placed on 37: Crotalus, Elachistodon, Suta
the Official List of Rejected Works in Zoology. H. Kaiser 40: Vermicella

5
 apers Covering Museum
P
Collections and/or Types

Uetz et al. (2019) provide background from a world perspec- Flores-Villela et al. (2016, Museo de Historia Natural Alfredo
tive on type material in collections, and a link to their file of Dugès).
the type material for all extant reptile species. Ganesh & Asokan (2010, Government Museum, Chennai).
Savage & McDiarmid (2017) evaluate all of Georgio Jan’s García-Díez & González-Fernández (2013, Museo Nacional
nomina, including nomenclatural availability and type material. de Ciencias Naturales).
Vanzolini & Myers (2015) illustrate and discuss the Wied Gonzales & Montaño-F (2005, Museo de Historia Natural
type material in the American Museum of Natural History “Noel Kempff Mercado”).
collection.
Guo et al. (2012a, 2016a, Chengdu Institute of Biology).
The following works cover museum collections:
Hamdan et al. (2013, Museu de Zoologia, Universidade
Andreone & Gavetti (2010, Museo Regionale di Scienze Federal da Bahia).
Naturali di Torino).
Kaya & Özuluğ (2017, Zoology Museum, Istanbul University).
Bauer & Wahlgren (2013, Linck Collection, Naturalenkabinett
Mekinić et al. (2015, Natural History Museum in Split).
Waldenburg).
Mlíkovský et al. (2011, National Museum, Prague).
Ceríaco et al. (2014b, Museu de História Natural da
Universidade do Porto). Moskvitin & Kuranova (2006, Zoological Museum, Tomsk
State University).
Céspedez, J. & Alvarez, B.B. (2005, Universidad Nacional del
Nordeste, Corrientes, Argentina [UNNEC]) Nistri (2010, Museo di Storia Naturale, University of Florence).
Conradie et al. (2019, Port Elizabeth Museum). Prudente et al. (2019, Museu Paraense Emílio Goeldi).
Doria (2010, Museo Civico di Storia Naturale “G. Doria”). Scali (2010, Museo Civico di Storia Naturale di Milano).
Dotsenko (2003, National Museum of Natural History, Scrocchi & Kretzchmar (2017, Fundacián Miguel Lillo).
Ukrainian Academy of Sciences). Serna-Botero & Ramírez-Castaño (2017, Museo de Historia
Džukić et al. (2017, Institute for Biological Research “Siniša Natural de la Universidad de Caldas).
Stanković,” University of Belgrade). Y. Werner & Shacham (2010, Hebrew University of Jerusalem).
Ferraro & Williams (2006, Museo de La Plata).

7
Classification

The following papers discuss snake classification in a broad Weinell & Brown (2018) use DNA sequence data to produce
phylogenetic sense, and/or at the suprageneric level. a phylogeny of the Lamprophiidae that resolves Philippine
taxa (Cyclocorus, Hologerrhum, Myersophis, Oxyrhabdium)
into a subfamily-level clade that they name Cyclocorinae.
Broad/Basal
Buhoma remains as the sister taxon to the remainder of the
Conrad (2008) presents alternative phylogenies and morpho- family aside from the Prosymninae. Micrelaps resolves as
logical diagnoses of the Squamata, including Serpentes, from the sister taxon to Psammodynastes plus the Lamprophiinae,
the Ophidia basal node through to Macrostomata. and Aparallactus forms a clade with Atractaspis and
Wiens et al. (2012) produce a phylogeny using 44 nuclear Homoroselaps to the exclusion of other Aparallactinae. Due
genes and representatives of then-recognized families, except to this latter arrangement, the authors suggest including the
Xenophidiidae. They find that the Scolecophidia are not Aparallactinae within the Atractaspidinae.
monophyletic. Garberoglio et al. (2019) produce a morphology-based phy-
Pyron et al. (2013) produce a phylogeny using numerous genes logeny that emphasizes relationships of certain pre-Serpentes
of 1262 snake species, establishing a new and subsequently snake taxa.
consistent arrangement of snake families and subfamilies. R.O. Goméz et al. (2019) produce a skeletal-based phylogeny
Palci et al. (2013b) use new osteological data to place hind- of early fossil snakes, focusing on the Madtsoiidae.
limbed Cretaceous snakes in a phylogeny in context with
other stem-group snakes. Madtsoiidae
Scanferla et al. (2013) produce a phylogeny of families, and
Rio & Mannion (2018) produce a phylogeny of the Madtsoiidae
primitive and/or monotypic genera to determine placement of
based on vertebral characters.
†Kataria.
K.T. Smith (2013) discusses classification and familial assign-
ments of some fossil taxa as follows: (1) †Eoanilius europae “Scolecophidia”
is probably a booid; (2) †Calamagras weigeli belongs to the Hedges et al. (2014) construct a molecular-based phylogeny
Ungaliophiinae; (3) †Paraplatyspondylia batesi is prob- of Typhlopidae, from which they erect four subfamilies, and
ably a species of †Platyspondylia; (4) †Ogmophis compactus new genera. They include data for 92 named and 60 yet-to-be
belongs to the Loxocemidae. described species.
Hsiang et al. (2015) discuss snake origins, and present a Pyron & Wallach (2014) produce a DNA sequence-based phy-
revised phylogeny. logeny of the Typhlopidae using 95 species, which conform to
Reeder et al. (2015) use morphological and molecular data to four subfamilies, and a revised arrangement of genera. Their
produce a phylogeny of Squamata that places snakes as sis- taxonomic conclusions are backed by data from internal and
ter taxon to the Mosasauria. Anomalepidiae are paraphyletic external morphology of nearly all recognized species.
with respect to Typhlopidae and Leptotyphlopidae, making Nagy et al. (2015) add genes and taxa to the gene matri-
the Scolecophidia non-monophyletic. These families are sis- ces studied by Hedges et al. (2014) and Pyron & Wallach
ters to remaining snakes, followed by Dinilysia and Najash, (2014), concluding that Antillotyphlops, Cubatyphlops,
with remaining snakes forming two clades. and Sundatyphlops are diagnosable, and that all native
Figueroa et al. (2016) present a ten gene-phylogeny using 1652 Madagascar species are assignable to Madatyphlops.
snake species, resulting in a modified resolution of suprage- Miralles et al. (2018) use 14 nuclear genes to examine relation-
neric snake classification. ships of the Scolecophidia. The resulting phylogeny reveals
Streicher & Wiens (2016) use 1.4 million base pairs from the Scolecophidia to be paraphyletic, with anomalepids
nuclear genes to produce a phylogeny using one or two rep- Liotyphlops and Typhlophis as sister taxa to the Alethinophidia.
resentatives from snake families. The phylogenies reveal
(1) a non-monophyletic Scolecophidia, (2) an association
Uropeltidae
between Aniliidae and Tropidophiidae, (3) and a clade
containing Casarea, Loxocemus, and Xenopeltis with the Pyron et al. (2016c) provide a revised phylogeny and a taxo-
Pythonidae. nomic summary of the Uropeltidae. Pseudotyphlops and
Zheng & Wiens (2016) produce a combination phylogeny Sri Lankan Uropeltis are included in Rhinophis.
based on numerous genes and taxa that is relatively similar to Cyriac & Kodandaramaiah (2017) use DNA sequence data to
the one produced by Wiens et al. (2012). produce a phylogeny of 32 described plus some undescribed
9
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