Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
TURTLES OF THE WORLD
Annotated Checklist and Atlas of Taxonomy, Synonymy,
Distribution, and Conservation Status (8th Ed.)
TuRTle TaxonoMy WoRking gRoup
andeRs g.J. Rhodin, John b. iveRson, RogeR bouR, uWe FRiTz,
aRThuR geoRges, h. bRadley shaFFeR, and peTeR paul van diJk
Chelonian ReseaRCh MonogRaphs, nuMbeR 7
Published by
Chelonian Research Foundation and Turtle Conservancy
in association with
IUCN/SSC Tortoise and Freshwater Turtle Specialist Group, Turtle Conservation Fund, Global Wildlife Conservation,
Conservation International, and International Union for Conservation of Nature / Species Survival Commission
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
CHELONIAN RESEARCH MONOGRAPHS
Contributions in Turtle and Tortoise Research
Series Editor
andeRs g.J. Rhodin
Chelonian Research Foundation, 168 Goodrich Street, Lunenburg, Massachusetts 01462 USA, and
Turtle Conservancy, 49 Bleecker Street, New York, New York 10012 USA [RhodinCRF@aol.com]
CHELONIAN RESEARCH MONOGRAPHS (ISSN 1088-7105) is an international peer-reviewed scientific publication series for monograph-length manuscripts, collected proceedings of symposia, edited compilations, and other longer
turtle-related research documents. The series accepts contributions dealing with any aspects of chelonian research, with
a preference for conservation or biology. Manuscripts or edited compilations dealing with conservation biology, systematic relationships, chelonian diversity, geographic distribution, natural history, ecology, reproduction, morphology and
natural variation, population status, husbandry, and human exploitation or conservation management issues are of special
interest. Bibliographic and other reference materials are also of interest. Submit proposals for publications directly to
A.G.J. Rhodin at the e-mail address above. Instructions for manuscript submission and detailed guidelines for authors are
available at the same address.The series is published on an occasional basis, from 1996 to 2016 by Chelonian Research
Foundation, and from 2017 on by Chelonian Research Foundation and Turtle Conservancy.
Published Issues in Series
1. The Galápagos Tortoises: Nomenclatural and Survival Status. 1996. By Peter C.H. PritCHard. 85 pp. ISBN: 0-96535400-8 (hard cover); 0-9653540-1-6 (soft cover).
2. Asian Turtle Trade: Proceedings of a Workshop on Conservation and Trade of Freshwater Turtles and Tortoises in
Asia. 2000. Edited by Peter Paul van dijk, Bryan l. Stuart, and anderS G.j. rHodin. 164 pp. ISBN: 0-9653540-2-4
(hard cover); 0-9653540-3-2 (soft cover).
3. Biology and Conservation of Florida Turtles. 2006. Edited by Peter a. Meylan. 376 pp. ISBN: 0-9653540-4-0 (hard
cover); 0-9653540-5-9 (soft cover).
4. Defining Turtle Diversity: Proceedings of a Workshop on Genetics, Ethics, and Taxonomy of Freshwater Turtles and
Tortoises. 2007. Edited by H. Bradley SHaffer, nanCy n. fitzSiMMonS, artHur GeorGeS, and anderS G.j. rHodin.
200 pp. ISBN 0-9653540-7-5 (hard cover); 0-9653540-8-3 (soft cover).
5. Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and
Freshwater Turtle Specialist Group. 2008–2016. Edited by anderS G.j. rHodin, Peter C.H. PritCHard, Peter Paul
van dijk, rayMond a. SauMure, kurt a. BuHlMann, joHn B. iverSon, and ruSSell a. MitterMeier. Installments 1–7,
1138 pp. to date. ISBN 0-9653540-9-1 (looseleaf).
6. Turtles on the Brink in Madagascar: Proceedings of Two Workshops on the Status, Conservation, and Biology of
Malagasy Tortoises and Freshwater Turtles. 2013. Edited by CHriStina M. CaStellano, anderS G.j. rHodin, MiCHael
oGle, ruSSell a. MitterMeier, Herilala randriaMaHazo, riCk HudSon, and riCHard e. lewiS. 184 pp. ISBN: 9780-9910368-0-6 (hardback), 978-0-9910368-1-3 (paperback).
7. Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status
(8th Ed.). 2017. turtle taxonoMy workinG GrouP [anderS G.j. rHodin, joHn B. iverSon, roGer Bour, uwe fritz,
artHur GeorGeS, H. Bradley SHaffer, and Peter Paul van dijk]. 292 pp. ISBN: 978-1-5323-5026-9 (online).
CHELONIAN RESEARCH MONOGRAPHS are available for purchase from Chelonian Research Foundation and Turtle Conservancy.
No overall series subscription rate is available, as individual monograph issues are priced separately. Contact either Turtle Conservancy
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Copyright © 2017 by Chelonian Research Foundation and Turtle Conservancy
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
TURTLES OF THE WORLD
Annotated Checklist and Atlas of Taxonomy, Synonymy,
Distribution, and Conservation Status (8th Ed.)
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
COVER AND PREFACE ILLUSTRATIONS
Front Cover: Left: Goode’s Thornscrub Tortoise, Gopherus evgoodei (Testudinidae), from Monte Mojino Reserve, Alamos,
Sonora, Mexico; photo by Eric V. Goode. Right: Ploughshare Tortoise, Astrochelys yniphora (Testudinidae), from Baly
Bay National Park, Madagascar; photo by Anders G.J. Rhodin.
Title Page: Radiated Tortoise, Astrochelys radiata (Testudinidae), from Madagascar; drawing by Stephen D. Nash.
Table of Contents: Left: Red-headed Amazon River Turtle, Podocnemis erythrocephala (Podocnemididae), from Rio Jufari,
Amazonas, Brazil; photo by Richard C. Vogt. Right: Euphrates Softshell Turtle, Rafetus euphraticus (Trionychidae), from
Balarood R., Khuzestan Prov., Iran; photo by Hanyeh Ghaffari.
Frontispiece: Left: Wood Turtle, Glyptemys insculpta (Emydidae), from Lunenburg, Worcester Co., Massachusetts, USA;
photo by Anders G.J. Rhodin. Right: Black-breasted Leaf Turtle, Geoemyda spengleri (Geoemydidae), from Tam Dao
National Park, Vietnam; photo by Flora Ihlow.
Back Cover: Top Left: Northern Snake-necked Turtle, Chelodina (Macrochelodina) rugosa (Chelidae), from Morehead
R., Western Prov., Papua New Guinea; photo by Arthur Georges. Top Right: Burmese Big-headed Turtle, Platysternon
megacephalum peguense (Platysternidae), from Phu Luang Wildlife Sanctuary, Loei Prov., Thailand; photo by Peter Paul van
Dijk. Bottom Left: Flatback Sea Turtle, Natator depressus (Cheloniidae), from Shoalwater Bay, Queensland, Australia;
photo by Carmen Pilcher. Bottom Right: Northern Giant Musk Turtle, Staurotypus triporcatus (Kinosternidae), from
Central Farms, Cayo Dist., Belize; photo by John B. Iverson.
RECOMMENDED CITATION
turtle taxonoMy workinG GrouP [rHodin, a.G.j., iverSon, j.B., Bour, r. fritz, u., GeorGeS, a., SHaffer, H.B., and van dijk,
P.P.]. 2017. Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status
(8th Ed.). In: Rhodin, A.G.J., Iverson, J.B., van Dijk, P.P., Saumure, R.A., Buhlmann, K.A., Pritchard, P.C.H., and Mittermeier, R.A.
(Eds.). Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater
Turtle Specialist Group. Chelonian Research Monographs 7:1–292. doi: 10.3854/crm.7.checklist.atlas.v8.2017.
CHelonian reSearCH MonoGraPHS. Series Edited by anderS G.j. rHodin.
voluMe nuMBer 7. Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation
Status (8th Ed.). In: Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC
Tortoise and Freshwater Turtle Specialist Group.
Published online 2 August 2017.
ISSN (monograph series): 1088-7105
ISBN (this volume): 978-1-5323-5026-9 (online)
Co-Published by Chelonian Research Foundation and Turtle Conservancy.
Copyright © 2017 by Chelonian Research Foundation, Lunenburg, Massachusetts, USA and Turtle Conservancy, New
York, New York, USA.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
TURTLES OF THE WORLD
Annotated Checklist and Atlas of Taxonomy, Synonymy,
Distribution, and Conservation Status (8th Ed.)
TuRTle TaxonoMy WoRking gRoup
andeRs g.J. Rhodin, John b. iveRson, RogeR bouR, uWe FRiTz,
aRThuR geoRges, h. bRadley shaFFeR, and peTeR paul van diJk
suppleMenT no. 1 oF Chelonian ReseaRCh MonogRaphs no. 5:
Conservation Biology of Freshwater Turtles and Tortoises:
A Compilation Project of the IUCN/SSC
Tortoise and Freshwater Turtle Specialist Group
ediTed by
andeRs g.J. Rhodin1,2, John b. iveRson3, peTeR paul van diJk2,4, RayMond a. sauMuRe5,
kuRT a. buhlMann6, peTeR C.h. pRiTChaRd7, and Russell a. MiTTeRMeieR8
1
Chelonian Research Foundation, Lunenburg, Massachusetts, USA; 2Turtle Conservancy, New York, New York, USA;
3
Earlham College, Richmond, Indiana, USA; 4Global Wildlife Conservation, Austin, Texas, USA;
5
Southern Nevada Water Authority, Las Vegas, Nevada, USA; 6University of Georgia, Aiken, South Carolina, USA;
7
Chelonian Research Institute, Oviedo, Florida, USA; 8Conservation International, Arlington, Virginia, USA
Chelonian ReseaRCh MonogRaphs
Number 7
2017
Chelonian Research Foundation and Turtle Conservancy
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
TURTLES OF THE WORLD
Annotated Checklist and Atlas of Taxonomy, Synonymy,
Distribution, and Conservation Status (8th Ed.)
Table oF ConTenTs
Abstract
...........................................................9
Introduction ...........................................................9
Methodology .......................................................10
TTWG Guidelines for Taxonomic Changes ....13
Taxonomic Changes ......................................14
Distributions..................................................15
GIS Maps ......................................................16
Conservation Status ......................................18
Results and Discussion ........................................19
Conservation Status ......................................19
Genetic Pollution ..........................................19
Request for Updates ............................................19
Acknowledgments ...............................................20
Photos and Photographers ..................................21
Checklist
.........................................................23
Linnaean Classification Content ..........................23
Alternative Phylocode Classification ..................23
Testudines
.........................................................24
Cryptodira .........................................................24
Chelydridae ............................................24
Chelydra ....................................24
Macrochelys ................................26
Chelonioidea .................................................27
Cheloniidae.............................................27
Carettinae .........................................27
Caretta ........................................27
Eretmochelys ...............................28
Lepidochelys ...............................29
Cheloniinae ......................................30
Chelonia ......................................30
Natator ........................................32
Dermochelyidae .....................................32
Dermochelys ...............................32
Kinosternoidea ..............................................34
Dermatemydidae ....................................34
Dermatemys ................................34
Kinosternidae .........................................34
Kinosterninae ...................................34
Kinosternon.................................34
Kinosternon or Cryptochelys ......35
Sternotherus ................................45
Staurotypinae or Staurotypidae........47
Claudius ......................................47
Staurotypus .................................48
Testudinoidea ................................................49
Emydidae ................................................49
Deirochelyinae .................................49
Chrysemys ...................................49
Deirochelys .................................51
Graptemys ...................................51
Malaclemys .................................57
Pseudemys...................................59
Trachemys ...................................62
Emydinae .........................................71
Clemmys ......................................71
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Emys ........................................ 72
Emys or Actinemys .................. 75
Emys or Emydoidea ................ 76
Glyptemys ............................... 77
Terrapene ................................ 77
Platysternidae ..................................... 82
Platysternon ............................ 82
Geoemydidae ...................................... 83
Geoemydinae ............................... 83
Batagur ....................................83
Cuora .......................................86
Cyclemys ..................................93
Geoclemys ................................96
Geoemyda ................................96
Hardella ...................................97
Heosemys .................................98
Leucocephalon .........................99
Malayemys .............................100
Mauremys...............................101
Melanochelys .........................107
Morenia ..................................109
Notochelys..............................109
Orlitia ....................................110
Pangshura ..............................110
Sacalia ...................................113
Siebenrockiella.......................114
Vijayachelys ...........................114
Rhinoclemmydinae .....................115
Rhinoclemmys ........................115
Testudinidae.......................................120
Aldabrachelys ........................120
Astrochelys .............................121
Centrochelys ..........................123
Chelonoidis ............................123
Chersina .................................132
Chersobius .............................132
Cylindraspis ...........................134
Geochelone ............................136
Gopherus................................137
Homopus ................................140
Indotestudo ............................140
Kinixys ...................................141
Malacochersus .......................145
Manouria ...............................145
Psammobates .........................146
Pyxis .......................................149
Stigmochelys ..........................150
Testudo ...................................151
(Testudo) ...........................151
(Agrionemys).....................155
(Chersine) .........................157
Trionychoidea ..........................................158
Carettochelyidae ................................158
Carettochelys .........................158
Trionychidae ......................................158
Cyclanorbinae .............................158
Cyclanorbis ............................158
Cycloderma ............................159
Lissemys .................................160
Trionychinae ...............................162
Amyda ....................................162
Apalone ..................................164
Chitra .....................................168
Dogania .................................169
Nilssonia ................................169
Palea ......................................171
Pelochelys ..............................172
Pelodiscus ..............................173
Rafetus ...................................175
Trionyx ...................................176
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Pleurodira ....................................................177
Chelidae .............................................177
Chelinae ......................................177
Acanthochelys ........................178
Chelus ....................................179
Mesoclemmys .........................180
Phrynops ................................183
Platemys .................................185
Rhinemys ................................186
Hydromedusinae .........................186
Hydromedusa .........................186
Chelodininae ...............................187
Chelodina ...............................187
(Chelodina) .......................187
(Macrochelodina) .............191
(Macrodiremys) .................193
Elseya .....................................193
(Elseya) .............................193
(Hanwarachelys)...............194
(Pelocomastes) ..................195
Elusor .....................................196
Emydura .................................197
Myuchelys ..............................200
Rheodytes ...............................201
Pseudemydurinae ........................202
Pseudemydura........................202
Pelomedusidae ...................................202
Pelomedusa ............................202
Pelusios ..................................206
Podocnemididae ................................214
Erymnochelys .........................214
Peltocephalus .........................214
Podocnemis ............................214
Regional Species Richness Maps .......................218
Annotations
....................................................221
Current 2017 Checklist ...................................221
Previous Checklist Annotations......................234
2007 Annotations .....................................234
2008 Annotations .....................................238
2009 Annotations .....................................239
2010 Annotations .....................................242
2011 Annotations .....................................246
2012 Annotations .....................................247
2014 Annotations .....................................251
Appendix – Distributional Data.........................256
Distribution Updates 2011 ..............................256
Distribution Updates 2012..............................256
Distribution Updates 2014..............................257
Literature Cited...................................................257
CRM 5 and CRM 7 Citations ............................290
TTWG, CBFTT, and TEWG Checklists ........290
CBFTT Species Accounts...............................290
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Conservation Biology of Freshwater Turtles and Tortoises:
of the
World:
Checklist
and AtlasTurtle
(8th Specialist
Ed.) – 2017
ATurtles
Compilation
Project
of theAnnotated
IUCN/SSC Tortoise
and Freshwater
Group
A.G.J. Rhodin, J.B. Iverson, P.P. van Dijk, R.A. Saumure, K.A. Buhlmann, P.C.H. Pritchard, and R.A. Mittermeier, Eds.
Chelonian Research Monographs (ISSN 1088-7105) No. 7, doi:10.3854/crm.7.checklist.atlas.v8.2017
© 2017 by Chelonian Research Foundation and Turtle Conservancy • Published 2 August 2017
Turtles of the World: Annotated Checklist and Atlas of Taxonomy,
Synonymy, Distribution, and Conservation Status (8th Ed.)
TuRTle TaxonoMy WoRking gRoup*
*Authorship of this article is by this working group of the IUCN SSC Tortoise and Freshwater Turtle Specialist Group,
which for the purposes of this document consisted of the following contributors:
andeRs g.J. Rhodin1,2, John b. iveRson3, RogeR bouR4, uWe FRiTz5,
aRThuR geoRges6, h. bRadley shaFFeR7, and peTeR paul van diJk2,8
1
Chairman Emeritus, IUCN SSC Tortoise and Freshwater Turtle Specialist Group,
Chelonian Research Foundation, 168 Goodrich St., Lunenburg, Massachusetts 01462 USA [rhodincrf@aol.com];
2
Turtle Conservancy, 49 Bleecker St., New York, New York 10012 USA;
3
Department of Biology, Earlham College, Richmond, Indiana 47374 USA [johni@earlham.edu];
4
Laboratoire des Reptiles et Amphibiens, Muséum National d’Histoire Naturelle, 75005 Paris, France [bour.roger@gmail.com];
5
Museum für Tierkunde, Senckenberg Dresden, A.B. Meyer Building, 01109 Dresden, Germany [uwe.fritz@senckenberg.de];
6
Institute for Applied Ecology, University of Canberra, ACT 2601, Australia [georges@aerg.canberra.edu.au];
7
Department of Ecology and Evolutionary Biology and La Kretz Center for California Conservation Science,
Institute of the Environment and Sustainability, University of California, Los Angeles, California 90095 USA [brad.shaffer@ucla.edu];
8
Deputy Chair, IUCN SSC Tortoise and Freshwater Turtle Specialist Group,
Global Wildlife Conservation, PO Box 129, Austin, Texas 78767 USA [ppvandijk@globalwildlife.org]
a bsTRaCT. – This is our 8th edition of an annotated checklist of all recognized and named
taxa of the world’s modern chelonian fauna, documenting recent changes and controversies
in nomenclature through early 2017, and including all primary synonyms, updated from 7
previous checklists (Turtle Taxonomy Working Group 2007b, 2009, 2010, 2011, 2012, 2014;
Rhodin et al. 2008). We provide an updated comprehensive listing of taxonomy, names, and
conservation status of all turtles and tortoises of the world, including detailed distribution
maps. We strive to record the most recent justified taxonomic assignment of taxa in a hierarchical framework, providing annotations, including alternative possible arrangements, for
some proposed changes. We provide common English names and detailed distributional data
for all taxa, listing occurrence by countries and many smaller political or geographic subunits
(states or regions), including indications of native, extirpated, and introduced (modern or
prehistoric) populations. We include current published and draft IUCN Red List status assessments for all turtles, as well as CITES listings. The diversity of turtles and tortoises in the
world that has existed in modern times (since 1500 AD) and currently generally recognized
as distinct and included in this checklist, now consists of 356 species. Of these, 60 are polytypic, representing 122 additional recognized subspecies, or 478 total taxa of modern turtles
and tortoises. Of these, 7 species and 3 subspecies, or 10 taxa (2.1%), have gone extinct. As
of the current IUCN 2017 Red List, 148 turtle species (60.4% of 245 species listed, 41.6% of
all 356 recognized modern species) are officially regarded as globally Threatened (Critically
Endangered [CR], Endangered [EN], or Vulnerable [VU]). We record additional draft Red
List assessments by the IUCN Tortoise and Freshwater Turtle Specialist Group (TFTSG)
of previously “unevaluated” species, and updated draft re-assessments of previously listed
species, allowing us to evaluate the overall current threat levels for all turtles and tortoises.
Of the 356 total species of turtles and tortoises, 114 (32.0%) are CR or EN, 179 (50.3%) are
Threatened (CR, EN, or VU), and 186 (52.2%) are Threatened or Extinct. If we provisionally
adjust for predicted threat rates of Data Deficient and Not Evaluated species, then ca. 59%
of all extant turtles are Threatened. These numbers and percentages of Threatened species
have increased since our last checklist. Turtles are among the most threatened of the major
groups of vertebrates, in general more than birds, mammals, cartilaginous or bony fishes,
or amphibians.
key W oRds. – Reptilia, Testudines, turtle, tortoise, chelonian, taxonomy, nomenclature, genera, species, subspecies, primary synonyms, suprageneric hierarchy, systematics, common
names, distribution, maps, introduced species, conservation status, IUCN Red List, CITES,
threatened species, extinction
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
The diversity of all turtles and tortoises (chelonians)
in the world that has existed in modern times (since 1500
AD), and currently generally recognized as distinct by
specialists in turtle taxonomy and systematics, consists
of approximately 356 species, of which 60 are polytypic,
with 122 additional recognized subspecies, or 478 total
taxa of modern chelonians. Of these, 8 species plus 3
subspecies, or 11 total taxa, of tortoises and freshwater
turtles have become extinct since 1500 AD (see Table 1),
leaving us currently with 348 species and 119 additional
subspecies, or 467 total taxa of living turtles and tortoises. Of all living turtle taxa, 7 species are marine turtles,
leaving 341 species and 460 total taxa of modern living
freshwater and terrestrial turtles and tortoises.
In this checklist we present a full taxonomic listing of
all recognized modern turtle and tortoise taxa, including
synonymized names and type localities, detailed distribution maps, and annotations concerning recently described
new taxa, nomenclatural and taxonomic updates, and significant taxon-related controversies or developments.
The 478 modern turtle and tortoise taxa we recognize
here are based on a synonymy of 1473 separate named
turtle and tortoise species and subspecies, including all
primary description names, secondary nomen novum replacement names, undescribed nomen nudum names, and
other nomenclaturally unavailable names. These names
also include those fossil taxa that have been synonymized
with modern taxa.
We also recognize 1 order, 2 suborders, 4 superfamilies, 14 families, 13 subfamilies, 94 genera (plus 5 potentially separate genera), and 6 subgenera of modern turtles, for a potential total of 139 supraspecific groupings.
These groups are based on 457 valid and synonymized
names, for a total listing here of 1930 taxonomic names
applied to all modern turtle taxa and groups.
As there is always some disagreement among experts
as to which taxa are distinct and valid, and at what systematic level or rank (species or subspecies), these numbers
are variable depending on the authorities presenting their
data or interpretations. For prior discussions and listings
of all recognized modern turtle taxa, with extensive annotations regarding areas of recent taxonomic change, instability, or controversy, see the previous publications by the
Turtle Taxonomy Working Group (TTWG 2007a,b, 2009,
2010, 2011, 2012, 2014), Rhodin et al. (2008), and the
turtle checklist produced for CITES by Fritz and Havaš
(2007). For a listing of all extinct Pleistocene and Holocene turtle and tortoise taxa, see our companion checklist
by the Turtle Extinctions Working Group (TEWG 2015).
METHODOLOGY
The Turtle Taxonomy Working Group (TTWG) functions under the auspices of the IUCN SSC Tortoise and
Freshwater Turtle Specialist Group (TFTSG), which
operates under the umbrella of the IUCN (International Union for Conservation of Nature) and its Species
Survival Commission (SSC). We first compiled our
checklist of modern turtle taxa in 2007 (TTWG 2007b),
and have previously updated it annually to reflect more
recent changes, as required by subsequent publications
with taxonomic novelties or proposed changes, as well
as adding primary synonyms for all recognized taxa, type
species and type locality designations, as well as distribution maps (Rhodin et al. 2008; TTWG 2009, 2010, 2011,
2012, 2014). This present checklist has taken three years
to update and produce because of extensive further expansion in content, and is now the 8th installment in this
series. It is current through approximately July 2017.
We list all primary and synonymized description
names, as well as all nomina nova and nomina nuda
names of which we are aware. We added nomen nudum
names in the previous checklist. We continue to exclude
most obvious ex errore names, especially the profusion of
recent egregious misspellings in modern literature (especially in the popular literature and other non-systematic
biological sciences). In addition, we do not list variations
in spelling of the two alternate patronymic endings (-ii
vs. -i), always using the original valid orthography.
Our listing of nomen novum names takes a broadly encompassing approach and lists both justified and unjustified subsequent emendations, including substantial name
changes caused by early writers’ occasional tendencies
to create new or “better” names that they felt were more
appropriate or more correct. Many early names were also
unjustifiably emended in order to try to comply with perceived rules about word constructions and the use of nonGreek vs. Greek letters, (e.g., c vs. k, as in Cinosternon
vs. Kinosternon, Cinixys vs. Kinixys). Occasionally, early
authors did not appear to remember what the previously
used names were, and simply came up with new spelling
variations, with these new names sometimes becoming
established in the literature for a while. This was especially true for the many names and spelling emendations
created and recorded by John Edward Gray between 1825
and 1874. Prior to the establishment of the International
Commission on Zoological Nomenclature in 1895, and
the publication of the first edition of the Code of Zoological Nomenclature in 1905, these kinds of new names
and changes were fairly common and we do not consider
them to be simple ex errore typographical errors, and
therefore, we have instead recorded many of them as nomina nova.
We include listings of subsequent new combination
names to reflect how taxa have been rearranged into new
genera or different specific or subspecific levels. The new
combination names are listed in lighter gray text following each associated primary name, arranged more or
less chronologically from oldest to most recently created
combinations, but without attributing authorship or date
of first use of the new combination. We have attempted to
list all known subsequent combination names, but these
listings may be incomplete. A few older ex errore misspelled names are included in these listings.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Table 1. Modern named freshwater turtles and tortoises that have
gone extinct since 1500 AD (7 species, 3 subspecies, 10 taxa),
with approximate or known extinction dates. For species that went
extinct during Holocene and Pleistocene times prior to 1500 AD,
see separate supplementary checklist and review (TEWG 2015)
and Fig. 1.
Kinosternidae
Kinosternon hirtipes megacephalum
Viesca Mud Turtle
Mexico (Coahuila); ca. 1970
Testudinidae
Aldabrachelys gigantea daudinii
Daudin’s Giant Tortoise
Seychelles (Mahé?); ca. 1850
Chelonoidis abingdonii
Pinta Giant Tortoise, Abingdon Island Giant Tortoise
Ecuador (Galápagos: Pinta [Abingdon]); 24 June 2012
Chelonoidis niger
Floreana Giant Tortoise, Charles Island Giant Tortoise
Ecuador (Galápagos: Floreana [Charles]); ca. 1850
Cylindraspis indica
Reunion Giant Tortoise
Réunion; ca. 1840
Cylindraspis inepta
Mauritius Giant Domed Tortoise
Mauritius (Mauritius); ca. 1735
Cylindraspis peltastes
Rodrigues Domed Tortoise
Mauritius (Rodrigues); ca. 1800
Cylindraspis triserrata
Mauritius Giant Flat-shelled Tortoise
Mauritius (Mauritius); ca. 1735
Cylindraspis vosmaeri
Rodrigues Giant Saddleback Tortoise
Mauritius (Rodrigues); ca. 1800
Pelomedusidae
Pelusios castaneus seychellensis
Seychelles Mud Turtle
Seychelles (Mahé); ca. 1950
Figure 1. While beyond the time-frame of extinctions for modern
turtles, it is worth noting the findings of White et al. (2010), who
documented the continued existence nearly into modern times of
an extraordinary giant tortoise, apparently of the extinct terrestrial horned family Meiolaniidae, on Efate Island, Vanuatu, in the
southwestern Pacific Ocean. The species persisted until as recently
as only 3100–2800 ybp (1150–850 BC) with a further calibrated
age of 2890–2760 ybp (940–810 BC) [see TEWG 2015]. White
et al. named it ?Meiolania damelipi and provided clear evidence
of human butchering and consumption of the species, further corroborated and expanded by Hawkins et al. (2016). This exploitation represented the final anthropogenic extinction event for this
spectacularly unique and evolutionarily distinct deep lineage of
giant terrestrial chelonians. For a more complete analysis of turtle
extinctions caused by humans and/or climate change during the
Pleistocene and Holocene, see our companion publication by the
Turtle Extinctions Working Group (2015). No complete skeletons
of ?M. damelipi are known, but the related Meiolania platyceps
from Lord Howe Island, Australia, which went extinct in the Late
Pleistocene, has been beautifully reconstructed (photo above) by
the American Museum of Natural History (Burke et al. 1983).
Original and synonymized taxon names (including
higher-category names) are listed using their original
spelling and genus-species combination as used by the
author at the time of first publication of the name. Our
synonymies for genus- and species-level taxa follow, to
our best efforts, the strict and established nomenclatural
rules established by the fourth edition of the International
Code of Zoological Nomenclature (ICZN 1999).
However, for the higher-level suprageneric categories used in this checklist, we have also provided some
synonyms and previously-used names for the same or included groupings whose usage may not necessarily correspond to nomenclatural guidelines under the ICZN. Since
the ICZN does not regulate names above the superfamily
rank, our listings of these names are intended to document historical use to aid understanding and resolving the
difficult questions of what names are most appropriately
used for these suprageneric categories and to what author
they should be attributed.
For example, the names we list under the Order-level
name for turtles (Testudines) are not all strict synonyms,
as some were proposed at different levels of groupings,
from “Family” to Order to various supra-ordinal categories. Many were utilized primarily for including various
fossil turtle-like ancestors in an expanded concept of
turtles, including some rank-free Phylocode names. The
names we list in other infra-ordinal suprageneric categories are not always strict synonyms either, as based on
nomenclatural acceptability or availability of the utilized
group name, but instead provide a partial historical record for names previously used for the same or similar
grouping.
Our checklist includes all currently recognized
named taxa (species and subspecies) of modern turtles
(extant after 1500 AD). By “currently recognized” we
mean those taxa that have not been demonstrably refuted or justifiably synonymized in published literature,
or whose description or recommended resurrection has
yet to receive wide community acceptance. We have attempted to describe all recent published taxonomic recommendations in our annotations, even though we have
not included all proposed changes in the checklist.
Since there are sometimes also different interpretations for some genera and polytypic species as to which
11
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Table 2. The top turtle-rich countries for all turtle species and taxa
(species and subspecies) per country, including tortoises, freshwater
and terrestrial turtles, and nesting sea turtles.
Species
1. USA, 62
2. Mexico, 49
3. Brazil, 36
4. China, 35
Ecuador, 35
6. Indonesia, 34
7. Colombia, 33
India, 33
9. Australia, 32
Vietnam, 32
11. Bangladesh, 30
Myanmar, 30
Thailand, 30
14. Malaysia, 24
15. Venezuela, 23
16. South Africa, 21
17. Congo (DRC), 19
18. Laos, 18
19. Guatemala, 16
Honduras, 16
Mozambique, 16
Nepal, 16
Papua New Guinea, 16
Tanzania, 16
25. Angola, 15
Cameroon, 15
Kenya, 15
Panama, 15
Peru, 15
Taxa (sp. & ssp.)
1. USA, 89
2. Mexico, 65
3. India, 41
4. Indonesia, 38
5. China, 37
6. Australia, 36
Brazil, 36
Vietnam, 36
9. Ecuador, 35
10. Colombia, 34
11. Thailand, 33
12. Myanmar, 31
13. Bangladesh, 30
14. Malaysia, 25
Venezuela, 25
16. South Africa, 23
17. Laos, 19
18. Congo (DRC), 18
Nepal, 18
20. Honduras, 17
Tanzania, 17
22. Guatemala, 16
Mozambique, 16
Panama, 16
Papua New Guinea, 16
Peru, 16
27. Angola, 15
Cameroon, 15
Costa Rica, 15
Nicaragua, 15
names are valid and whether to lump or split the contained taxa, we occasionally list recent alternative name
usages as ‘Xxxx or Yyyy’. Our most important criterion
for accepting proposed changes is that they be accompanied by adequate data and sound arguments justifying
the taxonomic revision. Consequently, some proposed
taxonomic changes from lists or publications with unsupported or untested revisions have not been incorporated. We also list and annotate recent systematic papers that do not necessarily commit nomenclatural or
taxonomic acts, but that present data on phylogenetic or
phylogeographic relationships that either serve to further support currently understood relationships, or are
indicators of distinct lineages or potential taxonomic
changes to come. Occasionally we also include annotations concerning dates of publication or other bibliographic considerations.
Currently recognized modern turtles and tortoises
(genera and terminal taxa, including species and subspecies) are listed in bold italics. Original description names
and synonymized names are in non-bold italicized text.
Higher suprageneric group-level names are listed in bold
non-italic text and are presented in an indented phylogenetic hierarchy. For competing alternative generic names,
we list them in phylogenetic order from most to least inclusive. All original names include authorship, year, and
page number.
As of this edition of the checklist, all described
genera include type species designations (original and
subsequent), and all described species include verbatim
original and subsequently restricted type localities.
This has been undertaken through a comprehensive reexamination of all original literature rather than relying
on secondary sources. As a result, many of these type
designations and localities are somewhat different from
those previously cited in Fritz and Havaš (2007).
Comments on names that have undergone recent
taxonomic change or phylogenetic analysis or are associated with instability or uncertainty or other changes are
indicated by superscript numbers that refer to annotations
at the end of this and earlier checklists. See the section on
Annotations at the end of the checklist for all detailed
explanations. A summary of all major taxonomic changes
in this checklist as compared to our previous one is provided in Table 3; all minor changes are only included in
the annotations.
Turtle taxa that have gone extinct within modern
times (since 1500 AD) are labeled in bold as Extinct, and
are also listed in Table 1. As of this checklist, this includes 7 species and 3 subspecies, or 10 taxa (2.1% of all
modern turtle taxa) that are extinct.
Turtle taxa that were originally described based on
Pleistocene or Holocene fossil, subfossil, or archeological material, but subsequently recognized as representing
extant taxa or synonymized with modern turtle taxa, are
included in the checklist and marked with a cross (†),
and include stratigraphic horizon and location data. Fossil taxa synonymized with extant polytypic species are
listed under the geographically most appropriate subspecies; however, such synonymizations may not be accurate
for some Early Pleistocene or older fossils which could
conceivably represent distinct chronospecies or extinct
subspecies. For further details on extinct fossil turtle
and tortoise taxa from the Pleistocene and Holocene, see
TEWG (2015).
Those modern species and subspecies for which
in-depth informational accounts have been published
in this TFTSG monograph series on Conservation Biology of Freshwater Turtles and Tortoises (CBFTT), are
indicated by a CBFTT Account heading, with interactive hyperlinks provided to the online published accounts
in dark blue typeface. We will gradually publish CBFTT
accounts for all non-marine turtle and tortoise species—
accounts also include recognized subspecies within the
account, but some subspecies have separate accounts
and are so indicated. As of December 2016 we have published 100 CBFTT accounts covering 130 turtle and tortoise taxa; these are all available online as downloadable
open-access doi-designated pdf’s on the TFTSG website
at www.iucn-tftsg.org/cbftt/.
The checklist includes English common names for
all taxa. We have tried to provide the most commonly
used names, although occasionally we have provided
two or more names. We do not support the practice of
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
TTWG Guidelines for Taxonomic Changes
Taxonomy is both a summary of scientific knowledge and a language for biological communication. As such, it is critical
that taxonomic changes be carefully considered and based on strong, comprehensive underlying data to ensure that changes
are stable and long-lasting. We fully recognize that taxonomy and the systematics research on which it is based, is a dynamic
field and that change is a sign of healthy science. However, we also recognize that taxonomic and nomenclatural stability are
of immense value to the wider community of biologists, conservationists, legislative authorities, and the public at large. Pauly
et al. (2009) argued that taxonomy should aim for stability and monophyly; in cases where these two objectives are in conflict,
well-supported monophyly prevails over stability. Given the dynamic nature of turtle taxonomy, we believe that a series of best
practices can and should be followed that should lead to changes that are stable, informative, and long-lasting. We summarize
these best practices both to identify many of the key points in our group discussions on newly-proposed name changes, and
as a set of considerations for authors who are considering new name changes. We hope the community finds them useful. For
additional discussion, see TTWG (2007a), Pauly et al. (2009), and Kaiser et al. (2013).
1. A proposed taxonomic change must meet the ICZN criteria for nomenclatural validity. Published names gain much greater
credibility by being published in a peer-reviewed scientific journal or equivalent publication standard. These standards include the
2012 emendations of the Code (ICZN 2012) regarding accepted
methods of electronic publication of new names.
2. Taxonomic changes above the species level should preferably be suggested and adopted only when a currently recognized
higher taxon is demonstrably non-monophyletic. We share the view
of the global systematics community that phylogeny should be reflected in higher taxonomic categories, and that changes should
be proposed to “fix” a non-monophyletic grouping. As discussed
in 5) below, non-monophyly should be based on multiple lines of
statistically well-supported evidence. As pointed out by Pauly et
al. (2009), the use of novel levels within a taxonomic hierarchy
(subgenera, supergenera, etc.) allows for the recognition of new/
previously known clades while still maintaining taxonomic stability within a group.
3. Taxonomic changes should incur the fewest possible name
changes while resulting in a final set of monophyletic taxa. We
share the view that taxonomic stability, and therefore the fewest
possible nomenclatural changes, is always a desirable outcome.
4. Avoid naming monotypic higher groups when possible.
As has been repeatedly stated in the literature, monotypic genera,
families, etc. provide only very limited information on group membership, and therefore are less informative than alternative schemes
where higher groups have multiple species within them. This may
imply merging / lumping, rather than splitting, to resolve issues of
non-monophyly. On the other hand, monotypic higher taxa emphasize the unique position of its contained (surviving) taxon. Monotypic higher taxa have been recognized among turtles for over two
centuries, and many (though not all) contain additional extinct taxa
as well as a single surviving species. We do not advocate eliminating traditionally recognized monotypic, and usually reciprocally
monophyletic,higher taxa (since that would lead to taxonomic destabilization), but caution against proliferation of monotypic higher
taxa.
5. Taxonomic arrangements that are supported by several
independent character sets, provide strong statistical support for
each, and report reasonable concordance between different datasets are more compelling than results from a single character set.
Independence in evolutionary studies is a complex concept. In systematics, independence means that characters are not constrained
to covary. For example, when multiple genetically independent
nuclear genes, or nuclear genes and morphological characters, imply the same phylogenetic relationships or species boundaries, they
presumably do so because both reflect the evolutionary history of
the contained lineages. However, two mitochondrial genes are far
less independent, since they are physically linked in the same non-
recombining piece of mtDNA, and natural selection, drift, or any
other process act simultaneously on that linked set of nucleotides.
Single character (e.g., only mtDNA, or only geographic distribution patterns) may reflect the history of the species, or they may reflect the history of that one character. We strongly recommend that
individual characters (each nuclear gene, composite set of mtDNA
data, morphological, behavioral, and other characters) be analyzed
separately to test for concordance among multiple independent data
sets.
6. Independent datasets may or may not provide convincing
evidence for monophyly, and thus for taxonomic changes. When
one dataset conflicts strongly with several other independent ones,
there may still be strong support for the hypothesis supported by
multiple independent data sets. However, character conflict may
often suggest that additional analyses or data are needed before
taxonomic changes should be endorsed and accepted.
7. Sampling should be comprehensive at the appropriate level. Broad taxon sampling for species trees, with multiple specimens
from across the geographic range of each taxon, can help avoid
spuriously high statistical support values for apparent clades (see
Spinks et al. 2013 for a recent chelonian example).
8. Species delimitation studies should include broad geographic sampling of all relevant taxa. Comprehensive geographic
sampling for each character from individuals across the ranges of
all species being considered is often critically important to correctly diagnose new species. We recognize that comprehensive geographic sampling may be difficult for rare species, but every effort
should be made to be as comprehensive as possible.
9. Studies that only evaluate a taxonomic or geographic subset
of the relevant group, or only make changes to some taxa without
evaluating the relevance of these changes to related taxa, are less
likely to be convincing and stand the test of time, and therefore are
less likely to be widely adopted. For example, a study that elevates
a particular subspecies to species rank, without examining variation among the remainder of the species is unlikely to be adopted
until further supporting and clarifying information is published.
10. The TTWG primarily reacts to taxonomic changes proposed in the published literature, although we also will take under
consideration publications that are under review but not yet published if they add additional information to a proposed change. Any
information that the TTWG members have access to can be used
to argue for or against adoption of a new taxonomic arrangement
proposed in a validly published publication, although in almost all
cases we rely on information that is either published or under review in a peer-reviewed journal. The TTWG will not use information from an as-yet unpublished study or manuscript to initiate a
taxonomic change. In very rare cases, the TTWG may decide to
make a new nomenclatural act, such as creating new nomenclatural
combinations.
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
designating “official” or “standard” common names for
species, as that is the domain for scientific names. Instead, common names tend to evolve and vary from
area to area and over time, as well as with language and
cultural context. However, in the field of conservation,
the use of reasonably widely recognized and appropriately descriptive common names is critically important
for communication purposes, and so we include English common names here. Though also important for the
global conservation community, and officially included
in IUCN Red List and CITES documentation, we do not
at this time include Spanish or French common names in
this checklist. Indigenous vernacular names for certain
species are often extensive and imprecise, and in general
we do not list such names here, although a few commonly-used ones are listed.
We are introducing representative photos of nearly
all species in this checklist, with the intention of adding images of all recognized taxa. We have started with
many photos previously published in our CBFTT species
accounts, as well as photos from our various Chelonian
Research Foundation publications: Chelonian Research
Monographs (CRM 1–6), our peer-reviewed journal
Chelonian Conservation and Biology (CCB), our informal Turtle and Tortoise Newsletter (TTN) and TurtleLog
(TL), the Turtle Conservation Fund (2002) (TCF) prospectus we co-published with Conservation International,
and the Top 25+ Endangered Turtles publication we coproduced with the Turtle Conservation Coalition (2011)
(TCC), all acknowledged as to sources. In addition, we
have included many photos from the authorship team as
well as many other contributing individuals. Photos are
preferentially of wild animals from known recorded localities, but a few are from captivity, and most extinct
species are from preserved museum specimens. We invite and urge our colleagues in the international turtle
community to consider submitting additional identified
quality portrait-style photos of missing taxa from known
localities to us for consideration of inclusion in the next
edition of this checklist.
Taxonomic Changes
A prime purpose of this checklist is to record taxonomic changes published in the literature, to evaluate the
strength of the data supporting those proposed changes,
and to recommend whether the community should adopt
or reject the proposed changes. It is important to note
that the recommendations we make here as the TTWG
regarding the validity or non-validity of any included or
excluded taxonomic names or systematic relationships
are not prescriptive, nor are they official recommendations by the TFTSG or the IUCN, as such matters are
generally best left to specialists working in these areas.
However, we have tried to be consistent in our listing of
what appear to be valid taxa and relationships, based on
criteria of published scientific descriptions and proposed
taxonomic changes accompanied by data and sound argumentation (TTWG 2007a).
Our hope is that through this process, the TTWG and
the TFTSG may help stabilize and guide the fluid state of
chelonian systematics and nomenclature, and provide a
standard reference source for updated taxonomy, systematic relationships, distributions, and conservation status
of all turtles and tortoises. The list should also provide an
impetus for ongoing and future work aimed at clarifying
and resolving areas of taxonomic disagreement and/or
uncertainty, as well as documented distribution patterns.
The very first checklist (TTWG 2007b) was compiled on the ‘last published revision’ principle, though reflecting some alternative arrangements through our use of
the ‘Xxxx or Yyyy’ arrangement. As the checklist has developed over the years and is increasingly adopted as the
taxonomic standard by other groups and entities (IUCN
Red List, Reptile Database, and others), and informs nomenclatural deliberations in CITES, ITIS, and other institutions, the TTWG author team has increasingly felt
a need to evaluate both the scientific merit and the wider
implications of adopting proposed taxonomic novelties.
Evaluations have always been on a case-by-case basis,
bringing the diverse perspectives of the authorship team
to bear on the merits of each proposed change. We have
considered drafting criteria for adoption or rejection, but
concluded that every case is unique, making it unrealistic and undesirable to rely on a single set of “rules”. Instead, we have formulated guidelines and considerations
of what increases (or decreases) the scientific credibility
of a proposed taxonomic novelty, and therefore the likelihood of its adoption into (or rejection from) the TTWG
turtle checklist (see inset on p. 13).
We have previously (TTWG 2007a) presented proactive guidelines for researchers proposing taxonomic
novelties; these remain valuable guidance also when
we evaluate new published names or arrangements. But
updating the checklist has required additional considerations, which we describe here. None of these are allor-nothing decisions; instead, almost every proposed
taxonomic novelty, and the underlying supporting data as
presented in the publication, falls somewhere on a continuum between ‘adopt unreservedly’ and ‘reject outright’.
The collective weight of evidence supporting any proposed change (availability of the name; strength and nature
of the supporting evidence; phylogenetic context; agreement with other studies; effect on taxonomic stability) is
deliberated by the TTWG team (often very extensively and
often with different philosophical views of the value of the
underlying evidence). In order to provide a more comprehensive and international approach to TTWG deliberations
and decisions, especially as regards issues of phylogenetic
analysis, we have recently expanded our previous TTWG
authorship team (Rhodin, Iverson, van Dijk, Shaffer, and
Bour) to now also include Arthur Georges and Uwe Fritz,
and we welcome them aboard. Our deliberations lead to
conclusions on whether to:
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
1) adopt fully, reject, or recommend modification of
a proposed taxonomic change,
2) include it as an ‘Xxxx or Yyyy’ arrangement, or
3) suspend adoption until additional, independent
supporting or alternative data are published.
It is important to note that decisions and recommendations within the TTWG are not always unanimous, and
our checklist is not necessarily reflective of the individual
taxonomic views or conclusions of all team members. In
the accompanying text box (see p. 13), we summarize our
guidelines and recommendations for making taxonomic
changes.
Nomenclatural and taxonomic changes often have
disruptive effects for legislation and other ‘users’ of
checklists. A degree of disruption is inevitable as phylogenetic knowledge accumulates; but we are more likely
to adopt proposed changes that have significant ‘disruptive’ effects on widely-used names if such changes are
strongly supported by robust data; in contrast, we are
inclined to suspend adoption of novel names and arrangements if they are based on weaker data sets or do not
greatly improve our overall phylogenetic understanding.
As an example, we would be reluctant to adopt a proposal
to transfer a single species out of an established genus
to form a new, monotypic genus, a move that would involve new names and combinations without significant
improvement of our understanding of the overall relationships of the group of species. We repeat our recommendation (TTWG 2007a) that taxonomy should not be
driven by politics or opportunism, and that the wider implications of taxonomic and nomenclatural decisions be
understood and carefully considered.
We have noted for many years that the ICZN (2012)
has emended its Code regarding accepted methods of
electronic publication of new names. The revision permits electronic publication after 2011 only after the
work (not the new name) is first registered in ZooBank
(http://zoobank.org/; The Official Registry of Zoological
Table 3. Summary of new or resurrected taxa (*) included in this 2017 checklist and major taxonomic changes from TTWG
2014. See the annotations for a full discussion of all these changes; minor changes associated only with overlooked or previously synonymized names or dates of authorship or other primarily nomenclatural changes are not listed here, but only in the
annotations. This table does not include added synonymized fossil taxa, nomina nuda, or names not considered valid in the
2014 checklist (i.e., newly added synonyms).
TTWG 2014 (335 species, 453 taxa)
TTWG 2017 (356 species, 478 taxa)
Change
Macrochelys temminckii
Macrochelys temminckii + Macrochelys suwanniensis *
(M. apalachicolae * = M. temminckii)
Kinosternon stejnegeri * (+ Kinosternon arizonense †)
taxon split, 1 new species described,
1 new species synonymized
taxon split, arizonense only fossil,
1 modern species resurrected
subspecies elevated to species
species changed to subspecies
subspecies reassigned
subspecies reassigned
taxon split, 1 species resurrected
taxon synonymized under occidentalis
taxon split, 1 new subspecies described
taxon split, 1 new species recognized,
and 5 subspecies (2 new, 2 resurrected)
recognized
taxon split, 1 new species described,
1 new species synonymized
taxon split, 1 new species described
taxon split, 3 previously synonymized
species resurrected
taxon split, 1 new species described
genus split into 2 genera
genus divided into 3 subgenera
Kinosternon arizonense
Kinosternon subrubrum steindachneri
Trachemys emolli
Trachemys ornata callirostris
Trachemys venusta panamensis
Emys or Actinemys marmorata
Emys orbicularis fritzjuergenobsti
Cuora aurocapitata
Cuora trifasciata
Malayemys subtrijuga
Chelonoidis porteri
Chelonoidis vicina
Gopherus morafkai
Homopus
Testudo
Amyda cartilaginea
Pelochelys cantorii
Chelodininae
Chelodina (Macrochelodina) expansa
Elseya
Elseya dentata
Elseya novaeguineae
Flaviemys
Flaviemys purvisi
Pelomedusa subrufa
Kinosternon steindachneri
Trachemys grayi emolli
Trachemys venusta callirostris
Trachemys grayi panamensis
E. or A. marmorata + E. or A. pallida *
Emys orbicularis occidentalis
Cuora aurocapitata aurocapitata + C. a. dabieshani *
Cuora trifasciata trifasciata + C. t. luteocephala *
+ Cuora cyclornata cyclornata * + C. c. meieri *
+ C. c. annamitica *
Malayemys subtrijuga + Malayemys khoratensis *
(M. isan * = M. khoratensis *)
Chelonoidis porteri + Chelonoidis donfaustoi *
Chelonoidis vicina + C. guntheri * + C. microphyes *
+ C. vandenburghi *
Gopherus morafkai + Gopherus evgoodei *
Homopus + Chersobius
Testudo (Testudo) + Testudo (Agrionemys)
+ Testudo (Chersine)
Amyda cartilaginea + A. cartilaginea cartilaginea,
+ A. c. maculosa * + Amyda c. or ornata ornata *
+ A. c. or o. phayrei *
[Pelochelys (Ferepelochelys)] + [Pelochelys clivepalmeri]
+ [Pelochelys telstraorum]
Chelodininae + Pseudemydurinae
[Chelodina (Supremechelys) expansa brisbaneensis]
+ [Chelodina (Supremechelys) duboisi]
Elseya (Elseya) + Elseya (Hanwarachelys)
+ Elseya (Pelocomastes)
Elseya (Elseya) dentata + E. (E.) flaviventralis *
Elseya (Hanwarachelys) novaeguineae + E. (H.) rhodini *
Myuchelys
Myuchelys purvisi
Pelomedusa subrufa + P. barbata * + P. galeata *
+ P. gehafie * + P. kobe * + P. neumanni * + P. olivacea *
+ P. schweinfurthi * + P. somalica * + P. variabilis *
taxon split, 1 new subspecies described,
1 species resurrected,
1 subspecies resurrected
unavailable names
subfamily split, 1 subfamily resurrected
unavailable names
genus divided into 3 subgenera
taxon split, 1 new species described
taxon split, 1 new species described
monotypic genus synonymized
generic allocation of taxon revised
taxon split, 6 new species described,
3 species resurrected
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Nomenclature). ZooBank must register the precise electronic archive where the work is to be published, as well
as the ISSN or ISBN of the work. In addition, amendments to the Code also clarify that preliminary electronic
versions of works due for publication on paper are unavailable, and that abstracts of meetings, presentation
texts and posters are unavailable for nomenclatural purposes, and preliminary electronic versions do not bring
forward the date of publication, unless the electronic
version meets the requirements for availability. Authors
intending to publish taxonomic papers in electronic archives are cautioned to read the text of International
Commission on Zoological Nomenclature (2012) carefully, and to follow the guidelines precisely, at risk of
having their work inadmissible.
Distributions
We summarize distributions for all taxa in the checklist, listing all nations and territories in which they occur
as native populations (see Table 2 for the top turtle-rich
nations). For several larger nations we also list political
or geographic subunits (e.g., states, provinces, regions,
or larger islands). We attempt to also indicate nations or
territories where species have been extirpated or where
they occur as non-native introduced or invasive species,
or where there are uncertainties as to occurrence.
For introductions, we attempt to distinguish between
two forms: (1) modern introductions (since ca. 1500 AD)
for those species that appear to have relatively well-established or potentially reproducing populations in extra-limital areas primarily as a result of relatively recent
trade for food or pets or planned conservation introductions (labeled “introduced” or “modern”), and (2) earlier
historic or prehistoric introductions for those species that
appear to have native populations, but where population
genetics studies find evidence of founder effects suggestive of possible introduction by humans, or other dispersal events, during the last ca. 2000–3000 years (labeled
“prehistoric introduction?”).
For freshwater and terrestrial turtles and tortoises,
we compiled native and introduced distributions and
locality records from a combination of multiple published and database sources. For native distributions
we used Iverson (1992) and Fritz and Havaš (2007) as
starting points, and then added data from other recent
literature by numerous authors, data from our extensive
database compiled from Iverson’s work by Buhlmann
et al. (2009), further data from our published CBFTT
species accounts, our TFTSG-organized IUCN Red
Listing workshops, and data from Iverson’s continuing
intensive compilation of distribution records from the
literature.
For introduced species, we used Kraus (2009) as a
starting point and have added data from other publications
and online sources and databases. We have attempted to
list introductions that are based on recorded populations
rather than just single animals (but not necessarily with
evidence of reproduction), but have not been rigorous in
this, as it can be difficult to determine what the actual
situation may be in each recorded case.
Finally, we also solicited and received input from
many members of the TFTSG for corrections and additions to all the native distributions and introductions
data. Despite this effort, it is likely that we have committed errors of omission or commission, and we request that any corrections or updates be brought to our
attention so that they can be included in future editions
of this checklist.
For sea turtles, we compiled distributions from a
combination of IUCN Red List data, CMS (Convention
on Migratory Species) listings, and the extensive listings
of nesting sites and foraging ranges that the IUCN/SSC
Marine Turtle Specialist Group (MTSG) includes in its
SWOT mapping application (The State of the World’s Sea
Turtles) (http://seaturtlestatus.org/learn/maps/all), generously supplied to us by the MTSG. Based on these data,
we list sea turtle distributions in three distributional categories: 1) nesting: native regularly nesting populations,
2) foraging: native permanently foraging or regularly migrating populations (but no evidence yet of regular nesting), and 3) vagrant: temporarily foraging or migrating
animals not necessarily considered native.
GIS Maps
In the previous edition of the checklist, we added simple GIS range distribution maps for all species. All maps
in this edition have now been enhanced and updated to
also include specific locality points upon which the ranges are based, as well as adding detailed color-coded altitudinal elevations. Nearly all distributional ranges have
been revised based on new data and improved geographic
analysis, in general restricting the presumed range somewhat tighter around our recorded localities. The locality
points have been obtained from a combination of data
from Iverson (1992), edited and corrected localities from
the EmySystem maps (http://emys.geo.orst.edu/), locality data that Iverson has continued to collect systematically since 1992, CBFTT species accounts, IUCN Red
Listing workshops, recent literature (although far from
complete) compiled mainly by Iverson and Rhodin, and
personal input from members of the TFTSG and other
specialists. Maps now also include color-coded ranges
for recognized subspecies.
Map production began with point locality datasets
from Iverson (1992), based on the many museum-held
voucher specimens and published records amassed by
John Iverson over the years and updated on the EmySystem (http://emys.geo.orst.edu/). These datasets were then
supplemented by newer data and converted into shapefiles and edited and corrected and updated for content by
Iverson, Ross Kiester, Tom Akre, Kurt Buhlmann, Peter
Paul van Dijk, Arthur Georges, Anders Rhodin, Russ
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Figure 2. Global species richness map for all tortoises and freshwater turtles. Composite map of all tortoise and freshwater turtle
distribution maps (349 species, 471 taxa) as presented in this checklist and atlas. Distribution shapefiles for all taxa in this map
are each shown in red at 80% transparency and stacked; lighest red color = one taxon, darkest red color = > 15 taxa. For enlarged
and more detailed regional species richness maps, see pp. 218–220 at the end of the taxonomic checklist.
Mittermeier, and Whit Gibbons, and analyzed by Buhlmann et al. (2009).
The original maps created this way were based on
constructing projected historical geographic ranges. This
was done by selecting GIS-defined hydrologic unit compartments (HUCs, at relatively coarse level 6 hydroshed
basins) with verified locality points, and then adding
HUCs that connected known point localities in the same
watershed or physiographic region and that had similar
habitats and elevations as the verified HUCs. As such,
these first maps represented assumed geographic ranges,
but generally somewhat larger than reality, and required
further verification and adjustment.
These distribution shapefiles were then further revised and formatted by Rhodin using ArcGIS Desktop
10.1 (www.esri.com) as part of the IUCN-associated BioFresh initiative (http://atlas.freshwaterbiodiversity.eu/),
using finer geographic scales (hydroshed basins at levels
10 or 12). This allowed elimination of many higher-altitude regions from the projected ranges (notably in areas
such as the Himalayan and Andean foothills and other
mountainous regions), while keeping lower altitude HUC
distributions in the same overall drainage basins, and in
general tightening up and reducing many of the projected
ranges. The maps have also been further revised through
input of data provided by authors of published CBFTT
accounts and participants in TFTSG-organized IUCN
Red Listing workshops, but still represent projected and
assumed historical ranges.
For some relatively cryptic, poorly known, or possibly questionable species, the ranges depicted in this
checklist are at best general approximations of their potential distributions. Species that fall into this category
include Pelodiscus axenaria, P. parviformis, Cuora
zhoui, Cyclemys enigmatica, Rafetus swinhoei, Mesoclemmys heliostemma, Chelodina gunaleni, C. kuchlingi,
and Emydura tanybaraga.
Other apparently widespread species with significant
documented phylogeographic differentiation in the
form of recognized subspecies or genetically-defined
lineages and evolutionarily significant units (ESUs) may
eventually warrant recognition as multiple taxa at the
species level. Some species that fall into this category
include Kinosternon hirtipes, K. integrum, K. scorpioides,
Terrapene carolina, Cuora amboinensis, Melanochelys
trijuga, Chelonoidis carbonarius, Testudo graeca, and
Phrynops geoffroanus.
Native populations and recorded specimens are
marked with yellow locality spots set in partially transparent distribution range polygons (using HUCs), using
red polygons at 50% transparency for species and nominate subspecies, and other colors at 30–50% transparency
for other subspecies. Populations that represent possible
prehistoric introductions (whether genetically verified
or hypothesized as such) or possible prehistoric or more
recent natural range extensions are also indicated with
yellow spots. A composite species richness map for all
tortoise and freshwater turtle taxa is depicted in Fig. 2
17
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
and enlarged regional detailed maps are included after the
checklist on pp. 218–221.
Apparently introduced populations near the native
range that are most likely of modern and recent historic
origin are generally included, and are indicated by orange
spots. Remote introduced populations are not included.
Questionable locality records near the native range (possibly misidentified and/or regional trade specimens) are
also indicated by orange spots. Extinct taxa (species or
subspecies) are designated by red spots.
It is critically important to note that the maps published here depict projected and presumed historical geographic ranges (defined as historical area of occupancy,
or AOO), as they are based on a combination of older
historical museum and literature data (Iverson 1992)
and more recent locality data, and do not in general reflect actual current areas of occupancy (AOO) of these
species. Most turtle and tortoise species have had their
historical ranges decrease considerably as a result of extensive habitat loss and degradation and/or overexploitation. For example, the ranges depicted here for Batagur
trivittata and Geochelone platynota from Myanmar, and
Psammobates geometricus from South Africa, show their
historical ranges, rather than their current ranges, which
have all been reduced by >90%. In general, all species
assessed as Critically Endangered or Endangered on the
IUCN Red List or TFTSG Draft Red List have had their
current AOO ranges greatly decreased from historical
extents.
Sea turtle maps were generated from GIS data generously supplied to us by the IUCN/SSC Marine Turtle
Specialist Group (MTSG) and SWOT (The State of the
World’s Sea Turtles), and show documented nesting sites
as yellow dots and generalized foraging distributions for
each species as shaded oceanic distributional ranges delimited as either regional management units (RMUs) or
distinctive population segments (DPSs) (see Seminoff et
al. 2015).
Conservation Status
We include current IUCN Red List conservation
status for all species. The status categorizations listed
here are current as of the IUCN Red List of Threatened
SpeciesTM, version 2017.1 (www.iucnredlist.org). The
TFTSG is the official global IUCN Red List Authority
responsible for continuously updating IUCN Red List assessments of all tortoises and freshwater turtles, and this
process is handled through multiple consensus-building
workshops and consultations.
As many species on the Red List need updating, either because their previous evaluations are more than
ten years old, or because of recent conservation status or
taxonomic changes, we have also included the results of
TFTSG Draft Red List assessments (through June 2017)
to indicate their current provisional status, which should
be released on the official IUCN Red List site in the
near future. In addition, many species that were determined by the TFTSG to be Least Concern in 1996 were
never formally listed (as per IUCN Red List protocol at
the time), but the original determinations as prepared at
that time are still available and are indicated here. For a
few species from the South African region we have also
added draft Red List assessments done in 2010 by the
South African Reptile Conservation Assessment (SARCA) committee, subsequently published by Hofmeyr et
al. (2014).
Finally, we include regulatory status listings on
CITES (Convention on International Trade in Endangered Species of Wild Fauna and Flora) Appendices I, II,
or III, current as of the 4 April 2017 listing (http://cites.
org/eng/app/appendices.php). As such, this document
brings together most important aspects of taxonomy,
names, distribution, and conservation status of all turtles
and tortoises of the world.
RESULTS AND DISCUSSION
Conservation Status
To assess and summarize the current conservation
status of turtles and tortoises in the broadest strokes, we
provide an update and analysis of the most current IUCN
Red List (www.iucnredlist.org), as well as provide provisional conservation status of species still under evaluation. The official determinations of conservation status of
turtles are provided to the IUCN Red List by the TFTSG,
which is continuously producing draft assessments for
previously unevaluated taxa as well as previously evaluated taxa needing updates (necessary every 10 years).
Knowing the overall conservation status and percentage of threatened species of turtles is important in understanding how seriously they are endangered, and how
they compare with other imperiled organisms. Mittermeier et al. (2015) provided an analysis of global Turtle
Hotspots as part of such an evaluation.
The current IUCN Red List (version 2017.1) formally lists 251 turtle species, including 7 separate subspecies
and 19 regional subpopulations, using a slightly different
taxonomy from the one presented in this checklist. Of the
251 species listed, 8 are Extinct (EX) [includes Pelusios
seychellensis, considered a subspecies in our checklist],
2 Extinct in the Wild (EW), 40 Critically Endangered
(CR), 44 Endangered (EN), 65 Vulnerable (VU), 34 Near
Threatened (NT), 1 Conservation Dependent (LR/cd; an
old category being phased out), 45 Least Concern (LC),
11 Data Deficient (DD), and 1 Not Assessed (NA).
By IUCN Red List protocol, Threatened species
are defined as those in the three categories of Critically Endangered, Endangered, and Vulnerable, meaning that 149 species are officially regarded as Threatened (59.4% of the 251 species listed), with 84 species
(33.5% of those listed) considered Critically Endangered or Endangered.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Of the 356 species recognized as distinct (or possibly distinct) in our checklist, 105 are not yet officially
listed on the IUCN Red List as species (although some
are listed as subspecies). Most of these apparently “unassessed” species have in fact already been evaluated by
the TFTSG, first in 1996, when Least Concern (LC) species were not formally listed (as some are now), and then
more recently through a series of draft assessments. Of
these species, the TFTSG evaluated 53 as Least Concern
in 1996 (J.L. Behler and C. Hilton-Taylor, in litt.), and
these are marked as such in this checklist.
Further status assessments have more recently been
accomplished through an ongoing series of regional IUCN Red Listing workshops held by the TFTSG.
These workshops have assessed both previously unevaluated species and updated older previously evaluated
species. Since 1999 the TFTSG has held Red Listing
workshops in or for Asia, Mexico, the Mediterranean,
India, Madagascar, Australia, New Guinea, USA, northern South America, southern South America, the Galápagos Islands, Asia a second time, Sub-Saharan Africa,
and India a second time. Although not yet official IUCN
Red List evaluations, we can use all these draft evaluations to determine overall threat rates to all turtles and
tortoises. The current assessments that are based on the
findings and results of these workshops, but have not
yet been finalized and published on the IUCN Red List,
are included in this checklist as ‘TFTSG Draft Red List’
status.
Combining the formal IUCN Red List assessments
with draft TFTSG status evaluations for previously unlisted species and draft updated assessments for currently
listed but outdated assessments, yields the following total current status numbers for all 356 species of turtles
and tortoises: 7 Extinct (EX), 1 Critically Endangered
(Possibly Extinct) [CR(PE)], 63 Critically Endangered
(CR), 50 Endangered (EN), 65 Vulnerable (VU), 38 Near
Threatened (NT), 81 Least Concern (LC), 35 Data Deficient (DD), and 16 Not Evaluated (NE). This yields 114
species (32.0%) that are Critically Endangered or Endangered, and 179 (50.3%) that are Threatened (Critically
Endangered, Endangered, or Vulnerable). If we also include Extinct species among the Threatened categories
(or more generally, “gone or nearly gone”), then 186 turtle species, or 52.2% of all currently recognized modern
turtle and tortoise species, are either already extinct or
threatened with extinction.
We can provisionally adjust these numbers to account
for Data Deficient and Not Evaluated species which may
also be Threatened. We follow the calculation method of
determining percentage of Threatened species utilized by
Hoffmann et al. (2010): the number of Threatened species
(179) is divided by the number of data-sufficient species
(305), i.e., the total number of species minus those Not
Evaluated (NE) and minus those that are Data Deficient
(DD). This assumes that DD and NE species will have the
same percentage of Threatened species as data-sufficient
species. Using this calculation methodology, 58.7% of all
assessed data-sufficient turtles and tortoises are Threatened, and 61.0% are Threatened or Extinct. For comparison, using the same methods, Hoffmann et al. (2010) determined that 41% of amphibians, 33% of cartilaginous
fishes, 25% of mammals, and 13% of birds were Threatened. Turtles were surpassed only by cycads, with 62% of
their 300+ species Threatened.
No matter how we analyze these various percentages
of threatened species, turtles and tortoises, with anywhere
from ca. 50–59% of all their modern species Threatened,
are among the most endangered of any of the major groups
of vertebrate species, more than birds (ca. 13%), mammals
(ca. 21–25%), cartilaginous and bony fishes (ca. 17–31%),
or amphibians (ca. 30–41%), and paralleled among the
larger vertebrate groups only by the primates (ca. 49%)
(www.iucnredlist.org, Hoffmann et al. 2010).
As part of the process of determining the relative
threatened status of the world’s tortoises and freshwater
turtles, the Turtle Conservation Coalition (2011) published a consensus listing of “Turtles in Trouble: The
World’s 25+ Most Endangered Tortoises and Freshwater
Turtles—2011”, which listed the top ca. 50 most endangered species at that time. This document has since become widely cited, especially as a basis for justifying and
supporting conservation grant proposals and action plans.
As such, continuing to evaluate changes to this Top 25+
list, both in terms of improved status for those species
benefitting from conservation efforts, and documenting
potentially deteriorating survival prospects for other species, will be critically important for future conservation
efforts for these highly threatened species. This continuing process is currently being undertaken by the TFTSG
in collaboration with other turtle conservation organizations, notably Turtle Conservancy, Turtle Survival Alliance, Global Wildlife Conservation, Wildlife Conservation Society, Conservation International, and Chelonian
Research Foundation.
Genetic Pollution
Aside from overt and highly impactful conservation
threats such as overexploitation and habitat destruction, the global turtle fauna is also increasingly facing
another insidious threat: genetic pollution caused by
human-facilitated hybridization and introgression from
introduced and invasive species (Rhymer and Simberloff 1996; Simison et al. 2013; Spencer et al. 2014;
García-Díaz et al. 2015; Nori et al. 2017). This is not
entirely new, but the current extent is unprecedented.
Some taxa have historically already been affected. This
is most probably true for Asian softshell turtles of the
genus Pelodiscus. These turtles have been farmed and
traded for centuries, with the corollary of translocating
different species and local genetic lineages, leading to
the admixture of different taxa and lineages in farms
and in the wild (Fritz et al. 2010b; Suzuki and Hikida
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
2014). Similarly, the historical introduction of Mauremys reevesii to Japan resulted in massive hybridization
with the native M. japonica (Suzuki et al. 2014). Another historical case of human-mediated admixture of
genetic lineages is known from European pond turtles
(Emys orbicularis). Here, the non-native populations
on the Balearic Islands, most probably introduced in
Roman times (Valenzuela et al. 2016), are of admixed
origin (Lenk et al. 1999). Another population with genetic signatures of an old or ancient introduction of E. o.
hellenica was discovered near Rome (Lenk et al. 1999;
Vamberger et al. 2015) within the range of another subspecies (E. o. galloitalica).
However, unlike in historical times, when only a few
turtle species were affected, genetic pollution has become
a major issue for nature conservation in recent years, facilitated by the massive pet and food trade and increased
human mobility. Today, genetic pollution is also caused
by well-meaning augmentation of endangered local turtle populations with genetically mismatched individuals
(typically, but not exclusively, from non-coordinated actions by turtle enthusiasts), the release of surplus or abandoned genetically divergent pet turtles, and also by largescale releases of confiscated turtle shipments, especially
in Southeast Asia.
Examples of restocking with mismatched genetic
individuals include endangered populations of E. orbicularis at the northern edge of its range (Fritz et al.
2004; Velo-Antón et al. 2011: genetic evidence for
restocking with several different subspecies), and in
southern France (Vamberger et al. 2015; Raemy et al.
2017: restocking with non-native E. o. hellenica instead
of native E. o. galloitalica), also northern edge populations of Mauremys leprosa in southern France (Palacios et al. 2015: restocking with M. l. saharica and
northern African M. l. leprosa instead of European M.
l. leprosa), of M. rivulata in Croatia (Vamberger et al.
2014: restocking with Cretan individuals), and of Testudo graeca in Doñana National Park in Spain (Graciá
et al. 2017b: restocking with non-native T. g. marokkensis from Morocco instead of T. g. graeca from Spain).
Examples of genetic pollution caused by abandoned pet
turtles include Chrysemys picta bellii from British Columbia introgressed by non-native subspecies (Jensen
et al. 2014b) and Antillean Trachemys introgressed by
red-eared sliders (Trachemys scripta elegans; Parham et
al. 2013). Also some of the above-mentioned cases for
European pond turtles refer at least partially to genetic
pollution by abandoned pet turtles. Hybridization in the
wild from released trade animals has been recorded in
Taiwan for Mauremys reevesii and M. sinensis (Fong
and Chen 2010).
This issue of potentially increasing genetic pollution needs to be kept in mind as conservationists devise
management plans designed to reinforce or restore dwindling or extirpated populations of turtles impacted by
overexploitation and habitat loss. The need to maintain
well-defined and relatively pure non-hybrid genetic lineages, subspecies, and species is important and needs to
be kept in focus.
Request for Updates
Please help the TTWG and the TFTSG keep this
Turtles of the World Checklist and Atlas up-to-date by
e-mailing any or all of us (addresses noted above) and
including pdf’s of any relevant articles about new taxonomic or distributional information and/or revisions
that should be included and annotated here in upcoming
checklists, whether you are an author on a paper providing updated information, or have become aware of data
that you believe should be included. Also please inform
us of any errors or discrepancies in any of our data, especially for geographic distributions (native or introduced)
in countries or states, and for cited references and names,
so that we may update or correct them as necessary.
For sea turtle distribution data, please also submit additions and corrections via the SWOT website. We want
this checklist and atlas to be as accurate, up-to-date, and
comprehensive as possible, and ask for your assistance to
help us accomplish this goal.
The maps published here all represent work in progress, and will continue to be updated and revised in future
checklists as we acquire new and improved locality data.
We strongly encourage and welcome our readers and professional colleagues, especially field-based turtle specialists and other enthusiasts, to inform us about proposed
corrections and changes to these maps, and to submit specific locality data on the presence or absence of species in
various locations for consideration of incorporation into
the next checklist.
We are especially interested in receiving new point
locality data from presumed range areas not well represented by locality points on our maps, especially from
gap areas in the presumed range, as well as verified range
extensions. Please help us improve and update these
maps by submitting any new locality information along
with geographic coordinates and the associated citations
to Rhodin at rhodincrf@aol.com.
aCknoWledgMenTs
We extend special thanks to James Parham, a major contributor and co-author on earlier versions of the
checklist, and a source of incisive initial input and analytic debate. Specific thanks also for extensive detailed
input from Holger Vetter, Peter Uetz, and Scott Thomson.
We thank various members of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group (TFTSG)
as well as many other individuals who have provided
pertinent data or references or localities for maps, or
helped in various ways with input or review of this and
earlier checklists: Rafael Acuña-Mesén, Kraig Adler,
Leandro Alcalde, Matthew Aresco, Mark Auliya, Ernst
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Baard, Sherif Baha El Din, Albert Bertolero, John Bickham, Torsten Blanck, Bill Branch, Ronald Brooks, Ortwin Bourquin, James Buskirk, Gisella Caccone, Patrick
Campbell, Linda Cayot, Eng Heng Chan, Laurent Chirio,
Marcelo de la Fuente, Jeffrey Dawson, Tomas Diagne,
Andrew DiMateo, David Emmett, Taylor Edwards,
Alejandro Fallabrino, Balazs Farkas, Bruno Ferronato,
Oscar Flores-Villela, Eugene Gaffney, Justin Gerlach,
Hanyeh Ghaffari, James Gibbs, Scott Gillingwater, Eric
Goode, Steve Gotte, Michael Granatosky, Ute Grimm,
Cris Hagen, Brian Henen, Craig Hilton-Taylor, Margaretha Hofmeyr, Marinus Hoogmoed, Brian Horne,
Flora Ihlow, Sixto Inchaustegui, Djoko Iskandar, Dale
Jackson, Fredric Janzen, Robert Jones, Ross Kiester,
Kenneth Krysko, Gerald Kuchling, Minh Le, Thomas
Leuteritz, Peter Lindeman, Jeffrey Lovich, Luca Luiselli, Roderic Mast, Peter Mikulíček, David Mifsud,
Russell Mittermeier, Robert Murphy, Simon Nemtzov,
Dario Ottonello, Vivian Páez, Olivier Pauwels, Fabio
Petrozzi, Hans-Dieter Philippen, Nicolas Pilcher, Peter
Praschag, Awal Riyanto, Jose Vicente Rueda-Almonacid, Raymond Saumure, Chuck Schaffer, Jason Schaffer, Hans-Hermann Schleich, Sabine Schoppe, Norman
Scott, Will Selman, Shi Haitao, Franco Souza, Phil
Spinks, Bryan Stuart, Travis Thomas, Robert Thomson,
Oguz Türkozan, Melita Vamberger, Richard Vogt, Bruce
Weissgold, and Reza Yadollahvand.
Distributional data on sea turtles from the IUCN/
SSC Marine Turtle Specialist Group (MTSG), SWOT
(The State of the World’s Sea Turtles), and Seminoff et
al. (2015) were gratefully received from Bryan Wallace,
Jeffrey Seminoff, and Kyle Van Houtan.
The Panaphil Foundation provided partial funding
support for PPvD, and HBS was partially supported by a
grant from the NSF (DEB-0817042). TFTSG-organized
IUCN Red Listing workshops and other redlisting efforts
have been generously supported by the Frankel Family
Foundation, Conservation International, Turtle Conservancy, Moore Family Foundation, George Meyer and
Maria Semple, Matt Frankel, Turtle Conservation Fund,
Chelonian Research Foundation, Galapagos Conservancy, U.S. Fish and Wildlife Service, Andrew Sabin Family
Foundation, and Mohamed bin Zayed Species Conservation Fund.
Initial map production and GIS-analysis by the
TFTSG and Chelonian Research Foundation was supported by the BioFresh Project of the IUCN Freshwater Biodiversity Unit, IUCN Species Programme, and
Forschungsverbund Berlin e.V., Leibniz-Institut für
Gewässerökologie und Binnenfischerei. We thank Kevin Smith, Savrina Carrizo, Will Darwall, Ackbar Joolia, Marcelo Tognelli, Adrian Hughes, and Diego Juffe
of IUCN and Melita Vamberger for welcome technical
assistance and helpful suggestions in our production of
maps. We thank Bryan Wallace, Rod Mast, and Ei Fujioka of SWOT and Duke University for help with sea turtle
maps. Ongoing support for map production and checklist
preparation has come from Chelonian Research Foundation (CRF), the TFTSG, and Turtle Conservancy (TC).
Publication costs have been shared by CRF, TC, TFTSG,
Turtle Conservation Fund, Global Wildlife Conservation,
and Surprise Spring Foundation.
phoTos and phoTogRapheRs
We also most gratefully acknowledge and very
much appreciate the use of the many turtle and tortoise
photos by the following photographers that enhance this
checklist. We include both previously unpublished and
republished photos from previous CRF publications,
with each acknowledged and (if republished) identified as to its original source: Conservation Biology
of Freshwater Turtles and Tortoises species accounts
(CBFTT), Chelonian Conservation and Biology journal
articles (CCB), Chelonian Research Monographs books
(CRM 1–6), Turtle and Tortoise Newsletter (TTN), TurtleLog (TL), Turtle Conservation Fund 2002 prospectus
(TCF), and the Turtle Conservation Coalition 2011 Top
25+ document (TCC). We strive to use primarily photos
from natural native settings and to include locality data
when available, although some photos are from captivity or the trade.
We include photos of 461 taxa (96.4% of all 478) by
188 contributors, but still lack photos of the following
17 taxa: Kinosternidae: Kinosternon hirtipes hirtipes;
Emydidae: Trachemys stejnegeri malonei, Trachemys
venusta panamensis, Trachemys venusta uhrigi; Geoemydidae: Rhinoclemmys punctularia flammigera;
Testudinidae: Testudo horsfieldii horsfieldii, Testudo
horsfieldii bogdanovi, Testudo horsfieldii kuznetzovi;
Trionychidae: Chitra chitra javanensis, Pelodiscus axenaria; Chelidae: Platemys platycephala melanonota,
Chelodina gunaleni, Chelodina mccordi roteensis; Pelomedusidae: Pelomedusa kobe, Pelomedusa schweinfurthi, Pelusios williamsi laurenti, and Pelusios williamsi lutescens.
We urge those of you who may have photos of any
of these missing taxa to submit them for consideration
of inclusion in the next checklist. We are also always
looking for better quality photos with locality data for
those taxa we have already illustrated; if you have photos that may offer such improvement, please submit
them for consideration.
Contributing Photographers. — Collette Adams,
Bhaba Amatya, Ben Anders, Matthew Aresco (3),
Mark Auliya (2), Roy C. Averill-Murray, Dinçer Ayaz,
Richard D. Bartlett (5), Chittaranjan Baruah (2), Aaron
S. Baxter, Rafael Bernhard, Albert Bertolero (2), S.
Bhupathy, Torsten Blanck (16), Roger Bour (9), Richard
C. Boycott (2), William R. Branch (5), Elizângela S.
Brito, Rafe M. Brown, Kurt A. Buhlmann (2), James R.
Buskirk (5), Alejandra Cadavid, Matt Cage, John Cann
(7), John L. Carr (3), Vinícius T. de Carvalho (3), Eng
Heng Chan, Tien-Hsi Chen (3), Laurent Chirio, B.C.
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Choudhury, Andrew T. Coleman, Marilyn Connell, Paul
Crow (3), Indraneil Das (11), Shekar Dattatri, Tui De
Roy/Roving Tortoise Photos (2), Anslem de Silva, V.
Deepak (2), David Dennis, Frank Deschandol, Bernard
Devaux, Tomas Diagne (7), Larry Ditto, C. Kenneth
Dodd, Jr. (2), Johannes Els, David Emmett (2), Kevin
Enge, Pablo Feliz, Vincenzo Ferri (5), Darren Fielder,
German Forero-Medina (2), Manuel Merchán Fornelino,
Alastair Freeman, Uwe Fritz (6), Carlos A. Galvis-Rizo,
Maren Gaulke (2), Saurav Gawan, Arthur Georges (8),
Justin Gerlach (3), David J. Germano, Hanyeh Ghaffari,
Paul M. Gibbons, Scott D. Gillingwater, James C.
Godwin (4), Eric V. Goode (4), Robert H. Goodman,
Jr., Wulf Haacke, Cris Hagen (2), Adrian Hailey,
Norbert Halasz (2), James H. Harding (6), Douglas
B. Hendrie (2), Judith Hirt, Kate Hodges, Margaretha
D. Hofmeyr, Brian D. Horne, Janet Hostetter, Jennifer
G. Howeth, Rick Hudson, Bonggi R. Ibarrondo, Flora
Ihlow (8), Alexander A. Inozemtsev, Iriomote Wildlife
Conservation Center, John B. Iverson (40), Dale R.
Jackson, S. Jayakumar, John Jensen, Carlos Alberto
Jimenez, Chris Johnson, Michael T. Jones (3), Robert
L. Jones, Nobuhiro Kawazoe, Rod Kennett, Gerald
Kuchling (11), Edgar Lehr, Chris Leone (3), Thomas
E.J. Leuteritz, Peter V. Lindeman (6), Victor J.T. Loehr
(8), Jeffrey E. Lovich (2), Gary Luciano, Luca Luiselli,
William E. Magnusson, Kevin Main, Barry Mansell
(4), Jérôme Maran (14), Peter May, Tomáš Mazuch
(3), William P. McCord, Timothy E.M. McCormack,
Suzanne E. McGaugh, Brian Mealey (2), Melvin
Mérida, Peter A. Meylan, Joseph C. Mitchell, Russell A.
Mittermeier (5), Asghar Mobaraki (2), Edward O. Moll,
Jiří Moravec, Robert H. Mount, Rahul Naik, Andreas
Nöllert (4), Michael Ogle, Annette Olsson, Vivian P.
Páez (2), Charles W. Painter, James F. Parham, Fred
Parker (2), Olivier S.G. Pauwels, Miguel Pedrono, John
Pemberton, Fabio Petrozzi, Hans-Dieter Philippen,
Pino Piccardo, Carmine Pilcher, Nicolas J. Pilcher,
Kalyar Platt, Peter Praschag (2), Peter C.H. Pritchard
(9), Michael Redmer, Renae Reed, Anders G.J. Rhodin
(20), Maurice Rodrigues (3), John Roe, Philip C. Rosen,
José Vicente Rueda-Almonacid, Daniel O. Santana,
Chuck Schaffer, Jason Schaffer (2), Hermann Schleich
(2), Fabian Schmidt, Sabine Schoppe (2), Gabriel H.
Segniagbeto, Shi Haitao (4), Shailendra Singh (3),
Pavel Široký (2), Sitha Som, Franco L. Souza, Craig B.
Stanford, Bev Steveson, James N. Stuart, Yik-Hei Sung,
Chris Tabaka, Washington Tapia, Robert C. Thomson,
Stanley E. Trauth (2), Claire Treilibs, Apostolis Trichas,
Oguz Türkozan, Melita Vamberger, Robert P. Van Dam,
Peter Paul van Dijk (13), Carla Van Ness, Sabine Vinke
(2), Thomas Vinke (2), Richard C. Vogt (7), Timothy
Walsh, Nikhil Whitaker, Dawn Wilson, Win Ko Ko,
Blair Witherington (3), Yuichirou Yasukawa (2), Zhou
Ting, Cassiano Zaparoli, Brian Zarate, Thomas Ziegler,
Anders Zimny, and Stephen M. Zozaya.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
CHECKLIST
ModeRn TuRTle and ToRToise Taxa exTanT sinCe 1500 ad
Phylogenetic Linnaean Classification of
Suprageneric Categories as used in this Checklist
TesTudines .................................... 356 spp., 478 taxa
• CRypTodiRa.............................. 263 spp., 373 taxa
__ChelydRidae ....................... 5 spp., 5 taxa
_Chelonioidea ............................. 7 spp., 7 taxa
__Cheloniidae........................ 6 spp., 6 taxa
___CaReTTinae .................. 4 spp., 4 taxa
___Cheloniinae ................ 2 spp., 2 taxa
__deRMoChelyidae ................ 1 sp., 1 taxon
_kinosTeRnoidea ...................... 28 spp., 39 taxa
__deRMaTeMydidae ................ 1 sp., 1 taxon
__kinosTeRnidae ................ 27 spp., 38 taxa
___kinosTeRninae ......... 24 spp., 35 taxa
___sTauRoTypinae ............. 3 spp., 3 taxa
_TesTudinoidea .................... 190 spp., 278 taxa
__eMydidae ........................ 53 spp., 91 taxa
___deiRoChelyinae ....... 42 spp., 66 taxa
___eMydinae ................. 11 spp., 25 taxa
__plaTysTeRnidae .................... 1 sp., 3 taxa
__geoeMydidae .................. 71 spp., 96 taxa
___geoeMydinae ........... 62 spp., 82 taxa
___RhinoCleMMydinae ... 9 spp., 14 taxa
__TesTudinidae................... 65 spp., 88 taxa
_TRionyChoidea ....................... 33 spp., 44 taxa
__CaReTToChelyidae ............. 1 sp., 1 taxon
__TRionyChidae.................. 32 spp., 43 taxa
___CyClanoRbinae............ 7 spp., 9 taxa
___TRionyChinae .......... 25 spp., 34 taxa
• pleuRodiRa ................................ 93 spp., 105 taxa
__Chelidae ......................... 58 spp., 65 taxa
___Chelinae .................. 20 spp., 21 taxa
___hydRoMedusinae ......... 2 spp., 2 taxa
___Chelodininae .......... 35 spp., 41 taxa
___pseudeMyduRinae ........ 1 sp., 1 taxon
__peloMedusidae ............... 27 spp., 32 taxa
__podoCneMididae ................. 8 spp., 8 taxa
* * * * * *
Alternative Phylogenetic Hierarchical
Phylocode Classification (1)
TesTudines
• pleuRodiRa
_peloMedusoides
__peloMedusidae
__podoCneMididae
_Chelidae
• CRypTodiRa
_duRoCRypTodiRa
__TesTudinoidea
___eMysTeRnia
____eMydidae
____plaTysTeRnidae
___TesTuguRia
____geoeMydidae
____TesTudinidae
__aMeRiChelydia
___ChelydRoidea
____kinosTeRnoidea
_____kinosTeRnidae
_____deRMaTeMydidae
____ChelydRidae
___Chelonioidea
____Cheloniidae
____deRMoChelyidae
_TRionyChia
__TRionyChidae
__CaReTToChelyidae
While the TTWG continues to prefer and adhere to the Linnaean classification presented here to the left (a system fundamental to, and compliant with, the International Code of
Zoological Nomenclature), we recognize that alternative, nonCode-compliant chelonian classification arrangements have
also been proposed, and include the above Phylocode classification scheme presented by Crawford et al. (2015) as a recent
example that has found adoption by some systematists.
23
24
Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
TesTudines Batsch 1788
(07:1, 10:4, 12:6) (1)
Testudinata Klein 1751:96 (invalid pre-Linnaean name)
Testudines Linnaeus 1758:194 (vernacular)
Testudinata Klein in Behn 1760:tab.gen.
Testudines Batsch 1788:437
Testudinea Batsch 1796:179
Cheloniens Brongniart 1800a:196 (vernacular)
Chelonii Latreille 1800:xi
Chelonia Ross and Macartney 1802:tab.iii
Cataphractae Link 1807:51
Testudinata Oppel 1811:3
Perostia Rafinesque 1814:66
Cataphracta Hemprich 1820:101
Chelonea Fleming 1822:268
Fornicata Haworth 1825:373
Chelynae Wagler 1828:861
Sterrichrotes Ritgen 1828:269
Chelonites Burmeister 1837:730
Chelonides Swainson 1839:112
Tylopoda Mayer 1849:197
Testudina Fry 1850:21
Chersemydes Strauch 1862:16
Rhynchochelones Dollo 1886:79
Cheloniae Hoffmann 1890:372
Testudoformes Chang 1957:50
Chelonomorpha Kuhn 1960:30
Casichelydia Gaffney 1975:4
Testudinomorpha Laurin and Reisz 1995:197
Pantestudines Joyce, Parham, and Gauthier 2004:996
CRypTodiRa Cope 1868b
1758], by original monotypy.
Devisia Ogilby 1905:11
Type species: Devisia mythodes Ogilby 1905 [= subjective synonym of
Testudo serpentina Linnaeus 1758], by original monotypy.
Chelydra acutirostris Peters 1862
South American Snapping Turtle
Vivian P. Páez / Colombia [Cali Zoo]
(08:20)
Cryptodères Duméril and Bibron 1834:354
Cryptodera Lichtenstein 1856:1 (08:20)
Cryptodira Cope 1868b:282
ChelydRidae Gray 1831d (09:3)
Chelydrae Gray 1831d:4
Chelydridae Swainson 1839:113
Chelydradae Gray 1869a:178
Chelydra Schweigger 1812 (07:2)
Chelydra Schweigger 1812:292
Type species: Chelydra serpentina Schweigger [= Testudo serpentina
Linnaeus 1758], by subsequent designation by Fitzinger (1843:29).
Cheliurus Rafinesque 1815:75 (nomen nudum)
Chelonura Fleming 1822:270 (senior homonym, not = Chelonura Rafinesque 1832)
Colombia (Antioquia, Caldas, Cauca, Chocó, Córdoba, Nariño,
Quindio, Valle del Cauca), Costa Rica, Ecuador, Honduras,
Nicaragua, Panama
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Near Threatened (South America
regional) (2011)
Chelydra serpentina acutirostris Peters 1862:627, Chelydra
acutirostris
Type locality: “Guayaquil” [Ecuador].
Chelydra rossignonii (Bocourt 1868)
Central American Snapping Turtle
Type species: Chelonura serpentina [= Testudo serpentina Linnaeus
1758], by original monotypy.
Ophichelone Jarocki 1822:21
Type species: Ophichelone serpentina [= Testudo serpentina Linnaeus
1758], by original monotypy.
Rapara Gray 1825:210
Type species: Rapara serpentina Gray [= Testudo serpentina Linnaeus
1758], by original monotypy.
Saurochelys Latreille 1825:92
Type species: Saurochelys “Tortue à longue queue” [= Testudo serpentina Linnaeus 1758], by original monotypy.
Cheliurus Rafinesque 1832:64
Type species: Cheliurus serpentina [= Testudo serpentina Linnaeus
1758], by original monotypy.
Emysaurus Duméril and Bibron 1835:348
Type species: Emysaura serpentina [= Testudo serpentina Linnaeus
Richard C. Vogt / Laguna Oaxaca, Rio Lacantun, Selva Lacandona, Chiapas, Mexico
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Belize, Guatemala, Honduras, Mexico (Campeche, Chiapas,
Oaxaca, Tabasco, Veracruz)
IUCN Red List: Vulnerable A2d (2007)
Emysaurus rossignonii Bocourt 1868:121, Chelydra rossignonii, Chelydra serpentina rossignonii
Type locality: “marais de Pansos, près le Rio Polochic
(Guatémala).”
Chelydra serpentina mexicanae Cope in Gray 1870c:64 (nomen nudum)
Chelydra serpentina (Linnaeus 1758) (08:5)
North American Snapping Turtle, Common Snapping Turtle
Matthew Aresco / CRM 3 / Leon Co., Florida
(orange dots = introduced)
Canada (Manitoba, New Brunswick, Nova Scotia, Ontario,
Québec, Saskatchewan), USA (Alabama, Arkansas,
Colorado, Connecticut, Delaware, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maine,
Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, Montana, Nebraska, New Hampshire,
New Jersey, New Mexico, New York, North Carolina,
North Dakota, Ohio, Oklahoma, Pennsylvania, Rhode
Island, South Carolina, South Dakota, Tennessee, Texas,
Vermont, Virginia, West Virginia, Wisconsin, Wyoming)
Introduced: China, Japan (mainland), Taiwan, USA (Arizona,
California, Nevada, Oregon, Washington)
IUCN Red List: Least Concern (2012); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix III (USA)
Testudo serpentina Linnaeus 1758:199, Chelydra serpentina,
Emys serpentina, Chelonura serpentina, Ophichelone
serpentina, Rapara serpentina, Saurochelys serpentina,
Chelidra serpentina, Cheliurus serpentina, Emysaurus
serpentina, Emysaura serpentina, Hydraspis (Chelydra)
serpentina, Emysaurus serpentinus, Chelydra serpentina
serpentina
Type locality: “Calidis regionibus.” Restricted to “Algiriae, Chinae
aquis dulcibus” by Linnaeus (1766:354); to “New Orleans, La.”
[Louisiana, USA] by Smith and Taylor (1950a:358, 1950b:21);
and to “vicinity of New York City” [New York, USA] by Schmidt
(1953:86).
Testudo serpentaria Wiedemann 1802:191 (nomen novum)
Chelydra lacertina Schweigger 1812:293 (senior homonym,
not = Gypochelys lacertina Agassiz 1857a), Chelydra
serpentina lacertina
Type locality: Not known. Restricted to “vicinity of New York City”
[New York, USA] by Schmidt (1953:86).
Testudo serrata Pennant in Gray 1830e:14 (nomen nudum
and junior synonym, not = Testudo serrata Daudin 1801 or
Testudo serrata Shaw 1802)
Testudo longicauda Shaw in Gray 1831d:36 (nomen nudum)
Chelydra emarginata Agassiz 1857a:417
Type locality: “Mobile and New Orleans.” Restricted to “Mobile”
[Alabama, USA] by Schmidt (1953:86).
Devisia mythodes Ogilby 1905:11
Type locality: “Fly River, British New Guinea” [Papua New
Guinea] [in error].
Chelydra laticarinata † Hay 1916a:72 (nomen suppressum,
ICZN 1986) [Pleistocene, USA (Florida)]
Chelydra sculpta † Hay 1916a:73 (nomen suppressum, ICZN
1986) [Pleistocene, USA (Florida)]
Chelydra osceola Stejneger 1918:89 (08:5) (nomen conservandum, ICZN 1986), Chelydra serpentina osceola
Type locality: “Clearwater, Pinellas County, Florida” [USA].
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Macrochelys Gray 1856a (07:3) (2)
Macrochelys Gray 1856a:200
Macrochelys temminckii (Troost in Harlan 1835) (09:4, 14:1) (2)
Western Alligator Snapping Turtle
Type species: Macrochelys temminckii [= Chelonura temminckii Troost
in Harlan 1835], by original monotypy.
Macroclemys Gray 1856b:48 (nomen novum)
Gypochelys Agassiz 1857a:248, 413
Type species: Gypochelys lacertina Agassiz 1857a [= subjective synonym of Chelonura temminckii Troost in Harlan 1835], by original
monotypy.
Macroclemmys Strauch 1862:35 (nomen novum)
Macrochelys suwanniensis Thomas, Granatosky, Bourque, Krysko,
Moler, Gamble, Suarez, Leone, Enge, and Roman 2014 (2)
Suwannee Alligator Snapping Turtle
James C. Godwin / CCB / lower Tallapoosa R., Elmore Co., Alabama / juvenile
Kevin Enge / White Springs, Suwannee R., Florida
USA (Florida, Georgia)
IUCN Red List: Not Evaluated
CITES: Appendix III (USA)
Macrochelys maxhoseri Hoser 2013:56 (unavailable name
pending ICZN decision; Rhodin et al. 2015) (14:1) (3, 4)
Macrochelys suwanniensis Thomas, Granatosky, Bourque,
Krysko, Moler, Gamble, Suarez, Leone, Enge, and Roman
2014:150
Type locality: “Santa Fe River and State Road 235, Alachua County,
Florida (29.87872ºN, 82.33619ºW...elev. 23 m)” [USA].
USA (Alabama, Arkansas, Florida, Georgia, Illinois, Indiana,
Iowa, Kansas, Kentucky, Louisiana, Mississippi, Missouri,
Oklahoma, Tennessee, Texas)
IUCN Red List: Vulnerable A1cd (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix III (USA)
Testudo planitia Gmelin 1789:1045 (junior homonym, not =
Testudo planitia Meuschen 1778; nomen suppressum, ICZN
1963), Chersine planitia
Type locality: “Surinami” [Suriname] [in error].
Chelonura temminckii Troost in Harlan 1835:158 (nomen
conservandum, ICZN 1963), Emysaurus temminckii, Macrochelys temminckii, Macroclemys temminckii, Chelydra
temminckii, Gypochelys temminckii, Macrochelys temminckii temminckii
Type locality: “a tributary stream of the Mississippi, which enters
that river above Memphis, in west Tennessee.” Emended to “Wolf
River, Shelby County, Tennessee, USA” by Bour (1987b:343).
Gypochelys lacertina Agassiz 1857a:414 (junior homonym,
not = Chelydra lacertina Schweigger 1812), Macrochelys
lacertina, Macroclemys lacertina
Macrochelys temminckii muscati Hoser 2013:55 (unavailable
name pending ICZN decision; Rhodin et al. 2015) (14:1) (3, 4)
Macrochelys apalachicolae Thomas, Granatosky, Bourque,
Krysko, Moler, Gamble, Suarez, Leone, Enge, and Roman
2014:151(2)
Type locality: “Apalachicola River, Gadsden County, Florida” [USA].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelonioidea Oppel 1811
Chelonii Oppel 1811:8
Chlonopteria Rafinesque 1814:66
Cheloniae Schmid 1819:14
Edigitata Haworth 1825:373
Oiacopodae Wagler 1828:861
Chelonidae Bonaparte 1831:64
Oeacopodes Burmeister 1837:731
Pterodactyli Mayer 1849:199
Chelonioidea Baur 1893b:673
Cheloniidae Oppel 1811 (09:5, 12:7, 12:8, 12:9)
Chelonii Oppel 1811:8 (partim)
Cheloniadae Gray 1825:212
Carettidae Gray 1825:212
Mydae Ritgen 1828:269
Chelonina Bonaparte 1831:64
Cheloniidae Cope 1868b:282
Cheloniadi Portis 1890:23
CaReTTinae Gray 1825 (12:7)
Carettidae Gray 1825:212
Carettinae Deraniyagala 1952:57
Caretta Rafinesque 1814 (12:8)
Caretta Rafinesque 1814:66
Type species: Caretta nasuta Rafinesque 1814 [= subjective synonym
of Testudo caretta Linnaeus 1758], by original monotypy.
Thalassochelys Fitzinger 1835:121
Type species: Thalassochelys caouana Fitzinger [= Testudo caouana
Lacepède 1788 (nomen suppressum) = Testudo caouana Bonnaterre
1789 = subjective synonym of Testudo caretta Linnaeus 1758], by
subsequent designation by Fitzinger (1843:30).
Caouana Cocteau and Bibron 1838:31
Type species: Chelonia (Caouana) cephalo [= Testudo cephalo Schneider 1783 = subjective synonym of Testudo caretta Linnaeus 1758],
by original monotypy.
Halichelys Fitzinger 1843:30
Type species: Thalassochelys (Halichelys) atra Fitzinger [= Caretta
atra Merrem 1820 = subjective synonym of Testudo caretta Linnaeus 1758], by original designation.
Eremonia Gray 1873i:408
Type species: Eremonia elongata Gray [= Caouana elongata Gray
1844 = subjective synonym of Testudo caretta Linnaeus 1758], by
original designation.
Caretta caretta (Linnaeus 1758) (10:5, 14:2) (5)
Loggerhead, Loggerhead Sea Turtle
Blair Witherington / CRM 3 / Atlantic Ocean off the Florida Keys, USA
(lines delimit Regional Management Units)
Nesting: Aruba, Australia (Queensland, Western Australia),
Bahamas, Bangladesh, Belize, Bermuda, Bonaire, Brazil
(Bahia, Espirito Santo, Rio de Janeiro, Sergipe), Cape
Verde, Cayman Islands, China, Colombia, Costa Rica,
Cuba, Curacao, Cyprus, Dominican Republic, Egypt,
France, Greece, Haiti, Honduras, Israel, Italy, Japan, Lebanon, Libya, Madagascar, Mauritania, Mexico, Montserrat,
Mozambique, Myanmar, New Caledonia, Oman, Panama,
Papua New Guinea (Trobriand Islands), Saint Lucia, Saint
Vincent and the Grenadines, Sierra Leone, South Africa,
Spain, Sri Lanka, Syria, Tunisia, Turkey, Turks and Caicos,
USA (Alabama, Florida, Georgia, Louisiana, Mississippi,
North Carolina, South Carolina, Texas), US Virgin Islands,
Vanuatu, Venezuela, Yemen
Foraging: Albania, Algeria, Anguilla, Antigua and Barbuda,
Argentina, Bahrain, Barbados, British Virgin Islands,
Canada, Chile, Comoros, Croatia, Djibouti, Dominica, Eritrea, Fiji, French Guiana, Gambia, Grenada, Guadeloupe,
Guinea-Bissau, Guatemala, Guyana, India, Indonesia, Iran,
Jamaica, Kenya, Malta, Martinique, Mauritius, Monaco,
Montenegro, Morocco, Namibia, Netherlands Antilles
(Bonaire, Saba, St. Eustatius), Nicaragua, North Korea,
Pakistan, Peru, Philippines, Portugal, Puerto Rico, Qatar,
Réunion, Saint Kitts and Nevis, Samoa, Saudi Arabia,
Senegal, Seychelles, Sint Maarten, Slovenia, Solomon
Islands, Somalia, South Korea, Sudan, Suriname, Taiwan,
Tanzania, Tonga, Trinidad and Tobago, United Arab
Emirates, Uruguay, USA (California, Hawaii, Oregon),
Vietnam, Western Sahara
Vagrant: Angola, Benin, Brunei, Cambodia, Cameroon, Congo
(DRC), Congo (ROC), Ecuador, El Salvador, Equatorial Guinea, Gabon, Ghana, Great Britain, Guinea, Iraq,
Ireland, Ivory Coast, Kuwait, Liberia, Malaysia, Maldives,
New Zealand, Nigeria, Thailand, Togo, Tuvalu
IUCN Red List: Global: Vulnerable A2b (2015); Previously:
Endangered (1996); Subpopulations: Mediterranean: Least
Concern (2015); North East Atlantic: Endangered B2ab(iii)
(2015); North East Indian Ocean: Critically Endangered D
(2015); North Pacific: Least Concern (2015); North West
Atlantic: Least Concern (2015); North West Indian Ocean:
Critically Endangered A4b (2015); South East Indian
Ocean: Near Threatened (2015); South Pacific: Critically Endangered A2b (2015); South West Atlantic: Least
Concern (2015); South West Indian Ocean: Near Threatened
(2015)
CITES: Appendix I, as Cheloniidae spp.
Testudo caretta Linnaeus 1758:197, Chelone caretta, Chelonia
caretta, Thalassochelys caretta, Talassochelys caretta, Caouana caretta, Caretta caretta, Caretta caretta caretta
Type locality: “insulas Americanas.” Restricted to “Mari Mediterraneo, Atlantico” by Schoepff (1793:70); to “Bermuda Islands” by
Smith and Taylor (1950a:315, 195b:16); and to “Bimini, British
27
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Bahamas” by Schmidt (1953:107).
Testudo marina Garsault 1764:pl.675 (10:5)
Testudo cephalo Schneider 1783:303, Caretta cephalo, Chelonia cephalo, Thalassochelys cephalo
Eretmochelys imbricata (Linnaeus 1766) (07:5, 09:9, 12:9, 14:4) (6)
Hawksbill Turtle, Hawksbill Sea Turtle
Type locality: Not designated. Restricted to “Charleston, South
Carolina” [USA] by Smith and Taylor (1950a:360).
Testudo caouana Lacepède 1788:95, synopsis[table] (09:6) (nomen
suppressum, ICZN 2005a)
Type locality: “les contrées chaudes du nouveau Continent.”
Testudo nasicornis Lacepède 1788:103, synopsis[table] (09:7)
(nomen suppressum, ICZN 2005a), Testudo caretta nasicornis, Caretta nasicornis
Type locality: “mers du nouveau Continent, voisines de l’équateur.”
Restricted to “Ascension Island” by Smith and Smith (1980:302).
Testudo caouana Bonnaterre 1789:20, Chelonia caouana,
Caretta caouana, Thalassochelys caouana
Testudo lauanna Meyer 1790:82 (09:8) (nomen novum et
oblitum)
Testudo gigas Walbaum in Donndorff 1798:35
Robert P. Van Dam / CCB / Mona Island, Puerto Rico
Type locality: Not designated. Restricted to “Westindischen Meere”
by Bechstein (1800:273).
Caretta nasuta Rafinesque 1814:66
Type locality: “Sicil.” [Sicily, Italy].
Caretta atra Merrem 1820:17, Chelonia (Thalassochelys)
atra, Thalassochelys atra, Thalassochelys (Halichelys)
atra, Halichelys atra
Type locality: “mari ad Insulam Adscensionis” [Ascension].
Testudo corianna Gray 1831d:53 (nomen novum)
Chelonia pelasgorum Valenciennes in Bory de Saint-Vincent
1833:planches, pl.6 (14:2)
Type locality: “sur la plage sablonneuse entre Arcadia et
l’embouchure de la Neda” [Greece].
Chelonia pelasgica Bibron and Bory de Saint-Vincent 1833:64
(nomen novum) (14:2)
Caouana elongata Gray 1844:53, Thalassochelys elongata,
Eremonia elongata
Type locality: Not designated. Restricted to “Ascension Island” by
Smith and Smith (1980:303).
Thalassochelys corticata Girard 1858:431
Type locality: “Madeira” [Canary Islands]. Restricted to “Funchal,
Madeira (Portugal)” by Cochran (1961:235).
Caretta gigas Deraniyagala 1933:66, Caretta caretta gigas
Type locality: “Ceylon” [Sri Lanka]. Restricted to “Gulf of Mannar...
Ceylon” [Sri Lanka] by Deraniyagala (1939:164).
Eretmochelys Fitzinger 1843 (09:5, 12:7, 12:8, 14:3)
Eretmochelys Fitzinger 1843:30
Type species: Chelonia (Eretmochelys) imbricata Cuvier [= Testudo
imbricata Linnaeus 1766], by original designation.
Herpysmostes Gistel 1868:145
Type species: Herpysmostes imbricatus [= Testudo imbricata Linnaeus
1766], by original monotypy.
Onychochelys Gray 1873i:397
Type species: Onychochelys kraussi Gray 1873i [= subjective synonym
of Testudo imbricata Linnaeus 1766], by original monotypy.
(lines delimit Regional Management Units)
Nesting: Anguilla, Antigua and Barbuda, Aruba, Australia,
Bahamas, Bangladesh, Barbados, Belize, Brazil, British
Indian Ocean Territory, British Virgin Islands, Cameroon,
Cayman Islands, China, Colombia, Congo (ROC), Costa
Rica, Cuba, Curacao, Dominica, Dominican Republic,
Ecuador, Egypt, El Salvador, Equatorial Guinea, Eritrea,
Fiji, French Guiana, French Southern Territories, Gabon,
Grenada, Guadeloupe, Guatemala, Guinea, Guinea-Bissau,
Guyana, Haiti, Honduras, India, Indonesia, Iran, Ivory
Coast, Jamaica, Japan, Kenya, Liberia, Madagascar,
Malaysia, Maldives, Martinique, Mexico (Campeche,
Yucatán), Micronesia, Montserrat, Mozambique, Netherlands Antilles (Bonaire, Sint Eustatius), Nicaragua, Oman,
Palau, Panama, Papua New Guinea, Philippines, Puerto
Rico, Qatar, Saint Kitts and Nevis, Saint Lucia, Saint Vincent and the Grenadines, Samoa, São Tomé and Príncipe,
Saudi Arabia, Senegal, Seychelles, Sint Maarten, Solomon
Islands, Sri Lanka, Suriname, Taiwan, Tanzania, Thailand,
Trinidad and Tobago, Turks and Caicos, USA (Florida,
Hawaii), US Virgin Islands, Vanuatu, Venezuela, Vietnam
Foraging: American Samoa, Argentina, Ascension, Bahrain, Benin, Bermuda, Brunei, Cambodia, Cook Islands, Djibouti,
French Polynesia, Gambia, Ghana, Guam, Iraq, Israel, Kuwait, Mauritania, Mauritius, Mayotte, Myanmar, Nigeria,
Northern Mariana Islands, Pakistan, Peru, Réunion, Sierra
Leone, Singapore, Society Islands, Somalia, South Africa,
Sudan, Togo, Tokelau, Tonga, Tuamotu, Tuvalu, United
Arab Emirates, Wallis and Futuna, Yemen
Vagrant: Algeria, Angola, Cape Verde, Chile, Comoros, Congo
(DRC), Kiribati, Marshall Islands, Morocco, Namibia,
Nauru, New Caledonia, North Korea, Pitcairn Island,
Portugal, South Korea, Spain, Uruguay
IUCN Red List: Critically Endangered A2bd (2008); Previously: Critically Endangered (1996)
CITES: Appendix I, as Cheloniidae spp.
Testudo imbricata Linnaeus 1766:350, Chelone imbricata,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelonia imbricata, Caretta imbricata, Eretmochelys
imbricata, Herpysmostes imbricatus, Herpysmostes
imbricata, Chelonius imbricatus, Eretmochelys imbricata
imbricata
Lepidochelys kempii (Garman 1880)
Kemp’s Ridley, Kemp’s Ridley Sea Turtle, Atlantic Ridley
Type locality: “Mari Americano, Asiatico.” Restricted to “Bermuda
Islands” by Smith and Taylor (1950a:315, 1950b:17), and to “Belize, British Honduras” by Schmidt (1953:106).
Testudo nasicornis Bonnaterre 1789:21 (09:7)
Chelonia radiata Cuvier 1829:14
Type locality: Not designated.
Chelonia pseudomydas Lesson 1831b:299
Type locality: “l’Océan atlantique.” Restricted to “Bermuda
Islands” by Smith and Taylor (1950a:315).
Chelonia pseudocaretta Lesson 1831b:302
Type locality: “l’Océan atlantique.” Restricted to “Bermuda
Islands” by Smith and Taylor (1950a:315).
Caretta bissa Rüppell 1835:4 (07:5, 09:9), Eretmochelys imbricata bissa
Blair Witherington / CRM 3 / Florida Bay, Monroe Co., Florida
Type locality: “im rothen Meere...Abyssinien” [Red Sea...Ethiopia].
Eretmochelys squamata Agassiz 1857a:382, Caretta squamata, Eretmochelys imbricata squamata
Type locality: “Singapore and Bengal, India.” Restricted to
“Singapore, Straits Settlements” by Smith and Taylor (1950a:315,
1950b:17).
Caretta squamosa Girard 1858:442 (nomen novum), Eretmochelys squamosa, Eretmochelys imbricata squamosa
Caretta rostrata Girard 1858:446
Type locality: “Feejee Islands” [Fiji].
Onychochelys kraussi Gray 1873i:398
Type locality: “Ocean, French Guiana.”
Lepidochelys Fitzinger 1843 (12:8)
Lepidochelys Fitzinger 1843:30
Type species: Thalassochelys (Lepidochelys) olivacea Eschscholtz
1829a [= Chelonia olivacea Eschscholtz 1829a], by original
designation.
Cephalochelys Gray 1873i:408
Type species: Cephalochelys oceanica Gray 1873i [= subjective
synonym of Chelonia olivacea Eschscholtz 1829a], by original
monotypy.
Colpochelys Garman 1880:124
Type species: Thalassochelys (Colpochelys) kempii Garman 1880, by
original monotypy.
Nesting: Mexico (Tamaulipas, Veracruz), USA (Texas)
Foraging: USA (Alabama, Connecticut, Delaware, Florida,
Georgia, Louisiana, Maryland, Massachusetts, Mississippi,
New Jersey, New York, North Carolina, Rhode Island,
South Carolina, Virginia)
Vagrant: Algeria, Anguilla, Bahamas, Bermuda, British Virgin
Islands, Canada, Cayman Islands, Cuba, France, Ireland,
Italy, Morocco, Portugal, Spain
IUCN Red List: Critically Endangered A1ab (1996)
CITES: Appendix I, as Cheloniidae spp.
Testudo viridisquamosa Lacepède 1788:92, synopsis[table]
(09:6)
(partim, nomen dubium et suppressum, ICZN 1963)
Testudo viridisquamosa Bonnaterre 1789:20 (partim, nomen
dubium)
Testudo bomarii Meyer 1790:82 (09:8) (partim, nomen dubium
et novum et oblitum)
Testudo mydas minor Suckow 1798:30 (partim, nomen
dubium et suppressum, ICZN 1963)
Type locality: “Amazonen-Flusse..[&]..Südsee..[&]..Cap Blanco
in Mexico.” Restricted to “the island of Blanquilla...West Indies”
[Venezuela] by Brongersma (1961:27).
Thalassochelys (Colpochelys) kempii Garman 1880:123 (nomen
conservandum, ICZN 1963), Lepidochelys kempii, Colpochelys kempii, Caretta kempii, Lepidochelys olivacea kempii
Type locality: “Gulf of Mexico.” Restricted to “Key West,
Monroe Co., Florida” [USA] by Smith and Taylor (1950a:358,
1950b:15).
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Lepidochelys olivacea (Eschscholtz 1829a) (7)
Olive Ridley, Olive Ridley Sea Turtle, Pacific Ridley
ICZN 1963)
Type locality: Not designated.
Chelonia olivacea Eschscholtz 1829a:15 (7), Chelonia caretta
olivacea, Caretta olivacea, Thalassochelys (Lepidochelys)
olivacea, Caouana olivacea, Lepidochelys olivacea,
Caretta caretta olivacea, Lepidochelys olivacea olivacea,
Caretta olivacea olivacea
Type locality: “Bai von Manilla” [Philippines].
Chelonia dussumierii Duméril and Bibron 1835:557 (nomen
novum), Lepidochelys dussumierii
Caouana ruppellii Gray 1844:53 (nomen nudum)
Type locality: “India?”
Chelonia subcarinata Rüppell in Gray 1844:53 (nomen
nudum)
Chelonia polyaspis Bleeker 1857b:239 (nomen nudum)
Nicolas J. Pilcher / Red Sea, nr. Assab, Eritrea
Type locality: “Batavia...Java” [Indonesia].
Chelonia dubia Bleeker in Gray 1864a:13 (nomen nudum)
Type locality: Not designated.
Cephalochelys oceanica Gray 1873i:408
Type locality: “West Coast of America – Mexico?”
Thalassiochelys tarapacona Philippi 1887:85, Thalassochelys tarapacona
Type locality: “Iquique...Chile.”
Thalassochelys controversa Philippi 1899:731
Type locality: “Quinteros...Chile.”
Caretta remivaga Hay 1908a:194, Lepidochelys olivacea
remivaga
(lines delimit Regional Management Units)
Nesting: Angola, Australia (Northern Territory), Bangladesh,
Benin, Brazil (Bahia, Espirito Santo, Sergipe), Brunei, Cameroon, Colombia, Congo (ROC), Costa Rica,
Ecuador, El Salvador, Equatorial Guinea, Eritrea, French
Guiana, Gabon, Ghana, Guatemala, Guinea-Bissau, Guyana, Honduras, India, Indonesia (Java, Papua), Iran, Ivory
Coast, Kenya, Liberia, Malaysia, Mexico (Baja California
Sur, Chiapas, Guerrero, Jalisco, Michoacán, Nayarit, Oaxaca, Sinaloa), Mozambique, Myanmar, Nicaragua, Oman,
Pakistan, Panama, Peru, São Tomé and Príncipe, Sierra
Leone, Sri Lanka, Suriname, Thailand, Togo, Trinidad and
Tobago, Vanuatu, Vietnam
Foraging: Bahrain, Cambodia, Cape Verde, China, Comoros,
Congo (DRC), Djibouti, Egypt, Gambia, Guinea, Iran,
Iraq, Israel (Southern), Kuwait, Liberia, Madagascar,
Maldives, Mauritius, New Caledonia, Nigeria, Papua
New Guinea, Philippines, Qatar, Saudi Arabia, Senegal,
Seychelles, Singapore, Solomon Islands, Somalia, Sudan,
Tanzania, Timor-Leste, United Arab Emirates, USA (Hawaii), Venezuela, Yemen
Vagrant: Antigua and Barbuda, Barbados, Canada, Chile, Cuba,
Dominica, Dominican Republic, Grenada, Guadeloupe,
Haiti, Jamaica, Japan, Marshall Islands, Martinique,
Mauritania, Micronesia, Morocco, Namibia, New Zealand,
North Korea, Puerto Rico, Saint Kitts and Nevis, Saint
Lucia, Saint Vincent and the Grenadines, South Africa,
South Korea, Spain, Taiwan, Uruguay, US Virgin Islands
IUCN Red List: Vulnerable A2bd (2008); Previously: Endangered (1996)
CITES: Appendix I, as Cheloniidae spp.
Testudo mydas minor Suckow 1798:30 (partim, nomen
dubium et suppressum, ICZN 1963)
Type locality: “Amazonen-Flusse..[&]..Südsee..[&]..Cap Blanco
in Mexico.” Restricted to “the island of Blanquilla...West Indies”
[Venezuela] by Brongersma (1961:27).
Chelonia multiscutata Kuhl 1820:78 (nomen suppressum,
Type locality: “Ventosa Bay, Gulf of Tehuantepec, Oaxaca,
Mexico.”
Cheloniinae Oppel 1811 (12:7)
Chelonii Oppel 1811:8 (partim)
Cheloniadae Gray 1825:212
Mydae Ritgen 1828:269
Cheloniidae Cope 1868b:282
Chelonia Brongniart 1800 (09:5, 12:8)
Chelonia Brongniart 1800b:89
Type species: Chelonia mydas [= Testudo mydas Linnaeus 1758], by
subsequent designation by Bell (1828c:516).
Chelone Brongniart 1805:610 (nomen novum)
Chelona Fleming 1828:149 (nomen novum)
Mydas Cocteau and Bibron 1838:22
Type species: Chelonia (Mydas) viridis [= Testudo viridis Schneider
1783 = subjective synonym of Testudo mydas Linnaeus 1758], by
tautonymy.
Mydasea Gervais 1843:457
Type species: Chelonia (Mydasea) mydas [= Testudo mydas Linnaeus
1758], by tautonymy.
Euchelonia Tschudi 1846:22
Type species: Chelonia (Euchelonia) midas Schweigger [= Testudo
mydas Linnaeus 1758], by original monotypy.
Megemys Gistel 1848:8 (nomen novum)
Euchelys Girard 1858:447
Type species: Euchelys macropus Girard [= Testudo macropus
Walbaum 1782 = subjective synonym of Testudo mydas Linnaeus
1758], by original monotypy.
Midas Herrera 1901:68 (nomen novum et suppressum, ICZN 1922)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelonia mydas (Linnaeus 1758) (07:4, 09:5, 12:9, 12:10) (8)
Green Turtle, Green Sea Turtle
Mariana Islands, Morocco, Nauru, New Zealand, Portugal,
Saint Helena, Slovenia, Solomon Islands, Spain, Tokelau,
Tuamotu, Tunisia, Tuvalu
IUCN Red List: Global: Endangered A2bd (2004); Previously:
Endangered (1996); Subpopulations: Hawaiian: Least
Concern (2012)
CITES: Appendix I, as Cheloniidae spp.
Testudo mydas Linnaeus 1758:197, Chelonia mydas, Chelone
mydas, Caretta mydas, Mydas mydas, Mydasea mydas,
Chelonia (Euchelonia) midas, Euchelonia mydas, Megemys mydas, Chelonia mydas mydas
Type locality: “insulas Pelagi: insulam Adscensionis.” Restricted to
“Insel Ascension” by Mertens and Müller (1928:23).
Blair Witherington / CRM 3 / Atlantic Ocean off Florida Keys, Florida, USA
Testudo macropus Walbaum 1782:112 (unavailable name),
Euchelys macropus, Chelone macropus
Testudo viridis Schneider 1783:299, Chelonia viridis,
Chelone viridis, Chelonia (Mydas) viridis, Mydas viridis,
Chelonia mydas viridis
Type locality: Not designated. Restricted to “Charleston, South
Carolina” [USA] by Smith and Taylor (1950a:360).
Testudo japonica Thunberg 1787:178, Chelonia japonica,
Chelonia mydas japonica
Type locality: “Japan.”
Testudo marina vulgaris Lacepède 1788:54, synopsis[table]
(09:6)
(nomen suppressum, ICZN 2005a)
Type locality: “contrées équatoriales.”
(lines delimit Distinctive Population Segments)
Nesting: American Samoa, Angola, Anguilla, Antigua and
Barbuda, Aruba, Ascension, Australia (Northern Territory,
Queensland, Western Australia), Bahamas, Bangladesh,
Barbados, Belize, Brazil, British Virgin Islands, Cayman Islands, China, Cocos (Keeling) Islands, Colombia,
Comoros, Congo (ROC), Costa Rica, Cuba, Curacao,
Cyprus, Dominica, Dominican Republic, Ecuador, Egypt, El
Salvador, Equatorial Guinea, Eritrea, French Guiana, French
Polynesia, French Southern Territories, Gambia, Grenada,
Guadeloupe, Guam, Guinea-Bissau, Guyana, Haiti, Honduras, India, Indonesia, Iran, Jamaica, Japan, Kenya, Lebanon,
Madagascar, Malaysia, Martinique, Mauritania, Mayotte,
Mexico (Baja California, Campeche, Michoacán, Quintana
Roo, Sinaloa, Sonora, Tabasco, Tamaulipas, Veracruz,
Yucatán), Micronesia, Montserrat, Mozambique, Myanmar,
Netherlands Antilles (Bonaire, Sint Eustatius), Nicaragua,
Oman, Pakistan, Panama, Papua New Guinea, Peru, Philippines, Puerto Rico, Réunion, Saint Kitts and Nevis, Saint
Lucia, Saint Vincent and the Grenadines, São Tomé and
Príncipe, Saudi Arabia, Senegal, Seychelles, Sint Maarten,
Sri Lanka, Suriname, Syria, Taiwan, Tanzania, Thailand,
Trinidad and Tobago, Turkey, Turks and Caicos, USA
(Florida, Hawaii), US Virgin Islands, Vanuatu, Venezuela,
Vietnam, Yemen
Foraging: Argentina, Bahrain, Benin, Bermuda, British Indian
Ocean Territory, Brunei, Cambodia, Cameroon, Cape Verde,
Chile, Christmas Island, Congo (DRC), Cook Islands,
Djibouti, Fiji, Gabon, Ghana, Greece, Guatemala, Guinea,
Iraq, Israel, Ivory Coast, Kuwait, Liberia, Libya, Maldives,
Marshall Islands, Mauritius, Namibia, New Caledonia, Nigeria, Niue, Palau, Qatar, Samoa, Sierra Leone, Singapore,
Somalia, South Africa, Sudan, Timor-Leste, Togo, Tonga,
United Arab Emirates, Uruguay, Wallis and Futuna, Western
Sahara
Vagrant: Algeria, Canada, Italy, Kiribati, Malta, Northern
Testudo viridisquamosa Lacepède 1788:92, synopsis[table] (09:6)
(partim, nomen dubium et suppressum, ICZN 1963)
Type locality: “la mer du Sud, auprès du cap Blanco, de la nouvelle
Espagne.” Restricted to “Bocas del Toro, Panama, Golf von
Mexico” by Wermuth (1956:405); and to “the island of Blanquilla...
West Indies” [Venezuela] by Brongersma (1961:27).
Testudo viridisquamosa Bonnaterre 1789:20 (partim, nomen
dubium)
Testudo macropus Gmelin 1789:1038
Type locality: Not designated.
Testudo bomarii Meyer 1790:82 (09:8) (partim, nomen dubium
et novum et oblitum)
Testudo chloronotus Bechstein 1800:107
Testudo rugosa Van-Ernest in Daudin 1801:37 (senior homonym, not = Testudo rugosa Shaw 1802)
Type locality: “la mer des Indes...près de la ligne à environ trois
dégrés des îles Maldives.”
Testudo cepediana Daudin 1801:50
Type locality: Not designated.
Chelonia virgata Schweigger 1812:291, Caretta virgata,
Chelonia (Mydas) virgata, Mydas virgata, Chelone virgata
Type locality: “mari sub zona torrida.” Restricted to “Bermuda
Islands” by Smith and Taylor (1950a:315).
Caretta cepedii Merrem 1820:18 (nomen novum)
Caretta esculenta Merrem 1820:18, Chelonia esculenta
Type locality: “Oceano Atlantico.”
Caretta thunbergii Merrem 1820:19 (nomen novum)
Type locality: “Japonia” [Japan].
Chelonia castanea Eschscholtz 1829a:11 (14:5) (7) (nomen
oblitum)
Type locality: “karaibischen Meere; küste von Surinam.”
Chelonia grisea Eschscholtz 1829a:13 (14:4) (7), Chelonia
griseam
Type locality: “kaspische Meer” [Caspian Sea] [in error].
Chelonia maculosa Cuvier 1829:13, Chelone maculosa
Type locality: Not designated. Restricted to “Ascension Island” by
Smith and Taylor (1950a:315).
Chelonia lachrymata Cuvier 1829:13
Type locality: Not designated.
Chelonia midas Wagler 1830b:133 (nomen novum)
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Chelonia bicarinata Lesson 1831b:301
Type locality: “l’Océan atlantique.”
Chelonia marmorata Duméril and Bibron 1835:546, Chelone
marmorata
Type locality: “l’île de l’Ascension.”
Chelonia formosa Girard 1858:456
Type locality: “Feejee Islands” [Fiji].
Chelonia tenuis Girard 1858:459
Type locality: “Honden Island, Paumotu Group; Tahiti and Eimo;
Rosa Island.” Rosa Island identified by Hirth (1980:1) as Rose
Atoll, American Samoa.
Chelonia albiventer Nardo 1864:1420
Type locality: “Adriatico...prossimata del porto di Malamocco”
[Italy].
Chelonia agassizii Bocourt 1868:122 (07:4), Chelonia mydas
agassizii
Type locality: “embouchure du Nagualate...Pacifique (Guatémala)”
[mouth of Rio Nagualate, Pacific coast of Guatemala].
Chelonia lata Philippi 1887:84
Type locality: “Valparaiso..[&]..Insel Chiloe” [Chile].
Chelonia mydas carrinegra Caldwell 1962:4
Type locality: “waters adjacent to Isla Angel de la Guarda, Bahia de
Los Angeles, central Gulf of California, Mexico.”
Testudo nigrita Tamayo 1962:358 (nomen nudum)
Natator McCulloch 1908 (12:8)
Natator McCulloch 1908:127
Type species: Natator tessellatus McCulloch 1908 [= subjective synonym of Chelonia depressa Garman 1880], by original monotypy.
Natator depressus (Garman 1880)
Flatback, Flatback Sea Turtle
“North Australia” by Loveridge (1934:261).
Natator tessellatus McCulloch 1908:127
Type locality: “near Port Darwin, North Australia.”
deRMoChelyidae Fitzinger 1843 (12:9)
Sphargidae Gray 1825:212
Sphargidina Bonaparte 1831:64
Dermatochelydae Fitzinger 1843:30
Athecae Cope 1871:235
Athecata Lydekker 1889:223
Dermochelyidae Lydekker 1889:223
Dermochelys Blainville 1816
Chelonias Rafinesque 1814:66 (nomen oblitum)
Chelyra Rafinesque 1815:74 (nomen nudum)
Dermochelys Blainville 1816:119 [“111”]
Type species: Dermochelys coriacea [= Testudo coriacea Vandelli
1761], by subsequent monotypy by Boulenger (1889:10), in accordance with ICZN Article 67.2.2.
Sphargis Merrem 1820:19
Type species: Sphargis mercurialis Merrem 1820 [= subjective synonym of Testudo coriacea Vandelli 1761], by original monotypy.
Coriudo Fleming 1822:271
Type species: Coriudo coriacea [= Testudo coriacea Vandelli 1761], by
original monotypy.
Scytina Wagler 1828:861 (nomen novum)
Dermochelis Cuvier 1829:14 (nomen novum)
Dermatochelys Wagler 1830b:133 (nomen novum) (9)
Chelyra Rafinesque 1832:64
Type species: Chelyra coriacea [= Testudo coriacea Vandelli 1761], by
original designation.
Dermochelys coriacea (Vandelli 1761) (12:9, 14:6, 14:7) (10)
Leatherback, Leatherback Sea Turtle
Carmen Pilcher / Shoalwater Bay, Queensland
Chris Johnson / CRM 3 / Juno Beach, Palm Beach Co., Florida, USA
(line delimits Regional Management Units)
Nesting: Australia (Northern Territory, Queensland, Western
Australia)
Foraging: Indonesia (Papua), Papua New Guinea (Southern)
Vagrant: Indonesia (Java, Lesser Sundas), Timor-Leste
IUCN Red List: Data Deficient (1996)
CITES: Appendix I, as Cheloniidae spp.
Chelonia depressa Garman 1880:124, Chelonia depressus,
Natator depressus, Natator depressa
Type locality: “East Indies and North Australia.” Restricted to
(lines delimit Regional Management Units)
Nesting: Angola, Anguilla, Antigua and Barbuda, Aruba,
Australia (Northern Territory), Bahamas, Bangladesh,
Barbados, Benin, Brazil, British Virgin Islands, Cameroon,
Colombia, Congo (ROC), Costa Rica, Cuba, Curacao,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Dominica, Dominican Republic, Ecuador, El Salvador,
Equatorial Guinea, French Guiana, Gabon, Ghana, Grenada, Guadeloupe, Guatemala, Guyana, Haiti, Honduras,
India, Indonesia (Java, Papua), Ivory Coast, Jamaica, Malaysia, Martinique, Mexico (Baja California Sur, Guerrero,
Jalisco, Michoacán, Oaxaca), Mozambique, Netherlands
Antilles (Bonaire, Sint Eustatius), Nicaragua, Panama,
Papua New Guinea (Northern), Puerto Rico, Saint Kitts
and Nevis, Saint Lucia, Saint Vincent and the Grenadines,
São Tomé and Príncipe, Sierra Leone, Sint Maarten, Solomon Islands, South Africa, Sri Lanka, Suriname, Thailand,
Togo, Trinidad and Tobago, USA (Florida), US Virgin
Islands, Vanuatu, Venezuela, Vietnam
Foraging: Albania, Algeria, Argentina, Belize, Brunei, Cambodia, Canada (British Columbia, New Brunswick, Newfoundland, Nova Scotia), Chile, China, Comoros, Congo
(DRC), Croatia, Cyprus, Egypt, Fiji, France, Gambia,
Great Britain, Greece, Guinea, Guinea-Bissau, Ireland,
Israel, Italy, Japan, Kenya, Kiribati, Lebanon, Liberia,
Libya, Madagascar, Malta, Marshall Islands, Mauritania,
Mauritius, Micronesia, Montenegro, Monaco, Morocco,
Myanmar, Namibia, Nauru, New Zealand, Nigeria, North
Korea, Palau, Peru, Philippines, Portugal, Russia, Samoa,
Senegal, Seychelles, Slovenia, South Korea, Spain, Syria,
Taiwan, Tanzania, Tonga, Tunisia, Turkey, Turks and
Caicos, Tuvalu, Uruguay, USA (Alaska, California, Connecticut, Delaware, Georgia, Hawaii, Maine, Maryland,
Massachusetts, New Jersey, New York, North Carolina,
Oregon, Rhode Island, South Carolina, Washington)
Vagrant: Bahrain, Denmark, Djibouti, Eritrea, Iceland, Iran,
Iraq, Kuwait, Maldives, Norway, Oman, Pakistan, Qatar,
Saudi Arabia, Somalia, Sudan, Sweden, United Arab Emirates, Yemen
IUCN Red List: Global: Vulnerable A2bd (2013); Subpopulations: East Pacific Ocean: Critically Endangered A2bd+4bd
(2013); Northeast Indian Ocean: Data Deficient (2013);
Northwest Atlantic Ocean: Least Concern (2013); Southeast
Atlantic Ocean: Data Deficient (2013); Southwest Atlantic
Ocean: Critically Endangered D (2013); Southwest Indian
Ocean: Critically Endangered C2a(ii) (2013); West Pacific
Ocean: Critically Endangered A2bd+4bd (2013)
CITES: Appendix I
Testudo coriacea Vandelli 1761:1 (senior homonym), Chelone coriacea, Chelonia coriacea, Dermochelys coriacea,
Coriudo coriacea, Scytina coriacea, Sphargis coriacea,
Dermatochelys coriacea, Dermochelys coriacea coriacea,
Chelyra coriacea
Type locality: “maris Tyrrheni oram in agro Laurentiano”
[Italy]. Restricted to “Palermo, Sicily” [Italy] by Smith and
Taylor (1950a:315, 1950b:13); to “la côte romaine (Italie), Mer
Tyrrhénienne, Méditerranée occidentale” [Italy] by Fretey and Bour
(1980:198); and to “Laurentum, between Lido di Ostia and Tor
Paterno, shore of the Tyrrhenian Sea, Italy” by Bour and Dubois
(1984:359).
Testudo coriacea Linnaeus 1766:350 (junior homonym)
Type locality: “Mari mediterraneo, Adriatico.” Restricted to “Palermo, Sicily” [Italy] by Smith and Taylor (1950a:315, 1950b:13).
Testudo arcuata Catesby 1771:40
Type locality: Not designated. Restricted to “Küsten-Gebiete von
Carolina und Florida” [USA] by Mertens and Wermuth (1955:386).
Testudo lyra Lacepède 1788:111, synopsis[table] (09:6) (nomen
suppressum, ICZN 2005a)
Type locality: “Méditerranée.”
Testudo lyra Bonnaterre 1789:22, Chelonia lyra
Testudo tuberculata Pennant in Schoepff 1801:123, Sphargis
tuberculata, Dermochelydis tuberculata
Type locality: Not designated. Restricted to “Palermo, Sicily” [Italy]
by Smith and Smith (1980:244).
Chelonias lutaria Rafinesque 1814:66
Type locality: “Sicil.” [Sicily, Italy].
Sphargis mercurialis Merrem 1820:19 (nomen novum et suppressum, ICZN 1956)
Type locality: “Mari mediterraneo et Oceano atlantico.”
Dermochelis atlantica LeSueur in Cuvier 1829:14 (nomen
nudum), Dermochelys atlantica, Dermatochelys atlantica
Dermatochelys porcata Wagler 1830c:explicatio tabularum
(nomen novum) (14:6) (9)
Testudo coriacea marina Ranzani 1832:3 (14:7)
Sphargis coriacea schlegelii Garman 1884:303, Dermochelys
schlegelii, Dermochelys coriacea schlegelii, Sphargis
schlegelii
Type locality: “Tropical Pacific and Indian Oceans.” Restricted
to “Guaymas, Sonora, Mexico” by Smith and Taylor (1950a:344,
1950b:13).
Sphargis angusta Philippi 1899:730, Dermatochelys angusta
Type locality: “cerca de Tocopilla” [Chile].
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kinosTeRnoidea Joyce, Parham, and Gauthier 2004
Kinosternoidea Joyce, Parham, and Gauthier 2004:1003
deRMaTeMydidae Gray 1870e
Dermatemydae Gray 1870e:714
Dermatemydidae Baur 1888b:595
Dermatemys Gray 1847
Dermatemys Gray 1847:55
Type species: Dermatemys mawii Gray 1847, by original monotypy.
Chloremys Gray 1870c:50
Type species: Chloremys abnormis [= Dermatemys abnormis Cope
1868a = subjective synonym of Dermatemys mawii Gray 1847], by
original monotypy.
Limnochelone Werner 1901b:297
Type species: Limnochelone micrura Werner 1901b [= subjective
synonym of Dermatemys mawii Gray 1847], by original monotypy.
Dermatemys mawii Gray 1847 (14:8)
Central American River Turtle
abnormis
Type locality: “Belize River, Yucatan” [Belize]. Restricted to
“Belize [city], British Honduras” by Smith and Taylor (1950a:316,
1950b:19); restriction reversed by Dunn and Stuart (1951:59).
Dermatemys salvinii Gray 1870c:50
Type locality: “Guatemala.”
Limnochelone micrura Werner 1901b:298
Type locality: “Mexico.” Restricted to “Alvarado, Veracruz,
Mexico” by Smith and Taylor (1950a:316, 1950b:19).
Dermatemys mawei Neill and Allen 1959:28 (nomen novum)
kinosTeRnidae Agassiz 1857a (14:9)
Cinosternoidae Agassiz 1857a:249
Kinosterna Gray 1869a:180
Kinosternidae Hay 1892:560
kinosTeRninae Agassiz 1857a (14:9)
Cinosternoidae Agassiz 1857a:249
Kinosternina Gray 1869a:180
Kinosterninae Lindholm 1929:277
Kinosternon Spix 1824 (14:9) (11)
Monoclida Rafinesque 1815:75 (nomen nudum)
Uronyx Rafinesque 1815:75 (nomen nudum)
Kinosternon Spix 1824:17 (nomen conservandum, ICZN 1989)
Type species: Kinosternon longicaudatum Spix 1824 [= subjective
synonym of Testudo scorpioides Linnaeus 1766], by subsequent
designation by Bell (1828c:515).
Kinosternum Bonaparte 1830:166 (nomen novum)
Cinosternon Wagler 1830b:137 (nomen novum)
Monoclida Rafinesque 1832:64
Melvin Mérida / CBFTT / TCC / Laguna el Tigre Reserve, Peten, Guatemala
Type species: Monoclida retziana Rafinesque 1832 [= subjective
synonym of Testudo scorpioides Linnaeus 1766], by original
monotypy.
Uronyx Rafinesque 1832:64
Type species: Uronyx scorpioides [= Testudo scorpioides Linnaeus
1766], by original monotypy.
Cinosternum Burmeister 1837:731 (nomen novum)
Swanka Gray 1844:32
Type species: Swanka scorpiodes Gray [= Testudo scorpioides Linnaeus 1766], by original monotypy.
Thyrosternum Agassiz 1857a:418, 427
Type species: Thyrosternum pensilvanica [= Testudo pensilvanica
Gmelin 1789 = subjective synonym of Testudo subrubra Bonnaterre 1789], by subsequent designation by Lindholm (1929:277).
Platythyra Agassiz 1857a:420, 429
(orange dots = introduced or trade)
Belize, Guatemala, Mexico (Campeche, Chiapas, Quintana
Roo, Tabasco, Veracruz)
CBFTT Account: Vogt, Polisar, Moll, and Gonzalez-Porter
(2011)
IUCN Red List: Critically Endangered A2abd+4d (2006); Previously: Endangered (1996)
CITES: Appendix II
Dermatemys mawii Gray 1847:55, Emys mawii, Dermatemys
mavei
Type locality: “South America.” Restricted to “Alvarado, Veracruz,
Mexico” by Smith and Taylor (1950a:346).
Emys berardii Duméril and Bibron in Duméril and Duméril
1851:11, Ptychemys berardii, Clemmys berardii, Dermatemys berardii
Type locality: “l’Amérique mérid., ...environs de Vera-Cruz” [Veracruz, Mexico].
Dermatemys abnormis Cope 1868a:120, Chloremys
Type species: Platythyra flavescens Agassiz 1857a, by original
monotypy.
Cinosternos Herrera 1901:35 (nomen novum et suppressum,
ICZN 1922)
Cryptochelys Iverson, Le, and Ingram 2013:933 (partim) (14:9) (10)
Type species: Cryptochelys leucostomum [= Cinosternon leucostomum
Duméril and Bibron in Duméril and Duméril 1851], by original
designation.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
35
Kinosternon abaxillare Baur in Stejneger 1925 (14:10) or
Kinosternon scorpioides abaxillare
Central Chiapas Mud Turtle
John B. Iverson / Río Cintalapa, Chiapas, Mexico
Belize, Guatemala, Mexico (Campeche, Chiapas, Tabasco,
Veracruz)
CBFTT Account: Iverson and Vogt (2011)
IUCN Red List: Near Threatened (1996)
Kinosternon scorpioides acuta Gray 1831d:34, Kinosternon
acutum, Cryptochelys acuta
Type locality: Not designated. Restricted to “C. America” [Central
America] by Gray (1844:33); to “British Honduras” [Belize] by
Schmidt (1941:476); and to “Cosamaloapan, Veracruz, Mexico” by
Smith and Taylor (1950a:347, 1950b:23).
Cinosternum berendtianum Cope 1865:189, Cinosternon
berendtianum, Kinosternon berendtianum
Type locality: “Tabasco” [Mexico].
Cinosternon effeldtii Peters 1873:603, Cinosternum effeldtii
Type locality: “angeblich aus Mexico (Veracruz).” Restricted
to “Cosamaloapan, Veracruz, Mexico” by Smith and Taylor
(1950a:347).
Guatemala, Mexico (Chiapas)
IUCN Red List: Not Evaluated
Kinosternon abaxillare Baur in Stejneger 1925:462,
Kinosternon scorpioides abaxillare, Kinosternon cruentatum abaxillare
Kinosternon alamosae Berry and Legler 1980
Alamos Mud Turtle
Type locality: “Tuxtla, Chiapas, Mexico”.
Kinosternon acutum Gray 1831d (14:9) or
Cryptochelys acuta
Tabasco Mud Turtle
John B. Iverson / Sonora, Mexico
John B. Iverson / CBFTT / Belize
Mexico (Sinaloa, Sonora)
IUCN Red List: Data Deficient (2007); Previously: Least Concern [Not Listed] (1996)
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Kinosternon alamose Pritchard 1979:556 (nomen suppressum, ICZN 1985c)
Kinosternon baurii Garman 1891
Striped Mud Turtle
Type locality: “vicinity of Alamos, Sonora, and the lower Rio
Yaqui” [Mexico].
Kinosternon alamosae Berry and Legler 1980:1 (nomen
conservandum, ICZN 1985c)
Type locality: “Rancho Carrizal, 7.2 km north and 11.5 km west of
Alamos, Sonora, Mexico (27º 05' N, 109º 03' W).”
Kinosternon angustipons Legler 1965 (14:9) or
Cryptochelys angustipons
Narrow-bridged Mud Turtle
Dawn Wilson / CRM 3 / Hillsborough Co., Florida
Renae Reed / Boca Tapada, Costa Rica
USA (Florida, Georgia, North Carolina, South Carolina,
Virginia)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
Cinosternum baurii Garman 1891:141, Kinosternon baurii,
Kinosternon baurii baurii
Type locality: “the island Key West...Florida..[&]..Cuba.” Restricted
to “Key West, Monroe County, Florida” [USA] by Uzzell and
Schwartz (1955:33).
Kinosternon bauri palmarum Stejneger 1925:463,
Kinosternon baurii palmarum
Type locality: “Royal Palm State Park, Dade County, Florida”
[USA]. Restricted to “Paradise Key, Dade County, Florida” [USA]
by Uzzell and Schwartz (1955:34).
Kinosternon chimalhuaca Berry, Seidel, and Iverson in Rogner
1996 (07:7, 14:11)
Jalisco Mud Turtle
Costa Rica, Nicaragua, Panama
IUCN Red List: Vulnerable B1+2c (1996)
Kinosternon angustipons Legler 1965:617, Cryptochelys
angustipons
Type locality: “Los Diamantes, Limón Province, Costa Rica.”
John B. Iverson / Rio Purificación, Jalisco, Mexico
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Kinosternon dunni Schmidt 1947 (14:9) or
Cryptochelys dunni
Dunn’s Mud Turtle
Mexico (Colima, Jalisco)
IUCN Red List: Least Concern (2007)
Kinosternon chimalhuaca Berry, Seidel, and Iverson in Rogner 1996:23
Kinosternon chimalhuaca Berry, Seidel, and Iverson
1997:331
German Forero-Medina / CBFTT / Rio Atrato, Chocó, Colombia
Type locality: “30 m southeast of Mexico Highway 80, 1.9 km northeast of Barra de Navidad, Jalisco, Mexico (19º 15' N, 104º 43' S).”
Kinosternon creaseri Hartweg 1934 (14:9) or
Cryptochelys creaseri
Creaser’s Mud Turtle
John B. Iverson / nr. Felipe Carillo Puerto, Quintana Roo, Mexico
Colombia (Chocó, Valle del Cauca [?])
CBFTT Account: Iverson, Carr, Castaño-Mora, Galvis-Rizo,
Rentería-Moreno, and Forero-Medina (2012)
IUCN Red List: Vulnerable B1+2c (1996)
TFTSG Draft Red List: Vulnerable (2011)
Kinosternon dunni Schmidt 1947:109, Cryptochelys dunni
Type locality: “Pizarro, Choco, Colombia.”
Mexico (Campeche, Quintana Roo, Yucatán)
IUCN Red List: Least Concern (2007)
Kinosternon creaseri Hartweg 1934:1, Cryptochelys creaseri
Type locality: “one mile south of the Hacienda, Chichen Itza,
Yucatan” [Mexico].
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Kinosternon durangoense Iverson 1979b (07:6)
Durango Mud Turtle
Veracruz?), USA (Arizona, Arkansas?, Colorado, Illinois,
Iowa, Kansas, Missouri, Nebraska, New Mexico, Oklahoma, Texas)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
Platythyra flavescens Agassiz 1857a:430, Cinosternon flavescens, Cinosternum flavescens, Kinosternum flavescens,
Kinosternon flavescens, Kinosternon flavescens flavescens
Type locality: “Texas, near San Antonio;...lower Rio Grande;...
Red River, Arkansas;...Camp Yuma;...Gila River” [USA]. Restricted to “Waco, McLennan County, Texas, USA” by Smith and
Taylor (1950a:362, 1950b:24), but rejected by Maslin (1959:22);
and to “Rio Blanco, near San Antonio, Texas” [USA] by Schmidt
(1953:89) and by lectotype designation by Iverson (1978:478).
John B. Iverson / Pedriceña, Durango, Mexico
Kinosternon flavescens spooneri Smith 1951:195,
Kinosternon spooneri
Type locality: “Henderson County State Forest, 7 miles north of
Oquawka, Illinois” [USA].
Kinosternon herrerai Stejneger 1925 (14:9) or
Cryptochelys herrerai
Herrera’s Mud Turtle
Mexico (Chihuahua, Coahuila, Durango)
IUCN Red List: Data Deficient (2007)
Kinosternon flavescens durangoense Iverson 1979b:219,
Kinosternon durangoense
Type locality: “8 km from Ceballos, in Lago de los Palomas,
Durango, Mexico.”
John B. Iverson / nr. Gutiérrez Zamora, Veracruz, Mexico
Kinosternon flavescens Agassiz 1857a (07:6)
Yellow Mud Turtle
John B. Iverson / Canadian R., Beaver Co., Oklahoma
Mexico (Hidalgo, San Luis Potosi, Tamaulipas, Veracruz)
IUCN Red List: Near Threatened (2007); Previously: Least
Concern [Not Listed] (1996)
Kinosternon herrerai Stejneger 1925:462, Cryptochelys
herrerai
Type locality: “Xochimilco, Valley of Mexico” [in error]. Restricted
to “La Laja, Veracruz, Mexico” by Smith and Taylor (1950a:349,
1950b:24); and to “vicinity of Tampico” [Tamaulipas, Mexico] by
Smith and Brandon (1968:54).
Mexico (Chihuahua, Coahuila, Nuevo Leon, Tamaulipas,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Kinosternon hirtipes Wagler 1830c (09:11) (9)
Rough-footed Mud Turtle
Kinosternon hirtipes chapalaense Iverson 1981
Lake Chapala Mud Turtle
John B. Iverson / Lago Zapotlan, Jalisco, Mexico
Mexico (Jalisco, Michoacán)
Kinosternon hirtipes chapalaense Iverson in Pritchard
1979:557 (nomen nudum)
Kinosternon hirtipes chapalaense Iverson 1981:51
Type locality: “Lake Chapala, 0.25 mile off Chapala, Jalisco,
Mexico [20º 18' N, 103º 12' W].”
Kinosternon hirtipes magdalense Iverson 1981
San Juanico Mud Turtle
(subspecies: hirtipes = red, chapalaense = purple, magdalense = orange,
megacephalum = pink, murrayi = brown, tarascense = green)
Mexico (Aguascalientes, Chihuahua, Coahuila, Distrito Federal,
Durango, Guanajuato, Jalisco, México, Michoacán, Morelos, Zacatecas), USA (Texas)
IUCN Red List: Least Concern (2007); Previously: Least Concern [Not Listed] (1996)
Kinosternon hirtipes hirtipes Wagler 1830c (09:11) (9)
Valley of Mexico Mud Turtle
Mexico (Distrito Federal, México, Morelos)
Cinosternon hirtipes Wagler 1830b:137 (nomen nudum) (9)
Cinosternon hirtipes Wagler 1830c:explicatio tabularum,
pl.V, figs.29-30 (09:11) (9), Clemmys (Cinosternon) hirtipes,
Kinosternum hirtipes, Kinosternon hirtipes, Cinosternum hirtipes, Thyrosternum hirtipes, Ozotheca hirtipes,
Kinosternon hirtipes hirtipes
John B. Iverson / Presa San Juanico, Michoacán, Mexico
Mexico (Michoacán)
Kinosternon hirtipes magdalense Iverson 1981:53
Type locality: “along the face of the dam at Presa San Juanico,
Michoacán [ca. 19º 50' N, 102º 40' W]” [Mexico].
Kinosternon hirtipes megacephalum Iverson 1981
(Extinct, ca. 1970)
Viesca Mud Turtle
Type locality: Not designated. Restricted to “Mexico” by Wagler
(1833:unpaginated); to “Mazatlán, Sinaloa” [Mexico] [in error] by
Smith and Taylor (1950a:343, 1950b:25); and to “lakes near Mexico
City” [Mexico] by Schmidt (1953:89).
John B. Iverson / 3.2 km SE Viesca, Coahuila, Mexico
Mexico (Coahuila [extinct])
Kinosternon hirtipes megacephalum Iverson 1981:52,
Kinosternon hirtipes megalocephala, Kinosternon
megacephalum
Type locality: “3.2 km SE Viesca [25º 21' N, 102º 48' W], Coahuila”
[Mexico].
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Kinosternon hirtipes murrayi Glass and Hartweg 1951
Mexican Plateau Mud Turtle
John B. Iverson / north of Durango, Durango, Mexico
Kinosternon integrum Le Conte 1854
Mexican Mud Turtle
John B. Iverson / Alamos, Sonora, Mexico
Mexico (Aguascalientes, Chihuahua, Coahuila, Durango,
Guanajuato, Jalisco, México, Michoacán, Zacatecas), USA
(Texas)
Kinosternon murrayi Glass and Hartweg 1951:50,
Kinosternon hirtipes murrayi
Type locality: “Harper Ranch, 37 miles south of Marfa, Presidio
County, Texas” [USA].
Kinosternon hirtipes tarascense Iverson 1981
Pátzcuaro Mud Turtle
Judith Hirt / Ichupio, Lago de Pátzcuaro, Michoacán, Mexico
Mexico (Michoacán)
Kinosternon hirtipes tarascense Iverson 1981:52
Type locality: “Lago de Pátzcuaro, adjacent to city of Pátzcuaro [19º
32' N, 101º 36' W]” [Michoacán, Mexico].
Mexico (Colima, Durango, Guanajuato, Guerrero, Hidalgo,
Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, Puebla,
San Luis Potosi, Sinaloa, Sonora, Tamaulipas, Zacatecas)
IUCN Red List: Least Concern (2007); Previously: Least Concern [Not Listed] (1996)
Kinosternum integrum Le Conte 1854:183, Cinosternum
integrum, Thyrosternum integrum, Thyrosternon integrum,
Cinosternon integrum, Kinosternon integrum, Swanka
integra, Cinosternum scorpioides integrum, Kinosternon
scorpioides integrum
Type locality: “Mexico.” Restricted to “Acapulco, Guerrero,
Mexico” by Smith and Taylor (1950a:331, 1950b:25).
Cinosternon rostellum Bocourt 1876a:391, Cinosternum
rostellum
Type locality: “Guanajuato” [Mexico].
Cinosternon guanajuatense Dugès 1888:107 (nomen nudum)
Cinosternum scorpioides integrum mexicana Siebenrock
1907:579 (unavailable name)
Type locality: “Mexiko.” Restricted to “Mazatlan” [Mexico] by
Smith and Smith (1980:115).
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Kinosternon leucostomum Duméril and Bibron in Duméril and
Duméril 1851 (14:9) or
Cryptochelys leucostoma
White-lipped Mud Turtle
and Duméril 1851:17, Kinosternum leucostomum,
Kinosternon leucostomum, Cinosternum leucostomum,
Thyrosternum leucostomum, Swanka leucostoma,
Kinosternon leucostomum leucostomum, Cryptochelys
leucostoma
Type locality: “N.-Orléans; Mexique; Rio-Sumasinta (Amér.
centr.).” Restricted to “Rio Usumacinta, El Peten, Guatemala” by
Schmidt (1941:488); and to “Cosamaloapam, Veracruz, Mexico” by
Smith and Taylor (1950a:347, 1950b:26).
Swanka maculata Gray 1869a:182
Type locality: “Mexico...Papalco Apoia; Vera Paz” [Mexico;
Guatemala]. Restricted to “Cosamaloapam, Veracruz, Mexico” by
Stejneger (1941:457).
Cinosternum brevigulare Günther 1885:17 (senior homonym,
not = Cinosternum brevigulare Cope 1885)
Type locality: “Mexico, Playa Vicente” [Veracruz].
Cinosternum cobanum Günther 1885:18, Cinosternon
cobanum
Type locality: “Guatemala, Coban; Cahabon.” Restricted to “Cobán,
Alta Verapaz, Guatemala” by Smith and Taylor (1950a:317,
1950b:25).
Kinosternon mopanum Neill 1965:117
Type locality: “Waha Leaf Creek, southern Stann Creek District,
British Honduras” [Belize].
Kinosternon leucostomum postinguinale Cope 1887
Southern White-lipped Mud Turtle
(subspecies: leucostomum = red, postinguinale = purple)
Belize, Colombia (Antioquia, Atlántico, Bolívar, Boyacá, Caldas, Cauca, Cesar, Chocó, Córdoba, Cundinamarca, Huila
[?], Magdalena, Nariño, Santander, Sucre, Tolima, Valle
del Cauca), Costa Rica, Ecuador, Guatemala, Honduras,
Mexico (Campeche, Chiapas, Oaxaca, Quintana Roo,
Tabasco, Veracruz, Yucatán [?]), Nicaragua, Panama, Peru
(Tumbes)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (South America regional)
(2011)
Kinosternon leucostomum leucostomum Duméril and Bibron in
Duméril and Duméril 1851
Northern White-lipped Mud Turtle
John B. Iverson / Turbo, Antioquia, Colombia
Colombia (Antioquia, Atlántico, Bolívar, Boyacá, Caldas,
Cauca, Cesar, Chocó, Córdoba, Cundinamarca, Huila [?],
Magdalena, Nariño, Santander, Sucre, Tolima, Valle del
Cauca), Costa Rica, Ecuador, Nicaragua, Panama, Peru
(Tumbes)
Cinosternum brevigulare Cope 1885:389 (junior homonym,
not = Cinosternum brevigulare Günther 1885), Cinosternon
brevigulare
Type locality: “Tierra Caliente of Costa Rica at Sipurio, on the east
coast.”
Cinosternum postinguinale Cope 1887:23 (nomen novum),
Kinosternon postinguinale, Kinosternon leucostomum
postinguinale
Type locality: “E. coast Costa Rica.”
Cinosternum spurrelli Boulenger 1913:1030, Kinosternon
spurrelli, Kinosternon leucostomum spurrelli
Type locality: “Choco, Colombia...Peña Lisa, Condoto, altitude 300
feet.”
John B. Iverson / Belize Distr., Belize
Belize, Guatemala, Honduras, Mexico (Campeche, Chiapas,
Oaxaca, Quintana Roo, Tabasco, Veracruz, Yucatán [?]),
Nicaragua
Cinosternon leucostomum Duméril and Bibron in Duméril
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Kinosternon oaxacae Berry and Iverson 1980
Oaxaca Mud Turtle
John B. Iverson / Puerto Angel, Oaxaca, Mexico
(subspecies: scorpioides = red, albogulare = purple, cruentatum = orange)
Mexico (Guerrero, Oaxaca)
IUCN Red List: Data Deficient (2007); Previously: Near Threatened (1996)
Kinosternon oaxacae Pritchard 1979:557 (nomen suppressum, ICZN 1985c)
Type locality: “vicinity of Pochutla, Oaxaca” [Mexico].
Kinosternon oaxacae Berry and Iverson 1980:313 (nomen
conservandum, ICZN 1985c)
Type locality: “11.6 km N. of Pochutla (San Pedro Pochutla), along
Mexican Hwy. 175 (ca. 235 m), Oaxaca, Mexico (15º46' N, 96º28'
W).”
Kinosternon scorpioides (Linnaeus 1766)
Scorpion Mud Turtle
Argentina (Formosa, Jujuy, Salta, Tucumán), Belize, Bolivia,
Brazil (Acre, Alagoas, Amapá, Amazonas, Bahia, Ceará,
Goiás, Maranhão, Mato Grosso, Minas Gerais, Pará,
Paraiba, Pernambuco, Piauí, Rio Grande do Norte, Rondônia, Sergipe, Tocantins), Colombia (Amazonas, Antioquia,
Arauca, Atlántico, Bolívar, Caldas, Caquetá, Casanare,
Cesar, Chocó, Córdoba, Guainía, Magdalena, Meta, Norte
de Santander, Putumayo, San Andrés, Sucre, Vaupés,
Vichada), Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Mexico (Campeche,
Chiapas, Oaxaca, Quintana Roo, Tabasco, Tamaulipas,
Veracruz, Yucatán), Nicaragua, Panama, Paraguay, Peru
(Amazonas, Huánuco, Loreto, Madre de Dios, Ucayali),
Suriname, Trinidad, Venezuela (Amazonas, Apure, Aragua, Bolívar, Cojedes, Falcón, Guárico, Lara, Monagas,
Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia)
CBFTT Account: Berry and Iverson (2011)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (South America regional)
(2011)
Kinosternon scorpioides scorpioides (Linnaeus 1766) (07:8)
Scorpion Mud Turtle
Frank Deschandol / French Guiana
Argentina (Formosa, Jujuy, Salta, Tucumán), Bolivia, Brazil
(Acre, Alagoas, Amapá, Amazonas, Bahia, Ceará, Goiás,
Maranhão, Mato Grosso, Minas Gerais, Pará, Paraiba, Pernambuco, Piauí, Rio Grande do Norte, Rondônia, Sergipe,
Tocantins), Colombia (Amazonas, Antioquia, Arauca,
Atlántico, Bolívar, Caldas, Caquetá, Casanare, Cesar,
Chocó, Córdoba, Guainía, Magdalena, Meta, Norte de
Santander, Putumayo, Sucre, Vaupés, Vichada), Ecuador,
French Guiana, Guyana, Panama, Paraguay, Peru (Amazonas, Huánuco, Loreto, Madre de Dios, Ucayali), Suriname,
Trinidad, Venezuela (Amazonas, Apure, Aragua, Bolívar,
Cojedes, Falcón, Guárico, Lara, Monagas, Portuguesa,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Sucre, Táchira, Trujillo, Yaracuy, Zulia)
Testudo scorpioides Linnaeus 1766:352, Emys scorpioides,
Chersine scorpioides, Terrapene scorpioidea, Cinosternon
scorpioidea, Kinosternon scorpioides, Uronyx scorpioides,
Terrapene scorpioides, Cinosternon scorpioides, Clemmys
(Cinosternon) scorpioidea, Kinosternum scorpioides, Cinosternum scorpioides, Thyrosternum scorpioides, Swanka
scorpiodes, Swanka scorpioides, Cinosternum scorpioides
scorpioides, Kinosternon scorpioides scorpioides
Type locality: “Surinami” [Surinam].
Testudo tricarinata Retzius in Schoepff 1792:9 (senior homonym, not = Testudo tricarinata Bory de Saint-Vincent
1804), Terrapene tricarinata, Clemmys tricarinata
Catalina [prehistoric or modern introduction?]), Costa
Rica, El Salvador, Honduras, Nicaragua, Panama
CBFTT Account: Forero-Medina and Castaño-Mora (2011)
Cinosternon albogulare Duméril and Bocourt 1870:24, Cinosternum albogulare, Kinosternon cruentatum albogulare,
Kinosternon scorpioides albogulare
Type locality: “S. Jose (Costa Rica).”
Kinosternon scorpioides cruentatum Duméril and Bibron in
Duméril and Duméril 1851
Red-cheeked Mud Turtle
Type locality: Not known. Restricted to “Surinam” by Fritz and
Havaš (2007:256).
Testudo retzii Daudin 1801:174 (nomen novum), Emys retzii,
Terrapene retzii
Type locality: Not known. Restricted to “Surinam” by Fritz and
Havaš (2007:256).
Kinosternon longicaudatum Spix 1824:17 (nomen conservandum, ICZN 1989), Cinosternon longicaudatum, Kinosternum longicaudatum, Cinosternum longicaudatum,
Thyrosternum longicaudatum, Swanka longicaudata
Type locality: “Brasiliam...ad campis aquosis” [Brazil].
Kinosternon brevicaudatum Spix 1824:18, Cinosternon
brevicaudatum, Kinosternum brevicaudatum, Cinosternum
brevicaudatum
Type locality: “Brasiliam...ad ripam fluminis Solimöens” [Brazil].
Kinosternon shavianum Bell 1825a:302, Cinosternon shavianum, Cinosternum shavianum, Thyrosternum shavianum
Type locality: Not known.
Monoclida retziana Rafinesque 1832:64 (nomen novum),
Testudo retziana
Cinosternon shawianum Bocourt 1876a:387 (nomen novum)
Cinosternum scorpioides integrum brasiliana Siebenrock
1907:579 (unavailable name)
Type locality: “Südamerika” [Brazil].
Kinosternon scorpioides pachyurum Müller and Hellmich
1936:100
Type locality: “Bolivien...Chaco...Villa Montes” [Bolivia].
Kinosternon scorpioides seriei Freiberg 1936:169 (07:8)
Type locality: “El Tabacal (Salta)” [Argentina].
Kinosternon panamensis Schmidt 1946:5
Type locality: “Panama Railroad, Canal Zone” [Panama].
Kinosternon scorpioides carajasensis Cunha 1970:1 (07:8)
Type locality: “compartimento da serra dos Carajás (serra Norte)
Pará” [Brazil].
John B. Iverson / CBFTT / Manuel, Tamaulipas, Mexico
Belize, Guatemala, Honduras, Mexico (Campeche, Chiapas,
Oaxaca, Quintana Roo, Tamaulipas, Veracruz, Yucatán)
Cinosternon cruentatum Duméril and Bibron in Duméril and
Duméril 1851:16, Kinosternum cruentatum, Kinosternon
cruentatum, Cinosternum cruentatum, Swanka cruentata,
Thyrosternum cruentatum, Kinosternon cruentatum cruentatum, Kinosternon scorpioides cruentatum
Type locality: “Amér. septentr.” Restricted to “San Mateo del Mar,
Oaxaca, Mexico” by Smith and Taylor (1950a:339, 1950b:23).
Kinosternum mexicanum Le Conte 1854:182, Cinosternum
mexicanum, Cinosternon mexicanum, Kinosternon mexicanum, Swanka mexicana
Type locality: “Mexico.” Restricted to “San Mateo del Mar, Oaxaca,
Mexico” by Smith and Taylor (1950a:339, 1950b:23).
Kinosternum triliratum Le Conte 1860:6, Cinosternon triliratum, Swanka trilirata, Cinosternum triliratum
Type locality: “Mexico.” Restricted to “San Mateo del Mar, Oaxaca,
Mexico” by Smith and Taylor (1950a:339, 1950b:23).
Kinosternon cruentatum consors Stejneger 1941:458
Type locality: “Cozumel Island, Yucatan, Mexico.”
Kinosternon sonoriense Le Conte 1854
Sonora Mud Turtle
Kinosternon scorpioides albogulare Duméril and Bocourt 1870
White-throated Mud Turtle
(subspecies: sonoriense = red, longifemorale = purple)
John B. Iverson / CBFTT / Rio Corobizi, Guanacaste, Costa Rica
Colombia (Archipiélago de San Andrés, Providencia y Santa
Mexico (Baja California, Chihuahua, Sonora), USA (Arizona,
California [extirpated], New Mexico)
IUCN Red List: Near Threatened (2013); Previously: Least
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Concern [Not Listed] (1996)
Kinosternon sonoriense sonoriense Le Conte 1854
Sonora Mud Turtle
Jeffrey E. Lovich / Montezuma Well, Yavapai Co., Arizona
Mexico (Chihuahua, Sonora), USA (Arizona, California
[extirpated], New Mexico)
Kinosternum sonoriense Le Conte 1854:184, Kinosternon
sonoriense, Cinosternum sonoriense, Thyrosternum sonoriense, Cinosternon sonoriense, Kinosternon sonoriense
sonoriense
Type locality: “Tucson...province of Sonora” [Mexico; now Arizona, USA].
Kinosternum henrici Le Conte 1860:4, Thyrosternum henrici,
Cinosternon henrici, Cinosternum henrici, Swanka henrici
USA (Florida)
IUCN Red List: Least Concern (2013), as part of Kinosternon
subrubrum
Cinosternum steindachneri Siebenrock 1906b:727,
Kinosternon steindachneri, Kinosternon subrubrum
steindachneri
Type locality: “Orlando in Florida” [USA].
Kinosternon stejnegeri (Hartweg 1938) (07:6, 09:10) (13)
[previously listed as Kinosternon arizonense] (07:6, 09:10)
Arizona Mud Turtle
Type locality: “New Mexico” [USA]. Data with holotype is “Gila
River, New Mexico”; incorrectly restricted to “vicinity of Las
Cruces” [New Mexico, USA] by Schmidt (1953:91).
Kinosternon sonoriense longifemorale Iverson 1981
Sonoyta Mud Turtle
John B. Iverson / nr. Sells, Pima Co., Arizona
John B. Iverson / Quitobaquito Springs, Organ Pipe National Monument, Arizona
Mexico (Sonora), USA (Arizona)
TFTSG Draft Red List: Critically Endangered (2016)
Kinosternon sonoriense longifemorale Iverson 1981:43
Type locality: “artificial pond fed by springs, Sonoyta, Sonora,
Mexico (31˚51' N, 112˚50' W).”
Kinosternon steindachneri Siebenrock 1906b (12:11, 14:12) (12)
Florida Mud Turtle
Mexico (Sonora), USA (Arizona)
IUCN Red List: Least Concern (2007), as Kinosternon
arizonense
Kinosternon flavescens stejnegeri Hartweg 1938:1 (12),
Kinosternon stejnegeri
Type locality: “Llano, Sonora...approximately midway between
Nogales and Hermosillo” [Mexico].
Previously listed as K. arizonense (now considered distinct
and extinct):
Kinosternon arizonense † Gilmore 1923:2 (07:6, 09:10) (12) [Pliocene–Pleistocene, USA (Arizona)], Kinosternon flavescens
arizonense
John B. Iverson / CRM 3 / CBFTT / Monroe Co., Florida
Type locality: “Benson Locality Quarry, two miles south of Benson,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Cochise County, Arizona” [USA].
Kinosternon subrubrum (Bonnaterre 1789) (09:6, 14:12)
Eastern Mud Turtle, Common Mud Turtle
Kinosternon pennsylvanicum Bell 1825a:304 (nomen
novum), Emys (Kinosternon) pennsylvanica, Kinosternum
pennsylvanicum, Cinosternon pennsylvanicum, Cinosternum pennsylvanicum, Cistudo pennsylvanica, Terrapene
pennsylvanica, Thyrosternum pennsylvanicum
Kinosternon (Kinosternon) doubledayii Gray 1844:33,
Kinosternon doubledayii, Kinosternum doubledayii, Cinosternum doubledayii, Cinosternon doubledayii
Type locality: “California” [in error]. Restricted to “vicinity of Philadelphia” [Pennsylvania, USA] by Schmidt (1953:90).
Kinosternon (Kinosternon) oblongum Gray 1844:33,
Kinosternon oblongum, Cinosternum oblongum
Type locality: “America.” Restricted to “vicinity of Philadelphia”
[Pennsylvania, USA] by Schmidt (1953:90).
Kinosternon punctatum Gray 1856a:198, Cinosternum
punctatum
Type locality: “North America.” Restricted to “East Florida” [USA]
by Gray (1856b:46).
(subspecies: subrubrum = red, hippocrepis = purple;
overlap = intergrades; orange dot = possibly introduced)
USA (Alabama, Arkansas, Delaware, Florida, Georgia, Illinois,
Indiana, Kentucky, Louisiana, Maryland, Mississippi, Missouri, New Jersey, New York, North Carolina, Oklahoma,
Pennsylvania, South Carolina, Tennessee, Texas, Virginia)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
Swanka fasciata Gray 1869a:183
Type locality: Not known. Restricted to “vicinity of Philadelphia”
[Pennsylvania, USA] by Schmidt (1953:90).
Kinosternon subrubrum hippocrepis Gray 1856a (14)
Mississippi Mud Turtle
Kinosternon subrubrum subrubrum (Bonnaterre 1789) (09:6)
Eastern Mud Turtle
Richard D. Bartlett / Mississippi or Louisiana
USA (Arkansas, Louisiana, Mississippi, Missouri, Oklahoma,
Texas)
Kinosternon hippocrepis Gray 1856a:198 (13), Cinosternum
hippocrepis, Cinosternon hippocrepis, Kinosternon subrubrum hippocrepis
Richard D. Bartlett / Apalachicola National Forest, Florida
USA (Alabama, Delaware, Florida, Georgia, Illinois,
Indiana, Kentucky, Maryland, Mississippi, New Jersey, New York, North Carolina, Pennsylvania, South
Carolina, Tennessee, Virginia)
Testudo subrubra Lacepède 1788:132, synopsis[table] (09:6)
(nomen suppressum, ICZN 2005a)
Type locality: “Pensylvanie” [USA]. Restricted to “vicinity of Philadelphia” [Pennsylvania, USA] by Schmidt (1953:90).
Testudo subrubra Bonnaterre 1789:27, Kinosternon subrubrum, Kinosternon subrubrum subrubrum
Testudo pensilvanica Gmelin 1789:1042, Emydes pensilvanica, Kinosternon pensilvanicum, Cinosternum
pensilvanicum
Type locality: “Pensilvaniae aquis stagnantibus” [USA]. Restricted
to “vicinity of Philadelphia” [Pennsylvania, USA] by Schmidt
(1953:90).
Emys pensylvanica Schweigger 1812:282 (nomen novum),
Terrapene pensylvanica, Cistuda pensylvanica, Sternotherus pensylvanica, Kinosternum pensylvanicum,
Cinosternon pensylvanicum, Clemmys (Cinosternon)
pensylvanica, Kinosternon pensylvanicum, Cinosternum
pensylvanicum
Type locality: “North America; New Orleans” [Louisiana, USA].
Kinosternon louisianae Baur 1893c:676, Cinosternum
louisianae
Type locality: “New Orleans, La.” [Louisiana, USA].
Sternotherus Bell in Gray 1825 (07:9) (10)
Sternothaerus Bell 1825a:305 (partim, nomen suppressum,
ICZN 1989)
Type species: Sternothaerus leachianus Bell 1825a [= subjective synonym of Emys castanea Schweigger 1812 = Pelusios castaneus],
by subsequent designation by Bell (1828c:515); not Sternothaerus
odoratus Bell [= Testudo odorata Latreille in Sonnini and Latreille
1801], by subsequent incorrect designation by Fitzinger (1843:290).
Sternotherus Bell in Gray 1825:211 [Bell 1825b] (nomen conservandum, ICZN 1989)
Type species: Sternotherus odoratus [= Testudo odorata Latreille in
Sonnini and Latreille 1801], by subsequent designation by Stejneger
(1902:237).
Aromochelys Gray 1856a:199
Type species: Aromochelys odorata [= Testudo odorata Latreille in
Sonnini and Latreille 1801], by subsequent designation by Strauch
(1862:38).
Ozotheca Agassiz 1857a:251,424
Type species: Ozotheca odorata [= Testudo odorata Latreille in Sonnini and Latreille 1801], by subsequent designation by Lindholm
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(1929:277).
Goniochelys Agassiz 1857a:420,423
Type species: Goniochelys triquetra Agassiz 1857a [= subjective
synonym of Aromochelys carinata Gray 1856a], by subsequent
designation by Lindholm (1929:277).
Sternotherus carinatus (Gray 1856a)
Razor-backed Musk Turtle
Robert C. Thomson / CBFTT / Pascagoula R., Mississippi
USA (Alabama)
CBFTT Account: Dodd (2008)
IUCN Red List: Critically Endangered A2bce+4bce (2013);
Previously: Vulnerable (1996)
Sternotherus depressus Tinkle and Webb 1955:53, Sternothaerus depressus, Sternotherus minor depressus,
Kinosternon depressum, Kinosternon depressus
Type locality: “Mulberry Fork of the Black Warrior River, 9 miles
east of Jasper, Walker County, Alabama, near the bridge crossing of
U.S. highway 78” [USA].
Sternotherus minor (Agassiz 1857a)
Loggerhead Musk Turtle
USA (Alabama, Arkansas, Louisiana, Mississippi, Oklahoma,
Texas)
CBFTT Account: Lindeman (2008)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
Aromochelys carinata Gray 1856a:199, Aromochelys carinatum, Aromochelys carinatus, Goniochelys carinata, Cinosternum carinata, Kinosternon carinatum, Sternotherus
carinata, Sternotherus carinata carinata, Kinosternon
carinata
Type locality: “North America, Louisiana” [USA]. Restricted to
“vicinity of New Orleans” [Louisiana, USA] by Schmidt (1953:87).
Goniochelys triquetra Agassiz 1857a:420,423
Type locality: “Lake Concordia, in Louisiana” [USA].
Sternotherus depressus Tinkle and Webb 1955 (07:10) (15)
Flattened Musk Turtle
(subspecies: minor = red, peltifer = purple;
overlap = intergrades; orange dot = introduced)
USA (Alabama, Florida, Georgia, Kentucky, Louisiana, Mississippi, Tennessee, Virginia)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
Sternotherus minor minor (Agassiz 1857a)
Loggerhead Musk Turtle
Timothy Walsh / CRM 3 / Marion Co., Florida
C. Kenneth Dodd, Jr. / CBFTT / TCC / Gurley Creek, Alabama
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
USA (Alabama, Florida, Georgia)
Goniochelys minor Agassiz 1857a:424, Aromochelys
minor, Sternotherus minor, Sternotherus carinatus minor,
Sternotherus minor minor, Sternothaerus minor minor,
Kinosternon minor, Kinosternon minor minor
Type locality: “neighborhood of Mobile;...Columbus, Georgia;...and
New Orleans” [USA]. Restricted to “Columbus, Georgia” [USA]
by Schmidt (1953:88).
Sternotherus minor peltifer Smith and Glass 1947 (16)
Stripe-necked Musk Turtle
Oklahoma, Pennsylvania, Rhode Island, South Carolina,
Tennessee, Texas, Vermont, Virginia, West Virginia,
Wisconsin)
IUCN Red List: Least Concern (2011); Previously: Least Concern [Not Listed] (1996)
Testudo odorata Latreille in Sonnini and Latreille 1801:122
(nomen conservandum, ICZN 1989), Emys odorata, Terrapene odorata, Cistuda odorata, Sternotherus odorata,
Sternothaerus odoratus, Kinosternum odoratum, Emys
(Kinosternon) odoratum, Kinosternon odoratum, Didicla
odorata, Staurotypus odoratus, Clemmys (Sternothaerus)
odorata, Cistudo odorata, Sternotherus odoratus, Aromochelys odorata, Aromochelys odoratum, Cinosternum
odoratum, Ozotheca odorata
Type locality: “les eaux dormantes de la Caroline” [USA].
Restricted to “vicinity of Charleston, South Carolina” [USA] by
Schmidt (1953:87).
Testudo glutinata Daudin 1801:194, Emys glutinata, Clemmys glutinata
Type locality: “les Etats-Unis d’Amérique” [USA]. Restricted to
“vicinity of Lancaster, Pennsylvania” [USA] by Schmidt (1953:87).
Terrapene boscii Merrem 1820:27, Sternothaerus boscii
C. Kenneth Dodd, Jr. / Buttahatchie R., Marion Co., Alabama
USA (Alabama, Georgia, Kentucky, Mississippi, Tennessee,
Virginia)
Sternotherus peltifer Smith and Glass 1947:22, Sternotherus
carinatus peltifer, Sternotherus minor peltifer, Sternothaerus minor peltifer, Kinosternon minor peltifer
Type locality: “Bassfield, Jefferson Davis County, 30 miles west of
Hattiesburg, Miss.” [Mississippi, USA].
Sternotherus odoratus (Latreille in Sonnini and Latreille 1801)
Musk Turtle, Stinkpot, Common Musk Turtle
Type locality: “America septentrionali.” Restricted to “vicinity of
Philadelphia” [Pennsylvania, USA] by Schmidt (1953:87).
Kinosternum guttatum Le Conte 1854:185, Cinosternum guttatum, Aromochelys guttata
Type locality: “Pennsylvania” [USA]. Restricted to “vicinity of
Philadelphia“ [Pennsylvania, USA] by Schmidt (1953:87).
Ozotheca tristycha Agassiz 1857a:392,425, Aromochelys
tristycha
Type locality: “Osage River, in Missouri, and in Williamson County, in Texas...near San Antonio,...Medina River, in Texas” [USA].
Restricted to “San Antonio” [Texas, USA] by Schmidt (1953:87).
Testudo glutinosa Agassiz 1857a:425 (nomen novum), Emys
glutinosa
sTauRoTypinae Gray 1869a (14:13) or
sTauRoTypidae
Staurotypina Gray 1869a:180
Staurotypinae Siebenrock 1907:531
Staurotypidae Bickham and Carr 1983:925
Claudius Cope 1865
Peter May / CRM 3 / Volusia Co., Florida
Claudius Cope 1865:187
Type species: Claudius angustatus Cope 1865, by original monotypy.
Staurosternon Duméril in Bocourt 1868:122
Type species: Claudius megalocephalus Bocourt 1868 [= subjective synonym of Claudius angustatus Cope 1865], by original
monotypy.
Claudius angustatus Cope 1865
Narrow-bridged Musk Turtle
Canada (Ontario, Québec), USA (Alabama, Arkansas, Connecticut, Delaware, Florida, Georgia, Illinois, Indiana,
Iowa, Kansas, Kentucky, Louisiana, Maine, Maryland,
Massachusetts, Michigan, Mississippi, Missouri, New
Hampshire, New Jersey, New York, North Carolina, Ohio,
John B. Iverson / Hattieville, Belize Dist., Belize
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IUCN Red List: Near Threatened (1996)
Staurotypus (Stauremys) salvinii Gray 1864c:127, Stauremys
salvinii, Staurotypus salvinii
Type locality: “Haumanchal, Guatemala” [= Huamúchil, Oaxaca,
Mexico].
Staurotypus marmoratus Fischer 1872:265
Type locality: “Mexico, Tejas.” Restricted to “Santa Efigenia,
Oaxaca, Mexico” by Smith and Taylor (1950a:339, 1950b:27).
Claudius severus Cope 1872:24, Staurotypus (Claudius)
severus
Type locality: “Santa Efigenia, on the western side of the Isthmus of
Tehuantepec, Mexico.”
Staurotypus biporcatus Gadow 1905:209 (nomen nudum)
Belize, Guatemala, Mexico (Campeche, Chiapas, Oaxaca,
Quintana Roo, Tabasco, Veracruz)
IUCN Red List: Near Threatened (1996)
Claudius angustatus Cope 1865:187, Claudius angustatum
Staurotypus triporcatus (Wiegmann 1828)
Northern Giant Musk Turtle
Type locality: “Tabasco, Mexico.”
Claudius megalocephalus Bocourt 1868:122
Type locality: “Mexico.” Restricted to “Tabasco, Mexico” by Smith
and Taylor (1950a:345).
Claudius macrocephalus Gray 1873d:69 (nomen novum)
Claudius megacephalus Boulenger 1889:33 (nomen novum)
Claudius agassizii Smith and Taylor 1950a:345 (nomen
nudum)
Staurotypus Wagler 1830b
Staurotypus Wagler 1830b:137
John B. Iverson / nr. Central Farms, Cayo Dist., Belize
Type species: Staurotypus triporcata [= Terrapene triporcata Wiegmann 1828], by original monotypy.
Stauremys Gray 1864c:127
Type species: Staurotypus (Stauremys) salvinii Gray 1864c, by original
monotypy.
Staurotypus salvinii Gray 1864c
Pacific Coast Giant Musk Turtle
John B. Iverson / Puerto Arista, Chiapas, Mexico
Belize, Guatemala, Honduras, Mexico (Campeche, Chiapas,
Oaxaca, Quintana Roo, Tabasco, Veracruz)
IUCN Red List: Near Threatened (1996)
Terrapene triporcata Wiegmann 1828:364, Staurotypus triporcata, Staurotypus triporcatus, Staurotypus
(Staurotypus) triporcatus, Emys (Kinosternon) triporcata, Kinosternon triporcatum, Clemmys (Staurotypus)
triporcata
Type locality: “Rio Alvarado” [Veracruz, Mexico].
Claudius pictus Cope 1872:26
Type locality: “Vera Paz” [Guatemala]. Emended to “Alta Verapaz”
[Guatemala] by Smith and Taylor (1950a:317, 1950b:27); and to “a
tributary of the Rio Polochic...Alta Verapaz, Guatemala” by Dunn
and Stuart (1951:59).
El Salvador, Guatemala, Mexico (Chiapas, Oaxaca)
Introduced: USA (Florida)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
TesTudinoidea Fitzinger 1826
Testudinoidea Fitzinger 1826:5
eMydidae Rafinesque 1815 (09:12) (17)
Chrysemys picta (Schneider 1783) (07:11, 10:6, 12:13) (18)
Painted Turtle
Emidania Rafinesque 1815:75
Emydes Schmid 1819:11
Emydidae Bell 1825a:302
Emydae Swainson 1839:113
Emididi Portis 1890:12
deiRoChelyinae Agassiz 1857a (09:12) (17)
Deirochelyoidae Agassiz 1857a:355
Deirochelyinae Gaffney and Meylan 1988:201
Chrysemys Gray 1844 (12:12)
Hydrochelys Wagler 1821:12 (12:12) (nomen oblitum)
Type species: Hydrochelys picta [= Testudo picta Schneider 1783], by
original monotypy.
Chrysemys Gray 1844:27
Type species: Emys (Chrysemys) picta Schweigger [= Testudo picta
Schneider 1783], by subsequent designation by Brown (1908:114).
Chrysemys dorsalis Agassiz 1857a (07:11, 10:6) (18) or
Chrysemys picta dorsalis
Southern Painted Turtle
(subspecies: picta = red, bellii = purple,
marginata = orange; overlap = intergrades)
James H. Harding / Reelfoot Lake, Tennessee
USA (Alabama, Arkansas, Illinois, Kentucky, Louisiana, Mississippi, Missouri, Oklahoma, Tennessee, Texas)
Introduced: USA (Florida)
IUCN Red List: Least Concern (2013), as Chrysemys picta
dorsalis
Chrysemys dorsalis Agassiz 1857a:439,440 (07:11, 10:6), Clemmys picta dorsalis, Chrysemys cinerea dorsalis, Chrysemys marginata dorsalis, Chrysemys bellii dorsalis,
Chrysemys picta dorsalis
Type locality: “Mississippi and Louisiana...Lake Concordia”
[USA]. Restricted to “vicinity of New Orleans” [Louisiana, USA]
by Schmidt (1953:100), but see Ernst (1967:133).
Canada (Alberta, British Columbia, Manitoba, New Brunswick,
Nova Scotia, Ontario, Québec, Saskatchewan), Mexico
(Chihuahua), USA (Alabama, Arizona, Colorado, Connecticut, Delaware, Georgia, Idaho, Illinois, Indiana, Iowa,
Kansas, Kentucky, Maine, Maryland, Massachusetts,
Michigan, Minnesota, Missouri, Montana, Nebraska, New
Hampshire, New Jersey, New Mexico, New York, North
Carolina, North Dakota, Ohio, Oklahoma, Oregon, Pennsylvania, Rhode Island, South Carolina, South Dakota,
Tennessee, Texas, Utah, Vermont, Virginia, Washington,
West Virginia, Wisconsin, Wyoming)
Introduced: Germany, Indonesia, Philippines, Spain, USA
(California)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
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Chrysemys picta picta (Schneider 1783) (07:11, 12:13)
Eastern Painted Turtle
Columbia River” [Oregon, USA].
Chrysemys nuttalii Agassiz 1857a:451 (nomen nudum)
Chrysemys nuttalii Agassiz 1857b:642 (nomen novum)
Type locality: “Minesota and westward to the junction of the Yellowstone and Missouri” [Minnesota, USA].
Chrysemys pulchra Gray 1873a:147
Type locality: “North America, Mississippi” [USA]. Restricted to
“upper Mississippi River” [USA] by Schmidt (1953:100).
Chrysemys timida Hay † 1908b:345 [Pleistocene, USA
(Nebraska)]
Type locality: “Equus beds of Sheridan County, Nebraska, not far
from the Niobrara River” [USA].
Chrysemys treleasei Hurter 1911:235
Type locality: “east side of the Mississippi River, in Madison, St.
Clair, and Monroe Counties, Ill.” [Illinois, USA].
Anders G.J. Rhodin / Acadia National Park, Bar Harbor, Mt. Desert Island, Maine
Canada (New Brunswick, Nova Scotia, Québec), USA (Alabama, Connecticut, Delaware, Georgia, Maine, Maryland,
Massachusetts, New Hampshire, New Jersey, New York,
North Carolina, Pennsylvania, Rhode Island, South Carolina, Vermont, Virginia)
Testudo picta Schneider 1783:348, Emys picta, Clemmys
picta, Terrapene picta, Emys (Chrysemys) picta, Chrysemys picta, Chrysemys picta picta, Pseudemys picta
Chrysemys picta marginata Agassiz 1857a
Midland Painted Turtle
Type locality: “England” [in error]. Restricted to “Lancaster, Pennsylvania” [USA] by Mittleman (1945:171); and to “vicinity of New
York City” [New York, USA] by Schmidt (1953:99) [in error according to Smith and Smith (1980:424)].
Testudo cinerea Bonnaterre 1789:25, Emys cinerea, Chrysemys cinerea, Chrysemys cinerea cinerea
Type locality: Not known. Restricted to “vicinity of Philadelphia”
[Pennsylvania, USA] by Schmidt (1953:99).
Chrysemys picta bellii (Gray 1830e) (10:7) (18)
Western Painted Turtle
Scott D. Gillingwater / Long Point, Norfolk Co., Ontario, Canada
Canada (Ontario, Québec), USA (Alabama, Illinois, Indiana, Kentucky, Michigan, New York, North Carolina,
Ohio, Pennsylvania, Tennessee, Vermont, Virginia, West
Virginia)
Chrysemys marginata Agassiz 1857a:262,439, Clemmys
marginata, Chrysemys marginata marginata, Chrysemys
bellii marginata, Chrysemys picta marginata
Type locality: “Racine, Wisconsin...Milwaukee, Wisconsin...Flint,
Michigan...Ann-Arbor, Michigan...Delphi, Indiana...Burlington,
Iowa” [USA]. Restricted to “northern Indiana” [USA] by Schmidt
(1953:99).
John B. Iverson / Oshkosh, Garden Co., Nebraska
Canada (Alberta, British Columbia, Manitoba, Ontario, Saskatchewan), Mexico (Chihuahua), USA (Arizona, Colorado,
Idaho, Illinois, Iowa, Kansas, Michigan, Minnesota, Missouri, Montana, Nebraska, New Mexico, North Dakota,
Oklahoma, Oregon, South Dakota, Texas, Utah, Washington, Wisconsin, Wyoming)
Emys bellii Gray 1830e:12 (10:7), Clemmys (Clemmys) bellii,
Emys (Chrysemys) bellii, Chrysemys bellii, Emys bellii,
Chrysemys cinerea bellii, Chrysemys marginata bellii,
Chrysemys bellii bellii, Chrysemys picta bellii
Type locality: Not known. Restricted to “West coast of North
America; British Columbia” by Gray (1873a:147); to “Manhattan,
Kans.” [Kansas, USA] by Smith and Taylor (1950b:34); and to
“Puget Sound, Washington” [USA] by Schmidt (1953:100).
Emys oregoniensis Harlan 1837:382, Chrysemys oregoniensis, Clemmys oregoniensis, Chrysemys oregonensis
Type locality: “fresh water ponds in the vicinity of the Oregon or
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Deirochelys Agassiz 1857a
51
Deirochelys reticularia chrysea Schwartz 1956a
Florida Chicken Turtle
Deirochelys Agassiz 1857a:252,441
Type species: Deirochelys reticulata Schweigger [= Testudo reticulata
Daudin 1801 = objective synonym of Testudo reticularia Latreille
in Sonnini and Latreille 1801], by original monotypy.
Hirochelys Beyer 1900:21 (nomen novum)
Deirochelys reticularia (Latreille in Sonnini and Latreille 1801)
Chicken Turtle
Barry Mansell / CRM 3 / Glades Co., Florida
USA (Florida)
Deirochelys reticularia chrysea Schwartz 1956a:476
Type locality: “5.8 miles east of Monroe Station, Collier County,
Florida” [USA].
Deirochelys reticularia miaria Schwartz 1956a
Western Chicken Turtle
(subspecies: reticularia = red, chrysea = purple, miaria = orange)
USA (Alabama, Arkansas, Florida, Georgia, Louisiana, Mississippi, Missouri, North Carolina, Oklahoma, South
Carolina, Texas, Virginia)
CBFTT Account: Buhlmann, Gibbons, and Jackson (2008)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Near Threatened (2011)
Deirochelys reticularia reticularia (Latreille in Sonnini and
Latreille 1801)
Eastern Chicken Turtle
John L. Carr / Ouachita Parish, Louisiana
USA (Arkansas, Louisiana, Mississippi, Missouri, Oklahoma,
Texas)
Deirochelys reticularia miaria Schwartz 1956a:486
Type locality: “College Station, Brazos County, Texas” [USA].
Graptemys Agassiz 1857a (12:14, 14:14) (19)
Graptemys Agassiz 1857a:252,436
Type species: Graptemys geographica [= Testudo geographica
LeSueur 1817], by subsequent designation by Stejneger and Barbour (1917:117).
Kurt A. Buhlmann / Virginia
USA (Alabama, Florida, Georgia, Louisiana, Mississippi,
North Carolina, South Carolina, Virginia)
Testudo reticularia Latreille in Sonnini and Latreille
1801:124, Emys reticularia, Clemmys reticularia, Deirochelys reticularia, Deirochelys reticularia reticularia
Neoclemmys Baur in Lindeman 2013:20 (nomen nudum)
Megaloclemmys Baur in Lindeman 2013:20 (nomen nudum)
Graptemys barbouri Carr and Marchand 1942 (19, 21)
Barbour’s Map Turtle
Type locality: “Caroline” [USA]. Restricted to “Charleston” [South
Carolina, USA] by Harper (1940:711); to “vicinity of Charleston,
South Carolina” [USA] by Schmidt (1953:104); and to “9 miles
northwest of Charleston, Charleston County, South Carolina”
[USA] by neotype designation by Schwartz (1956a:466).
Testudo reticulata Daudin 1801:144 (nomen novum), Emys
reticulata, Clemmys (Clemmys) reticulata, Deirochelys
reticulata, Hirochelys reticulata, Chrysemys reticulata,
Chrysemys reticulatus
Type locality: “Caroline” [USA]. Restricted to “vicinity of Charleston, South Carolina” [USA] by Schmidt (1953:104).
David Dennis / CRM 3 / Florida
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Graptemys ernsti Lovich and McCoy 1992 (19, 21)
Escambia Map Turtle
James C. Godwin / CBFTT / Yellow R., Covington Co., Alabama
USA (Alabama, Florida, Georgia)
IUCN Red List: Vulnerable A2bcde (2013); Previously: Near
Threatened (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys barbouri Carr and Marchand 1942:98, Malaclemys barbouri
Type locality: “Chipola River north of Marianna, Jackson County,
Florida” [USA].
Graptemys caglei Haynes and McKown 1974 (19, 20)
Cagle’s Map Turtle
USA (Alabama, Florida)
CBFTT Account: Lovich, Godwin, and McCoy (2011)
IUCN Red List: Near Threatened (2013); Previously: Near
Threatened (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys ernsti Lovich and McCoy 1992:300, Graptemys
pulchra ernsti
Type locality: “Conecuh River, 1 mile upstream from County Road
4 Bridge, 14 km east of East Brewton, Escambia County, Alabama,
USA.”
John B. Iverson / Texas
USA (Texas)
IUCN Red List: Endangered A2c+4c; B2ab(iii) (2013); Previously: Vulnerable (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys caglei Haynes and McKown 1974:143
Type locality: “Guadalupe River, 8 km NW Cuero, DeWitt Co.,
Texas” [USA].
Graptemys flavimaculata Cagle 1954 (14:15) (19)
Yellow-blotched Map Turtle, Yellow-blotched Sawback
Peter V. Lindeman / Merrill, Mississippi
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Testudo geographica LeSueur 1817:86, Emys geographica,
Terrapene geographica, Clemmys (Clemmys) geographica, Clemmys geographica, Graptemys geographica,
Malacoclemmys geographica, Malacoclemmys geographicus, Malaclemys geographica, Malaclemys geographicus,
Malaclemmys geographicus, Graptemys geographicus
Type locality: “marsh, on the borders of Lake Erie” [USA]. Restricted
to “peninsula of Presque Isle and adjacent Presque Isle Bay in Erie
County, Pennsylvania” [USA] by Lindeman (2009:97).
Emys lesueurii Gray 1830e:12 (08:18, 10:7), Graptemys lesueurii,
Malacoclemmys lesueurii, Malaclemys lesueurii, Malaclemys lesueurii lesueurii
Type locality: “North America.”
USA (Mississippi)
CBFTT Account: Selman and Jones (2011)
IUCN Red List: Vulnerable A2bce+4ce (2013); Previously:
Endangered (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys flavimaculata Cagle 1954:167, Graptemys oculifera flavimaculata, Malaclemys flavimaculata
Emys megacephala Holbrook 1836:51
Type locality: “Cumberland river...[&]...in the neighbourhood of Nashville, Tennessee” [USA].
Emys macrocephala Agassiz 1857a:436 (nomen novum)
Graptemys gibbonsi Lovich and McCoy 1992 (10:8) (19)
Pascagoula Map Turtle
Type locality: “Pascagoula River, 13 miles S.W. of Lucedale,
George Co., Mississippi” [USA]. Emended to “Pascagoula River at
Old Benndale Crossing (T3S, R8W, Sec. 1), George County” [Mississippi, USA] by Cliburn (1971:17).
Graptemys geographica (LeSueur 1817) (08:18, 12:14) (19)
Northern Map Turtle, Common Map Turtle
Jeffrey E. Lovich / CBFTT / Chickasawhay R., nr. Leakesville, Mississippi
John B. Iverson / Dewart Lake, Kosciusko Co., Indiana
USA (Mississippi)
CBFTT Account: Lovich, Selman, and McCoy (2009)
IUCN Red List: Endangered A2bce+4ce (2013); Previously:
Near Threatened (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys gibbonsi Lovich and McCoy 1992:302, Graptemys pulchra gibbonsi
Type locality: “Chickasawhay River, Leakesville, Greene Co., Mississippi, USA.”
Canada (Ontario, Québec), USA (Alabama, Arkansas, Delaware, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky,
Louisiana, Maryland, Michigan, Minnesota, Mississippi,
Missouri, New Jersey, New York, North Carolina, Ohio,
Oklahoma, Pennsylvania, Tennessee, Vermont, Virginia,
West Virginia, Wisconsin)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix III (USA), as Graptemys spp.
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Graptemys nigrinoda Cagle 1954 (19, 22)
Black-knobbed Map Turtle, Black-knobbed Sawback
James C. Godwin / CBFTT / Alabama
USA (Louisiana, Mississippi)
CBFTT Account: Jones and Selman (2009)
IUCN Red List: Vulnerable B2ab(iii) (2013); Previously: Endangered (1996)
CITES: Appendix III (USA), as Graptemys spp.
Malacoclemmys oculifera Baur 1890a:262, Graptemys oculifera, Malaclemys lesueurii oculifera, Graptemys pseudogeographica oculifera, Graptemys oculifera oculifera,
Malaclemys oculifera
Type locality: “Mandeville, La.” [Louisiana, USA]. Emended to
“Pearl River, 26 miles east of Mandeville” [Louisiana, USA] by
Cagle (1953b:138).
USA (Alabama, Mississippi)
CBFTT Account: Blankenship, Butterfield, and Godwin (2008)
IUCN Red List: Least Concern (2013); Previously: Near Threatened (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys nigrinoda Cagle 1954:173 (22), Graptemys
oculifera nigrinoda, Graptemys nigrinoda nigrinoda,
Malaclemys nigrinoda
Graptemys ouachitensis Cagle 1953a (12:14, 12:15, 14:16) (19) or
Graptemys ouachitensis ouachitensis
Ouachita Map Turtle
Type locality: “Black Warrior River, above Lock 9, 17.5 miles SSW
of Tuscaloosa, Tuscaloosa County, Alabama” [USA].
Graptemys nigrinoda delticola Folkerts and Mount 1969:677 (22)
Type locality: “Hubbard’s Landing on Tensaw Lake, 2.6 air miles
SW of Latham, Baldwin County, Alabama” [USA].
Graptemys oculifera (Baur 1890a) (19)
Ringed Map Turtle, Ringed Sawback
Peter V. Lindeman / Paris Landing State Park, Henry Co., Tennessee
Robert L. Jones / CCB / Pearl R., Madison Co., Mississippi
USA (Alabama, Arkansas, Georgia (?), Illinois, Indiana, Iowa,
Kansas, Kentucky, Louisiana, Minnesota, Mississippi,
Missouri, Ohio, Oklahoma, Tennessee, Texas, West Virginia, Wisconsin)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix III (USA), as Graptemys spp.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Graptemys pseudogeographica ouachitensis Cagle 1953a:10,
Malaclemys pseudogeographica ouachitensis, Graptemys
ouachitensis, Graptemys ouachitensis ouachitensis
Type locality: “Ouachita River, four miles northeast of Harrisonburg, Louisiana” [USA].
Graptemys pearlensis Ennen, Lovich, Kreiser, Selman, and
Qualls 2010 (10:8) (19)
Pearl River Map Turtle
Cris Hagen / CCB / Pearl River, Mississippi
USA (Louisiana, Mississippi)
CBFTT Account: Ennen, Lovich, and Jones (2016)
IUCN Red List: Endangered A1bcde+A4bcde (2013)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys pearlensis Ennen, Lovich, Kreiser, Selman, and
Qualls 2010:104
55
Texas, Wisconsin)
Introduced: USA (Florida, Virginia)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys pseudogeographica pseudogeographica (Gray 1831d)
(08:19)
False Map Turtle
James H. Harding / Indiana
USA (Illinois, Indiana, Iowa, Kansas, Kentucky, Minnesota,
Missouri, Nebraska, North Dakota, South Dakota, Tennessee, Wisconsin)
Introduced: USA (Florida, Virginia)
Emys pseudogeographica Gray 1831d:31, Clemmys pseudogeographica, Graptemys pseudogeographica, Malacoclemmys pseudogeographicus, Malaclemys pseudogeographica, Malaclemys pseudogeographicus, Graptemys
pseudogeographicus, Graptemys pseudogeographica
pseudogeographica, Malaclemys pseudogeographica
pseudogeographica
Type locality: Not designated. Restricted to “Etats-Unis, Indiana,
rivière Wabash, entre MontVernon et Chaumetown (= Shawneetown)...près du confluent de la Wabash et de l’Ohio” [USA] by
Bour and Dubois (1983:45).
Graptemys pseudogeographica kohnii (Baur 1890a)
Mississippi Map Turtle
Type locality: “Mississippi, Copiah County, Pearl River at State
Highway 28, near Georgetown” [USA].
Graptemys pseudogeographica (Gray 1831d) (12:14, 12:15) (19, 23)
False Map Turtle
Stanley E. Trauth / Strawberry R., Lawrence Co., Arkansas
(subspecies: pseudogeographica = red, kohnii = purple;
overlap = intergrades)
USA (Arkansas, Illinois, Indiana, Iowa, Kansas, Kentucky,
Louisiana, Minnesota, Mississippi, Missouri, Nebraska,
North Dakota, Oklahoma, South Dakota, Tennessee,
USA (Arkansas, Kansas, Kentucky, Louisiana, Mississippi,
Missouri, Oklahoma, Tennessee, Texas)
Malacoclemmys kohnii Baur 1890a:263, Graptemys kohnii,
Malaclemys lesueurii kohnii, Graptemys pseudogeographica kohnii, Malaclemys kohnii
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Type locality: “Bayou Lafourche, La.; Bayou Teche, St. Martinsville, La.” [Louisiana, USA].
Graptemys pulchra Baur 1893c (14:17) (19)
Alabama Map Turtle
James C. Godwin / CBFTT / Tallapoosa R., Elmore Co., Alabama
USA (Louisiana, Texas)
IUCN Red List: Least Concern (2013), as part of Graptemys
ouachitensis
CITES: Appendix III (USA), as Graptemys spp.
Malacoclemmys intermedia Baur in Beyer 1900:21 (14:19) (nomen nudum), Graptemys intermedia
Graptemys pseudogeographica sabinensis Cagle 1953a:2,
Malaclemys pseudogeographica sabinensis, Graptemys
ouachitensis sabinensis, Graptemys sabinensis
Type locality: “Sabine River, eight miles southwest of Negreet,
Louisiana” [USA].
Graptemys versa Stejneger 1925 (19)
Texas Map Turtle
USA (Alabama, Georgia, Mississippi)
CBFTT Account: Lovich, Godwin, and McCoy (2014)
IUCN Red List: Near Threatened (2013); Previously: Least
Concern (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys pulchra Baur 1893c:675, Malacoclemmys pulchra, Malaclemys lesueurii pulchra, Malaclemys pulchra,
Graptemys pulchra pulchra
Type locality: “Montgomery, Alabama” [USA].
Graptemys alabamensis Baur in Lindeman 2013:20 (14:17) (nomen nudum)
Graptemys grandis Baur in Lindeman 2013:20 (14:17) (nomen
nudum)
Peter V. Lindeman / CBFTT / Live Oak Creek, Gillespie Co., Texas
Graptemys sabinensis Cagle 1953a (07:12, 12:15, 14:18) (19) or
Graptemys ouachitensis sabinensis
Sabine Map Turtle
Peter V. Lindeman / CCB / nr. Estherwood, Acadia Parish, Louisiana
USA (Texas)
CBFTT Account: Lindeman, Stuart, and Killebrew (2016)
IUCN Red List: Least Concern (2013); Previously: Near Threatened (1996)
CITES: Appendix III (USA), as Graptemys spp.
Graptemys pseudogeographica versa Stejneger 1925:463,
Graptemys versa, Malaclemys versa
Type locality: “Austin, Texas” [USA].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Malaclemys Gray 1844
Malaclemys Gray 1844:28
Type species: Malaclemys concentrica [= Testudo concentrica Shaw
1802 = subjective synonym of Testudo terrapin Schoepff 1793], by
original monotypy.
Malacoclemmys Agassiz 1857a:392,437 (nomen novum)
Euchyloclemmys Sclater 1858:292 (nomen novum)
Euchyloclemys Sclater in Gray 1863c:181 (nomen novum)
Malaclemmys Gray 1870c:41 (nomen novum)
Malaclemys terrapin (Schoepff 1793) (11:5) (24)
Diamondback Terrapin
USA (Connecticut, Delaware, Massachusetts, New Jersey,
New York, North Carolina, Maryland, Rhode Island,
Virginia)
Testudo terrapin Schoepff 1793:64, Emys terrapin, Clemmys
terrapin, Malaclemys terrapin, Malacoclemmys terrapin,
Malaclemys terrapin terrapin
Type locality: “America septentrionali...in foris Philadelphiae...et...
aquis subdulcibus Insulae Longae” [USA]. Restricted to “probably Delaware Bay” [Delaware and New Jersey, USA] by Hay
(1905:16); and to “coastal waters of Long Island” [New York, USA]
by Schmidt (1953:95).
Testudo concentrica Shaw 1802:43, Emys concentrica,
Malaclemys concentrica, Malaclemmys concentrica,
Malaclemmys centrata concentrica, Malaclemys centrata
concentrica, Malaclemys terrapin concentrica
Type locality: “North America...sold in the markets at Philadelphia”
[Pennsylvania, USA]. Restricted to “probably Delaware Bay” [Delaware or New Jersey, USA] by Hay (1905:16).
Testudo ocellata Link 1807:52
Type locality: “Nord-Amerika.” Restricted to “Philadelphia markets” [Pennsylvania, USA] by Schmidt (1953:96).
Emys concentrica polita Gray 1830e:11 (10:7)
Type locality: Not designated.
Testudo palustris Le Conte 1830:113 (nomen novum and junior
homonym, not = Testudo palustris Gmelin 1789), Emys
palustris, Malacoclemmys palustris
Type locality: “New-York to Florida, and even in the West Indies, in
salt water” [USA].
Emys macrocephalus Gray 1844:26 (junior homonym, not =
Emys macrocephala Spix 1824), Emys macrocephala
Type locality: “America?” Restricted to “Philadelphia markets”
[Pennsylvania, USA] by Schmidt (1953:96).
Malaclemys terrapin centrata (Latreille in Sonnini and Latreille
1801) (11:5)
Carolina Diamondback Terrapin
(subspecies: terrapin = red, centrata = purple, littoralis = orange,
macrospilota = pink, pileata = brown, rhizophorarum = green,
tequesta = gray; orange dot = trade)
Bermuda, USA (Alabama, Connecticut, Delaware, Florida,
Georgia, Louisiana, Maryland, Massachusetts, Mississippi,
New Jersey, New York, North Carolina, Rhode Island,
South Carolina, Texas, Virginia)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II
Malaclemys terrapin terrapin (Schoepff 1793)
Northern Diamondback Terrapin
Carla Van Ness / CRM 3 / Duval Co., Florida
Bermuda, USA (Georgia, Florida, North Carolina, South
Carolina)
Testudo centrata Latreille in Sonnini and Latreille 1801:145,
Emys centrata, Clemmys (Clemmys) centrata, Malaclemmys centrata, Malaclemys centrata, Malaclemys centrata
centrata, Malaclemmys terrapin centrata, Malaclemys
terrapin centrata
Type locality: “les grands marais de la Caroline” [USA]. Restricted
to “neighborhood of Charleston, South Carolina” [USA] by Hay
(1905:14).
Emys concentrica livida Gray 1831d:27, Emys livida
Joseph C. Mitchell / Fisherman Island, Northampton Co., Virginia
Type locality: “America Boreali.” Restricted to “vicinity of Charleston, South Carolina” [USA] by Schmidt (1953:96).
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Malaclemys terrapin littoralis Hay 1905
Texas Diamondback Terrapin
Aaron S. Baxter / Nueces Bay, Nueces Co., Texas
USA (Texas)
Malaclemmys littoralis Hay 1905:18, Malaclemys centrata
littoralis, Malaclemys pileata littoralis, Malaclemys terrapin littoralis
Type locality: “Rockport, Texas” [USA].
Malaclemys terrapin macrospilota Hay 1905
Ornate Diamondback Terrapin
Malaclemys terrapin rhizophorarum Fowler 1906
Mangrove Diamondback Terrapin
Brian Mealey / CRM 3 / Monroe Co., Florida
USA (Florida)
Malaclemys tuberculifera Gray 1844:29 (nomen oblitum)
Type locality: “California” [in error]. Restricted to “Philadelphia
market” [Pennsylvania, USA] by Schmidt (1953:96) [in error];
shown to be from the Florida Keys [Florida, USA] by Ernst and
Hartsell (2000:887), but they did not formally restrict the type
locality.
Malaclemmys littoralis rhizophorarum Fowler 1906:112,
Malaclemmys terrapin rhizophorarum, Malaclemys terrapin rhizophorarum
Type locality: “Boca Grande Key, Florida” [USA].
Malaclemys terrapin fordorum Wood 1994:1 (nomen nudum)
Malaclemys terrapin tequesta Schwartz 1955
Eastern Florida Diamondback Terrapin
Brian Mealey / CRM 3 / Monroe Co., Florida
USA (Florida)
Malaclemmys macrospilota Hay 1905:16, Malaclemys
centrata macrospilota, Malaclemys pileata macrospilota,
Malaclemys terrapin macrospilota
Type locality: “Charlotte Harbor, Florida” [USA].
Malaclemys terrapin pileata (Wied 1865)
Mississippi Diamondback Terrapin
Richard D. Bartlett / CRM 3 / Martin Co., Florida
USA (Florida)
Malaclemys terrapin tequesta Schwartz 1955:158
Type locality: “Miami Beach, Dade County, Florida” [USA].
Andrew T. Coleman / CCB / Bayou Caddy, Hancock Co., Mississippi
USA (Alabama, Florida, Louisiana, Mississippi, Texas)
Emys pileata Wied 1865:17, Malaclemmys pileata, Malaclemys centrata pileata, Malaclemys pileata pileata, Malaclemys terrapin pileata
Type locality: “Sümpfen mit salzigem Wasser an der Mündung
des Mississippi bei New-Orleans” [Louisiana, USA]. Emended to
“New Orleans, Louisiana” [USA] by Hay (1905:17).
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Pseudemys Gray 1856a (09:13, 12:16, 14:20)
Pseudemys Gray 1856a:197
Type species: Pseudemys concinna [= Testudo concinna LeConte
1830], by subsequent designation by Baur (1893a:221).
Ptychemys Agassiz 1857a:252,431
Type species: Ptychemys concinna [= Testudo concinna LeConte
1830], by subsequent designation by Brown (1908:114).
Nectemys Agassiz 1857b:642 (nomen novum)
Pseudemys alabamensis Baur 1893a
Alabama Red-bellied Cooter
Robert H. Mount / CCB / CBFTT / Alabama
(orange dots = waifs on offshore islands)
USA (Alabama, Mississippi)
CBFTT Account: Leary, Dobie, Mann, Floyd, and Nelson
(2008)
IUCN Red List: Endangered B1+2c (1996)
TFTSG Draft Red List: Endangered (2011)
Pseudemys alabamensis Baur 1893a:224, Pseudemys rubriventris alabamensis, Chrysemys (Pseudemys) alabamensis, Chrysemys rubriventris alabamensis
Type locality: “Mobile bay, Ala.” [Alabama, USA].
Pseudemys alabamiensis Beyer 1900:20 (nomen nudum)
Pseudemys concinna (Le Conte 1830) (09:13) (25)
River Cooter
(subspecies: concinna = red, suwanniensis = purple;
orange dot = introduced suwanniensis)
USA (Alabama, Arkansas, Florida, Georgia, Illinois, Indiana,
Kansas, Kentucky, Louisiana, Mississippi, Missouri, North
Carolina, Ohio, Oklahoma, South Carolina, Tennessee,
Texas, Virginia, West Virginia)
CBFTT Account: Ward and Jackson (2008)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
Pseudemys concinna concinna (Le Conte 1830) (07:13, 09:13, 10:9) (25)
Eastern River Cooter
Peter A. Meylan / CBFTT / CRM 3 / Spring Creek, Decatur Co., Georgia
USA (Alabama, Arkansas, Florida, Georgia, Illinois, Indiana,
Kansas, Kentucky, Louisiana, Mississippi, Missouri, North
Carolina, Ohio, Oklahoma, South Carolina, Tennessee,
Texas, Virginia, West Virginia)
Testudo concinna Le Conte 1830:106 (junior homonym, not
= Emys concinna Guérin 1829 [nomen oblitum]) (25), Emys
(Testudo) concinna, Terrapene concinna, Clemmys (Clemmys) concinna, Pseudemys concinna, Ptychemys concinna,
Chrysemys concinna, Pseudemys floridana concinna,
Pseudemys concinna concinna, Chrysemys concinna
concinna
Type locality: “rivers of Georgia and Carolina, where the beds are
rocky...never...below Augusta on the Savannah, or Columbia on
the Congaree” [USA]. Restricted to “vicinity of Columbia, South
Carolina” [USA] by Schmidt (1953:101).
Emys annulifera Gray 1830e:12 (10:7), Trachemys annulifera
Type locality: Not known. Restricted to “Columbia, South Carolina” [USA] by Schmidt (1953:101).
Emys hieroglyphica Holbrook 1836:47, Pseudemys hieroglyphica, Ptychemys hieroglyphica, Clemmys hieroglyphica, Chrysemys hieroglyphica, Pseudemys floridana hieroglyphica, Pseudemys concinna hieroglyphica, Chrysemys
concinna hieroglyphica
Type locality: “Cumberland river” [probably Tennessee, USA].
Emys mobilensis Holbrook 1838a:53, Ptychemys mobilensis,
Clemmys mobilensis, Pseudemys mobilensis, Chrysemys
mobilensis, Pseudemys floridana mobilensis, Pseudemys
concinna mobilensis, Chrysemys concinna mobilensis
Type locality: “Alabama...in the neighbourhood of Mobile” [USA].
Emys labyrinthica Duméril and Bibron in Duméril and
Duméril 1851:13, Clemmys labyrinthica, Pseudemys
labyrinthica, Chrysemys labyrinthica
Type locality: “Wabash-River (États-Unis)” [Indiana, USA].
Ptychemys hoyi Agassiz 1857a:433, Pseudemys concinna
hoyi, Pseudemys floridana hoyi, Chrysemys floridana hoyi
Type locality: Not designated. Restricted to “vicinity of Springfield,
Missouri” [USA] by Schmidt (1953:101).
Emys orthonyx Wied 1865:23
Type locality: “südlichen Gewässern bei New-Orleans” [Louisiana,
USA].
Pseudemys vioscana Brimley 1928:66
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Type locality: “Lake Des Allemands, La.” [Louisiana, USA].
Pseudemys elonae Brimley 1928:67
Type locality: “a pond in Guilford County, North Carolina, not far
from Elon College, in the Cape Fear drainage” [USA].
Pseudemys concinna metteri Ward 1984:34
Type locality: “Old Fort Cobb, Caddo County, Oklahoma” [USA].
Pseudemys concinna suwanniensis Carr 1937 (07:15, 09:13)
Suwannee Cooter
floridana; Previously: Least Concern [Not Listed] (1996)
Testudo floridana Le Conte 1830:100, Terrapene floridana,
Emys floridana, Clemmys (Clemmys) floridana, Pseudemys floridana, Chrysemys floridana, Pseudemys floridana
floridana, Chrysemys floridana floridana, Pseudemys
concinna floridana
Type locality: “St. John’s river of East Florida” [USA]. Emended
to “lower reaches of the St. John’s River (Duval County), Florida”
[USA] by Bour (2003:540).
Pseudemys gorzugi Ward 1984 (07:16, 12:16, 14:19)
Rio Grande Cooter
Dale R. Jackson / CBFTT / CRM 3 / Wakulla River, Wakulla Co., Florida
USA (Florida, Georgia)
Pseudemys floridana suwanniensis Carr 1937:4, Pseudemys
concinna suwanniensis, Chrysemys concinna suwanniensis, Pseudemys suwanniensis
Type locality: “Suwannee River at Manatee Springs, Levy-Dixie
County line, Florida” [USA].
Charles W. Painter / CBFTT / Black River, Eddy Co., New Mexico
Pseudemys floridana (Le Conte 1830) (07:14, 09:13, 10:9) or
Pseudemys concinna floridana or
Pseudemys floridana floridana
Coastal Plain Cooter
(orange dots = introduced or misidentified)
John Jensen / CRM 3 / Okaloosa Co., Florida
Mexico (Chihuahua [?], Coahuila, Nuevo Leon, Tamaulipas),
USA (New Mexico, Texas)
CBFTT Account: Pierce, Stuart, Ward, and Painter (2016)
IUCN Red List: Near Threatened (2013); Previously: Near
Threatened (1996)
Pseudemys concinna gorzugi Ward 1984:29, Pseudemys
gorzugi
Type locality: “3 1/2 mi. W Jimenez, Río San Diego, Coahuila,
México, 850 feet altitude.”
(orange dots = possibly introduced)
USA (Alabama, Georgia, Florida, Mississippi, North Carolina,
South Carolina, Virginia)
IUCN Red List: Least Concern (2013), as Pseudemys concinna
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
61
Pseudemys nelsoni Carr 1938a (09:13)
Florida Red-bellied Cooter
(orange dot = introduced)
Gary Luciano / CBFTT / Big Cypress National Preserve, Collier Co., Florida
USA (Florida)
IUCN Red List: Least Concern (2013), as Pseudemys
peninsularis
Pseudemys floridana persimilis † Hay 1916a:71 (nomen
dubium et oblitum) [Pleistocene, USA (Florida)]
Type locality: “Vero, St. Lucie County, Florida” [USA].
Pseudemys floridana peninsularis Carr 1938b:105, Chrysemys floridana peninsularis, Pseudemys peninsularis
Type locality: “Crystal Springs, Pasco County, Florida” [USA].
Pseudemys rubriventris (Le Conte 1830)
Northern Red-bellied Cooter
(orange dot = possibly introduced)
USA (Florida, Georgia)
Introduced: British Virgin Islands (Tortola), USA (Texas)
CBFTT Account: Jackson (2010)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
Deirochelys floridana † Hay 1908b:346 (nomen dubium)
[Pleistocene, USA (Florida)]
Type locality: “Hillsboro County, Florida...Peace Creek beds”
[USA].
Trachemys jarmani † Hay 1908b:351 (nomen dubium) [Late
Pleistocene, USA (Florida)], Pseudemys jarmani
Type locality: “Hillsboro County, Florida...Peace Creek beds”
[USA].
James H. Harding / Virginia
Pseudemys nelsoni Carr 1938a:307, Pseudemys rubriventris
nelsoni, Chrysemys (Pseudemys) nelsoni, Chrysemys
rubriventris nelsoni
Type locality: “Fellsmere, Indian River County, Florida” [USA].
Pseudemys peninsularis Carr 1938b (07:17, 09:13, 10:9) or
Pseudemys floridana peninsularis
Peninsula Cooter
(orange dot = possible occurrence)
Richard D. Bartlett / CRM 3 / Lee Co., Florida
USA (Delaware, Maryland, Massachusetts, New Jersey, North
Carolina, Pennsylvania, Virginia, West Virginia)
Introduced: South Korea
IUCN Red List: Near Threatened (2013); Previously: Near
Threatened (1996)
Testudo rubriventris Le Conte 1830:101, Terrapene rubriventris, Emys rubriventris, Clemmys (Clemmys) rubriventris,
Chrysemys rubriventris, Pseudemys rubriventris, Pseudemys rubiventris, Pseudemys rubriventris rubriventris,
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Chrysemys rubriventris rubriventris
Type locality: “rivers from New-Jersey to Virginia, chiefly...in such
as are rocky; in the Delaware, near Trenton” [USA]. Restricted
to “in the Delaware, near Trenton” [New Jersey, USA] by Baur
(1893a:224).
Emys irrigata Bell in Duméril and Bibron 1835:276, Emys
irrigita
Type locality: “la partie septentrionale du Nouveau-Monde.”
Restricted to “vicinity of Trenton, New Jersey” [USA] by Schmidt
(1953:103).
Emys rivulata Gray 1844:22 (junior homonym, not = Emys
rivulata Valenciennes in Bory de Saint-Vincent 1833)
Type locality: “N. America.” Restricted to “vicinity of Trenton, New
Jersey” [USA] by Schmidt (1953:103).
Pseudemys extincta † Hay 1908b:356 (nomen dubium) [Early
Pleistocene, Blancan, USA (Florida)]
and Smith 1980:434). Rhodin and Carr (2009:14) demonstrated that
Testudo scabra Linnaeus 1758 was synonymous with Rhinoclemmys
punctularia [= Testudo punctularia Daudin 1801] and declared T.
scabra a nomen oblitum, validating Lindholm’s designation.
Callichelys Gray 1863c:179,181
Type species: Callichelys ornata [= Emys ornata Gray in Griffith and
Pidgeon 1830], by original designation.
Redamia Gray 1870c:35
Type species: Redamia olivacea [= Emys olivacea Gray 1856b =
subjective synonym of Pseudemys stejnegeri Schmidt 1928], by
original monotypy.
Trachemys adiutrix Vanzolini 1995 (11:6) (27) or
Trachemys dorbigni adiutrix
Maranhao Slider
Type locality: “Hillsboro County, Florida...probably the Peace
Creek formation” [USA].
Pseudemys rubriventris bangsi Babcock 1937:293, Chrysemys rubriventris bangsi, Pseudemys bangsi
Type locality: “Boot Pond, Plymouth, Massachusetts” [USA].
Corrected to “Gunner’s Exchange Pond, Plymouth, Massachusetts”
[USA] by Graham (1991:1).
Pseudemys texana Baur 1893a (12:16)
Texas Cooter
Jérôme Maran / Santo Amaro, Maranhão, Brazil
James H. Harding / Texas
Brazil (Maranhão, Piauí)
IUCN Red List: Endangered B1+2c (1996)
TFTSG Draft Red List: Near Threatened (2011)
Trachemys adiutrix Vanzolini 1995:111, Trachemys dorbigni
adiutrix
Type locality: “Brasil: Maranhão: Santo Amaro, 02º33' S, 43º14' W”
[Brazil]
Trachemys callirostris (Gray 1856b) (11:6) (26) or
Trachemys venusta callirostris
Colombian Slider
USA (Texas)
IUCN Red List: Least Concern (2013); Previously: Least Concern [Not Listed] (1996)
Pseudemys texana Baur 1893a:223, Chrysemys texana,
Pseudemys floridana texana, Pseudemys concinna texana,
Chrysemys concinna texana
Type locality: “San Antonio, Texas” [USA].
Trachemys Agassiz 1857a (07:18, 09:14, 11:6, 14:21, 14:22, 14:23) (26)
Trachemys Agassiz 1857a:252,434
Type species: Trachemys scabra [= Testudo scabra Linnaeus 1758],
by subsequent designation by Brown (1908:114), but its uncertain
identity led Lindholm (1929:280) to designate Trachemys troosti
[= Emys troosti Holbrook 1836] as the type species (see also Smith
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
(subspecies: callirostris = red, chichiriviche = purple;
orange dot = trade or introduced)
Colombia (Antioquia, Atlántico, Bolívar, Cesar, Córdoba,
Cundinamarca, La Guajira, Magdalena, Santander, Sucre),
Venezuela (Carabobo, Falcón, Yaracuy, Zulia)
CBFTT Account: Bock, Páez, and Daza (2010)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Vulnerable (2011)
Trachemys decorata (Barbour and Carr 1940) (14:21)
Hispaniolan Slider
Trachemys callirostris callirostris (Gray 1856b) (07:19, 11:6) (26) or
Trachemys venusta callirostris
Colombian Slider
Pablo Feliz / Cabral, Barahona, Dominican Republic / male
Vivian P. Páez / CBFTT / Colombia
Colombia (Antioquia, Atlántico, Bolívar, Cesar, Córdoba,
Cundinamarca, La Guajira, Magdalena, Santander, Sucre),
Venezuela (Zulia)
Emys callirostris Gray 1856b:25, Callichelys callirostris,
Pseudemys callirostris, Chrysemys ornata callirostris,
Pseudemys scripta callirostris, Pseudemys ornata callirostris, Chrysemys callirostris, Chrysemys scripta callirostris,
Trachemys scripta callirostris, Trachemys callirostris,
Trachemys ornata callirostris, Trachemys dorbigni callirostris, Trachemys callirostris callirostris
Type locality: “America.” Restricted to “Unterlauf des Rio Magdalena” [Colombia] by Müller (1940:109).
Trachemys callirostris chichiriviche (Pritchard and Trebbau
1984) (07:19, 11:6) (26) or
Trachemys venusta chichiriviche
Venezuelan Slider
Dominican Republic, Haiti
IUCN Red List: Vulnerable B1+2c (1996)
Pseudemys decorata Barbour and Carr 1940:409, Pseudemys
terrapen decorata, Chrysemys (Trachemys) decorata,
Chrysemys terrapen decorata, Trachemys decorata,
Trachemys stejnegeri decorata
Type locality: “Fond Parisien, Haiti.”
Trachemys decussata (Bell in Griffith and Pidgeon 1830) (14:22)
Cuban Slider
Peter C.H. Pritchard / CBFTT / central coastal Venezuela
Venezuela (Carabobo, Falcón, Yaracuy)
Pseudemys scripta chichiriviche Pritchard and Trebbau
1984:191, Trachemys scripta chichiriviche, Trachemys
ornata chichiriviche, Trachemys callirostris chichiriviche
Type locality: “Lago de Tacarigua, Edo. Falcón, Venezuela (68º15'
W, 11º4' N).”
(subspecies: decussata = red, angusta = purple;
orange dots = introduced angusta)
Cayman Islands [historic introduction?], Cuba, Jamaica [prehistoric introduction?]
IUCN Red List: Least Concern [Not Listed] (1996)
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Trachemys decussata decussata (Bell in Griffith and Pidgeon 1830)
(08:17, 14:22)
Trachemys decussata angusta (Barbour and Carr 1940) (14:22)
Western Cuban Slider
Eastern Cuban Slider
Vincenzo Ferri / Cuba [captivity]
Cuba, Jamaica [prehistoric introduction?]
Testudo rugosa Shaw 1802:28 (partim, nomen dubium and
junior homonym, not = Testudo rugosa Van-Ernest in
Daudin 1801), Emys rugosa, Trachemys rugosa, Clemmys
rugosa, Pseudemys rugosa, Pseudemys rugosa rugosa,
Pseudemys terrapen rugosa, Chrysemys terrapen rugosa,
Trachemys terrapen rugosa
Type locality: Not designated. Restricted to “Rio Jobabo drainage in
eastern Cuba” by Mittleman (1947:176).
Emys decussata Bell in Griffith and Pidgeon 1830:76 (08:17),
Ptychemys decussata, Clemmys decussata, Pseudemys
decussata, Pseudemys decussata decussata, Pseudemys rugosa decussata, Pseudemys terrapen decussata,
Chrysemys (Trachemys) decussata, Chrysemys decussata
decussata, Chrysemys terrapen decussata, Trachemys
decussata, Trachemys decussata decussata
Type locality: Not designated. Restricted to “North America”
by Gray (1830e:11); to “America Boreali” by Gray (1831d:28);
to “Cuba, exclusive of the drainage systems of the Rio Jobabo
and the Caribbean slope of Pinar del Rio Province” by Mittleman (1947:176); and to “Westindien” by Mertens and Wermuth
(1955:366).
James H. Harding / Grand Cayman Island
Cayman Islands [historic introduction?], Cuba
Pseudemys decussata angusta Barbour and Carr 1940:402,
Pseudemys rugosa angusta, Pseudemys terrapen angusta,
Chrysemys terrapen angusta, Trachemys decussata angusta, Trachemys decorata angusta
Type locality: “Taco River, Pinar del Rio, Cuba.”
Pseudemys granti Barbour and Carr 1941:59, Pseudemys
terrapen granti, Pseudemys decussata granti, Pseudemys
stejnegeri granti, Chrysemys terrapen granti, Chrysemys
decussata granti, Trachemys decussata granti, Trachemys
granti, Trachemys stejnegeri granti
Type locality: “Grand Cayman” [Cayman Islands].
Trachemys dorbigni (Duméril and Bibron 1835) (07:20, 11:6) (27) or
Trachemys dorbigni dorbigni
D’Orbigny’s Slider
Emys vermiculata Gray 1844:25
Type locality: “Tropical America.” Restricted to “Westindien” by
Mertens and Wermuth (1955:366).
Emys jamao Duméril 1861b:435 (nomen nudum)
Emys jamao Vilaró 1867a:121
Type locality: Not designated. [Cuba].
Emys gnatho Vilaró 1867b:204
Type locality: Not designated. [Cuba].
Jérôme Maran / Brazil
Pseudemys decussata plana Barbour and Carr 1940:405,
Pseudemys terrapen plana, Chrysemys terrapen plana,
Trachemys decussata plana
Type locality: “Rio Jobabo, Western Oriente, Cuba.”
(orange dots = introduced)
Argentina (Buenos Aires, Corrientes, Entre Rios), Brazil (Rio
Grande do Sul, Santa Catarina), Uruguay
Introduced: Brazil (Bahia, Goiás, Minas Gerais, Paraná, Rio de
Janeiro, São Paulo, Sergipe, Tocantins)
IUCN Red List: Least Concern [Not Listed] (1996)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
TFTSG Draft Red List: Least Concern (2011)
Emys dorbigni Duméril and Bibron 1835:272, Clemmys
dorbigni, Pseudemys dorbigni, Chrysemys (Trachemys)
dorbigni, Pseudemys scripta dorbigni, Pseudemys dorbigni dorbigni, Chrysemys dorbigni dorbigni, Chrysemys
scripta dorbigni, Trachemys scripta dorbigni, Trachemys
dorbigni, Trachemys dorbigni dorbigni
Type locality: “Buenos-Ayres” [Argentina].
Clemmys (Rhinoclemmys) orbignyi Fitzinger 1835:124 (nomen novum), Emys orbignyi
Clemmys dorbignyi Boulenger 1886b:424 (nomen novum),
Chrysemys dorbignyi, Emys dorbignyi, Pseudemys
dorbignyi, Pseudemys dorbignyi dorbignyi, Pseudemys
scripta dorbignyi, Chrysemys scripta dorbignyi, Trachemys scripta dorbignyi, Trachemys dorbignyi, Trachemys
dorbignyi dorbignyi
Pseudemys dorbignyi brasiliensis Freiberg 1969:301 (07:20),
Pseudemys dorbigni brasiliensis, Pseudemys scripta brasiliensis, Chrysemys dorbigni brasiliensis, Chrysemys scripta brasiliensis, Trachemys scripta brasiliensis, Trachemys dorbigni
brasiliensis, Trachemys dorbignyi brasiliensis
Type locality: “río Guaíba, Porto Alegre, Brasil.”
Trachemys gaigeae (Hartweg 1939)
Big Bend Slider
65
Trachemys nebulosa gaigeae, Trachemys scripta gaigeae,
Trachemys gaigeae, Trachemys ornata gaigeae, Trachemys gaigeae gaigeae
Type locality: “Boquillas, Rio Grande River, Brewster County,
Texas” [USA].
Trachemys gaigeae hartwegi (Legler 1990) (07:18)
Nazas Slider
John B. Iverson / CBFTT / Río Nazas, Preza Francisco Zarco, Durango, Mexico
Mexico (Coahuila, Durango)
Pseudemys scripta hartwegi Legler 1990:89, Chrysemys
scripta hartwegi, Trachemys scripta hartwegi, Trachemys
ornata hartwegi, Trachemys nebulosa hartwegi, Trachemys gaigeae hartwegi
Type locality: “Río Nazas, 1.2 km east of Presa Lázaro Cardenas,
Durango, Mexico.”
Trachemys grayi (Bocourt 1868) (07:18, 10:10, 12:6) (26, 28)
Western Meso-American Slider
(subspecies: gaigeae = red, hartwegi = purple)
Mexico (Chihuahua, Coahuila, Durango), USA (New Mexico,
Texas)
CBFTT Account: Stuart and Ward (2009)
IUCN Red List: Vulnerable A2ce+4ce (2013); Previously:
Vulnerable (1996)
Trachemys gaigeae gaigeae (Hartweg 1939) (07:18)
Big Bend Slider
(orange dots = possible misidentified)
Costa Rica, El Salvador, Guatemala, Honduras, Mexico (Chiapas, Oaxaca), Nicaragua, Panama
IUCN Red List: Not Evaluated
Trachemys grayi grayi (Bocourt 1868) (07:18, 10:10, 12:6) (26, 28)
Gray’s Slider, Tehuantepec Slider
James N. Stuart / CBFTT / Socorro Co., New Mexico
Mexico (Chihuahua, Coahuila), USA (New Mexico, Texas)
Pseudemys scripta gaigeae Hartweg 1939:1, Pseudemys
gaigeae, Chrysemys scripta gaigeae, Chrysemys gaigeae,
Shi Haitao / Mexico [captivity]
El Salvador, Guatemala, Mexico (Chiapas, Oaxaca)
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Emys grayi Bocourt 1868:121 (senior homonym, not = Emys
grayi Günther 1869), Callichelys grayi, Chrysemys grayi,
Pseudemys grayi, Pseudemys ornata grayi, Pseudemys
scripta grayi, Chrysemys scripta grayi, Trachemys scripta
grayi, Trachemys grayi, Trachemys ornata grayi, Trachemys venusta grayi; Trachemys grayi grayi
Mexico (Baja California Sur, Sinaloa, Sonora)
IUCN Red List: Not Evaluated
Trachemys nebulosa nebulosa (Van Denburgh 1895) (07:18)
Baja California Slider
Type locality: “l’embouchure du Nagualate, dans le Pacifique
(Guatémala).”
Callichelys concinna Gray 1873a:148 (28)
Type locality: “San Mateo, Tehuantepec” [Oaxaca, Mexico].
Emys umbra Bocourt 1876b:26 (nomen novum), Pseudemys
umbra, Clemmys umbra, Chrysemys umbra, Pseudemys
scripta umbra
Trachemys grayi emolli (Legler 1990) (07:18, 11:6, 14:23) (26)
Nicaraguan Slider
James R. Buskirk / Cadeje, Baja California Sur, Mexico
Mexico (Baja California Sur)
Chrysemys nebulosa Van Denburgh 1895:84, Chrysemys
ornata nebulosa, Pseudemys ornata nebulosa, Pseudemys
nebulosa, Pseudemys scripta nebulosa, Chrysemys scripta
nebulosa, Trachemys scripta nebulosa, Trachemys dorbigni nebulosa, Trachemys ornata nebulosa, Trachemys
nebulosa, Trachemys nebulosa nebulosa
Type locality: “Mainland abreast of San José Island, Lower California...Los Dolores, L.C.” [Baja California Sur, Mexico].
Shi Haitao / No data [captivity]
Costa Rica, El Salvador, Honduras, Nicaragua
Pseudemys scripta emolli Legler 1990:91, Trachemys scripta
emolli, Trachemys ornata emolli, Trachemys emolli,
Trachemys venusta emolli, Trachemys grayi emolli
Trachemys nebulosa hiltoni (Carr 1942) (07:18)
Fuerte Slider
Type locality: “Río Tepetate, 2.5 km northeast of Granada,
Granada Province, Nicaragua.”
Trachemys grayi panamensis McCord, Joseph-Ouni, Hagen,
and Blanck 2010 (10:10, 11:6) (26)
Panamanian Slider
Costa Rica, Panama
Trachemys venusta panamensis McCord, Joseph-Ouni,
Hagen, and Blanck 2010:46, Trachemys grayi panamensis
Type locality: “Chiva-Chiva Road (trail), 1 km from Gaillané (Gaillard) Highway (Fort Clayton entrance), north of Miraflores Lake,
Pacific-side Panama Canal Zone, Panamá Province, Panama.”
Trachemys nebulosa (Van Denburgh 1895) (07:18)
Baja California Slider
Philip C. Rosen / San Miguel Zapotitlán, Rio Fuerte, Sinaloa, Mexico
Mexico (Sinaloa, Sonora)
Pseudemys scripta hiltoni Carr 1942:1, Pseudemys concinna
hiltoni, Chrysemys scripta hiltoni, Chrysemys gaigeae hiltoni, Trachemys scripta hiltoni, Trachemys ornata hiltoni,
Trachemys nebulosa hiltoni
Type locality: “Guirocoba about 28 miles southeast of Alamos,
Sonora, Mexico, at an elevation of approximately 1,485 feet.”
(subspecies: nebulosa = red, hiltoni = purple)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Trachemys ornata (Gray in Griffith and Pidgeon 1830) (07:18, 07:19, 10:10,
11:6) (26)
Ornate Slider
Tennessee, Texas, Virginia, West Virginia)
Introduced: Multiple global locations, most apparently Trachemys scripta elegans (see below)
IUCN Red List: Least Concern (2013); Previously: Near Threatened (1996)
Trachemys scripta scripta (Thunberg in Schoepff 1792) (09:15)
Yellow-bellied Slider
James R. Buskirk / Presidios, Sinaloa, Mexico
Kurt A. Buhlmann / Savannah River Site, Aiken Co., South Carolina
(orange dots = possibly introduced)
Mexico (Jalisco, Nayarit, Sinaloa)
Possibly Introduced: Mexico (Guerrero?, Michoacán?)
IUCN Red List: Vulnerable B1ab(iii)+2ab(iii) (2007)
Emys ornata Gray in Griffith and Pidgeon 1830:76 [Gray
1830c], Clemmys (Clemmys) ornata, Callichelys ornata,
Pseudemys ornata, Chrysemys ornata, Chrysemys ornata
ornata, Pseudemys ornata ornata, Pseudemys scripta ornata, Chrysemys scripta ornata, Trachemys scripta ornata,
Trachemys ornata, Trachemys ornata ornata
Type locality: Not designated. Restricted to “South America”
by Gray (1830e:12); and to “America Meridionali...Mazetland”
[Mazatlán, Sinaloa, Mexico] by Gray (1831d:30).
Trachemys scripta (Thunberg in Schoepff 1792) (09:15)
Pond Slider, Common Slider
USA (Alabama, Florida, Georgia, North Carolina, South
Carolina, Virginia)
Introduced: South Korea, USA (Florida)
Testudo scripta Thunberg in Schoepff 1792:16 (09:15) (nomen
conservandum, ICZN 1985b), Emys scripta, Trachemys
scripta, Chrysemys scripta, Pseudemys scripta, Chrysemys
scripta scripta, Chrysemys palustris scripta, Pseudemys
scripta scripta, Trachemys scripta scripta
Type locality: Not known. Restricted to “Charleston, South Carolina” [USA] by Schmidt (1953:102).
Testudo serrata Daudin 1801:148 (senior homonym, not =
Testudo serrata Shaw 1802), Emys serrata, Terrapene serrata, Clemmys (Clemmys) serrata, Pseudemys serrata
Type locality: “la Caroline” [South Carolina, USA].
Emys occipitatis Gray in Griffith and Pidgeon 1830:75 [Gray
1830c]
Type locality: Not designated.
Emys vittata Gray 1830e:11 (10:7)
Type locality: “North America?”
Emys euglypha † Leidy 1889:97 (nomen dubium) [Pleistocene, uSa (Florida)], Trachemys euglypha, Pseudemys
euglypha
Type locality: “Florida...Arcadia, on Peace Creek” [USA].
Trachemys sculpta † Hay 1908b:351 (nomen dubium) [Pleistocene, uSa (Florida)], Pseudemys sculpta
Type locality: “Hillsboro County, Florida...probably the Peace Creek
beds” [USA].
Trachemys delicata † Hay 1916a:66 (nomen dubium) [Pleistocene, uSa (Florida)], Pseudemys delicata
Type locality: “Near Labelle, Lee County, Florida” [USA].
(subspecies: scripta = red, elegans = purple, troostii = orange;
overlap = intergrades; orange dots = introduced elegans)
Mexico (Nuevo Leon, Tamaulipas), USA (Alabama, Arkansas,
Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Mississippi, Missouri, Nebraska, New
Mexico, North Carolina, Ohio, Oklahoma, South Carolina,
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Trachemys scripta elegans (Wied 1839) (25)
Red-eared Slider
Pseudemys scripta petrolei, Trachemys scripta petrolei
Type locality: “Harden Co., Texas” [USA].
Trachemys lineata Gray 1873a:147
Type locality: “North America.” Restricted to “New Harmony, Posey
County, Indiana” [USA] by Schmidt (1953:103).
Pseudemys bisornatus † Cope 1878:228 [Pleistocene, USA
(Texas)], Pseudemys bisornata, Chrysemys scripta bisornata, Pseudemys scripta bisornata, Trachemys scripta
bisornata, Trachemys bisornata
Type locality: “South-western Texas” [USA]. Restricted to “Atascosa County, Texas” [USA] by Hay (1908b:354).
Trachemys trulla † Hay 1908b:355 (nomen dubium) [Pleistocene, uSa (Texas)], Pseudemys trulla
Type locality: “Hardin County, Texas” [USA].
Matthew Aresco / CRM 3 / Leon Co., Florida
Mexico (Nuevo Leon, Tamaulipas), USA (Alabama, Arkansas, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Mississippi, Missouri, Nebraska, New
Mexico [eastern], Ohio, Oklahoma, Tennessee, Texas,
West Virginia)
Introduced: Argentina, Australia (New South Wales,
Queensland, Victoria), Austria, Bahamas, Bahrain, Belgium, Bermuda, Bulgaria, Brazil, British Virgin Islands,
Cambodia, Canada (Ontario), Cayman Islands, Chile,
China (Hong Kong), Colombia, Cyprus, Czech Republic,
Denmark, Dominican Republic, Ecuador, Egypt, Finland,
France, French Polynesia, Germany, Great Britain, Greece,
Guadeloupe, Guam, Guyana, Honduras, Hungary, Indonesia (Java, Kalimantan, Papua, Sulawesi, Sumatra), Iran
(Mazandaran, Tehran), Ireland, Israel, Italy, Japan (mainland, Ryukyu Archipelago), Latvia, Malaysia (Peninsular,
East), Martinique, Mexico, Micronesia, Myanmar, Netherlands, Netherlands Antilles, New Zealand, Nicaragua,
Northern Mariana Islands [Saipan], Palau, Panama, Philippines (Cebu, Luzon, Mindanao), Poland, Portugal, Puerto
Rico, Réunion, Russia, Saudi Arabia, Seychelles (Mahé),
Singapore, Sint Maarten, Slovakia, Slovenia, South Africa,
South Korea, Spain (Balearic Islands, Continental), Sri
Lanka, Suriname, Sweden, Switzerland, Taiwan, Thailand,
Trinidad, Turkey, USA (Arizona, California, Colorado,
Connecticut, Delaware, Florida, Hawaii, Maine, Maryland, Massachusetts, Michigan, New Jersey, New Mexico
[western], New York, North Carolina, Ohio, Oregon,
Pennsylvania, South Carolina, Virginia, Washington), US
Virgin Islands, Vietnam
Emys concinna Guérin 1829:pl.1,f.3 (nomen oblitum and
senior homonym, not = Emys concinna Le Conte 1830) (25)
Trachemys scripta troostii (Holbrook 1836)
Cumberland Slider
James H. Harding / Tennessee
USA (Tennessee, Virginia)
Introduced: Latvia
Emys troostii Holbrook 1836:55, Trachemys troostii, Clemmys troostii, Pseudemys troostii, Chrysemys troostii,
Pseudemys scripta troostii, Pseudemys troostii troostii,
Chrysemys scripta troostii, Trachemys scripta troostii
Type locality: “Cumberland river” [Tennessee, USA].
Emys cumberlandensis Holbrook 1840:55
Type locality: “Tennessee...Cumberland river” [USA].
Trachemys stejnegeri (Schmidt 1928) (14:22)
Central Antillean Slider
Type locality: Not designated.
Emys elegans Wied 1839:176,213, Trachemys elegans,
Clemmys elegans, Pseudemys elegans, Chrysemys
elegans, Chrysemys scripta elegans, Chrysemys palustris
elegans, Pseudemys troostii elegans, Pseudemys scripta
elegans, Trachemys scripta elegans
Type locality: “Gegend von Harmony...in dem Wabasch und
Fox-River” [Indiana, USA]. Emended to “in Fox Rivers bei NewHarmony aus einem nebenflusse des Wabasch” [Indiana, USA] by
Wied (1865:41).
Emys holbrookii Gray 1844:23, Trachemys holbrookii
Type locality: “N. America. Louisiana” [USA].
Emys sanguinolenta Gray 1856b:26,pl.15,f.1
Type locality: Not designated.
Emys petrolei † Leidy 1868:176 [Late Pleistocene, Rancholabrean, USA (Texas)], Pseudemys petrolei, Chrysemys
petrolei, Chrysemys scripta petrolei, Trachemys petrolei,
(subspecies: stejnegeri = red, malonei = purple, vicina = orange)
Bahamas (Inagua), Dominican Republic, Haiti, Puerto Rico
Introduced: Dominica, Guadeloupe
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Near Threatened (2011)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Trachemys stejnegeri stejnegeri (Schmidt 1928)
Puerto Rican Slider
69
Trachemys taylori (Legler 1960) (07:18, 12:17)
Cuatro Cienegas Slider
James R. Buskirk / Cuatro Ciénegas, Coahuila, Mexico
Anders G.J. Rhodin / Fajardo, Puerto Rico
USA (Puerto Rico)
Emys olivacea Gray 1856b:30 (junior homonym, not = Emys
olivacea Schweigger 1812), Clemmys olivacea, Redamia
olivacea, Chrysemys olivacea
Type locality: “N. America?”
Pseudemys stejnegeri Schmidt 1928:147, Pseudemys palustris stejnegeri, Pseudemys stejnegeri stejnegeri, Pseudemys terrapen stejnegeri, Pseudemys decussata stejnegeri,
Chrysemys decussata stejnegeri, Chrysemys stejnegeri,
Chrysemys terrapen stejnegeri, Trachemys stejnegeri,
Trachemys stejnegeri stejnegeri
Type locality: “San Juan, Porto Rico” [Puerto Rico, USA].
Trachemys stejnegeri malonei (Barbour and Carr 1938)
Inagua Slider
Bahamas (Inagua)
Pseudemys malonei Barbour and Carr 1938:76, Pseudemys
palustris malonei, Pseudemys terrapen malonei, Chrysemys malonei, Chrysemys terrapen malonei, Trachemys
stejnegeri malonei, Trachemys malonei
Type locality: “ponds near Northwest Point, Great Inagua Island,
B.W.I.” [Bahamas].
Mexico (Coahuila)
IUCN Red List: Endangered A4e, B1ab(iii,v)+2ab(iii,v) (2007)
Pseudemys scripta taylori Legler 1960:75, Chrysemys scripta
taylori, Chrysemys gaigeae taylori, Chrysemys taylori,
Trachemys scripta taylori, Trachemys nebulosa taylori,
Trachemys ornata taylori, Trachemys taylori
Type locality: “16 km. S Cuatro Ciénegas, Coahuila, México.”
Trachemys terrapen (Bonnaterre 1789) (09:6, 14:22)
Jamaican Slider
Trachemys stejnegeri vicina (Barbour and Carr 1940)
Dominican Slider
Uwe Fritz / Montego Bay, Jamaica
Peter Paul van Dijk / Vuelta Larga, Dominican Republic
Dominican Republic, Haiti
Pseudemys stejnegeri vicina Barbour and Carr 1940:408,
Pseudemys terrapen vicina, Pseudemys decussata vicina,
Chrysemys decussata vicina, Chrysemys stejnegeri vicina,
Chrysemys terrapen vicina, Trachemys stejnegeri vicina
Type locality: “Sanchez, San Domingo” [Dominican Republic].
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Bahamas (Cat Island, Eleuthera [prehistoric introduction?]),
Jamaica
IUCN Red List: Vulnerable B1+2c (1996)
Testudo terrapen Lacepède 1788:129, synopsis[table] (09:6)
(nomen suppressum, ICZN 2005a)
Trachemys venusta venusta (Gray 1856b) (07:18, 10:10, 11:6, 12:18, 14:23) (26)
Meso-American Slider
Type locality: “aux Antilles, & particulièrement à la Jamaïque”
[Jamaica].
Testudo terrapen Bonnaterre 1789:30, Pseudemys terrapen,
Pseudemys terrapen terrapen, Chrysemys (Trachemys)
terrapen, Chrysemys terrapen, Chrysemys terrapen terrapen, Trachemys terrapen, Trachemys terrapen terrapen
Type locality: “la Jamaïque” [Jamaica].
Testudo palustris Gmelin 1789:1041 (senior homonym, not
= Testudo palustris Le Conte 1830), Trachemys palustris,
Pseudemys palustris, Chrysemys scripta palustris, Pseudemys
palustris palustris
Type locality: “Jamaicae aquis stagnantibus” [Jamaica].
Testudo fasciata Suckow 1798:40 (senior homonym, not =
Testudo fasciata Daudin 1801)
Type locality: “Amboina, und besonders in Nordamerika zu Carolina” [in error]. Restricted to “Jamaica” by Seidel (1988:23).
Testudo rugosa Shaw 1802:28 (partim, nomen dubium and
junior homonym, not = Testudo rugosa Van-Ernest in
Daudin 1801), Emys rugosa, Clemmys rugosa, Chrysemys
scripta rugosa, Pseudemys rugosa
Type locality: Not designated. Restricted to “Rio Jobabo drainage in
eastern Cuba” by Mittleman (1947:176).
Emys rugosa livida Gray 1831d:30
Type locality: “America septentrionali?”
Pseudemys felis Barbour 1935:205, Pseudemys palustris
felis, Pseudemys terrapen felis, Chrysemys decussata felis,
Chrysemys felis, Chrysemys terrapen felis, Trachemys terrapen felis, Trachemys felis
Vincenzo Ferri / Mexico [captivity]
Belize, Guatemala, Mexico (Campeche, Chiapas, Oaxaca,
Quintana Roo, Tabasco, Tamaulipas, Veracruz)
Emys venusta Gray 1856b:24 (12:18), Callichelys venusta,
Pseudemys scripta venusta, Chrysemys scripta venusta,
Trachemys scripta venusta, Trachemys ornata venusta,
Trachemys venusta, Trachemys venusta venusta
Type locality: “Southern States of America; Honduras.” Restricted
to “Honduras” by lectotype designation by Smith and Smith
(1980:495).
Emys valida Le Conte 1860:7, Clemmys valida
Type locality: “Honduras.”
Emys (Clemmys) salvini Günther 1885:4, Pseudemys salvini
Type locality: “Guatemala.”
Trachemys venusta cataspila (Günther 1885) (07:18, 10:10, 11:6) (26)
Huastecan Slider
Type locality: “Tea Bay, Cat Island, Bahamas.”
Trachemys venusta (Gray 1856b) (07:18, 10:6, 10:10, 11:6, 12:18, 14:23) (26)
Eastern Meso-American Slider
James R. Buskirk / Aldama, Tamaulipas, Mexico
Mexico (San Luis Potosi, Tamaulipas, Veracruz)
Emys ventricosa Gray 1856b:28 (nomen suppressum, ICZN
1985b), Pseudemys ventricosa
Type locality: Not known.
(subspecies: venusta = red, cataspila = purple,
iversoni = blue, uhrigi = brown; orange dots = introduced)
Belize, Colombia (Antioquia, Chocó), Costa Rica, El Salvador,
Guatemala, Honduras, Mexico (Campeche, Chiapas,
Oaxaca, Quintana Roo, San Luis Potosi, Tabasco, Tamaulipas, Veracruz, Yucatán), Nicaragua, Panama
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (South America regional)
(2011)
Emys (Clemmys) cataspila Günther 1885:4 (nomen conservandum, ICZN 1985b), Pseudemys cataspila, Chrysemys
ornata cataspila, Pseudemys scripta cataspila, Pseudemys
ornata cataspila, Chrysemys scripta cataspila, Trachemys
scripta cataspila, Trachemys ornata cataspila, Trachemys
venusta cataspila
Type locality: “Mexico.” Restricted to “Alvarado, Veracruz, Mexico” [in error] by Smith and Taylor (1950a:346, 1950b:32); and to
“Tampico, Tamaulipas” [Mexico] by Smith and Smith (1980:486).
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Trachemys venusta iversoni McCord, Joseph-Ouni, Hagen, and
Blanck 2010 (10:10, 11:6) (26)
Yucatan Slider
Trachemys scripta yaquia, Trachemys dorbigni yaquia,
Trachemys ornata yaquia, Trachemys yaquia
Type locality: “Río Mayo, Conicarit, Sonora, México (27º14' N,
109º06' W).”
eMydinae Rafinesque 1815 (17)
Emidania Rafinesque 1815:75
Emydidae Bell 1825a:302
Emydinae Cope 1870b:123
Clemmys Ritgen 1828
Chelopus Rafinesque 1815:75 (nomen nudum)
Clemmys Ritgen 1828:270
John B. Iverson / Cobá, Quintana Roo, Mexico
Mexico (Quintana Roo, Yucatán)
Trachemys venusta iversoni McCord, Joseph-Ouni, Hagen,
and Blanck 2010:45
Type locality: “Cenote on the north side of the highway, 13.8 km
east of Buctzotz, Yucatán, Mexico.”
Trachemys venusta uhrigi McCord, Joseph-Ouni, Hagen, and
Blanck 2010 (10:10, 11:6, 14:23) (26)
Uhrig’s Slider
Colombia (Antioquia, Chocó), Costa Rica, Honduras, Nicaragua, Panama
Testudo panama Perry 1810:[unpaginated], pl.33 (12:18) (nomen
oblitum et dubium)
Type locality: “countries of South America, adjoining to the Isthmus
of Panama.”
Type species: Clemmys punctata [= Testudo punctata Schoepff 1792
= subjective synonym of Testudo guttata Schneider 1792], by
subsequent designation by Baur (1892:43).
Chelopus Rafinesque 1832:64
Type species: Chelopus punctatus [= Testudo punctata Schoepff 1792
= subjective synonym of Testudo guttata Schneider 1792], by
original monotypy.
Nanemys Agassiz 1857a:252,442
Type species: Nanemys guttata [= Testudo guttata Schneider 1792], by
original monotypy.
Melanemys Shufeldt 1919:157
Type species: Melanemys guttatus [= Testudo guttata Schneider 1792],
by subsequent designation by Dunn (1920:8).
Clemmys guttata (Schneider 1792) (29)
Spotted Turtle
Trachemys venusta uhrigi McCord, Joseph-Ouni, Hagen, and
Blanck 2010:43
Type locality: “Río Chamelecón drainage 3 km south of San Pedro
Sula, northwestern Caribbean coastal Honduras.”
Trachemys yaquia (Legler and Webb 1970) (07:18)
Yaqui Slider
Barry Mansell / CRM 3 / Seminole Co., Florida
James R. Buskirk / Rio Mayo, below Presa Mocuzari, Sonora, Mexico
Mexico (Sonora)
IUCN Red List: Vulnerable B1ab(iii)+2ab(iii) (2007)
Pseudemys scripta yaquia Legler and Webb 1970:158,
Chrysemys scripta yaquia, Pseudemys ornata yaquia,
Canada (Ontario), USA (Connecticut, Delaware, Florida,
Georgia, Illinois, Indiana, Maine, Maryland, Massachusetts, Michigan, New Hampshire, New Jersey, New York,
North Carolina, Ohio, Pennsylvania, Rhode Island, South
Carolina, Vermont, Virginia, West Virginia)
IUCN Red List: Endangered A2cde+4ce (2013); Previously:
Vulnerable (1996)
CITES: Appendix II
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Testudo guttata Schneider 1792:264, Emys guttata, Geoclemys guttata, Nanemys guttata, Clemmys guttata, Geoclemmys guttata, Chelopus guttatus, Melanemys guttatus
Type locality: Not designated. Restricted to “Philadelphia, Pennsylvania” [USA] by Mittleman (1945:171); and to “vicinity of
Philadelphia, Pennsylvania” [USA] by Schmidt (1953:91).
Testudo punctata Schoepff 1792:25 (junior homonym, not
= Testudo punctata Lacepède 1788 or Testudo punctata
Bonnaterre 1789), Emys punctata, Clemmys punctata, Terrapene punctata, Chelopus punctatus
Type locality: “in paludosis Americae septentrionalis... Philadelphiam” [Pennsylvania, USA].
Testudo anonyma Schneider in Schoepff 1792:25 (nomen
nudum)
Geoclemmys sebae Gray 1869a:188
Type locality: Not designated. Restricted to “vicinity of Philadelphia” [Pennsylvania, USA] by Schmidt (1953:91).
Emys Duméril 1805 (07:21, 09:16, 10:11, 10:12, 11:7, 14:24)
Emydes Brongniart 1805:27 (nomen suppressum, ICZN 1995b)
Emys Duméril 1805:76 (10:11) (nomen conservandum, ICZN
1995b)
Type species: Emys europaea Schweigger [= Testudo europaea
Schneider 1783 = subjective synonym of Testudo orbicularis Linnaeus 1758], by subsequent designation by Fitzinger (1843:29).
Hydrone Rafinesque 1814:66
Type species: Hydrone orbicularis [= Testudo orbicularis Linnaeus
1758], by subsequent designation by Loveridge and Williams
(1957:201).
Emyda Rafinesque 1815:75 (nomen novum and senior homonym, not = Emyda Gray 1830e)
Lutremys Gray 1844:31
Type species: Cistudo (Lutremys) europaea [= Testudo europaea
Schneider 1783 = subjective synonym of Testudo orbicularis Linnaeus 1758], by original monotypy.
Emys orbicularis (Linnaeus 1758)
European Pond Turtle
(subspecies: orbicularis = red, eiselti = purple, galloitalica = blue,
hellenica = brown, ingauna = pink, occidentalis = green, persica = tourmaline;
unassigned E. orbicularis sensu lato = gray (Algeria, Tunisia, southern Turkey);
overlap = intergrades; orange dots = introduced; red dots = extirpated)
Albania, Algeria, Armenia, Austria, Azerbaijan, Belarus,
Belgium (extirpated), Bosnia and Herzegovina, Bulgaria,
Croatia, Czech Republic (extirpated, reintroduced),
Denmark (extirpated, reintroduced), Estonia (extirpated),
France (Continental, Corsica [prehistoric introduction?]),
Georgia, Germany, Greece, Hungary, Iran, Italy (Continental, Sardinia [prehistoric introduction]), Kazakhstan,
Kosovo, Latvia, Lithuania, Luxembourg (extirpated),
Macedonia, Moldova, Montenegro, Morocco, Netherlands
(extirpated), Poland, Portugal, Romania, Russia, Serbia,
Slovakia, Slovenia, Spain (Continental), Switzerland (extirpated, reintroduced), Syria, Tunisia, Turkey, Turkmenistan, Ukraine
Introduced: Spain (Balearic Islands)
IUCN Red List: Global: Near Threatened (1996); Regional: Europe: Near Threatened (2004); European Union: Vulnerable A2bcde (2004)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Emys orbicularis orbicularis (Linnaeus 1758) (09:17) (30, 31)
European Pond Turtle
Type locality: “Gräfve af Brågarps pastorat...Skåne..[&]..Götha
kanal, Östergöthland...vid Nordskogsvägen nära intill Svartjordshålan” [Gräfve in Brågarp’s Parish...Scania..[&]..Göta Canal,
East Götaland...along Nordskogs Road near Svartjordshålan]
[Sweden]. Restricted to “Schonen, Südschweden” [Scania (Skåne),
Sweden] by Fritz (1992:67).
Testudo (Emys) canstadiensis † Plieninger 1847:208 (31) [Pleistocene, Germany]
Type locality: “Württemberg...Cannstadt” [Germany].
Cistudo anhaltina † Giebel 1866a:1 [Holocene, subfossil,
Germany], Emys anhaltina
Uwe Fritz / Dnieper R. Delta, Ukraine
Austria, Belarus, Belgium (extirpated), Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic (extirpated,
reintroduced), Denmark (extirpated, reintroduced), Estonia
(extirpated), France, Georgia, Germany, Hungary, Italy,
Kazakhstan, Kosovo, Latvia, Lithuania, Luxembourg
(extirpated), Moldova, Netherlands (extirpated), Poland,
Romania, Russia, Serbia, Slovakia, Slovenia, Switzerland
(extirpated), Turkey, Ukraine
Introduced: Spain (Balearic Islands)
Testudo orbicularis Linnaeus 1758:198 (nomen conservandum, ICZN 1995b), Hydrone orbicularis, Emys orbicularis, Emys orbicularis orbicularis
Type locality: “meridionalibus Europae.” Restricted to “Mecklenburgisch-Pommersche Seenplatte” [Germany] by Fritz (1992:67),
and by neotype designation by Fritz (1994:65).
Testudo lutaria Linnaeus 1758:198, Emydes lutaria, Hydrone
lutaria, Emys lutaria, Clemmys (Clemmys) lutaria, Cistudo
lutaria
Type locality: “Italia, Oriente.” Restricted to “MecklenburgischPommersche Seenplatte” [Germany] by Fritz (1992:67).
Testudo terrestris Garsault 1764:pl.675 (10:13) (nomen oblitum
and senior homonym, not = Testudo terrestris Fermin 1765
or Testudo terrestris Forskål 1775)
Type locality: Not designated.
Type locality: “Latdorf...Nord-deutschland” [Germany].
Emended to “Lattorf vid Bernberg a. d. Saale” [Germany] by Kurck
(1917:23); and to “Latdorf bei Bernburg an der Saale, Deutschland”
[Germany] by Fritz (1995:227).
Emys lutaria taurica Mehnert 1890:537
Type locality: “Ufer des Dnjepr, einige Meilen von seiner Ausmündung” [Ukraine].
Emys europaea sparsa Dürigen 1897:14
Type locality: Not designated. Restricted to “Ungarn” [Hungary] by
Fritz (1992:67).
Emys europaea punctata Dürigen 1897:15
Type locality: Not designated. Restricted to “Mecklenburg” [Germany] by Fritz (1992:67).
Emys europaea concolor Dürigen 1897:15
Type locality: Not designated. Restricted to “Mark Brandenburg”
[Germany] by Fritz (1992:67).
Emys orbicularis aralensis Nikolsky 1915:24
Type locality: “Lac. Aral” [Kazakhstan].
Emys orbicularis luteofusca Fritz 1989:145 (09:17)
Type locality: “See-Ebene westlich von Ereğli, Provinz Konya,
Türkei” [Turkey].
Emys orbicularis colchica Fritz 1994:61 (09:17)
Type locality: “Batumi (Batum)” [Georgia].
Emys orbicularis eiselti Fritz, Baran, Budak, and Amthauer
1998
Eiselt’s Pond Turtle, Turkish Pond Turtle
Testudo europaea Schneider 1783:323, Emys europaea,
Terrapene europaea, Cistuda europaea, Cistudo europaea,
Cistudo (Lutremys) europaea, Lutremys europaea
Type locality: “meisten Ländern von Europa bis in Preussen” [Germany]. Restricted to “Frankfurt an der Oder” [Germany] by Fritz
(1992:67).
Testudo pulchella Schoepff 1801:113 (senior homonym, not =
Emys pulchella sensu Schweigger 1812), Emys pulchella
Type locality: Not known. Restricted to “Mecklenburgisch-Pommersche Seenplatte” by Fritz (1992:67).
Terrapene europea Bell 1825a:308 (nomen novum)
Emys turfa † Meyer 1835:67 [Holocene, subfossil, Germany], Cistudo lutaria turfa
Type locality: “Enkheim unweit Frankfurt” [Germany].
Clemmys schlotheimii † Fitzinger 1835:127 (30) (nomen nudum)
[Pleistocene, Germany]
Type locality: Not designated. Restricted to “Burgtonna”
[Thuringia, Germany] by lectotype designation by Karl and Paust
(2014:156).
Trionyx schlotheimii † Fitzinger 1835:128 (30) (nomen nudum)
[Pleistocene, Germany]
Type locality: Not designated. Restricted to “Burgtonna”
[Thuringia, Germany] by lectotype designation by Karl and Paust
(2014:156).
Emys fossilis † Nilsson 1841:206 (nomen nudum), Emys
lutaria fossilis
Emys lutaria borealis † Nilsson 1841:208 [Holocene, Boreal
(Atlantic), subfossil, Sweden]
Dinçer Ayaz / Samandağ Distr., Antakya, Turkey
Syria, Turkey
Emys orbicularis eiselti Fritz, Baran, Budak, and Amthauer
1998b:113
Type locality: “14 km NE of Fevzipaşa (about 450 m above sea
level), Vilayet Gaziantep” [Turkey].
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Emys orbicularis galloitalica Fritz 1995 (10:14)
Franco-Italian Pond Turtle
Type locality: “Dalmatien und Cephallonia” [Croatia and Greece].
Emys europaea maculosa Dürigen 1897:15
Type locality: Not designated. Restricted to “Dalmatien” [Croatia]
by Fritz (1992:68).
Emys orbicularis ingauna Jesu, Piombo, Salvidio, Lamagni,
Ortale, and Genta 2004 (10:14)
Ligurian Pond Turtle
Uwe Fritz / Plaine des Maures, France
France (Continental, Corsica [prehistoric introduction?]), Italy
(Continental, Sardinia [prehistoric introduction]), Spain
(Continental)
Introduced: Spain (Balearic Islands)
Emys orbicularis (galloitalica) capolongoi Fritz 1995:204
(10:14)
, Emys orbicularis capolongoi
Type locality: “Olbia (Sardinien)” [Italy].
Emys orbicularis (galloitalica) lanzai Fritz 1995:211 (10:14),
Emys orbicularis lanzai
Type locality: “Conca-Mündung bei Fontea, unweit Santa Lucia di
Porto-Vecchio (Korsika)” [France].
Emys orbicularis (galloitalica) galloitalica Fritz 1995:217,
Emys orbicularis galloitalica
Type locality: “5 km östlich Collobrières, Département Var, Südfrankreich” [France].
Pino Piccardo / Albenga, Italy
Italy (Continental)
Emys orbicularis ingauna Jesu, Piombo, Salvidio, Lamagni,
Ortale, and Genta 2004:139, Emys (Emys) orbicularis
ingauna
Type locality: “Peagna (Comune di Ceriale), Provincia di Savona
(Regione Liguria, Italia)” [Italy].
Emys orbicularis occidentalis Fritz 1993 (32)
Western Pond Turtle, Spanish Pond Turtle, Magreb Pond
Turtle
Emys orbicularis hellenica Valenciennes in Bory de Saint-Vincent
1833 (12:19, 14:25)
Hellenic Pond Turtle
Uwe Fritz / Ifrane, Morocco
Uwe Fritz / Kefalinia, Greece / male
Albania, Bosnia and Herzegovina, Croatia, Greece, Italy,
Kosovo, Macedonia, Montenegro, Serbia, Slovenia
Emys hellenica Valenciennes in Bory de Saint-Vincent
1833:planches, pl.8 (14:25), Cistuda hellenica, Emys orbicularis hellenica
Type locality: Not designated. Restricted to “plaine de Nisi que
baigne le Pamisus au coeur de la Messénie” [Peloponnes, Greece]
by Bibron and Bory de Saint-Vincent (1833:61).
Emys iberica Valenciennes in Bory de Saint-Vincent
1833:planches, pl.9 (14:25)
Type locality: Not designated. Restricted to “Morée, où
l’embouchure de l’Eurotas en est remplie, ainsi que le principal ruisseau de l’île de Tine” [Greece] by Bibron and Bory de Saint-Vincent
(1833:61).
Emys antiquorum Bory de Saint-Vincent 1835:Atlas, pl.9
[corrigenda] (14:25) (nomen novum et nomen nudum)
Type locality: Not designated.
Emys (Emys) hofmanni Fitzinger 1835:123 (12:19) (nomen
novum), Emys orbicularis hoffmanni, Cistudo hoffmanni
Emys orbicularis atra Werner 1897:15
Morocco, Portugal, Spain
Emys orbicularis occidentalis Fritz 1993:136 (32)
Type locality: “Lagune von Medhiya unweit Kenitra, Marokko”
[Morocco].
Emys orbicularis fritzjuergenobsti Fritz 1993:132 (09:18) (32)
Type locality: “Castellón de la Plana, Spanien” [Spain].
Emys orbicularis hispanica Fritz, Keller, and Budde
1996:132 (09:18)
Type locality: “Doñana, Huelva” [Spain].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Emys orbicularis persica Eichwald 1831 (07:23, 07:24, 09:19) (33)
Eastern Pond Turtle, Persian Pond Turtle
James F. Parham / CRM 4 / Shahrbijar, Gilan Prov., Iran
Armenia, Azerbaijan, Georgia, Iran (Ardabil, Gilan, Golestan,
Mazandaran), Russia (Dagestan), Turkmenistan
Introduced: Iran (Alborz, Teheran)
Emys europaea persica Eichwald 1831:196 (33), Emys europaea persicae, Emys orbicularis persica
Type locality: “provincia Masanderan” [Mazandaran Province,
Caspian Sea, Iran].
Emys europaea iberica Eichwald 1831:196 (09:19) (33), Emys
europaea ibericae, Emys orbicularis iberica
Type locality: Not designated. Restricted to “in Iberiae convallibus
paludosis et fluviis, Cyrum amnem petentibus” [in marshy Iberian
valleys and rivers, Kura River creeks] [Georgia and Azerbaijan] by
Eichwald (1840:47).
Emys orbicularis orientalis Fritz 1994:72
Type locality: “Bandar-e-Anzali (Enzeli), Prov. Gilan, Iran.”
Emys orbicularis kurae Fritz 1994:78
(09:19)
Type locality: “Bank (Bank Promisl) an der Kura-Mündung, Aserbaidshan” [Azerbaijan].
Italy (Sicily)
IUCN Red List: Data Deficient (2009)
Emys trinacris Fritz, Fattizzo, Guicking, Tripepi, Pennisi,
Lenk, Joger, and Wink 2005a:364
Type locality: “Lago Gian Fenaro, below the pass of Pizzo
Laminaria approximately 1400 m above sea level, Monte Nebrodi,
Sicily” [Italy]. Emended to “Laghetto Gianferraro...14.497241 E,
37.951625 N; Elevation: 1007 m a.s.l.” by Marrone et al. (2016:60).
Emys Duméril 1805 or
Actinemys Agassiz 1857a (07:22, 09:16, 10:12, 11:7, 14:24) (35)
Actinemys Agassiz 1857a:252,444
Type species: Actinemys marmorata [= Emys marmorata Baird and
Girard 1852], by original designation.
Emys marmorata (Baird and Girard 1852) (07:22, 10:15) (35) or
Actinemys marmorata
Northern Pacific Pond Turtle, Northern Western Pond Turtle
Emys orbicularis ssp. indet. (14:26)
Testudo purgotii † Ceselli 1846:24 (nomen oblitum) [Late
Pleistocene, Italy]
Type locality: “Viterbo...Viterbesi...acque Caje” [Italy].
Emys maior † Portis 1890:16 (14:26) (nomen dubium) [Late
Pliocene to Early Pleistocene, Villafranchian, Italy]
Type locality: “Poderaccio sotto Persignano nella Valle Superiore
dell’Arno” [Italy].
Emys latens † Portis 1890:16 (14:26) (nomen dubium) [Late
Pliocene to Early Pleistocene, Villafranchian, Italy]
Type locality: “Colombajolo presso S. Giovanni” [Italy].
David J. Germano / CBFTT / Fresno, San Joaquin Valley, California
Emys tigris Salvator 1897:280 (nomen nudum)
Type locality: “Mercadal, Insel Menorca” [Balearic Islands, Spain].
Emys trinacris Fritz, Fattizzo, Guicking, Tripepi, Pennisi, Lenk,
Joger, and Wink 2005 (34)
Sicilian Pond Turtle
Melita Vamberger / Laghetto Gorgo, Sicily, Italy
Canada (?) (British Columbia), USA (California, Nevada,
75
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Oregon, Washington)
Introduced: Australia (New South Wales)
CBFTT Account: Bury and Germano (2008)
IUCN Red List: Vulnerable A1cd (1996)
TFTSG Draft Red List: Vulnerable (2011)
Emys marmorata Baird and Girard 1852:177, Actinemys
marmorata, Clemmys marmorata, Geoclemmys marmorata, Chelopus marmoratus, Melanemys marmorata,
Clemmys marmorata marmorata, Actinemys marmorata marmorata, Emys marmorata, Emys marmorata
marmorata
marmorata pallida, Emys marmorata pallida, Emys pallida, Actinemys pallida
Type locality: “Lower Coyote Creek, near Alamitos, Orange
County, California” [USA].
Emys Duméril 1805 or
Emydoidea Gray 1870c (07:21, 09:16, 10:12, 11:7, 14:24) (36)
Emydoidea Gray 1870c:19
Type species: Emydoidea blandingii [= Cistuda blandingii Holbrook
1838b], by original monotypy.
Neoemys Lindholm 1929:282 (nomen novum)
Type locality: “Puget Sound” [Washington, USA].
Emys nigra Hallowell 1854:91 (senior homonym, not = Emys
nigra Blyth 1856)
Type locality: “Posa Creek, Lower California” [Kern County,
California, USA].
Emys blandingii (Holbrook 1838b) (36) or
Emydoidea blandingii
Blanding’s Turtle
Clemmys wosnessenskyi Strauch 1862:114, Geoclemmys
wosnessenskyi
Type locality: “Rio Sacramento in Californien” [USA].
Clemmys hesperia † Hay 1903:238 [Pliocene, USA
(Oregon)]
Type locality: “Rattlesnake beds, Rattlesnake Creek, Oregon”
[USA].
Emys pallida (Seeliger 1945) (07:22, 10:15) (35) or
Actinemys pallida
Southern Pacific Pond Turtle, Southern Western Pond Turtle
Janet Hostetter / CBFTT / Weaver Dunes, Minnesota
Robert H. Goodman, Jr. / Camp Pendleton, San Diego Co., California
Canada (Nova Scotia, Ontario, Québec), USA (Illinois, Indiana,
Iowa, Maine, Massachusetts, Michigan, Minnesota,
Missouri, Nebraska, New Hampshire, New York, Ohio,
Pennsylvania, South Dakota, Wisconsin)
CBFTT Account: Congdon, Graham, Herman, Lang, Pappas,
and Brecke (2008)
IUCN Red List: Endangered A2cde+4ce (2013); Previously:
Near Threatened (1996)
CITES: Appendix II
Testudo flava Lacepède 1788:135, synopsis[table] (09:6) (nomen suppressum, ICZN 1963)
Type locality: “Amérique..[&]..l’isle de l’Ascension.” Restricted to
“Amerika” by Wermuth (1956:407).
Testudo flava Bonnaterre 1789:26 (nomen oblitum)
Mexico (Baja California), USA (California)
CBFTT Account: Bury and Germano (2008) [as part of A.
marmorata]
IUCN Red List: Vulnerable A1cd (1996), as part of A.
marmorata
TFTSG Draft Red List: Vulnerable (2011), as part of A.
marmorata
Clemmys marmorata pallida Seeliger 1945:158, Actinemys
Type locality: “Amérique, l’ile de l’Ascension.”
Testudo meleagris Shaw 1793:147 (nomen suppressum,
ICZN 1963), Lutremys meleagris, Emys meleagris
Type locality: “America” [USA].
Cistuda blandingii Holbrook 1838b:35 (nomen conservandum, ICZN 1963), Cistudo blandingii, Emys blandingii,
Emydoidea blandingii, Neoemys blandingii
Type locality: “Fox river, a tributary of the Illinois” [Illinois, USA].
Emys twentei † Taylor 1943:250 [Pleistocene, USA (Kansas)]
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “north side of the Cimarron river, 13 miles southwest
of Meade, Meade County, Kansas (Loc. No. 7, XI Ranch)” [USA].
Glyptemys muhlenbergii (Schoepff 1801)
Bog Turtle
Glyptemys Agassiz 1857a (07:21)
Calemys Agassiz 1857a:252,443
Type species: Calemys muhlenbergii [= Testudo muhlenbergii Schoepff
1801], by original designation.
Glyptemys Agassiz 1857a:252,443
Type species: Glyptemys inculpta [= Testudo insculpta Le Conte 1830],
by original designation.
Glyptemys insculpta (Le Conte 1830)
Wood Turtle
Brian Zarate / TCC / Sussex Co., New Jersey
Anders G.J. Rhodin / Lunenburg, Worcester Co., Massachusettts
Canada (New Brunswick, Nova Scotia, Ontario, Québec), USA
(Connecticut, Delaware, Iowa, Maine, Maryland, Massachusetts, Michigan, Minnesota, New Hampshire, New
Jersey, New York, Pennsylvania, Rhode Island, Vermont,
Virginia, West Virginia, Wisconsin)
IUCN Red List: Endangered A2cd+4c (2013); Previously:
Vulnerable (1996)
CITES: Appendix II
Emys pulchella sensu Schweigger 1812:303 (junior homonym,
not = Testudo pulchella Schoepff 1801 [= Emys pulchella]),
Geoclemys pulchella, Glyptemys pulchella
Testudo insculpta Le Conte 1830:112, Clemmys (Clemmys)
insculpta, Clemmys insculpta, Emys insculpta, Glyptemys
insculpta, Chelopus insculptus, Calemys insculpta
Type locality: “the northern states” [USA]. Restricted to “vicinity of
New York City” [New York, USA] by Schmidt (1953:92).
Emys speciosa Gray 1830e:10
(10:7)
Type locality: “North America?” Restricted to “America Boreali,
New Jersey” [USA] by Gray (1831d:26).
Emys inscripta Gray 1831d:26 (nomen novum)
Emys speciosa levigata Gray 1831d:26
Type locality: “America Boreali, New Jersey” [USA].
USA (Connecticut, Delaware, Georgia, Maryland, Massachusetts, New Jersey, New York, North Carolina, Pennsylvania, South Carolina, Tennessee, Virginia)
IUCN Red List: Critically Endangered A2cd+4ce (2013); Previously: Endangered (1996)
CITES: Appendix I
Testudo muhlenbergii Schoepff 1801:132, Emys muhlenbergii, Emys mühlenbergii, Chersine muhlenbergii,
Terrapene muhlenbergii, Clemmys (Clemmys) muhlenbergii, Clemmys muhlenbergii, Geoclemys muhlenbergii,
Calemys mühlenbergii, Calemys muhlenbergii, Geoclemmys muhlenbergii, Chelopus muhlenbergii, Melanemys
muhlenbergii, Glyptemys muhlenbergii
Type locality: “Pensylvaniae rivulis” [USA]. Restricted to “Lancaster, Pennsylvania” [USA] by Stejneger and Barbour (1917:114).
Emys biguttata Say 1825:212 (10:16)
Type locality: “United States.” Restricted to “vicinity of Philadelphia” [Pennsylvania, USA] by Schmidt (1953:92).
Emys bipunctata Say in Gray 1830e:10 (nomen novum)
Emys fusca LeSueur in Gray 1831d:25 (nomen nudum)
Clemmys nuchalis Dunn 1917:624
Type locality: “side of Yonahlossee Road, about 3 miles from Linville, North Carolina...altitude, 4200 feet” [USA].
Terrapene Merrem 1820 (14:27)
Didicla Rafinesque 1815:75 (nomen nudum)
Terrapene Merrem 1820:27
Type species: Terrapene clausa [= Testudo clausa Gmelin 1789 = subjective synonym of Testudo carolina Linnaeus 1758], by subsequent
designation by Bell (1828c:514).
Cistuda Fleming 1822:270
Type species: “Box tortoise”, by original designation.
Didicla Rafinesque 1832:64
Type species: Didicla clausa [= Testudo clausa Gmelin 1789 =
77
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
subjective synonym of Testudo carolina Linnaeus 1758], by original
designation.
Cistudo Duméril and Bibron 1835:207 (nomen novum) (10:17)
Pyxidemys Fitzinger 1835:123
Type species: Pyxidemys clausa [= Testudo clausa Gmelin 1789 = subjective synonym of Testudo carolina Linnaeus 1758], by subsequent
designation by Fitzinger (1843:29).
IUCN Red List: Vulnerable A2bcde+4bcde (2013); Previously:
Near Threatened (1996)
CITES: Appendix II, as Terrapene spp.
Terrapene carolina carolina (Linnaeus 1758) (11:8, 14:27) (37, 38)
Eastern Box Turtle, Woodland Box Turtle
Emyoides Gray 1844:27
Type species: Emys (Emyoides) kinosternoides [= Emys kinosternoides
Gray 1830e = subjective synonym of Testudo carolina Linnaeus
1758], by original monotypy.
Onychotria Gray 1849:17
Type species: Cistudo (Onychotria) mexicana Gray 1849, by original
monotypy.
Pariemys Cope 1895:757
Type species: Pariemys bauri [= Terrapene bauri Taylor 1895], by
original monotypy.
Toxaspis Cope 1895:757
Type species: Toxaspis major [= Cistudo major Agassiz 1857a], by
original monotypy.
Cistudos Herrera 1901:36 (nomen novum et suppressum, ICZN
1922)
Terrapene carolina (Linnaeus 1758) (11:8, 14:27) (37)
Eastern Box Turtle, Common Box Turtle
Peter Paul van Dijk / CBFTT / Colesville, Montgomery Co., Maryland
Canada (Ontario [extirpated]), USA (Alabama, Connecticut,
Delaware, Georgia, Illinois, Indiana, Kentucky, Maine,
Maryland, Massachusetts, Michigan, Mississippi, New
Hampshire, New Jersey, New York, North Carolina, Ohio,
Pennsylvania, Rhode Island, South Carolina, Tennessee,
Virginia, West Virginia)
Testudo carolina Linnaeus 1758:198 (senior homonym, not
= Testudo carolina Le Conte 1830), Terrapene carolina,
Testudo carolinina, Terrapene carolinensis, Emys (Cistuda)
carolinae, Terrapene carolinae, Cistuda carolina, Cistudo
carolina, Cistudo carolinensis, Terrapene carolina carolina
Type locality: “Carolina” [USA]. Restricted to “vicinity of Charleston,
South Carolina” [USA] by Schmidt (1953:93).
Testudo carinata Linnaeus 1758:198, Terrapene carinata,
Cistudo carinata
Type locality: “Calidis regionibus.” Restricted to “vicinity of Charleston, South Carolina” [USA] by Schmidt (1953:93).
Testudo brevicaudata Lacepède 1788:169, synopsis[table] (09:6)
(nomen suppressum, ICZN 2005a), Testudo brevicauda
Type locality: “Caroline” [USA].
Testudo incarcerata Bonnaterre 1789:29
Type locality: “Philadelphie...L’Amérique septentrionale” [Pennsylvania, USA].
Testudo incarceratostriata Bonnaterre 1789:29
Type locality: “L’Amérique septentrionale.” Restricted to “vicinity
of Philadelphia” [Pennsylvania, USA] by Schmidt (1953:93).
Testudo clausa Gmelin 1789:1042, Emydes clausa, Emys
clausa, Didicla clausa, Terrapene clausa, Cistudo clausa,
Cinosternon clausum, Emys (Pyxidemys) clausa, Pyxidemys clausa, Cinosternum clausum
(subspecies: carolina = red, bauri = purple, major = blue,
mexicana = pink, triunguis = brown, yucatana = green;
overlap = intergrades, red dots = extirpated)
Canada (Ontario [extirpated]), Mexico (Campeche, Nuevo
León, Quintana Roo, San Luis Potosi, Tamaulipas, Veracruz, Yucatán), USA (Alabama, Arkansas, Connecticut,
Delaware, Florida, Georgia, Illinois, Indiana, Kansas,
Kentucky, Louisiana, Maine, Maryland, Massachusetts,
Michigan, Mississippi, Missouri, New Hampshire, New
Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Rhode Island, South Carolina, Tennessee, Texas,
Virginia, West Virginia)
CBFTT Account: Kiester and Willey (2015)
Type locality: “America septentrionali.” Restricted to “vicinity of
Philadelphia” [Pennsylvania, USA] by Schmidt (1953:94).
Testudo virgulata Latreille in Sonnini and Latreille 1801:100,
Emys virgulata, Terrapene virgulata
Type locality: “les grands bois de la Caroline” [USA]. Restricted to
“Charleston, South Carolina” [USA] by Schmidt (1953:94).
Emys schneideri Schweigger 1812:317, Emys (Pyxidemys)
schneideri
Type locality: Not known. Restricted to “vicinity of Philadelphia”
[Pennsylvania, USA] by Schmidt (1953:94).
Monoclida kentukensis Rafinesque 1822:5 (nomen suppressum, ICZN 1984)
Type locality: “United States...Kentucky.”
Didicla erythrops Rafinesque 1822:5 (nomen nudum)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “United States.”
Terrapene maculata Bell 1825a:309, Terrapene carolina
maculata
Type locality: Not known. Restricted to “vicinity of Philadelphia”
[Pennsylvania, USA] by Schmidt (1953:94).
Terrapene nebulosa Bell 1825a:310, Terrapene carolina
nebulosa
79
(Florida)]
Type locality: “Two miles west of Melbourne, Brevard County,
Fla.” [Florida, USA].
Terrapene carolina major (Agassiz 1857a) (11:8, 14:27) (37)
Gulf Coast Box Turtle
Type locality: Not known. Restricted to “vicinity of Philadelphia”
[Pennsylvania, USA] by Schmidt (1953:94).
Testudo irregulata Daudin in Gray 1830e:7 (10:7) (nomen
nudum)
Emys (Cistuda) carolinae fusca Gray 1830e:7 (10:7), Emys
carolinae fusca, Cistuda carolinae fusca
Type locality: “North America.”
Emys kinosternoides Gray 1830e:12 (10:7), Emys (Emyoides)
kinosternoides, Terrapene kinosternoides
Type locality: Not known. Restricted to “vicinity of Philadelphia”
[Pennsylvania, USA] by Schmidt (1953:94).
Emys cinosternoides Duméril and Bibron 1835:303 (nomen novum), Cistudo carolina cinosternoides, Cistudo
cinosternoides, Terrapene cinosternoides
Cistudo pickeringi Duméril 1855:199 (nomen nudum)
Cistudo virginea Agassiz 1857a:260,445
Type locality: “New England, and westward as far as Michigan, and
southward as far as the Carolinas” [USA]. Restricted to “vicinity of
Cambridge, Massachusetts” [USA] by Schmidt (1953:94).
Cistudo eurypygia † Cope 1870b:124 [Pleistocene, USA
(Maryland)], Terrapene eurypygia
Type locality: “Oxford Neck in Talbot Co., Maryland” [USA].
Toxaspis anguillulatus † Cope 1899:196 [Pleistocene, USA
(Pennsylvania)], Terrapene anguillulatus
Type locality: “Port Kennedy, Upper Merion Township, Montgomery County, Pennsylvania” [USA].
Testudo munda † Hay 1920:86 [Pleistocene, USA
(Tennessee)]
Type locality: “Whitesburg, Hamblen County, Tennessee” [USA].
Terrapene carolina bauri Taylor 1895 (11:8, 14:27) (37) or
Terrapene bauri
Florida Box Turtle
Richard D. Bartlett / CRM 3 / Liberty Co., Florida
USA (Alabama, Florida, Georgia, Louisiana, Mississippi,
Texas)
Cistudo major Agassiz 1857a:445, Cistudo carolina major,
Terrapene major, Toxaspis major, Terrapene carolina
major
Type locality: “Mobile..[&]..Florida. Restricted to “Mobile” [Alabama, USA] by Schmidt (1953:94).
Cistudo marnochii † Cope 1878:229 [Pliocene–Pleistocene,
USA (Texas)], Terrapene marnochii
Type locality: “South-western Texas” [USA]. Restricted to
“the Equus beds of Atascosa County, Texas” [USA] by Hay
(1908b:362).
Terrapene putnami † Hay 1906:30 (11:8, 14:28) [Pliocene? to Late
Pleistocene?, USA (Florida)], Terrapene carolina putnami
Type locality: “Alifia River, Florida, about a mile from its mouth...
into Tampa Bay” [USA].
Terrapene canaliculata † Hay 1907:850 [Pliocene–Early
Pleistocene, USA (Georgia)]
Type locality: “Whitemarsh Island or Skedaway Island, Georgia...
southeast of Savannah” [USA].
Terrapene formosa † Hay 1916a:57 [Late Pleistocene, USA
(Florida)]
Type locality: “Ocala, Florida” [USA].
Terrapene antipex † Hay 1916a:58 [Late Pleistocene, USA
(Florida)]
Type locality: “Vero, St. Lucie County, Florida” [USA].
Terrapene carolina mexicana (Gray 1849) (07:25, 14:27) (37) or
Terrapene mexicana mexicana or
Terrapene mexicana
Mexican Box Turtle
Michael T. Jones / CBFTT / peninsular Florida
USA (Florida)
Terrapene bauri Taylor 1895:576, Pariemys bauri, Cistudo
bauri, Terrapene carolina bauri
Type locality: “Florida” [USA]. Restricted to “Orlando, Florida”
[USA] by Schmidt (1953:94).
Terrapene innoxia † Hay 1916a:61 [Pleistocene, USA
(Florida)]
Type locality: “Vero, St. Lucie County, Florida” [USA].
Trachemys nuchocarinata † Hay 1916a:70 (nomen dubium)
[Pleistocene, uSa (Florida)]
Type locality: “Florida Coast Line Canal, 20 miles north of St.
Augustine” [Florida, USA].
Terrapene singletoni † Gilmore 1927:1 [Pleistocene, USA
Collette Adams / CBFTT / Mexico
Mexico (Nuevo León, San Luis Potosi, Tamaulipas, Veracruz)
Cistudo (Onychotria) mexicana Gray 1849:17, Onychotria
mexicana, Cistudo mexicana, Cistudo carolina mexicana,
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Chelopus mexicanus, Terrapene mexicana, Terrapene
mexicana mexicana, Terrapene carolina mexicana
Type locality: “Mexico.” Restricted to “Tampico, Tamaulipas”
[Mexico] by Müller (1936:112).
Terrapene goldmani Stejneger 1933:119
Type locality: “Chijol or Chijoles, southeastern corner of San Louis
Potosi, Mexico.”
Terrapene carolina triunguis (Agassiz 1857a) (11:8, 14:27) (37) or
Terrapene mexicana triunguis or
Terrapene triunguis
Three-toed Box Turtle
Mexico (Campeche, Quintana Roo, Yucatán)
Cistudo yucatana Boulenger 1895b:330, Terrapene yucatana, Terrapene mexicana yucatana, Terrapene carolina
yucatana
Type locality: “Mexico...North Yucatan.” Restricted to “Chichen
Itzá, Yucatán, Mexico” by Smith and Taylor (1950a:351, 1950b:35).
Terrapene coahuila Schmidt and Owens 1944 (14:27)
Coahuilan Box Turtle
Jennifer G. Howeth / CBFTT / TCC / Cuatro Ciénegas, Coahuila, Mexico
Michael T. Jones / CBFTT / Missouri
USA (Alabama, Arkansas, Illinois, Kansas, Louisiana, Mississippi, Missouri, Oklahoma, Texas)
Cistudo triunguis Agassiz 1857a:279,445, Cistudo carolina
triunguis, Terrapene triunguis, Onychotria triunguis, Terrapene carolina triunguis, Terrapene mexicana triunguis
Type locality: “Louisiana...Mississippi...New Orleans...Osage
River...Georgia” [USA]. Restricted to “New Orleans, Louisiana”
[USA] by Schmidt (1953:94).
Terrapene whitneyi † Hay 1916b:8 [Pleistocene, USA
(Texas)]
Type locality: “Austin, Texas” [USA].
Terrapene bulverda † Hay 1920:133 [Pleistocene, USA
(Texas)]
Type locality: “near Bulverde, Bexar County, Texas” [USA].
Terrapene impressa † Hay 1924:245 [Pleistocene, USA
(Texas)]
Type locality: “Munson’s Shoals, 2 or 3 miles below Pittbridge...
Texas” [USA].
Terrapene llanensis † Oelrich 1953:35 [Late Pleistocene,
Sangamonian, USA (Kansas)]
Type locality: “tributary of Shorts Creek...Lone Tree Arroyo, Meade
County, Kansas” [USA].
Mexico (Coahuila)
CBFTT Account: Howeth and Brown (2011)
IUCN Red List: Endangered A2c+4c, B1ab(i,ii,iii,iv,v)+2b(i,ii,
iii,iv,v) (2007); Previously: Endangered (1996)
CITES: Appendix I
Terrapene coahuila Schmidt and Owens 1944:101, Terrapene ornata coahuila, Terrapene coahuilae
Type locality: “Cuatro Cienegas, Coahuila” [Mexico].
Terrapene nelsoni Stejneger 1925 (14:27)
Spotted Box Turtle
Terrapene carolina yucatana (Boulenger 1895b) (07:25, 14:27) (37) or
Terrapene mexicana yucatana or
Terrapene yucatana
Yucatan Box Turtle
(subspecies: nelsoni = red, klauberi = purple)
Michael T. Jones / CBFTT / Yucatán, Mexico
Mexico (Chihuahua, Jalisco, Nayarit, Sinaloa, Sonora)
CBFTT Account: Buskirk and Ponce-Campos (2011)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
IUCN Red List: Data Deficient (1996)
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II, as Terrapene spp.
Terrapene nelsoni nelsoni Stejneger 1925
Southern Spotted Box Turtle
81
Mexico (Chihuahua, Coahuila, Sonora), USA (Arizona, Arkansas, Colorado, Illinois, Indiana, Iowa, Kansas, Louisiana,
Missouri, Nebraska, New Mexico, Oklahoma, South
Dakota, Texas, Wisconsin, Wyoming)
IUCN Red List: Near Threatened (2013); Previously: Near
Threatened (1996)
CITES: Appendix II, as Terrapene spp.
Terrapene ornata ornata (Agassiz 1857a) (12:20, 14:27)
Ornate Box Turtle, Western Box Turtle
John B. Iverson / CBFTT / Nayarit, Mexico
Mexico (Jalisco, Nayarit, Sinaloa)
Terrapene nelsoni Stejneger 1925:463, Terrapene nelsoni
nelsoni
Type locality: “Pedro Pablo, Tepic, Mexico; 2500 feet altitude.”
Jeffrey E. Dawson / Barton Co., Kansas
Terrapene nelsoni klauberi Bogert 1943
Northern Spotted Box Turtle
USA (Arkansas, Colorado, Illinois, Indiana, Iowa, Kansas,
Louisiana, Missouri, Nebraska, New Mexico, Oklahoma,
South Dakota, Texas, Wisconsin, Wyoming)
Cistudo ornata Agassiz 1857a:392,445, Terrapene ornata,
Terrapene ornata ornata, Terrapene carolina ornata
Type locality: “Upper Missouri...and...Iowa” [USA]. Restricted to
“Council Bluffs, Pottawatomie County, Iowa, USA” [in error] by
Smith and Taylor (1950a:358, 1950b:36); and to “junction of the
Platte and Missouri River” [Nebraska, USA] [in error] by Schmidt
(1953:95); and corrected to “Burlington, Des Moines County, Iowa”
[USA] by lectotype designation by Smith and Smith (1980:587).
Terrapene ornata cimarronensis Cragin 1894:37
Type locality: “Red beds country of the Cimarron Basin” [Kansas,
USA].
Matt Cage / CBFTT / nr. Yecora, Sonora, Mexico
Mexico (Chihuahua, Sinaloa, Sonora)
Terrapene klauberi Bogert 1943:2, Terrapene nelsoni
klauberi
Type locality: “Rancho Guirocoba, approximately eighteen miles
southeast of Alamos, Sonora, Mexico.”
Terrapene longinsulae † Hay 1908c:166 (12:20) [Upper Miocene or Lower Pliocene to possibly Pleistocene, USA
(Kansas)], Terrapene ornata longinsulae
Type locality: “Long Island, Phillips County, Kansas” [USA].
Terrapene ornata luteola Smith and Ramsey 1952 (14:27)
Desert Box Turtle
Terrapene ornata (Agassiz 1857a) (12:20, 14:27)
Ornate Box Turtle, Western Box Turtle
Peter Paul van Dijk / Portal, Arizona
Mexico (Chihuahua, Coahuila, Sonora), USA (Arizona, New
Mexico, Texas)
Terrapene ornata luteola Smith and Ramsey 1952:45
Type locality: “17 miles south of Van Horn, Culberson County,
Texas” [USA].
(subspecies: ornata = red, luteola = purple; overlap = intergrades)
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plaTysTeRnidae Gray 1869a (07:26)
Platysternidae Gray 1869a:208
Platysternon megacephalum peguense Gray 1870c (39)
Burmese Big-headed Turtle
Platysternon Gray 1831c
Platysternon Gray 1831c:106
Type species: Platysternon megacephalum Gray 1831c, by original
monotypy.
Platysternum Agassiz 1846:297 (nomen novum)
Platysternon megacephalum Gray 1831c (39)
Big-headed Turtle
Peter Paul van Dijk / CRM 2 / TCF / CCB / Phu Luang Wildlife Sanctuary, Loei Prov., Thailand
Cambodia, China (Hainan), Laos, Myanmar, Thailand,
Vietnam
Platysternon peguense Gray 1870c:70, Platysternon megacephalum peguense
Type locality: “Pegu” [Myanmar].
Platysternon megacephalum vogeli Wermuth 1969:374
Type locality: “Provinz Chiang Mai, Nordwest-Thailand”
Platysternon megacephalum tristernalis Schleich and Gruber
1984:68 (39)
Type locality: “zwischen Mung Lun und Simao, Ostufer des Mekongflusses, südliches Yünnan (VR China).” [Menglun, Lancang
Jiang, Yunnan].
(subspecies: megacephalum = red, peguense = purple, shiui = blue;
overlap = intergrades; orange dots = probable trade or introduced)
Cambodia, China (Anhui, Fujian, Guangdong, Guangxi,
Hainan, Hong Kong, Hunan, Jiangxi, Yunnan, Zhejian),
Laos, Myanmar, Thailand, Vietnam
IUCN Red List: Endangered A1d+2d (2000); Previously: Data
Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix I, as Platysternidae spp.
Platysternon megacephalum shiui Ernst and McCord 1987 (39)
Vietnamese Big-headed Turtle
Platysternon megacephalum megacephalum Gray 1831c (07:27) (39)
Chinese Big-headed Turtle
Torsten Blanck / Vietnam [captivity]
China (?) (Guangxi, Hainan), Vietnam
Platysternon megacephalum shiui Ernst and McCord
1987:626
Type locality: “vicinity of Langson, Langson Province, Vietnam
(26°50' N, 106°45' E).” GPS coordinates in error, corrected here to
21°50' N, 106°45' E.
Yik-Hei Sung / Hong Kong, China
China (Anhui, Fujian, Guangdong, Guangxi, Hong Kong,
Hunan, Jiangxi, Zhejian), Vietnam
Platysternon megacephalum Gray 1831c:107, Emys
megacephala, Platysternon megacephalus, Platysternon
megacephalum megacephalum
Type locality: “Chinâ.” Restricted to “S. China” by Boulenger
(1889:48).
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
geoeMydidae Theobald 1868a (07:29, 09:20, 12:21)
Geoemydidae Theobald 1868a:vi
Batagurina Gray 1869a:185
Bataguridae Gray 1870c:17
Batagur affinis (Cantor 1847) (08:9)
Southern River Terrapin
geoeMydinae Theobald 1868a (12:21)
Geoemydidae Theobald 1868a:vi
Batagurina Gray 1869a:185
Bataguridae Gray 1870c:17
Batagur Gray 1856b (07:30, 08:9)
Trionyx (Tetraonyx) Gray 1830e:19 (10:7) (junior homonym, not =
Tetraonyx Latreille 1809 [= Coleoptera])
Type species: Trionyx (Tetraonyx) cuvieri Gray 1830e [= subjective
synonym of Emys baska Gray 1830d], by original monotypy.
Tetronyx Lesson 1832:pl.7 (nomen novum et oblitum)
Type species: Tetronyx longicollis [= Tetraonyx longicollis Lesson
1831b = subjective synonym of Emys baska Gray 1830d], by
original monotypy.
Batagur Gray 1856b:35
Type species: Batagur baska [= Emys baska Gray 1830d], by subsequent designation by Smith (1931:134).
Batagur (Kachuga) Gray 1856b:35
Type species: Batagur (Kachuga) lineata [= Emys kachuga Gray
1831a], by tautonymy. Not Kachuga trilineata Gray 1869a [=
subjective synonym of Emys trivittata Duméril and Bibron 1835],
by subsequent erroneous designation by Smith (1931:124).
Kachuga (Batagurella) Gray 1869a:200
Type species: Kachuga (Batagurella) peguensis Gray 1869a [= subjective synonym of Emys trivittata Duméril and Bibron 1835], by
original monotypy.
Dongoka Gray 1869a:202
Type species: Dongoka hardwickii [= Kachuga (Dongoka) hardwickii
Gray 1869a = subjective synonym of Emys dhongoka Gray 1832b],
by subsequent designation by Lindholm (1929:278).
(subspecies: affinis = red, edwardmolli = purple)
Cambodia, Indonesia (Sumatra), Malaysia (Peninsular),
Singapore (extirpated, reintroduced), Thailand, Vietnam
(extirpated)
CBFTT Account: Moll, Platt, Chan, Horne, Platt, Praschag,
Chen, and van Dijk (2015)
IUCN Red List: Critically Endangered A2bcd+4bcd (2016);
Previously: Critically Endangered, as part of Batagur
baska (2000).
CITES: Appendix I
Batagur affinis affinis (Cantor 1847) (08:9, 09:21)
Western Malay River Terrapin
Dhongoka Gray 1870c:57 (nomen novum)
Type species: Dhongoka hardwickii [= Kachuga (Dongoka) hardwickii
Gray 1869a = subjective synonym of Emys dhongoka Gray 1832b],
by subsequent monotypy.
Callagur Gray 1870c:53
Type species: Callagur picta [= Batagur picta Gray 1862b = subjective
synonym of Emys borneoensis Schlegel and Müller 1845], by
original monotypy.
Cantorella Gray 1870c:58
Type species: Cantorella affinis [= Tetraonyx affinis Cantor 1847], by
original monotypy.
Dhougoka Gray 1873j:52 (nomen novum)
Cachuga Lydekker 1889:123 (nomen novum)
Edward O. Moll / CBFTT / Perak R., Malaysia / female
Indonesia (Sumatra), Malaysia (Peninsular), Singapore (extirpated, reintroduced), Thailand
Tetraonyx affinis Cantor 1847:6, Batagur affinis, Kachuga
affinis, Kachuga (Dongoka) affinis, Cantorella affinis,
Batagur affinis affinis
Type locality: “sea off Pinang…along the sea-shore of Pinang..[&]..
estuaries and rivers on the Peninsula” [Malaysia].
Batagur siebenrocki † Jaekel 1911:76 [Pleistocene, Pithecanthropus Trinil Beds, Indonesia (Java)]
Type locality: “Pithecanthropus-schichten...Java...Trinil”
[Indonesia].
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Batagur affinis edwardmolli Praschag, Holloway, Georges, Päckert, Hundsdörfer, and Fritz 2009 (09:21)
Eastern Malay River Terrapin
baska, Tetronyx baska, Batagur (Batagur) baska, Batagur
baska, Tetraonyx baska, Batagur baska baska
Type locality: “India.”
Emys batagur Gray 1830e:9 (10:7), Clemmys (Clemmys)
batagur, Tetraonyx batagur, Batagur batagur, Batagur
batagur batagur
Type locality: Not designated. Restricted to “India” by Gray
(1831d:24).
Trionyx (Tetraonyx) cuvieri Gray 1830e:19 (10:7), Trionyx
cuvierii
Type locality: Not designated. Restricted to “l’Irrawady, fleuve
du royaume de Pégu” [Myanmar] by Temminck and Schlegel
(1834:43).
Tetraonyx longicollis Lesson 1831b:297, Clemmys longicollis, Tetronyx longicollis
Type locality: “le fleuve de l’Irravaddy au Pégou” [Myanmar].
Eng Heng Chan / CBFTT / TCC / Setiu R., Terengganu, Malaysia / male in breeding color
Cambodia, Malaysia (Peninsular), Thailand (extirpated),
Vietnam (extirpated)
Batagur affinis edwardmolli Praschag, Holloway, Georges, Päckert, Hundsdörfer, and Fritz 2009a:64 (09:21)
Type locality: “Sre Ambel River system, Koh Kong Province,
Cambodia.”
Batagur baska (Gray 1830d) (07:31, 08:9)
Northern River Terrapin
Peter Praschag / Satkhira Distr., Sundarbans Bangladesh / male in breeding color
Bangladesh, India (Odisha, West Bengal), Myanmar, Thailand
(extirpated?)
CBFTT Account: Moll, Platt, Platt, Praschag, and van Dijk
(2009)
IUCN Red List: Critically Endangered A1cd (2000); Previously:
Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix I
Emys baska Gray 1830d:pl.75, Testudo baska, Emys batagur
Emys tetraonyx Temminck and Schlegel 1834:43 (10:18) (nomen
novum)
Tetraonyx lessonii Duméril and Bibron 1835:338 (nomen
novum), Hydraspis (Tetronyx) lessonii
Batagur baska ranongensis Nutaphand 1979:181 (07:31),
Batagur ranongensis, Batagur batagur ranongensis
Type locality: “mouth of rivers in Ranong Province” [Thailand].
Batagur borneoensis (Schlegel and Müller 1845) (07:30)
Painted Terrapin
Douglas B. Hendrie / TCC / Perak, Malaysia / male in breeding color
Brunei, Indonesia (Kalimantan, Sumatra), Malaysia (Peninsular,
East), Thailand
IUCN Red List: Critically Endangered A1bcd (2000); Previously: Critically Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Batagur spp.
Emys borneoensis Schlegel and Müller 1845:30, Clemmys
borneoensis, Callagur borneoensis, Batagur borneoensis
Type locality: “Borneo” [East Malaysia or Kalimantan, Indonesia].
Batagur picta Gray 1862b:204, Callagur picta, Tetraonyx
pictus, Callagur pictus
Type locality: “Borneo, Sarawak” [East Malaysia].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Clemmys grayi Strauch 1865:88 (nomen novum)
Kachuga major Gray 1873c:300
Type locality: “India?” Emended to “India” by Gray (1873j:51).
Batagur kachuga (Gray 1831a) (07:30)
Red-crowned Roofed Turtle
Kachuga brookei Bartlett 1895a:29
Type locality: “Borneo” [East Malaysia or Kalimantan, Indonesia].
Batagur dhongoka (Gray 1832b) (07:30)
Three-striped Roofed Turtle
Saurav Gawan / Chambal R., Uttar Pradesh, India / male in breeding color
Indraneil Das / Bihar, India
(orange dot = trade)
Bangladesh, India (Assam, Bihar, Madhya Pradesh, Rajasthan,
Uttar Pradesh, West Bengal), Nepal
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Near Threatened (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Batagur spp.
Emys dhongoka Gray 1832b:pl.60, Batagur (Kachuga) dhongoka, Batagur dhongoka, Clemmys dhongoka, Kachuga
dhongoka
Type locality: “India.” Restricted to “N. India” by Smith
(1931:130).
Emys duvaucelii Duméril and Bibron 1835:334, Batagur
duvaucelii
Type locality: “Bengale” [Bangladesh or India].
Kachuga (Dongoka) hardwickii Gray 1869a:202, Kachuga
hardwickii, Dhongoka hardwickii, Dhougoka hardwickii
Type locality: “Nepal.”
Batagur durandi † Lydekker 1885:192 [Late Pliocene (Pinjor) to Early Pleistocene (Tatrot), Siwaliks, India (Punjab)]
Type locality: “Siwalik Hills” [Punjab, India].
Bangladesh, India (Bihar, Madhya Pradesh, Punjab, Uttar
Pradesh, West Bengal), Nepal
IUCN Red List: Critically Endangered A1cd (2000); Previously:
Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Batagur spp.
Emys lineata Gray 1830e:9 (10:7) (nomen oblitum), Clemmys
(Clemmys) lineata, Batagur (Kachuga) lineata, Batagur
lineatus, Kachuga lineata
Type locality: “India.”
Emys kachuga Gray 1831a:pl.74, Batagur kachuga, Kachuga
kachuga
Type locality: “India.” Restricted to “N. India” by Smith
(1931:131).
Batagur ellioti Gray 1862b:264, Batagur elliotti, Clemmys
ellioti
Type locality: “Southern India, River Kistna” [Krishna River,
Andhra Pradesh, India] [in error].
Kachuga fusca Gray 1870c:56 (partim)
Type locality: “India.”
Batagur bakeri † Lydekker 1885:190 [Late Pliocene (Pinjor)
to Early Pleistocene (Tatrot), Siwaliks, India (Punjab)]
Type locality: “Siwalik Hills” [Punjab, India].
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Batagur trivittata (Duméril and Bibron 1835) (07:30)
Burmese Roofed Turtle
trifasciatus Bell 1825a], by original monotypy.
Cuora amboinensis (Riche in Daudin 1801) (12:23) (41)
Southeast Asian Box Turtle
Rick Hudson / CRM 5 / TCC / Myanmar [Yadanabon Zoo, Mandalay] / male in breeding color
(subspecies: amboinensis = red, couro = purple, kamaroma = orange,
lineata = pink, overlap = intergrades (Palawan);
orange dot = introduced or trade)
Myanmar
IUCN Red List: Endangered A1c (2000); Previously: Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Batagur spp.
Emys trivittata Duméril and Bibron 1835:331, Batagur trivittata, Kachuga trivittata
Type locality: “Bengale.” [India or Bangladesh]
Kachuga (Batagurella) peguensis Gray 1869a:200, Kachuga
peguensis
Bangladesh, Bhutan, Brunei, Cambodia, India (Arunachal
Pradesh, Assam, Nagaland, Nicobar Islands), Indonesia
(Java, Kalimantan, Lesser Sundas, Moluccas, Sulawesi,
Sumatra, Timor), Laos, Malaysia (Peninsular, East),
Myanmar, Philippines (Bohol, Cebu, Leyte, Luzon,
Mindanao, Mindoro, Negros, Palawan, Panay, Samar, Sulu
Archipelago), Thailand, Timor-Leste (?), Vietnam
CBFTT Account: Schoppe and Das (2011)
IUCN Red List: Vulnerable A1d+2d (2000); Previously: Near
Threatened (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II, as Cuora spp.
Cuora amboinensis amboinensis (Riche in Daudin 1801) (12:23)
East Indian Box Turtle
Type locality: “India.” Restricted to “Pegu” [Myanmar] by Gray
(1873j:50).
Kachuga trilineata Gray 1869a:200, Kachuga (Kachuga)
trilineata
Type locality: “India.” Restricted to “Pegu” [Myanmar] by Boulenger (1889:56).
Kachuga fusca Gray 1870c:56 (partim)
Type locality: “India.”
Batagur iravadica Anderson 1879:736, Batagur iravadicus,
Clemmys iravadica
Type locality: “Pegu..[&]..Bhamô in Upper Burma..[&].. throughout the Irawady” [Myanmar].
Cuora Gray 1856a (07:32, 12:22) (40)
Cuora Gray 1856a:198
Type species: Cuora amboinensis [= Testudo amboinensis Riche in
Daudin 1801], by subsequent designation by Stejneger (1907:503).
Cistoclemmys Gray 1863e:175
Type species: Cistoclemmys flavomarginata Gray 1863e, by original
monotypy.
Pyxidea Gray 1863e:175
Type species: Pyxidea mouhotii [= Cyclemys mouhotii Gray 1862a], by
original monotypy.
Cuora (Pyxiclemmys) Gray 1863e:176
Type species: Cuora (Pyxiclemmys) trifasciata [= Sternothaerus
Sabine Schoppe / CBFTT / Sulawesi, Indonesia
Indonesia (Moluccas, Sulawesi), Philippines (Bohol, Cebu,
Leyte, Luzon, Mindanao, Mindoro, Negros, Palawan,
Panay, Samar)
Testudo melanocephala Van-Ernest in Daudin 1801:128 (nomen oblitum), Emys melanocephala, Clemmys (Clemmys)
melanocephala
Type locality: “une des îles Moluques ou Philippines” [Moluccas,
Indonesia, or Philippines]. Restricted to “Moluccas” [Indonesia] by
Fritz and Havaš (2007:214).
Testudo amboinensis Riche in Daudin 1801:309, Emys
amboinensis, Terrapene amboinensis, Kinosternon amboinense, Cistuda amboinensis, Cuora amboinensis, Cistudo
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
amboinensis, Cyclemys amboinensis, Cuora amboinensis
amboinensis
Type locality: “Amboine” [Ambon, Moluccas, Indonesia].
Cuora amboinensis lineata McCord and Philippen 1998
Burmese Box Turtle
Emys melanogaster Bleeker in Gray 1864a:12 (nomen
nudum)
Type locality: “Batchian and Boero” [Bacan and Buru, Moluccas,
Indonesia]. Restricted to “Borneo” [in error] by Gray (1873j:21).
Emys hypselonotus Bleeker in Gray 1864a:12 (nomen nudum), Emys hypsilonotus
Type locality: “Batchian and Boero” [Bacan and Buru, Moluccas,
Indonesia]. Restricted to “Borneo” [in error] by Gray (1873j:22).
Emys gastrotaenia Bleeker in Gray 1873j:21 (nomen nudum)
Type locality: “Borneo.” Locality likely in error, probably “Batchian
and Boero” [Bacan and Buru, Moluccas, Indonesia].
Cuora amboinensis couro (Lechenault in Schweigger 1812)
Indonesian Box Turtle
Hans-Dieter Philippen / CBFTT / Myanmar
Myanmar
Cuora amboinensis lineata McCord and Philippen 1998:54
Type locality: “Myitkyina, Kachin Province, Myanmar (Burma).”
Cuora aurocapitata Luo and Zong 1988 (12:22) (42)
Yellow-headed Box Turtle
Torsten Blanck / nr. Jakarta, Java, Indonesia
Indonesia (Java, Lesser Sundas, Sumatra, Timor), Timor-Leste (?)
Emys couro Lechenault in Schweigger 1812:315, Terrapene
couro, Cuora amboinensis couro
Type locality: “Java” [Indonesia].
Terrapene bicolor Bell 1826:485
Type locality: “Americâ septentrionali” [in error].
Emys (Cistuda) amboinensis leveriana Gray 1830e:7 (10:7),
Cistuda amboinensis leveriana
Type locality: “Java and Penang” [Indonesia and Peninsular
Malaysia].
Cuora amboinensis kamaroma Rummler and Fritz 1991 (12:23)
Malayan Box Turtle
(subspecies: aurocapitata = red, dabieshani = purple;
orange dots = probable trade)
China (Anhui, Henan, Hubei, Zhejiang)
IUCN Red List: Critically Endangered A1d+2d (2000); Previously: Data Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Cuora spp.
Cuora aurocapitata aurocapitata Luo and Zong 1988 (12:22) (42)
Eastern Yellow-headed Box Turtle
Indraneil Das / Malaysia (East) [Borneo]
Bangladesh, Bhutan, Brunei, Cambodia, India (Arunachal
Pradesh, Assam, Nagaland, Nicobar Islands), Indonesia
(Kalimantan), Malaysia (Peninsular, East), Laos, Myanmar
(?), Philippines (Palawan [?], Sulu Archipelago [?]), Thailand, Vietnam
Cuora amboinensis kamaroma Rummler and Fritz 1991:39
Type locality: “circa 50 km nördlich von Bangkok, Thailand.”
Torsten Blanck / Jing County, Anhui, China
China (Anhui, Zhejiang)
Cuora aurocapitata Luo and Zong 1988:13, Cuora pani
aurocapitata, Pyxiclemmys aurocapitata, Pyxiclemmys
pani aurocapitata, Cuora aurocapitata aurocapitata
Type locality: “Nanling County, Anhui” [China].
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Cuora aurocapitata dabieshani Blanck, Protiva, Zhou, Li, Crow,
and Tiedemann 2017 (42)
Western Yellow-headed Box Turtle
Cuora cyclornata Blanck, McCord, and Le 2006a:133 (07:36, 09:23, 12:22)
(40, 42)
Vietnamese Three-striped Box Turtle
Torsten Blanck / Anhui, China [captivity]
China (Anhui, Henan, Hubei)
Cuora aurocapitata dabieshani Blanck, Protiva, Zhou, Li,
Crow, and Tiedemann 2017:16
Type locality: “Anhui province, China.”
Cuora bourreti Obst and Reimann 1994 (07:35, 09:22, 12:22)
Bourret’s Box Turtle
(subspecies: cyclornata = red, annamitica = purple, meieri = orange;
orange dot = trade)
China (Guangxi), Laos, Vietnam
IUCN Red List: Critically Endangered A1d+2d (2000) (as part
of Cuora trifasciata); Previously: Endangered (1996) (as
part of C. trifasciata)
TFTSG Draft Red List: Critically Endangered (2016)
CITES: Appendix II, as Cuora spp.
Cuora cyclornata cyclornata Blanck, McCord, and Le 2006a:133
(07:36, 09:23, 12:22) (40, 42)
Southern Vietnamese Three-striped Box Turtle
Torsten Blanck / TCC / Vietnam
Torsten Blanck / Vietnam [captivity]
Laos, Vietnam
Cuora cyclornata cyclornata Blanck, McCord, and Le
2006a:133 (07:36, 09:23, 12:22)
Type locality: “Phong Nha Ke Bang Nat. Res., Quang Binh Province, central Vietnam.”
Laos, Vietnam
IUCN Red List: Critically Endangered A2bd+4bd (2016);
Previously: Critically Endangered, as part of Cuora galbinifrons (2000)
CITES: Appendix II, as Cuora spp.
Cuora galbinifrons serrata Iverson and McCord 1992b:434
(07:33)
(partim, hybrid)
Cuora galbinifrons bourreti Obst and Reimann 1994:135, Cistoclemmys galbinifrons bourreti, Cuora bourreti, Cistoclemmys
bourreti
Cuora cyclornata annamitica Blanck, Protiva, Zhou, Li, Crow,
and Tiedemann 2017 (42)
Central Vietnamese Three-striped Box Turtle
Type locality: “Linh-Cam (heute Ha-Tinh), in Mittel-Vietnam”.
Emended to “Bach Ma, Thua Thien Hue Province, Central Vietnam” by Fritz et al. (2002:71).
Torsten Blanck / Vietnam [captivity]
Laos (?), Vietnam
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Cuora cyclornata annamitica Blanck, Protiva, Zhou, Li,
Crow, and Tiedemann 2017:12
Type locality: “Vietnam, Nghe An Province, Tan Ky district, near
Ky Son Village.”
Cuora flavomarginata flavomarginata (Gray 1863e)
Yellow-margined Box Turtle
Cuora cyclornata meieri Blanck, McCord, and Le 2006a:142 (07:36,
09:23, 12:22) (40, 42)
Northern Vietnamese Three-striped Box Turtle
Tien-Hsi Chen / CBFTT / Nantou, Taiwan
Torsten Blanck / Vietnam [captivity]
China (Guangxi), Vietnam
Cuora cyclornata meieri Blanck, McCord, and Le
2006a:142 (07:36, 09:23, 12:22)
Type locality: “Tam Dao, Provinz Vinh Phuc, Nord-Vietnam.”
Cuora flavomarginata (Gray 1863e) (08:21, 11:9, 12:22) (43)
Yellow-margined Box Turtle
China (Anhui, Fujian, Henan, Hubei, Hunan, Jiangsu, Jiangxi,
Zhejiang), Taiwan
Cistoclemmys flavomarginata Gray 1863e:175, Cuora
flavomarginata, Terrapene flavomarginata, Cyclemys
flavomarginata, Cyclemys flavomarginata flavomarginata,
Cuora flavomarginata flavomarginata, Geoemyda flavomarginata, Cistoclemmys flavomarginatus, Cistoclemmys
flavomarginata flavomarginata
Type locality: “China; Formosa...district of Tamsuy, N.W. Formosa”
[Taiwan]. Restricted to “Tamsui, Formosa” [Taiwan] by Stejneger
(1907:503).
Cyclemys flavomarginata sinensis Hsü 1930:3 (07:34, 08:21), Cuora flavomarginata sinensis, Cistoclemmys flavomarginata
sinensis
Type locality: “Künshan Island, Tungting Lake, Central China.”
[Junshan Dao, Dongting Lake, Hunan, China].
Terrapene culturalia † Yeh 1961:59 [Holocene, Neolithic,
subfossil, Lung-shan Period (Ying Dynasty), China (Shandong)], Emydoidea culturalia
Type locality: “Dawenkou, Taian, Shantung” [Shandong, China].
Cuora flavomarginata evelynae Ernst and Lovich 1990 (08:21, 11:9)
Ryukyu Yellow-margined Box Turtle
Iriomote Wildlife Conservation Center / CBFTT / Iriomotejima, Japan
Japan (Ryukyu Archipelago)
Cuora evelynae Ernst and Lovich 1990:31, Cuora flavomarginata evelynae, Cistoclemmys flavomarginata evelynae
Type locality: “Ishigaki Shima, Ryukyu Islands, Japan.”
(subspecies: flavomarginata = red, evelynae = purple;
orange dots = introduced or trade)
China (Anhui, Fujian, Henan, Hubei, Hunan, Jiangsu, Jiangxi,
Sichuan, Zhejiang), Japan (Ryukyu Archipelago), Taiwan
CBFTT Account: Ota, Yasukawa, Fu, and Chen (2009)
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Cuora spp.
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Cuora galbinifrons Bourret 1940 (07:35, 09:22, 12:22, 12:24)
Indochinese Box Turtle
Douglas B. Hendrie / TCF / TCC / Vietnam
Cuora mccordi Ernst 1988 (12:22)
McCord’s Box Turtle
Torsten Blanck / TCC / Guangxi, China [captivity]
(red = possible range; orange dots = trade)
China (Guangxi, Hainan), Laos, Vietnam
IUCN Red List: Critically Endangered A2bd+4bd (2016); Previously: Critically Endangered (2000), Near Threatened
(1996)
CITES: Appendix II, as Cuora spp.
Cuora galbinifrons Bourret 1940:11, Cistoclemmys galbinifrons, Cuora galbinifrons galbinifrons, Cistoclemmys
galbinifrons galbinifrons
Type locality: “Tam-Dao..[&]..Bach-Ma (Annam)..[&]..Linh-Cam
(Ha-Tinh, Nord Annam)” [Tonkin..[&]..Annam..[&]..Ha-Tinh]
[Vietnam]. Restricted to “Tam-Dao, Nord-Vietnam” by Obst and
Reimann (1994:136).
Cyclemus flavomarginata hainanensis Li 1958:234, Cuora
hainanensis, Cyclemys flavomarginata hainanensis, Cistoclemmys hainanensis, Cuora flavomarginata hainanensis,
Cistoclemmys flavomarginata hainanensis, Cuora galbinifrons hainanensis, Cyclemys flavomarginatus hainanensis, Cistoclemmys galbinifrons hainanensis
China (Guangxi)
IUCN Red List: Critically Endangered A1d+2d (2000); Previously: Data Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Cuora spp.
Cuora mccordi Ernst 1988:466, Cistoclemmys mccordi
Type locality: “highland near Paise, Guangxi Province, China
(23º54'N, 106º37'E)” [in error]. Restricted to “Yunnan Province,
west of Paise, Guangxi Province, China” by McCord and Iverson
(1991:414).
Cuora mouhotii (Gray 1862a) (07:32, 12:22)
Keeled Box Turtle
Type locality: In Chinese [“Dali village, Mt. Diaoluo, Linshui
County, Hainan Island, China”].
Cuora galbinifrons serrata Iverson and McCord 1992b:434
(07:33)
(partim, hybrid)
Type locality: “100 km east of Tungfang at Tainhfien in central
Hainan Island, China.”
(subspecies: mouhotii = red, obsti = purple;
overlap = intergrades; orange dots = trade)
Bangladesh, Bhutan, China (Guangxi, Hainan, Yunnan),
India (Arunachal Pradesh, Assam, Manipur, Meghalaya,
Mizoram, Nagaland), Laos, Myanmar, Thailand (?),
Vietnam
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
CBFTT Account: Das, McCormack, van Dijk, Hoang, and
Struijk (2016)
IUCN Red List: Endangered A1d+2d (2000); Previously: Least
Concern [Not Listed] (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Cuora spp.
91
Cuora pani Song 1984 (12:22)
Pan’s Box Turtle
Cuora mouhotii mouhotii (Gray 1862a)
Northern Keeled Box Turtle
Torsten Blanck / TCC / China [captivity]
Indraneil Das / CBFTT / China [captivity]
Bangladesh, Bhutan, China (Guangxi, Hainan, Yunnan),
India (Arunachal Pradesh, Assam, Manipur, Meghalaya,
Mizoram, Nagaland), Laos, Myanmar, Thailand (?),
Vietnam
Cyclemys mouhotii Gray 1862a:157, Pyxidea mouhotii,
Cyclemys mohouti, Pyxidea mouboti, Emys mouhotii,
Geoemyda mouhotii, Pyxidea mouhotii mouhotii, Cuora
mouhotii, Cuora mouhotii mouhotii
Type locality: “Lao Mountains, in Siam.” Restricted to “Luang
Prabang, Laos (19º54' N, 102º8' O)” by lectotype designation by
Fritz et al. (1998:40).
Cuora galbinifrons serrata Iverson and McCord 1992b:434
(07:33)
(partim, hybrid)
Type locality: “100 km east of Tungfang at Tainhfien in central
Hainan Island, China.”
Cuora mouhotii obsti (Fritz, Andreas, and Lehr 1998)
Southern Keeled Box Turtle
(orange dots = trade)
China (Henan, Hubei, Shaanxi, Sichuan)
IUCN Red List: Critically Endangered A1d+2d (2000); Previously: Data Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Cuora spp.
Cuora pani Song 1984:330, Cuora pani pani, Pyxiclemmys
pani pani
Type locality: “Xujiaba (alt. 420 m) of Pingli County in Shaanxi
Province” [China].
Cuora chriskarannarum Ernst and McCord 1987:624
Type locality: “Ta Lau Shan, Yunnan Province, China (23º30' N,
102º25' E).”
Cuora picturata Lehr, Fritz, and Obst 1998 (07:35, 09:22, 12:22)
Southern Vietnam Box Turtle
Torsten Blanck / Phu Yen Prov., Vietnam / male
Laos, Vietnam
Pyxidea mouhotii obsti Fritz, Andreas, and Lehr 1998a:35,
Cuora mouhotii obsti
Type locality: “Umgebung von Phú Lôc, Annam (Vietnam), 16º16'
N, 107º56' O).”
Torsten Blanck / TCC / Vietnam
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China (Fujian, Guangdong, Guangxi, Hong Kong)
Sternothaerus trifasciatus Bell 1825a:305, Emys (Cistuda)
trifasciata, Emys trifasciata, Cistuda trifasciata, Cistudo
trifasciata, Cuora trifasciata, Pyxidemys trifasciata, Terrapene trifasciata, Cyclemys trifasciata, Cuora (Pyxiclemmys) trifasciata, Pyxiclemmys trifasciata, Cuora trifasciata
trifasciata
Type locality: Not known. Restricted to “Luofo Shan Mountains,
Guangdong, China” by Blanck et al. (2006:40).
Mauremys iversoni Pritchard and McCord 1991:140 (07:33) (partim, hybrid)
Type locality: “People’s Republic of China: Fujian Province: vicinity
of Nanping (26º38' N, 118º10' E).”
Clemmys guangxiensis Qin 1992:60 (14:29) (partim, hybrid)
Vietnam
IUCN Red List: Critically Endangered A2bd+4bd (2016);
Previously: Critically Endangered, as part of Cuora galbinifrons (2000)
CITES: Appendix II, as Cuora spp.
Cuora galbinifrons picturata Lehr, Fritz, and Obst 1998:7,
Cistoclemmys galbinifrons picturata, Cuora picturata,
Cistoclemmys picturata
Type locality: “Nanning, Guangxi” [China].
Cuora trifasciata luteocephala Blanck, Protiva, Zhou, Li, Crow,
and Tiedemann 2017 (42)
Hainan Three-striped Box Turtle
Type locality: “südliches Annam (Vietnam).”
Cuora trifasciata (Bell 1825a) (07:36, 09:23, 12:22) (40, 42)
Chinese Three-striped Box Turtle, Golden Coin Turtle
Torsten Blanck / Hainan, China [captivity]
China (Hainan)
Sacalia pseudocellata Iverson and McCord 1992a:426 (07:33)
(partim, hybrid)
Type locality: “between Tungfang [19º03' N, 108º56' E] and
Kancheng [18º51' N, 108º37' E; ca. 48 km from Tungfang], western
Hainan Island, China.”
Ocadia philippeni McCord and Iverson 1992:13 (07:33) (partim,
hybrid)
(subspecies: trifasciata = red, luteocephala = purple; orange dots = trade)
China (Fujian, Guangdong, Guangxi, Hainan, Hong Kong)
IUCN Red List: Critically Endangered A1d+2d (2000); Previously: Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Cuora spp.
Cuora trifasciata trifasciata (Bell 1825a) (07:36, 09:23, 12:22) (40, 42)
Chinese Three-striped Box Turtle
Type locality: “near Kancheng [18º51' N, 108º37' E; = 48 km from
Tungfang (19º03' N, 108º56' E)], western Hainan Island, China.”
Cuora trifasciata luteocephala Blanck, Protiva, Zhou, Li,
Crow, and Tiedemann 2017:14
Type locality: “China, Hainan Province, Dan County, 300m
elevation.”
Cuora yunnanensis (Boulenger 1906) (07:37, 12:22)
Yunnan Box Turtle
Zhou Ting, William P. McCord, Torsten Blanck / TCC / Yunnan, China [captivity]
Paul Crow / TCF / TCC / Hong Kong, China
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Cyclemys Bell 1834 (07:38, 08:6, 09:24)
Cyclemys Bell 1834:17
Type species: Cyclemys orbiculata Bell 1834 [= subjective synonym of
Emys dentata Gray 1831d], by original designation.
Cyclemis Tirant 1884:156 (nomen novum)
Cyclemys atripons Iverson and McCord 1997 (44)
Western Black-bridged Leaf Turtle
(red = possible range; dots = possible native or trade)
China (Sichuan, Yunnan)
IUCN Red List: Critically Endangered B2ab(ii,iii,v), D (2010);
Previously: Extinct (2000), Data Deficient (1996)
CITES: Appendix II, as Cuora spp.
Cyclemys yunnanensis Boulenger 1906a:567, Cuora yunnanensis, Pyxiclemmys yunnanensis
Type locality: “Yunnan fu..[&].. Tongchuan fu” [Kunming Shi..
[&].. Dongchuan Shi, Yunnan, China]; restricted to “vicinity of
Zhongping (Huize) City (26°42'N, 103°30'E) in Huize County,
northeastern Yunnan Province, China (= Tongchuan Fu)” by Blanck
et al. (2006b:31).
Flora Ihlow / Trat, Cambodia
Cuora zhoui Zhao in Zhao, Zhou, and Ye 1990 (12:22)
Zhou’s Box Turtle
(orange dots = uncertain identification or possible hybrids)
Torsten Blanck / TCC / Guangxi, China? [captivity]
Cambodia, Thailand
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Near Threatened (2011)
CITES: Appendix II, as Cyclemys spp.
Cyclemys atripons Iverson and McCord 1997:632, Cyclemys
atripons atripons, Cyclemys pulchristriata atripons
Type locality: “Thailand, Krat [= Trat], Kao [= Mt.] Kuap (= Khao
Kuap).”
Cyclemys dentata (Gray 1831d) (08:7)
Asian Leaf Turtle
(red = possible range; yellow dot = possible native; orange dots = trade)
China (Guangxi?, Yunnan?), Vietnam
IUCN Red List: Critically Endangered A1d+2d (2000); Previously: Data Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Cuora spp.
Cuora zhoui Zhao in Zhao, Zhou, and Ye 1990:213, Pyxiclemmys zhoui
Type locality: “market at Nanning, Guangxi Zhuang Autonomous
Region” [China].
Cuora pallidicephala McCord and Iverson 1991:414
Type locality: “Wuting [= Wuding: 25º26'N, 102º21'W] or Yuanmow [25º41'N, 101º54'W], Yunnan Province, China.”
Maren Gaulke / Kota Pinang, Sumatra, Indonesia
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Brunei, Indonesia (Bali, Java, Kalimantan, Sumatra), Malaysia
(Peninsular, East), Philippines (Palawan, Sulu Archipelago), Singapore, Thailand
IUCN Red List: Near Threatened (2000); Previously: Least
Concern [Not Listed] (1996)
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II, as Cyclemys spp.
Emys hasseltii Boie in Fitzinger 1826:45 (nomen nudum),
Clemmys (Clemmys) hasseltii
Type locality: “Asia, Insula Java” [Indonesia].
Emys dhor Gray 1830e:8
dhor
(10:7)
(nomen oblitum), Cyclemys
Malaysia (Peninsular, East), Singapore, Indonesia (Java, Kalimantan, Sumatra), Thailand
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II, as Cyclemys spp.
Cyclemys enigmatica Fritz, Guicking, Auer, Sommer, Wink,
and Hundsdörfer 2008b:381
Type locality: “Padang, Sumatra” [Indonesia].
Cyclemys fusca Fritz, Guicking, Auer, Sommer, Wink, and
Hundsdörfer 2008 (08:6)
Myanmar Brown Leaf Turtle
Type locality: “India.” Restricted to “Bengal..[and]..Java” by Gray
(1831d:20); and to “Java” [Indonesia] by lectotype designation by
Fritz et al. (1997:188).
Emys dentata Gray 1831d:errata[btw 78-79] (nomen novum),
Cistudo (Cyclemys) dentata, Cistudo dentata, Cyclemys
dentata, Cyclemys dentata dentata
Type locality: “Bengal..[and]..Java.” Restricted to “Java” [Indonesia]
by Smith (1931:80), and by lectotype designation by Fritz et al.
(1997:188).
Cyclemys orbiculata Bell 1834:17, Emys orbiculata, Emys
(Cyclemys) orbiculata, Cistudo orbiculata
Type locality: “Indiâ.” Restricted to “Java” [Indonesia] by neotype
designation by Fritz et al. (1997:188).
Cistudo diardii Duméril and Bibron 1835:227 (nomen novum), Emys diardii
Peter Paul van Dijk / Myanmar [trade]
Type locality: “Bengale et l’ile de Java.”
Cyclemys ovata Gray 1863e:178
Type locality: “Sarawak” [East Malaysia].
Cyclemys bellii Gray 1863e:179
Type locality: Not known.
Cyclemys enigmatica Fritz, Guicking, Auer, Sommer, Wink, and
Hundsdörfer 2008 (08:6)
Enigmatic Leaf Turtle
India (Nagaland, Manipur), Myanmar
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II, as Cyclemys spp.
Cyclemys fusca Fritz, Guicking, Auer, Sommer, Wink, and
Hundsdörfer 2008b:383
Type locality: “Kachin State, Myanmar.”
Maren Gaulke / Rantau Prapat, Sumatra, Indonesia
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95
Cyclemys gemeli Fritz, Guicking, Auer, Sommer, Wink, and
Hundsdörfer 2008 (08:6)
Assam Leaf Turtle
Peter Praschag / Dimapur, Nagaland, India
Cambodia, China (?) (Yunnan [?]), Laos, Myanmar, Thailand,
Vietnam
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II, as Cyclemys spp.
Cyclemys oldhamii Gray 1863e:178, Cyclemis oldhami
Type locality: “Mergui..[&]..Siam.” Restricted to “Mergui” [Myanmar] by Smith (1931:81) and by lectotype designation by Fritz et al.
(1997:196).
Cyclemys dhor shanensis Annandale 1918:67, Cyclemys
shanensis, Cyclemys shanensis shanensis
Type locality: “Fort Stedman on the Inlé Lake, altitude 3,000 feet..
[&]..He-Ho plain 800 feet higher” [Myanmar].
Bangladesh, Bhutan, India (Arunachal Pradesh, Assam, Bihar,
Manipur, Meghalaya, Mizoram, Nagaland, Uttar Pradesh),
Myanmar, Nepal
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II, as Cyclemys spp.
Cyclemys gemeli Fritz, Guicking, Auer, Sommer, Wink, and
Hundsdörfer 2008b:384
Type locality: “street from Tezpur to Arunachal Pradesh, 5 km to
border of Arunachal Pradesh, Jia Bhoroli River Region, Assam,
India.”
Geoemyda tcheponensis Bourret 1939:7, Cyclemys tcheponensis, Cyclemys dentata tcheponensis, Cyclemys shanensis tcheponensis
Type locality: “Haute Sé-Bang-Hien, (centre de la Chaîne annamitique)” [Laos].
Cyclemys tiannanensis Kou 1989:193
Type locality: “Nanliang, Mengla County, Xishuangbanna of Yunnan
Province, alt. 620 m” [China]. Probable market specimen according
to Fritz et al. (1997:209).
Cyclemys pulchristriata Fritz, Gaulke, and Lehr 1997 (45)
Eastern Black-bridged Leaf Turtle
Cyclemys oldhamii Gray 1863e (08:8) (44)
Southeast Asian Leaf Turtle
Edgar Lehr / Vietnam [trade]
Thomas Ziegler / NW of Ky Thuong, Vietnam
Cambodia, Vietnam
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IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II, as Cyclemys spp.
Cyclemys pulchristriata Fritz, Gaulke, and Lehr 1997:203,
Cyclemys atripons pulchristriata
Type locality: “Phuc-Son, Annam” [Vietnam].
Geoclemys Gray 1856b
Geoclemys Gray 1856b:17
Type species: Geoclemys hamiltonii [= Emys hamiltonii Gray 1830e],
by subsequent designation by Stejneger (1907:496).
Geoclemmys Cope 1865:186 (nomen novum)
Geoclemys hamiltonii (Gray 1830e) (10:7)
Spotted Pond Turtle, Black Pond Turtle
Sind” [Pakistan].
Clemmys palaeindica † Lydekker 1885:178 (nomen novum)
[Late Pliocene (Pinjor) to Early Pleistocene (Tatrot), Siwaliks, India (Punjab)], Damonia palaeindica
Type locality: “Siwalik Hills” [Punjab, India].
Geoclemys sivalensis † Tewari and Badam 1969:555 [Lower
Pleistocene, Upper Siwaliks, India (Haryana)]
Type locality: “Upper Siwaliks...1 km south-east of Quranwalla...6
km northeast of Chandigarh Lake...Panjore, India.” [Haryana,
India].
Geoemyda Gray 1834b (07:39)
Geoemyda Gray 1834b:100 (nomen conservandum, ICZN 1985a)
Type species: Geoemyda spengleri [= Testudo spengleri Gmelin 1789],
by original designation.
Geoemys Bonaparte 1836:6 (nomen novum)
Geomyda Blyth 1856:714 (nomen novum)
Nicoria Gray 1856b:17
Type species: Nicoria spengleri [= Testudo spengleri Gmelin 1789], by
original monotypy.
Geoemyda japonica Fan 1931
Ryukyu Black-breasted Leaf Turtle
Indraneil Das / CBFTT / Kaziranga National Park, Assam, India
Yuichirou Yasukawa / CBFTT / Okinawajima, Japan
Bangladesh, India (Assam, Bihar, Jammu, Meghalaya, Punjab,
Rajasthan, Uttar Pradesh, West Bengal), Nepal, Pakistan
CBFTT Account: Das and Bhupathy (2010)
IUCN Red List: Vulnerable A1d+2d (2000); Previously: Near
Threatened (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix I
Emys hamiltonii Gray 1830e:9 (10:7), Clemmys (Clemmys)
hamiltonii, Clemmys hamiltonii, Geoclemys hamiltonii,
Damonia hamiltonii, Geoclemmys hamiltonii, Geoclemys
hamoltoni
Type locality: “India.”
Emys guttata Gray 1831b:pl.76
Type locality: “India.”
Emys piquotii Lesson 1831a:120
Type locality: “le Gange” [India].
Emys picquotii Lesson in Duméril and Bibron 1835:316 (nomen novum)
Emys hamiltonoides † Falconer and Cautley in Lydekker
1880:21 (nomen nudum) [Late Pliocene (Pinjor) to Early
Pleistocene (Tatrot), Siwaliks, India (Punjab)], Damonia
hamiltonoides
Melanochelys pictus Murray 1884:107
Type locality: “in the Sind "Doro," in the Kushmore Talooka, Upper
(orange dots = introduced)
Japan (Ryukyu Archipelago)
CBFTT Account: Yasukawa and Ota (2008)
IUCN Red List: Endangered A1ce, B1+2c (2000); Previously:
Endangered (1996)
CITES: Appendix II
Geoemyda spengleri japonica Fan 1931:148, Geoemyda
japonica, Geoemyda japonicus
Type locality: “Japan and other Pacific Islands.” Restricted to
“Nawa (?Naha), Okinawajima Is., Japan” by Yasukawa et al.
(1992:149).
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Geoemyda spengleri (Gmelin 1789) (09:25)
Black-breasted Leaf Turtle
Hardella Gray 1870c
Hardella Gray 1870c:58
Type species: Hardella thurjii [= Emys thurjii Gray 1831d], by subsequent designation as Hardella thurgi by Günther (1871:70).
Hardella thurjii (Gray 1831d) (07:40)
Crowned River Turtle
Flora Ihlow / Tam Dao National Park, Vietnam
Indraneil Das / CBFTT / National Chambal Sanctuary, Madhya Pradesh, India
China (Guangdong, Guangxi, Hainan, Hunan [?], Jiangxi),
Laos, Vietnam
CBFTT Account: Yasukawa and Ota (2010)
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Testudo spengleri Gmelin 1789:1043 (nomen conservandum,
ICZN 1985a), Emys spengleri, Geoemyda spengleri,
Clemmys (Clemmys) spengleri, Clemmys spengleri, Nicoria spengleri, Geoemyda spengleri spengleri
Type locality: Not designated. Originally indicated as coming from
“vermuthlich...Ostindien” by Walbaum (1785:129); and restricted to
“probably...from the East Indies” by Pope (1935:36).
Testudo serrata Shaw 1802:51 (junior homonym, not =
Testudo serrata Daudin 1801)
Type locality: Not designated.
Testudo tricarinata Bory de Saint-Vincent 1804:308 (junior
homonym, not = Testudo tricarinata Retzius in Schoepff
1792)
Type locality: “Bourbon...mare d’Arzule” [Réunion] [in error].
Geoemyda spengleri sinensis Fan 1931:146
Type locality: “Loshiang and Kutchen” [Luoxiang and Guchen,
Guangxi, China].
Bangladesh, India (Assam, Bihar, Madhya Pradesh, Meghalaya,
Punjab, Uttar Pradesh, West Bengal), Nepal, Pakistan
CBFTT Account: Das and Bhupathy (2009a)
IUCN Red List: Vulnerable A1cd+2cd (2000); Previously: Near
Threatened (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Emys thuryi Gray 1830e:8 (10:7) (nomen oblitum)
Type locality: “Bengal” [Bangladesh or India].
Emys thurjii Gray 1831d:22 (nomen novum), Testudo thurjii,
Hardella thurjii, Hardella thurjii thurjii
Type locality: “India.” Restricted to “Ganges and Brahmaputra
systems” [India] as Hardella thurgii by McDowell (1964:255).
Emys flavonigra Lesson 1831a:120
Type locality: “le Gange” [India].
Clemmys (Clemmys) thurgii Fitzinger 1835:123 (nomen
novum), Clemmys thurgii, Testudo thurgii, Emys thurgii,
Emys thurgi, Batagur thurgii, Hardella thurgii, Batagur
(Hardella) thurgii
Kachuga oldhami Gray 1869a:200, Kachuga (Kachuga)
oldhami
Type locality: “India.”
Hardella indi Gray 1870c:58 (07:40), Hardella thurjii indi
Type locality: “Indus River” [Pakistan].
Batagur falconeri † Lydekker 1885:187 [Late Pliocene (Pinjor) to Early Pleistocene (Tatrot), Siwaliks, India (Punjab)],
Hardella falconeri
Type locality: “Siwalik Hills” [Punjab, India].
Batagur cautleyi † Lydekker 1885:194 [Late Pliocene (Pinjor) to Early Pleistocene (Tatrot), Siwaliks, India (Punjab)],
Type locality: “Siwalik Hills” [Punjab, India].
Clemmys watsoni † Lydekker 1886:541 [Late Pliocene to Early
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Pleistocene, Siwaliks, India (Gujarat)]
Type locality: “Siwaliks of Perim Island, Gulf of Cambay, India”
[Gujarat, India].
Heosemys depressa (Anderson 1875)
Arakan Forest Turtle
Geoemyda pilgrimi † Prasad and Satsangi 1967 [Late Pliocene to Early Pleistocene, Siwaliks, Haritalyanagar, India
(Himachal Pradesh)]
Type locality: “Tatrot beds east of Chakrana in Bilaspur District,
Himachal Pradesh, India.”
Heosemys Stejneger 1902
Heosemys Stejneger 1902:238
Type species: Heosemys spinosa [= Emys spinosa Gray 1831d], by
original designation.
Brian D. Horne / TCC / Myanmar
Hieremys Smith 1916:50
Type species: Hieremys annandalii [= Cyclemys annandalii Boulenger
1903a], by original monotypy.
Heosemys annandalii (Boulenger 1903a) (07:41)
Yellow-headed Temple Turtle
Sitha Som / TL / Tonle Sap, Cambodia
Bangladesh, Myanmar
IUCN Red List: Critically Endangered A2cd, B1+2c (2000); Previously: Critically Endangered (1996)
TFTSG Draft Red List: Critically Endangered or Endangered
(2011)
CITES: Appendix II
Geoëmyda depressa Anderson 1875:284, Geoemyda depressa, Heosemys depressa
Type locality: “Arakan” [Myanmar]. Restricted to “Akyab, near
Arakan” [Myanmar] by Smith (1931:95).
Geomyda arakana Theobald 1876:vii, Geoemyda arakana
Type locality: “Akyab” [Myanmar].
Heosemys grandis (Gray 1860d)
Giant Asian Pond Turtle
Cambodia, Laos, Malaysia (Peninsular), Thailand, Vietnam
IUCN Red List: Endangered A1cd+2d (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Critically Endangered or Endangered
(2011)
CITES: Appendix II
Cyclemys annandalii Boulenger 1903a:142 (07:41), Hieremys
annandalii, Heosemys annandalii
Type locality: “Kampong Jalor” [= Yala, Yala Prov., Thailand].
Hieremys annandalei Smith 1916:50 (nomen novum), Cyclemys annandalei
Jérôme Maran / Vietnam
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Emys spinosa Gray 1830e:8 (nomen novum), Geoemyda
spinosa, Geomyda spinosa, Clemmys (Clemmys) spinosa,
Clemmys spinosa, Heosemys spinosa
Type locality: “Penang” [Peninsular Malaysia].
Leucocephalon McCord, Iverson, Spinks, and Shaffer 2000
Leucocephalon McCord, Iverson, Spinks, and Shaffer 2000:21
Type species: Leucocephalon yuwonoi McCord, Iverson, and Boeadi
1995, by original designation.
Leucocephalon yuwonoi (McCord, Iverson, and Boeadi 1995) (07:42)
Sulawesi Forest Turtle
Cambodia, Laos, Malaysia (Peninsular), Myanmar, Thailand,
Vietnam
IUCN Red List: Vulnerable A1d+2cd (2000); Previously: Near
Threatened (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Emys siamensis Gray in Günther 1860:114 (nomen nudum)
Type locality: “Siam” [Thailand].
Geoemyda grandis Gray 1860d:218, Geomyda grandis,
Clemmys grandis, Heosemys grandis
Cris Hagen / CBFTT / TCC / Sulawesi, Indonesia / male
Type locality: “Cambojia” [Cambodia].
Heosemys spinosa (Bell in Gray 1830a) (12:25)
Spiny Turtle
(orange dots = trade)
Peter Paul van Dijk / Perlis State Park, Perlis, Malaysia
Indonesia (Sulawesi)
CBFTT Account: Hagen, Platt, and Innis (2009)
IUCN Red List: Critically Endangered A1cd+2cd, C1 (2000); Previously: Data Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Geoemyda yuwonoi McCord, Iverson, and Boeadi 1995:311,
Heosemys yuwonoi, Leucocephalon yuwonoi
Type locality: “near Gorontalo [0º33' N, 123º05' E] on the Minahassa Peninsula, northern Sulawesi [Celebes], Indonesia.”
Brunei, Indonesia (Sumatra, Kalimantan), Malaysia (Peninsular,
East), Myanmar, Philippines (Sulu Archipelago [TawiTawi]), Singapore, Thailand
IUCN Red List: Endangered A1bcd (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Emys spinosae Bell in Gray 1830a:pl.77 (nomen oblitum)
Type locality: “Penang” [Peninsular Malaysia].
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Malayemys Lindholm 1931 (46)
Damonia Gray 1869a:193 (junior homonym, not = Damonia
Robineau-Desvoidy 1847 [= Diptera])
Malayemys macrocephala (Gray 1859) (07:43) (46)
Malayan Snail-eating Turtle
Type species: Damonia macrocephala [= Geoclemmys macrocephala
Gray 1859], by subsequent designation by Stejneger (1907:496).
Malayemys Lindholm 1931:30 (nomen novum)
Type species: Malayemys macrocephala [= Geoclemmys macrocephala Gray 1859], in accordance with ICZN Article 67.8, not
Malayemys subtrijuga [= Emys subtrijuga Schlegel and Müller
1845], as by original designation by Lindholm (1931:30).
Malayemys khoratensis Ihlow, Vamberger, Flecks, Hartmann, Cota,
Makchai, Meewattana, Dawson, Kheng, Rödder, and Fritz
2016 (46, 47)
Khorat Snail-eating Turtle
Flora Ihlow / Tap Than, Thailand
Flora Ihlow / Sikhio, Thailand
(orange dots = possible trade or introduced)
Laos, Thailand
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Not Evaluated
Malayemys khoratensis Ihlow, Vamberger, Flecks, Hartmann,
Cota, Makchai, Meewattana, Dawson, Kheng, Rödder, and
Fritz 2016:16 (46, 47)
Type locality: “Udon Thani, Udon Thani Province, Thailand
(17.36555°N, 102.81427°E)”
Malayemys isan Sumontha, Brophy, Kunya, Wiboonatthapol,
and Pauwels 2016:2 (46, 47)
Type locality: “Ban Na Klang (17°14'48.728"N, 102°12'32.479"E),
Na Klang Sub-district, Na Klang District, Nong Bua Lamphu
Province, northeastern Thailand.”
Malaysia (Peninsular), Thailand
Possibly Introduced or Native: Indonesia (Sumatra) (prehistoric
or historic?)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Vulnerable (2011); Previously: Least
Concern [Not Listed] (1996), as part of Malayemys
subtrijuga
CITES: Appendix II
Geoclemmys macrocephala Gray 1859:479, Clemmys macrocephala, Emys macrocephala, Damonia macrocephala,
Geoclemys macrocephala, Malayemys macrocephala
Type locality: “Siam” [Thailand]. Restricted to “Thanyaburi,
Pathum Thani Province, Thailand (Chao Phraya River Basin; approx. 50 km NNE of Bangkok; 14.017 N, 100.733 E)” by Brophy
(2004:75).
Emys megacephala Gray 1870c:44 (nomen nudum), Damonia megacephala
Malayemys subtrijuga (Schlegel and Müller 1845) (46)
Mekong Snail-eating Turtle
Flora Ihlow / Prek Toal, Tonle Sap Lake, Cambodia
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chinemys Smith 1931:116
Type species: Chinemys reevesi [= Emys reevesii Gray 1831d)], by
original monotypy.
Annamemys Bourret 1940:15
Type species: Annamemys merkleni Bourret 1940 [= subjective
synonym of Cyclemys annamensis Siebenrock 1903a], by original
monotypy.
Mauremys annamensis (Siebenrock 1903a) (07:44, 14:29) (49)
Vietnamese Pond Turtle, Annam Pond Turtle
(orange dots = possibly native or introduced or trade)
Cambodia, Laos, Thailand (?), Vietnam
Possibly Introduced or Native: Indonesia (Java, Sumatra?)
(prehistoric or historic?)
IUCN Red List: Vulnerable A1d+2d (2000); Previously: Least
Concern [Not Listed] (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II
Emys herrmanni Schweigger 1812:311 (nomen dubium),
Clemmys (Clemmys) hermanni
Jeffrey E. Dawson / CBFTT / Vietnam [captivity]
Type locality: Not known.
Emys subtrijuga Schlegel and Müller 1845:30 (46), Damonia
subtrijuga, Geoclemys subtrijuga, Malayemys subtrijuga
Type locality: “Java” [Indonesia]. Possibly introduced, but perhaps
native (46).
Cistudo gibbosa Bleeker 1857b:239 (nomen nudum)
Type locality: “Batavia...Java” [Indonesia].
Emys nuchalis Blyth 1863:82, Bellia nuchalis
Type locality: “Java?” [Indonesia].
Damonia crassiceps Gray 1870c:43 (48)
Type locality: “China.”
Damonia oblonga Gray 1871c:367
Type locality: “Batavia” [Java, Indonesia].
Mauremys Gray 1869b (07:44, 09:26)
Mauremys Gray 1869b:500
Type species: Mauremys fuliginosa [= Emys fuliginosus Gray 1860c
= subjective synonym of Emys leprosa Schoepff in Schweigger
1812], by subsequent designation by Lindholm (1929:281).
Ocadia Gray 1870c:35
Type species: Ocadia sinensis [= Emys sinensis Gray 1834a], by
original monotypy.
Emmenia Gray 1870c:38
Type species: Emmenia grayi [= Emys grayi Günther 1869 = subjective synonym of Mauremys caspica siebenrocki Wischuf and Fritz
in Fritz and Wischuf 1997 = subjective synonym of Testudo caspica
Gmelin 1774], by original monotypy.
Eryma Gray 1870c:44 (junior homonym, not = Eryma Meyer
1840 [= Crustacea †] or Eryma Albers 1854 [= Gastropoda] or Eryma Förster 1868 [= Hymenoptera])
Type species: Eryma laticeps [= Emys laticeps Gray 1854a = subjective synonym of Emys leprosa Schoepff in Schweigger 1812], by
original monotypy.
Cathaiemys Lindholm 1931:29
Type species: Cathaiemys mutica [= Emys mutica Cantor 1842b], by
original designation.
Pseudocadia Lindholm 1931:30
Type species: Pseudocadia anyangensis [= Testudo anyangensis † Ping
1930], by original designation.
Vietnam
CBFTT Account: McCormack, Dawson, Hendrie, Ewert,
Iverson, Hatcher, and Goode (2014)
IUCN Red List: Critically Endangered A1d+2d (2000); Previously: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Cyclemys annamensis Siebenrock 1903a:341, Cuora (Cyclemys) annamensis, Cuora annamensis, Annamemys annamensis, Mauremys annamensis, Cathaiemys annamensis
Type locality: “Annam (Phuc-Son)” [Vietnam].
Annamemys merkleni Bourret 1940:15
Type locality: “Annam...Fai-Fo” [Vietnam]
Clemmys guangxiensis Qin 1992:60 (14:29) (partim, hybrid),
Mauremys guangxiensis
Type locality: “Nanning, Guangxi” [China]. [in error?]
Ocadia glyphistoma McCord and Iverson 1994:53 (07:33)
(partim, hybrid)
Type locality: “near the Vietnam border southwest of Nanning,
Guangxi Province, China.” [in error?]
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Mauremys caspica (Gmelin 1774) (09:27, 12:26) (50)
Caspian Turtle, Caspian Terrapin
Mauremys japonica (Temminck and Schlegel 1838) (10:18, 14:30)
Japanese Pond Turtle
Asghar Mobaraki / Maharloo Lake, Shiraz, Fars, Iran
Yuichirou Yasukawa / CBFTT / Shiga, Honshu, Japan
Armenia, Azerbaijan, Bahrain, Georgia, Iran (Ardabil, Bushehr,
Chahar Mahal Va Bakhtiari, East Azarbaijan, Esfahan,
Fars, Gilan, Golestan, Ilam, Kermanshah, Khuzestan,
Kordestan, Lorestan, Mazandaran, West Azarbaijan), Iraq,
Russia (Dagestan), Saudi Arabia (Eastern), Syria, Turkey,
Turkmenistan
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Testudo caspica Gmelin 1774:59,pl.10, Emys caspica, Clemmys caspica, Clemmys (Clemmys) caspica, Terrapene
caspica, Clemmys caspica caspica, Mauremys caspica,
Mauremys caspica caspica
Type locality: “Schamachie...bei Bach Pusahat” [Pirsagat, Shemakha, Azerbaijan]. Emended to “Hircaniae aquis dulcibus” by
Gmelin (1789:1042); and to “les bords de la mer Caspienne...dans
les eaux douces de l’Hircanie” [Azerbaijan] by Daudin (1801:124).
Testudo ecaudata Daudin 1801:125 (nomen novum and
senior homonym, not = Testudo ecaudata Pallas 1814)
Type locality: “les bords de la mer Caspienne...dans les eaux douces
de l’Hircanie” [Azerbaijan].
Emys caspia Rüppell in Gray 1830e:9 (nomen novum),
Testudo caspia, Clemmys caspia, Clemmys caspia caspia
Type locality: “Caspian Sea.”
Emys grayi Günther 1869:504 (junior homonym, not = Emys
grayi Bocourt 1868), Emmenia grayi
Type locality: “Bussora” [Basrah, Iraq].
Mauremys caspica siebenrocki Wischuf in Maran 1996:17
(nomen nudum)
Mauremys caspica schiras Wischuf in Maran 1996:17 (nomen nudum)
Mauremys caspica ventrimaculata Wischuf and Fritz
1996:114
Type locality: “Tang-e Karam (Tang-i-Kerim), 70 Meilen östlich
von Schiraz, Iran (29º 03' N, 53º 38' E).”
Mauremys caspica siebenrocki Wischuf and Fritz in Fritz and
Wischuf 1997:240
Type locality: “Kerbela, Irak” [Iraq].
Japan (Honshu, Kyoshu, Shikoku)
CBFTT Account: Yasukawa, Yabe, and Ota (2008)
IUCN Red List: Near Threatened (2000); Previously: Least
Concern [Not Listed] (1996)
CITES: Appendix II
Emys palustris var. Japon Temminck and Schlegel
1834:pls.8-9 (14:30) (invalid vernacular name)
Emys vulgaris var. japonaise Temminck and Schlegel 1834:54
(14:30)
(invalid vernacular name)
Emys vulgaris japonica Temminck and Schlegel 1838:139
(10:18, 14:30)
, Emys japonica, Emys caspica japonica, Clemmys japonica, Mauremys japonica, Ocadia japonica
Type locality: “Japon” [Japan].
Testudo margaritifera Schlegel in Gray 1856b:11 (nomen
nudum)
Type locality: “Japon” [Japan].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Mauremys leprosa (Schoepff in Schweigger 1812) (51)
Mediterranean Pond Turtle, Spanish Terrapin
103
Emys marmorea Spix 1824:13, Clemmys marmorea
Type locality: “Brasil” [in error]. Restricted to “southern Spain” by
Fritz and Havaš (2007:230).
Clemmys sigriz Michahelles 1829:1296, Terrapene sigriz,
Emys sigriz, Clemmys caspica sigriz
Type locality: “paludibus Hispaniae meridionalis” [marshes of
southern Spain].
Emys vulgaris Gray 1830e:9 (10:7)
Type locality: “South of Europe...Spain.” Validated by lectotype
designation by Fritz and Wischuf (1997:249).
Emys sigritzii Michahelles in Gray 1831d:24 (nomen novum)
Emys laticeps Gray 1854a:134, Clemmys laticeps, Eryma
laticeps
Type locality: “West Africa, River Gambia” [Gambia]; possibly
in error. Emended to “Tétouan, Morocco” by Fritz and Havaš
(2007:230) without explanation.
Emys fuliginosus Gray 1860c:232, Emys fuliginosa, Clemmys
fuliginosa, Mauremys fuliginosa
Type locality: “Egypt?...North Africa?” Emended to “southern
Spain” by Fritz and Havaš (2007:230) without explanation.
Mauremys laniaria Gray 1869b:499
Type locality: Not known. Restricted to “southern Spain” by Fritz
and Havaš (2007:230).
Emys flavipes Gray 1869c:643
Type locality: Not known. Restricted to “southern Spain” by Fritz
and Havaš (2007:230).
Emys fraseri Gray 1869c:643 (partim, nomen dubium)
Type locality: “North Africa.” Restricted to “Lake Tetzara, Algiers”
[Algeria] by Gray (1873a:146).
Mauremys leprosa atlantica Schleich 1996a:32 (07:45)
(subspecies: leprosa = red, saharica = purple,
overlap = intergrades; orange dots = possibly introduced)
Algeria, France, Libya, Mauritania (prehistoric introduction?),
Mali (prehistoric introduction?), Morocco, Niger (prehistoric introduction?), Portugal, Spain (Continental), Tunisia
IUCN Red List: Global: Least Concern [Not Listed] (1996);
Regional: Europe: Vulnerable A2ac+3c (2004)
TFTSG Draft Red List: Vulnerable (2011)
Mauremys leprosa leprosa (Schoepff in Schweigger 1812) (07:45)
Mediterranean Pond Turtle
Type locality: “Larache, Nord-Marokko” [Morocco].
Mauremys leprosa erhardi Schleich 1996a:36 (07:45)
Type locality: “NNE Taza, N 35º 25' 32.8, W 02º 52' 11.5, Nordost
Marokko” [Morocco].
Mauremys leprosa marokkensis Schleich 1996a:40 (07:45)
Type locality: “ca. 2 km NE Tahanaoute / Kreuzung S 501, wenige
km vor Aguelmouss; S-Marrakech, N 31º 24' 54.0, W 7º 49' 44.4,
Zentral-Morokko” [Morocco].
Mauremys leprosa wernerkaestlei Schleich 1996a:44 (07:45)
Type locality: “Wasserlauf im Schnittpunkt von Oued Serou, Oued
Oum de Oumbia. Oued Oum er Rbia, S-Khenifra, N 32º 44' 55.1,
W 5º 41' 10.7, ca. 90 m NN, Zentral-Morokko” [Morocco].
Mauremys leprosa saharica Schleich 1996 (07:45)
Saharan Pond Turtle
Albert Bertolero / Flix, Catalonia, Spain
Algeria, France, Morocco, Portugal, Spain (Continental)
Emys leprosa Schoepff in Schweigger 1812:298, Clemmys (Clemmys) leprosa, Clemmys leprosa, Emys caspica
leprosa, Clemmys caspica leprosa, Mauremys caspica
leprosa, Mauremys leprosa, Mauremys leprosa leprosa
Type locality: Not known. Restricted to “Südspanien” [Spain] by
Mertens and Müller (1928:22).
Emys lutescens Schweigger 1812:302, Clemmys (Clemmys)
lutescens, Clemmys lutescens
Type locality: Not known. Restricted to “southern Spain” by Fritz
and Havaš (2007:230).
Andreas Nöllert / Bou-Jerif, Morocco
Algeria, Libya, Mauritania (prehistoric introduction?), Mali
(prehistoric introduction?), Morocco, Niger (prehistoric
introduction?), Tunisia
Emys fraseri Gray 1869c:643 (partim, nomen dubium)
Type locality: “North Africa.” Restricted to “Lake Tetzara, Algiers”
[Algeria] by Gray (1873a:146).
Mauremys leprosa saharica Schleich 1996a:45 (07:45)
Type locality: “Fort Bou Jerif, Goulmime, Südwest-Marokko”
[Morocco].
Mauremys leprosa zizi Schleich 1996a:48 (07:45)
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Type locality: “Oued Ziz, Meski bis Erfoud, Süd-Marokko”
[Morocco].
Mauremys leprosa vanmeerhaeghei Bour and Maran 1999:42
(07:45)
Type locality: “Mare amont dans l’oasis de Sidi El Mehdaoui
(29º30 N, 8º00 W)...Province de Tata (ca. 30 km de Tata), Maroc...
affluent de...l’oued Draa” [Morocco].
Mauremys mutica (Cantor 1842b) (49)
Yellow Pond Turtle
Vietnam
Emys muticus Cantor 1842b:482, Emys mutica, Clemmys
mutica, Damonia mutica, Geoclemys mutica, Cathaiemys
mutica, Mauremys mutica, Mauremys mutica mutica,
Cathaiemys mutica mutica
Type locality: “Chusan...Island” [Zhoushan Island, Zhejiang,
China]. Cited as “Canton” [in error] by many authors, corrected by
Iverson and McCord (1989:23).
Clemmys schmackeri Boettger 1894:129
Type locality: “China, wahrscheinlich Hainan.”
Annamemys grochovskiae Dao 1957:1214, Mauremys
grochovskiae
Type locality: In Russian [“forest near Vinh-Linh, Quang Tri Province, central Vietnam”].
Mauremys iversoni Pritchard and McCord 1991:140 (07:33) (partim,
hybrid)
Type locality: “People’s Republic of China: Fujian Province: vicinity
of Nanping (26º38' N, 118º10' E).”
Clemmys guangxiensis Qin 1992:60 (14:29) (partim, hybrid),
Mauremys guangxiensis
Type locality: “Nanning, Guangxi” [China].
Mauremys pritchardi McCord 1997:555 (07:33) (partim,
hybrid)
Type locality: “Lashio, Myanmar (97º14' E, 22º56' N).”
Mauremys mutica kami Yasukawa, Ota, and Iverson 1996
Ryukyu Yellow Pond Turtle
John B. Iverson / Ryukyus, Japan
(subspecies: mutica = red; kami = purple)
China (Anhui, Fujian, Guangdong, Guangxi, Hainan, Hubei, Hunan, Jiangsu, Jiangxi, Yunnan, Zhejiang), Japan
(Ryukyu Archipelago), Taiwan, Vietnam
Introduced: Japan (mainland)
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Japan (Ryukyu Archipelago)
Mauremys mutica kami Yasukawa, Ota, and Iverson
1996:311, Cathaiemys mutica kami
Type locality: “Okawa, Ishigakijima Is., the Yaeyama Group,
Ryukyu Archipelago, Japan.”
Mauremys nigricans (Gray 1834a)
Chinese Red-necked Turtle, Red-necked Pond Turtle
Mauremys mutica mutica (Cantor 1842b)
Yellow Pond Turtle
Ben Anders / CBFTT / China [trade]
Tien-Hsi Chen / Taiwan
China (Anhui, Fujian, Guangdong, Guangxi, Hainan, Hubei,
Hunan, Jiangsu, Jiangxi, Yunnan, Zhejiang), Taiwan,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
China (Fujian [?], Guangdong, Guangxi, Hainan [?]), Vietnam (?)
CBFTT Account: Anders and Iverson (2012)
IUCN Red List: Endangered A1d+2d (2000); Previously: Data
Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Emys nigricans Gray 1834a:53, Emys nigracans, Clemmys
nigricans, Damonia nigricans, Chinemys nigricans, Mauremys nigricans
Type locality: “Chinâ prope Canton” [Gangzhou, Guangdong].
Geoclemys kwangtungensis Pope 1934:1, Clemmys kuangtungensis, Chinemys kwangtungensis
Type locality: “Lofaoshan, Kwangtung, 300–400 m. altitude” [Mt.
Luofu, Guangdong, China].
Geoclemys palaeannamitica † Bourret 1941a:10 [Holocene,
Neolithic, subfossil, Vietnam], Geoclemmys palaeannamitica, Chinemys palaeannamitica
Type locality: “la grotte de Dong-Giao, appelée ‘Thung Gianh’... à
moins de 1 km de la voie ferrée, vers le N.W., au S.W. de la station
de Dong-Giao, tout près de la frontière entre Annam et Tonkin”
[Vietnam].
(native populations = red, introduced populations = gray;
orange dots = introduced and probable trade)
China (Anhui, Fujian, Gansu, Guangdong, Guizhou, Henan,
Hong Kong, Hubei, Hunan, Jiangsu, Jiangxi, Shangdong,
Shanxi, Shaanxi, Sichuan, Zhejiang), Japan (prehistoric
introduction), North Korea, South Korea, Taiwan (prehistoric introduction)
Introduced: Indonesia (Timor), Japan, Palau, Philippines, Taiwan, Timor-Leste
CBFTT Account: Lovich, Yasukawa, and Ota (2011)
IUCN Red List: Endangered A2bcd+4bcd (2011); Previously:
Endangered (2000), Least Concern [Not Listed] (1996)
CITES: Appendix III (China)
Emys reevesii Gray 1831d:73, Clemmys (Clemmys) reevesii,
Clemmys reevesii, Geoclemys reevesii, Geoclemmys
reevesii, Damonia reevesii, Geoclemys reevesii reevesii,
Chinemys reevesi, Chinemys reevesii, Mauremys reevesii
Type locality: “China.”
Emys vulgaris picta Schlegel 1844:127
Type locality: “Japan.”
Mauremys reevesii (Gray1831d) (07:46) (52)
Reeves’ Turtle, Chinese Three-keeled Pond Turtle
Emys japonica Duméril and Bibron in Duméril and Duméril
1851:8 (nomen novum)
Type locality: “Japon” [Japan].
Damonia unicolor Gray 1873e:78, Clemmys unicolor, Damonia reevesii unicolor, Geoclemys reevesii unicolor
Type locality: “China...Shanghai.”
Geoclemys grangeri Schmidt 1925:1, Geoclemys reevesii
grangeri, Chinemys grangeri
Type locality: “Yenchingkao, Wanhsien, Szechwan, 1500 feet
altitude” [Yanjinggou, Sichuan, China].
Geoclemys paracaretta Chang 1929:1, Geoclemys
papacaretta
Nobuhiro Kawazoe / CBFTT / Honshu, Japan
Type locality: “Fuchow” [Fuzhou Shi, Fujian, China].
Chinemys megalocephala Fang 1934:158, Mauremys megalocephala, Chinemys megacephala
Type locality: “hill-sides of the vicinity of Nanking city” [Nanjing
Shi, Jiangsu, China].
Chinemys macrocephala Bourret 1941c:140 (nomen novum)
Chinemys pani † Tao 1985:45 [Pleistocene, Chi-Ting,
Taiwan]
Type locality: “Tsochen, Tsai-liao-chi, Tainan, Taiwan.”
Mauremys pritchardi McCord 1997:555 (07:33) (partim, hybrid)
Type locality: “Lashio, Myanmar (97º14' E, 22º56' N).”
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Mauremys rivulata (Valenciennes in Bory de Saint-Vincent 1833)
(14:25) (50, 53)
Western Caspian Turtle, Balkan Terrapin
di Cattaro, Süd-Dalmatien” [Croatia] by Wermuth and Mertens
(1961:60).
Clemmys caspica cretica Mertens 1946:115, Mauremys
caspica cretica, Mauremys rivulata cretica
Type locality: “Rapaniana, Kreta” [Crete, Greece].
Mauremys sinensis (Gray 1834a) (07:44)
Chinese Stripe-necked Turtle
Apostolis Trichas / CBFTT / Crete, Greece
John B. Iverson / No data [trade]
Albania, Bosnia and Herzegovina, Bulgaria, Croatia, Cyprus,
Greece, Israel, Jordan, Kosovo (?), Lebanon, Macedonia,
Montenegro, Palestine (West Bank), Syria, Turkey
Introduced: Latvia
CBFTT Account: Mantziou and Rifai (2014)
IUCN Red List: Global: Not Evaluated; Regional: Europe:
Least Concern (2004)
TFTSG Draft Red List: Least Concern (2011)
Emys pannonica Michahelles in Bonaparte 1831:87 (nomen
nudum)
Emys rivulata Valenciennes in Bory de Saint-Vincent
1833:planches,pl.9 (14:25) (senior homonym, not = Emys
rivulata Gray 1844), Clemmys caspica rivulata, Mauremys
caspica rivulata, Mauremys rivulata, Mauremys rivulata
rivulata, Emmenia rivulata
Type locality: Not designated. Restricted to “Morée...Modon”
[Greece] by Bory de Saint-Vincent (1836:110), and emended
to “Umgebung von Modon, Morea, Griechenland” [Greece] by
Mertens and Müller (1928:22).
Emys tristrami Gray 1869a:190, Emys tristram, Mauremys
rivulata tristrami
Type locality: “Holy Land” [Israel or Jordan]. Restricted to “Jabook
River” [Zarqa River, Jordan] by Gray (1873j:35); and to “Yarmuk
River” [Israel or Jordan] by Boulenger (1889:104).
Emys caspica arabica Gray 1870c:36, Emys arabica
Type locality: “Arabia Petraea?” Restricted to “Mt. Carmel” [Israel]
by Gray (1873j:34).
Emys pannonica Gray 1870c:36
Type locality: “Xantos” [Xanthos, Greece].
Clemmys caspica orientalis Bedriaga 1881:335
Type locality: “Umgebung von Athen und von Nauplia, ferner auf
den Inseln Milo, Mykonos, Syra, Tinos, Siphnos und Seriphos”
[Greece].
Clemmys caspica obsoleta Schreiber 1912:946
Type locality: “südlichen Dalmatien...Bocche di Cattaro..[&].. Halbinsel Sabbioncello..[&]..Menge” [Croatia]. Restricted to “Bocche
China (Fujian, Guangdong, Guangxi, Hainan, Zhejiang), Taiwan, Vietnam
Introduced: South Korea, USA (Florida)
IUCN Red List: Endangered A1cd (2000); Previously: Near
Threatened (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix III (China)
Emys sinensis Gray 1834a:53, Graptemys sinensis, Clemmys sinensis, Ocadia sinensis, Ocadia sinensis sinensis,
Mauremys sinensis
Type locality: “Chinâ.” Restricted to “China; Canton” [Gangzhou,
Guangdong] by Gray (1856b:21).
Emys bennettii Gray 1844:21, Clemmys bennettii
Type locality: “North America?” [in error]. Restricted to “China?”
by Gray (1856b:22).
Testudo anyangensis † Ping 1930:217 [Holocene, Neolithic,
subfossil, China (Henan)], Ocadia anyangensis, Pseudocadia anyangensis, Mauremys anyangensis
Type locality: “ancient ruins in Annyang Hsien, Honan Province”
[Anyang Shi, Henan, China].
Ocadia sinensis changwui † Tao 1988:233 [Late Pleistocene,
Taiwan]
Type locality: “Penghu Channel in the Taiwan Strait” [Taiwan].
Ocadia philippeni McCord and Iverson 1992:13 (07:33) (partim,
hybrid)
Type locality: “near Kancheng [18º51'N, 108º37'E; = 48 km from
Tungfang (19º03'N, 108º56'E)], western Hainan Island, China.” [in
error?]
Ocadia glyphistoma McCord and Iverson 1994:53 (07:33) (partim,
hybrid)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “near the Vietnam border southwest of Nanning,
Guangxi Province, China.” [in error?]
Melanochelys trijuga (Schweigger 1812)
Indian Black Turtle
Melanochelys Gray 1869a
Melanochelys Gray 1869a:187
Type species: Melanochelys trijuga [= Emys trijuga Schweigger 1812],
by original monotypy.
Geomyda (Chaibassia) Theobald 1876:6
Type species: Geomyda (Chaibassia) tricarinata [= Geomyda tricarinata Blyth 1856], by original monotypy.
Melanochelys tricarinata (Blyth 1856)
Tricarinate Hill Turtle, Three-keeled Land Turtle
Indraneil Das / CBFTT / Dehradun, Uttarakhand, India
(subspecies: trijuga = red, coronata = purple, edeniana = orange,
indopeninsularis = pink, parkeri = brown, thermalis = green;
overlap = intergrades; orange dots = introduced or trade)
Bangladesh, Bhutan, India (Arunachal Pradesh, Assam, Bihar,
Jharkhand, Uttar Pradesh, Uttarakhand, West Bengal),
Nepal
CBFTT Account: Das (2009)
IUCN Red List: Vulnerable B1+2c (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix I
Geomyda tricarinata Blyth 1856:714, Geomyda (Chaibassia)
tricarinata, Geoemyda tricarinata, Chaibassia tricarinata,
Nicoria tricarinata, Melanochleys tricarinata
Bangladesh, China (?) (Yunnan), India (Andhra Pradesh,
Assam, Bihar, Gujarat, Karnataka, Kerala, Maharashtra,
Meghalaya, Mizoram, Tamil Nadu, Uttar Pradesh, West
Bengal), Myanmar, Nepal, Pakistan (?), Sri Lanka
Introduced: British Indian Ocean Territory (Chagos Archipelago), Maldives
CBFTT Account: Das and Bhupathy (2009b)
IUCN Red List: Near Threatened (2000); Previously: Data
Deficient (1996)
TFTSG Draft Red List: Near Threatened (2011)
CITES: Appendix II
Melanochelys trijuga trijuga (Schweigger 1812)
Indian Black Turtle
Type locality: “Central India (Chaibása).” Restricted to “Chybassa,
in the District of Singhbhum, Chota Nagpur, Bengal” [Jharkhand,
India] by Anderson (1879:717).
Geoemyda carinata Blyth in Jerdon 1870:69 (nomen novum)
Chaibassia theobaldi Anderson 1879:718
Type locality: “Bishnath Plain, near Tezpur, in Assam, close to the
Brahmaputra” [Assam, India].
Nicoria tricarinata sivalensis † Lydekker 1889:100 [Late
Pliocene (Pinjor) to Early Pleistocene (Tatrot), Siwaliks,
India (Punjab)], Nicoria sivalensis
Type locality: “Siwalik Hills” [Punjab, India].
S. Jayakumar / CBFTT / Anaikatti Hills, Coimbatore, Western Ghats, Tamil Nadu, India
India (Andhra Pradesh, Gujarat, Karnataka, Maharashtra,
Tamil Nadu), Pakistan (?)
Emys trijuga Schweigger 1812:310, Clemmys (Clemmys)
trijuga, Clemmys trijuga, Melanochelys trijuga, Nicoria
trijuga, Geoemyda trijuga, Geoemyda trijuga trijuga,
Melanochelys trijuga trijuga
Type locality: “insula Java” [in error].
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Emys belangeri Lesson 1831b:291
Type locality: “Bengale..[&]..Carnate” [India]. Restricted to “Bengal” [India] by Smith (1931:97).
Melanochelys trijuga indopeninsularis (Annandale 1913)
Bengal Black Turtle
Emys trijuga maderaspatana Anderson 1879:729, Emys
trijuga madraspatana
Type locality: “Madras” [Chennai, Tamil Nadu, India].
Geoemyda trijuga plumbea Annandale 1915a:192
Type locality: “Coorg on the east side of the Western Ghats considerably east and a little south of the Madras district” [Karnataka,
India].
Melanochelys trijuga coronata (Anderson 1879)
Cochin Black Turtle
Bhaba Amatya / CBFTT / Rupa Lake, Pokhara, Nepal
Bangladesh, India (Assam, Bihar, Jharkhand, Meghalaya,
Mizoram, Uttar Pradesh, West Bengal), Nepal
Bellia sivalensis † Theobald 1877:44 (nomen dubium) [Late
Pliocene to Early Pleistocene, Siwaliks, India (Punjab)],
Clemmys sivalensis
Type locality: “Siwaliks in the Punjab...half a mile south of Jaba (a
village 6 miles south-west from Jhand)” [Himachal Pradesh, India].
S. Bhupathy / CBFTT / Kumarakom, Kerala, India
India (Kerala)
Emys trijuga coronata Anderson 1879:729, Nicoria trijuga
coronata, Geoemyda trijuga coronata, Melanochelys
trijuga coronata
Type locality: “Southern India...Travancore” [Kerala, India].
Melanochelys trijuga edeniana Theobald 1876 (07:47)
Burmese Black Turtle
Clemmys hydaspica † Lydekker 1885:172 (nomen dubium)
[Late Pliocene to Early Pleistocene, Siwaliks, India
(Punjab)]
Type locality: “Siwaliks of the Jhelam district in the Punjab” [Uttarakhand, India].
Clemmys theobaldi † Lydekker 1885:173 (nomen dubium)
[Late Pliocene to Early Pleistocene, Siwaliks, India (Punjab)], Bellia theobaldi, Mauremys theobaldi
Type locality: “Siwaliks of Jhand in the Punjab” [Himachal Pradesh,
India].
Clemmys punjabiensis † Lydekker 1885:175 (nomen dubium)
[Late Pliocene to Early Pleistocene, Siwaliks, India
(Punjab)]
Type locality: “Siwaliks of the Punjab” [Punjab, India].
Geoemyda indopeninsularis Annandale 1913:71, Geoemyda
trijuga indopeninsularis, Melanochelys trijuga indopeninsularis, Melanochelys edeniana indopeninsularis
Type locality: “Singhbhum district of Chota Nagpur..[&]..Dharwar
district in...Bombay Presidency...at Devikop, 26 miles south of...
Dharwar” [India]. Restricted to “Singhbhum District, Chota Nagpur” [Jharkhand, India] by Smith (1931:99).
Melanochelys trijuga parkeri Deraniyagala 1939 (54)
Parker’s Black Turtle
Indraneil Das / CBFTT / Myanmar
China (?) (Yunnan), Myanmar
Melanochelys edeniana Theobald 1876:12, Nicoria trijuga
edeniana, Geoemyda trijuga edeniana, Melanochelys
trijuga edeniana, Emys trijuga edeniana, Melanochelys
edeniana edeniana
Type locality: “Arakan, Pegu, and Tenasserim..[&]..near Tounghu”
[Myanmar].
Emys trijuga burmana Anderson 1879:723
Type locality: “Bhamô..[&]..Moulmein..[&]..Khyouk-Phyoo in
Arracan” [Myanmar]. Restricted to “Bhamo, Burma” [Myanmar]
by Smith (1931:98).
Geoemyda trijuga wiroti Reimann in Nutaphand
1979:177, Melanochelys trijuga wiroti, Melanochelys
edeniana wiroti
Type locality: “Thai-Burmese border area (Tak and Mae Hong Son
Provinces)” [Thailand].
Indraneil Das / CBFTT / Sri Lanka
Sri Lanka
Melanochelys trijuga parkeri Deraniyagala 1939:269, Geoemyda trijuga parkeri
Type locality: “Polonnaruva, N.-C.P., ...Ceylon” [Sri Lanka].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Melanochelys trijuga thermalis (Lesson 1830)
Sri Lanka Black Turtle
Threatened (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix I
Emys ocellata Duméril and Bibron 1835:329, Batagur
(Kachuga) ocellata, Batagur ocellata, Clemmys ocellata,
Morenia ocellata
Type locality: “Bengale” [India or Bangladesh] [in error]. Restricted
to “Irrawaddy (Ayeyarwady) river delta, Myanmar” by lectotype
designation by Bour (2009:41).
Emys berdmorei Blyth 1859:281, Emys berdmorii, Batagur
berdmorii, Batagur berdmorei, Kachuga berdmorei,
Kachuga berdmoorei, Kachuga (Dongoka) berdmoorei,
Morenia berdmorei
Type locality: “Arakan or Tenasserim” [Myanmar].
Peter Paul van Dijk / CBFTT / nr. Bentota, Sri Lanka
India (Tamil Nadu), Sri Lanka
Emys thermalis Lesson 1830:86, Clemmys thermalis, Nicoria
trijuga thermalis, Geoemyda trijuga thermalis, Melanochelys trijuga thermalis
Morenia petersi Anderson 1879
Indian Eyed Turtle
Type locality: “eaux thermales de Cannia, près Trinquemalé à
Ceylan,” [Sri Lanka].
Emys sebae Gray 1831d:75, Emys seba, Geoclemys seba,
Geoclemys sebae, Melanochelys sebae, Emys trijuga
sebae
Type locality: Not designated. Restricted to “Ceylon” [Sri Lanka]
by Gray (1844:19).
Shailendra Singh / CBFTT / Ganga R., Uttar Pradesh, India
Morenia Gray 1870c
Morenia Gray 1870c:62
Type species: Morenia berdmorei [= Emys berdmorei Blyth 1859 =
subjective synonym of Emys ocellata Duméril and Bibron 1835],
by subsequent designation by Lindholm (1929:279).
Morenia ocellata (Duméril and Bibron 1835)
Burmese Eyed Turtle
Indraneil Das / CBFTT / Yangon, Myanmar
Bangladesh, India (Assam, Bihar, Uttarakhand, Uttar Pradesh,
West Bengal), Nepal
CBFTT Account: Das and Sengupta (2010)
IUCN Red List: Vulnerable A1cd+2d (2000); Previously: Near
Threatened (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II
Batagur (Morenia) petersi Anderson 1879:761, Batagur
petersi, Morenia petersi
Type locality: “Huzurapur in the Jessore District..[&]..Furreedpore..
[&]..Dacca” [Uttar Pradesh, India..[&]..Bangladesh].
Notochelys Gray 1863e
Notochelys Gray 1863e:177 (senior homonym, not = Notochelys
Owen 1882 [= Testudines: Protostegidae †])
Type species: Notochelys platynota [= Emys platynota Gray 1834a],
by original monotypy.
China (?) (Yunnan), Myanmar
CBFTT Account: Das (2010)
IUCN Red List: Vulnerable A1cd+2cd (2000); Previously: Near
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Notochelys platynota (Gray 1834a)
Malayan Flat-shelled Turtle
Orlitia borneensis Gray 1873b (12:27)
Malaysian Giant Turtle
Jérôme Maran / Kalimantan, Indonesia [Borneo]
Sabine Schoppe / No data [trade]
(orange dots = probable trade or misidentified)
Brunei, Indonesia (Java, Kalimantan, Sumatra), Malaysia (Peninsular, East), Myanmar, Singapore, Thailand
IUCN Red List: Vulnerable A1cd+2cd (2000); Previously: Data
Deficient (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II
Emys platynota Gray 1834a:54, Emys platynotha, Cyclemys
platynota, Notochelys platynota
Type locality: “Indiâ Orientali.” Restricted to “Sumatra” [Indonesia]
by Gray (1835:pl.57).
Cistudo bankanensis Bleeker in Gray 1864a:12
Indonesia (Kalimantan, Sumatra), Malaysia (Peninsular, East)
IUCN Red List: Endangered A1d+2d (2000); Previously: Near
Threatened (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Cistudo borneënsis Bleeker 1857a:473 (nomen nudum),
Cistudo borneensis
Type locality: “Sintang aan de Kapoeas...Borneo” [Sintang, Kapuas
River, Kalimantan, Indonesia].
Orlitia borneensis Gray 1873b:157, Bellia borneensis
Type locality: “Borneo” [Kalimantan, Indonesia].
Clemmys (Heteroclemmys) gibbera Peters 1875:622, Clemmys gibbera, Heteroclemmys gibbera
Type locality: “Pulo Matjan (Borneo)” [Kalimantan, Indonesia].
Type locality: “Banka Island” [Sumatra, Indonesia].
Hardella baileyi Bartlett 1895b:83, Brookeia baileyi
Type locality: “Borneo” [Kalimantan, Indonesia].
Adelochelys crassa Baur 1896:319
Cyclemys giebelii Hubrecht 1881:45
Orlitia Gray 1873b
Orlitia Gray 1873b:156
Type species: Orlitia borneensis Gray 1873b, by original monotypy.
Clemmys (Heteroclemmys) Peters 1875:622
Type species: Clemmys (Heteroclemmys) gibbera Peters 1875 [=
subjective synonym of Orlitia borneensis Gray 1873b], by original
monotypy.
Brookeia Bartlett 1896:113
Type species: Brookeia baileyi [= Hardella baileyi Bartlett 1895b =
subjective synonym of Orlitia borneensis Gray 1873b], by original
monotypy.
Adelochelys Baur 1896:319
Type species: Adelochelys crassa Baur 1896 [= subjective synonym of
Orlitia borneensis Gray 1873b], by original monotypy.
Liemys Boulenger 1897a:468
Type species: Liemys inornata Boulenger 1897a [= subjective synonym of Orlitia borneensis Gray 1873b], by original monotypy.
Type locality: “Batang Lupar” [Sarawak, East Malaysia].
Type locality: Not known.
Liemys inornata Boulenger 1897a:469
Type locality: “Lobuk Antu district, Borneo” [Sarawak, East
Malaysia].
Batagur signatus † Jaekel 1911:77 [Pleistocene, Pithecanthropus Trinil Beds, Indonesia (Java)]
Type locality: “Pithecanthropus-schichten...Java...Trinil”
[Indonesia].
Pangshura Gray 1856b (07:48)
Batagur (Pangshura) Gray 1856b:36
Type species: Kachuga (Pangshura) tecta [= Emys tectum Gray
1830b], by subsequent designation by Lindholm (1929:278).
Cuchoa Gray 1870c:61
Type species: Cuchoa tentoria [= Emys tentoria Gray 1834a], by
subsequent designation by Lindholm (1929:278).
Jerdonella Gray 1870c:61
Type species: Jerdonella sylhetensis [= Pangshura sylhetensis Jerdon
1870], by original monotypy.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Emia Gray 1870c:61
Type species: Emia smithii [= Batagur smithii Gray 1863g], by original
monotypy.
111
Pangshura smithii pallidipes (Moll 1987)
Pale-footed Roofed Turtle
Pangshura smithii (Gray 1863g)
Brown Roofed Turtle
Paul Crow / India [trade]
India (Bihar, Uttar Pradesh), Nepal
Kachuga smithii pallidipes Moll 1987:8, Pangshura smithii
pallidipes
Type locality: “Gandak River, Bherihari Wildlife Sanctuary, Bettiah
(West Champaran) District, Bihar” [India].
(subspecies: smithii = red, pallidipes = purple;
overlap = intergrades)
Bangladesh, India (Assam, Bihar, Punjab, Uttar Pradesh),
Nepal, Pakistan
IUCN Red List: Near Threatened (2000); Previously: Least
Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern or Near Threatened
(2011)
CITES: Appendix II, as Pangshura spp.
Pangshura sylhetensis Jerdon 1870
Assam Roofed Turtle, Sylhet Roofed Turtle
Pangshura smithii smithii (Gray 1863g)
Brown Roofed Turtle
Indraneil Das / CBFTT / Kulsi River, Kamrup District, Assam, India
Hermann Schleich / Koshi R., Nepal
Bangladesh, India (Assam, Bihar, Punjab, Uttar Pradesh),
Nepal, Pakistan
Batagur smithii Gray 1863g:253, Pangshura smithii, Clemmys smithii, Emia smithii, Kachuga smithii, Kachuga
smithii smithii, Pangshura smithii smithii
Type locality: “North-western India: Punjab; River Chenab” [Punjab, India, or Pakistan].
Bangladesh, Bhutan (?), India (Arunachal Pradesh, Assam,
Meghalaya, Mizoram, Nagaland), Myanmar (?)
CBFTT Account: Das, Sengupta, and Praschag (2010)
IUCN Red List: Endangered B1+2c (2000); Previously: Data
Deficient (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Pangshura spp.
Pangshura sylhetensis Jerdon 1870:69, Jerdonella sylhetensis, Kachuga sylhetensis
Type locality: “the stream that runs from the Terria Ghat at the foot
of the Khasi Hills” [India; now Bangladesh]. Emended to “Sylhet, a
stream at the foot of the Khasi Hills” [originally Assam, India; now
Sylhet, Bangladesh] by Boulenger (1889:58).
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Pangshura tecta (Gray 1830b)
Indian Roofed Turtle
Pangshura tentoria (Gray 1834a)
Indian Tent Turtle
Paul Crow / Bangladesh [trade]
(subspecies: tentoria = red, circumdata = purple, flaviventer = blue;
unassigned P. tentoria sensu lato = gray; overlap = intergrades;
orange dot = possibly erroneous tentoria restricted type locality)
Bangladesh, India (Andhra Pradesh, Assam, Bihar, Chhattisgarh,
Maharashtra, Manipur, Odisha, Rajasthan, Uttar Pradesh,
Uttarakhand, West Bengal), Nepal
IUCN Red List: Least Concern (2000); Previously: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
CITES: Appendix II, as Pangshura spp.
Bangladesh, India (Arunachal Pradesh, Assam, Bihar, Gujarat,
Haryana, Madhya Pradesh, Meghalaya, Odisha, Punjab,
Rajasthan, Uttar Pradesh, Uttarakhand, West Bengal), Nepal,
Pakistan
IUCN Red List: Least Concern (2000); Previously: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Near Threatened (2011)
CITES: Appendix I
Emys tectum Gray 1830b:pl.72, Emys tecta, Clemmys
(Clemmys) tecta, Clemmys tecta, Batagur (Pangshura)
tecta, Batagur tecta, Clemmys tectum, Pangshura tecta,
Pangshura tectum, Kachuga tectum, Kachuga (Pangshura) tecta, Kachuga tectum tectum, Kachuga tecta, Kachuga tecta tecta
Type locality: “India.”
Testudo dura Hamilton in Gray 1831d:23 (nomen nudum)
Type locality: “India.”
Testudo katuya Hamilton in Gray 1831d:23 (nomen nudum)
Type locality: “India.”
Testudo pangshure Hamilton in Gray 1831d:23 (nomen
nudum), Testudo pangshura
Type locality: “India.”
Testudo khagraskata Hamilton in Gray 1831d:23 (nomen
nudum)
Type locality: “India.”
Emys trigibbosa Lesson 1831a:121
Type locality: “le Gange” [India].
Emys namadicus † Theobald 1860:295 (nomen nudum) [Tertiary, Nerbudda, India], Emys nomadicus, Emys namadica
Type locality: “near Moar Domur...Nerbudda Valley” [Madhya
Pradesh, India].
Pangshura dura Gray 1869a:205
Type locality: Not designated.
Pangshura ventricosa Gray 1870c:60
Type locality: “India.” Restricted to “Assam” [India] by Boulenger
(1889:59).
Pangshura tentoria tentoria (Gray 1834a) (07:49)
Indian Tent Turtle
Peter Paul van Dijk / India [captivity, Madras Crocodile Bank Trust]
India (Andhra Pradesh, Chhattisgarh, Maharashtra, Odisha)
Emys tentoria Gray 1834a:54, Batagur (Pangshura) tentoria,
Batagur tentoria, Clemmys tentoria, Pangshura tentoria,
Pangshura tentori, Cuchoa tentoria, Kachuga tectum tentoria, Kachuga tecta tentoria, Kachuga tentoria tentoria,
Pangshura tentoria tentoria
Type locality: “Indiae Orientalis regione Dukhun” [India]. Emended
to “Deccan” [India] by Boulenger (1889:59); and restricted to
“Dhond, Poona Dist.” [Daund, Pune Dist., Maharashtra, India; in
error?] by Smith (1931:128).
Pangshura tentorium Gray 1869a:205 (nomen novum)
Emys (Pangshura) tectum intermedia Blanford 1870:339,
Kachuga intermedia, Kachuga tectum intermedia
Type locality: “Chappa and Korba in Biláspúr, on the Hasdo River,
a tributary of the upper Mahanaddi...above Sambhalpúr” [Chhattisgarh, India].
Pangshura leithii Gray 1870c:60
Type locality: “River Poonah” [Maharashtra, India; in error, trade?].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
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Pangshura tentoria circumdata (Mertens 1969)
Pink-ringed Tent Turtle
Nikhil Whitaker / Uttar Pradesh, India
India (Bihar, Rajasthan, Uttar Pradesh, Uttarakhand), Nepal
Kachuga tecta circumdata Mertens 1969a:24, Kachuga
tentoria circumdata, Pangshura tentoria circumdata
Type locality: “Meerut, Indien” [Uttar Pradesh, India].
Pangshura tentoria flaviventer Günther 1864 (07:49)
Yellow-bellied Tent Turtle
China (Fujian, Guangdong, Hong Kong, Hunan, Jiangxi)
IUCN Red List: Endangered A1d+2d (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Cistuda bealei Gray 1831d:71, Emys (Pyxidemys) bealei,
Emys bealei, Clemmys bealei, Cistudo bealei, Sacalia
bealei, Mauremys bealei, Sacalia bealei bealei
Type locality: “China.”
Emys bealii Gray 1834a:54 (nomen novum), Clemmys bealii,
Sacalia bealii, Cistudo bealii, Clemmys bealii bealii
Sacalia quadriocellata (Siebenrock 1903a) (08:24)
Four-eyed Turtle
Hermann Schleich / Koshi R., Nepal
Bangladesh, India (Bihar, Uttar Pradesh, West Bengal), Nepal
Pangshura flaviventer Günther 1864:35, Kachuga tecta flaviventer, Pangshura flaviventus, Kachuga tentoria flaviventer, Pangshura tentoria flaviventer
Type locality: “Bengal” [India]. Emended to “India” by Boulenger
(1889:59).
Cuchoa flaviventris Gray 1870c:61 (nomen novum)
John B. Iverson / No data [trade]
Sacalia Gray 1870c
Sacalia Gray 1870c:35
Type species: Sacalia bealii [= Emys bealii Gray 1834a = Cistuda
bealei Gray 1831d], by original monotypy.
Sacalia bealei (Gray 1831d)
Beale’s Eyed Turtle
John B. Iverson / No data [trade]
China (Guangdong, Guangxi, Hainan), Laos, Vietnam
IUCN Red List: Endangered A1d+2d (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Clemmys bealii quadriocellata Siebenrock 1903a:336 (senior
homonym), Clemmys quadriocellata, Clemmys quadriocellata, Clemmys bealei quadriocellata, Sacalia quadriocellata, Sacalia bealei quadriocellata, Sacalia quadriocellata quadriocellata
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Type locality: “Annam” [Vietnam]. Restricted to “Annam, PhucSon” [Vietnam] by Siebenrock (1909a:482).
Clemmys bealii quadriocellata Li 1958:235 (junior
homonym)
Type locality: “small stream of Dali village of Mt. Diaoluo,
Lingshui Co., Hainan Island, China” [in Chinese].
Clemmys beali insulensis Adler 1962:135 (nomen novum)
(08:24)
Sacalia pseudocellata Iverson and McCord 1992a:426 (07:33)
(partim, hybrid)
Type locality: “between Tungfang [19º03'N, 108º56'E] and
Kancheng [18º51'N, 108º37'E; ca. 48 km from Tungfang], western
Hainan Island, China.” [in error?]
Sacalia quadriocellata insularis Artner 2003:xviii (nomen
novum)
Siebenrockiella Lindholm 1929
Bellia Gray 1869a:197 (junior homonym, not = Bellia MilneEdwards 1848 [= Crustacea] or Bellia Bate 1851 [=
Crustacea] or Bellia Tutt 1902 [= Lepidoptera])
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Emys crassicollis Gray 1830e:8 (10:7), Clemmys (Clemmys)
crassicollis, Clemmys crassicollis, Bellia crassicollis,
Orlitia crassicollis, Siebenrockiella crassicollis
Type locality: “Sumatra” [Indonesia].
Emys nigra Blyth 1856:713 (junior homonym, not = Emys
nigra Hallowell 1854)
Type locality: “valley of the Tenasserim” [Myanmar].
Emys crassilabris Gray in Theobald 1876:10 (nomen novum),
Bellia crassilabris
Pangshura cochinchinensis Tirant 1884:159, Kachuga
cochinchinensis
Type locality: “Cochinchine” [Vietnam].
Siebenrockiella leytensis (Taylor 1920) (07:50)
Palawan Forest Turtle, Philippine Forest Turtle
Type species: Bellia crassicollis [= Emys crassicollis Gray 1830e], by
original monotypy.
Siebenrockiella Lindholm 1929:280 (nomen novum)
Type species: Siebenrockiella crassicollis [= Emys crassicollis Gray
1830e], by original designation.
Siebenrockiella (Panyaenemys) Diesmos, Parham, Stuart, and
Brown 2005:38
Type species: Siebenrockiella (Panyaenemys) leytensis [= Heosemys
leytensis Taylor 1920], by original designation.
Siebenrockiella crassicollis (Gray 1830e) (10:7)
Black Marsh Turtle
Rafe M. Brown / TCC / Palawan, Philippines
Jérôme Maran / No data [captivity]
(orange dot = erroneous type locality)
Philippines (Palawan [not Leyte])
CBFTT Account: Diesmos, Buskirk, Schoppe, Diesmos, Sy,
and Brown (2012)
IUCN Red List: Critically Endangered A2d, B1+2c (2000);
Previously: Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2015)
CITES: Appendix II
Heosemys leytensis Taylor 1920:131, Geoemyda leytensis,
Siebenrockiella leytensis, Siebenrockiella (Panyaenemys)
leytensis
Type locality: “Cabalian, southern Leyte, P. I.” [Philippines] [in
error]. Restricted to “northern Palawan in the Province of Palawan,
Philippines” by Diesmos et al. (2012:3).
Cambodia, Indonesia (Java, Kalimantan, Sumatra), Laos,
Malaysia (Peninsular, East [?]), Myanmar, Singapore,
Thailand, Vietnam
IUCN Red List: Vulnerable A1cd+2cd (2000); Previously: Least
Concern [Not Listed] (1996)
Vijayachelys Praschag, Schmidt, Fritzsch, Müller, Gemel, and Fritz 2006 (07:51)
Vijayachelys Praschag, Schmidt, Fritzsch, Müller, Gemel, and
Fritz 2006:156
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type species: Vijayachelys silvatica [= Geoemyda silvatica Henderson
1912], by original designation.
Rhinoclemmys annulata (Gray 1860b)
Brown Wood Turtle
Vijayachelys silvatica (Henderson 1912)
Cochin Forest Cane Turtle
John L. Carr / Isla Palma, Valle del Cauca, Colombia
V. Deepak / CBFTT / Anamalai Hills, Western Ghats, Tamil Nadu, India
India (Karnataka, Kerala, Tamil Nadu)
CBFTT Account: Deepak, Praschag, and Vasudevan (2014)
IUCN Red List: Endangered B1+2c (2000); Previously: Endangered (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Geoemyda silvatica Henderson 1912:217, Heosemys silvatica, Vijayachelys silvatica
Type locality: “South India...near Kavalai in the Cochin State
Forests...an elevation of about 1500 feet” [Kerala, India].
RhinoCleMMydinae Gray 1873j (12:21)
Rhinoclemmyina Gray 1873j:27
Rhinoclemminae Le and McCord 2008:763
Rhinoclemmydinae Turtle Taxonomy Working Group 2012:274
Rhinoclemmys Fitzinger 1835 (09:20, 12:21, 14:31)
Chemelys Rafinesque 1815:75 (nomen nudum)
Chemelys Rafinesque 1832:64 (nomen suppressum, ICZN
1985a)
Type species: Chemelys verrucosa [= Testudo verrucosa Suckow 1798
= subjective synonym of Testudo punctularia Daudin 1801], by
original monotypy.
Clemmys (Rhinoclemmys) Fitzinger 1835:115 (nomen conservandum, ICZN 1985a)
Type species: Geoemyda (Rhinoclemmys) dorsata [= Testudo dorsata
Schoepff 1801 = subjective synonym of Testudo punctularia
Daudin 1801], by subsequent designation as Emys dorsata sensu
Schweigger 1812 by Lindholm (1929:283).
Rhinoclemys Gray 1863c:182 (nomen novum)
Rhinoclemys (Callopsis) Gray 1863c:183
Type species: Rhinoclemys (Callopsis) annulata [= Geoclemmys annulata Gray 1860b], by original monotypy.
(orange dot = possible trade)
Colombia (Antioquia, Cauca, Chocó, Córdoba, Nariño, Valle
del Cauca), Costa Rica, Ecuador, Honduras, Nicaragua,
Panama
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Data Deficient (South America regional)
(2011)
Geoclemmys annulata Gray 1860b:231, Clemmys annulata,
Rhinoclemys annulata, Rhinoclemys (Callopsis) annulata,
Chelopus annulatus, Rhinoclemmys annulata, Nicoria
annulata, Geoemyda annulata, Callopsis annulata
Type locality: “Esmeraldas, ...western coast of Ecuador.”
Chelopus gabbii Cope 1876:153, Emys gabbii, Nicoria gabbii, Geoemyda gabbii, Rhinoclemmys gabbii
Type locality: “Costa Rica.” Restricted to “Sipurio, Costa Rica” by
Dunn (1930:32).
Testudo mercatoria Vaillant 1911:47 (nomen nudum)
Type locality: “Amérique centrale.”
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Rhinoclemmys areolata (Duméril and Bibron in Duméril and Duméril
1851)
Furrowed Wood Turtle
John B. Iverson / CBFTT / nr. Paamul, Quintana Roo, Mexico
Colombia (Norte de Santander), Venezuela (Mérida, Táchira,
Trujillo, Zulia)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Vulnerable (2011)
Geoemyda punctularia diademata Mertens 1954:4, Callopsis
punctularia diademata, Rhinoclemmys punctularia diademata, Rhinoclemmys diademata
Type locality: “Maracay, Venezuela.” Considered in error by
Pritchard (1979:182); and considered in error for “Maracaibo”
[Venezuela] by Pritchard and Trebbau (1984:172).
Rhinoclemmys funerea (Cope 1876)
Black Wood Turtle
Belize, Guatemala, Honduras, Mexico (Campeche, Chiapas,
Quintana Roo, Tabasco, Veracruz, Yucatán)
CBFTT Account: Vogt, Platt, and Rainwater (2009)
IUCN Red List: Near Threatened (2007); Previously: Least
Concern [Not Listed] (1996)
Emys areolata Duméril and Bibron in Duméril and Duméril
1851:10, Malaclemmys concentrica areolata, Clemmys
areolata, Malaclemys concentrica areolata, Chelopus
areolatus, Nicoria punctularia areolata, Geoemyda
punctularia areolata, Geoemyda areolata, Rhinoclemmys
areolata, Callopsis areolata
Manuel Merchán Fornelino / Costa Rica
Type locality: “Province du Petén (Amér. centr.).” Restricted to “La
Libertad, El Petén, Guatemala” by Smith and Taylor (1950a:318;
1950b:30); and to “Flores...El Petén, La Libertad...Guatemala” by
Dunn and Stuart (1951:60).
Rhinoclemmys diademata (Mertens 1954)
Maracaibo Wood Turtle
(orange dot = possible trade)
Costa Rica, Honduras, Nicaragua, Panama
IUCN Red List: Near Threatened (1996)
Chelopus funereus Cope 1876:154, Emys funerea, Geoemyda
funerea, Geoemyda punctularia funerea, Rhinoclemmys
funerea, Callopsis funerea
Type locality: “Costa Rica...Limon.”
Carlos A. Galvis-Rizo / Colombia [captivity]
Geoemyda costaricensis Kanberg 1930:162
Type locality: “Costa Rica.”
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Rhinoclemmys melanosterna (Gray 1861b) (14:31)
Colombian Wood Turtle
Uwe Fritz / El Silencio, Antioquia, Colombia
Colombia (Cauca, Chocó, Nariño, Valle del Cauca), Ecuador
CBFTT Account: Carr and Giraldo (2009)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Near Threatened (2011)
Nicoria nasuta Boulenger 1902a:53, Geoemyda nasuta,
Geoemyda punctularia nasuta, Callopsis punctularia nasuta, Rhinoclemmys punctularia nasuta, Callopsis nasuta,
Rhinoclemmys nasuta
Type locality: “North-western Ecuador...Bulún, 160 feet, and from
the Rio Durango, 350 feet.” Emended to “San Francisco de Pulún,
Río Bogota” [Colombia] by Carr and Almendariz (1990:92).
Rhinoclemmys pulcherrima (Gray 1856b)
Painted Wood Turtle
Colombia (Antioquia, Atlántico, Bolívar, Boyacá, Caldas, Cauca, Cesar, Chocó, Córdoba, Cundinamarca, La Guajira,
Magdalena, Nariño, Santander, Sucre, Valle del Cauca),
Ecuador, Panama
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Geoclemmys melanosterna Gray 1861b:205, Clemmys
melanosterna, Rhinoclemmys melanosterna, Rhinoclemys
melanosterna, Nicoria punctularia melanosternum, Geoemyda punctularia melanosternum, Geoemyda punctularia melanosterna, Callopsis punctularia melanosterna,
Rhinoclemmys punctularia melanosterna
Type locality: “Gulf of Darien: Cherunha” [Colombia]. Restricted to
“River Buonaventura..[and]..Chirambira, Gulf of Darien” [Colombia] by Boulenger (1889:124); to “Chirambira bei Buenaventura,
Columbien” [Colombia] by Mertens and Wermuth (1955: 352),
and to “Punta Chirambira en el Delta del Río San Juan...al norte
de Buenaventura en la Costa del Pacífico, Chocó” [Colombia] by
Medem (1958:21).
Rhinoclemmys nasuta (Boulenger 1902a)
Large-nosed Wood Turtle
(subspecies: pulcherrima = red, incisa = purple,
manni = orange, rogerbarbouri = pink)
Costa Rica, El Salvador, Guatemala, Honduras, Mexico (Chiapas, Colima, Guerrero, Jalisco, Nayarit, Oaxaca, Sinaloa,
Sonora), Nicaragua
IUCN Red List: Least Concern [Not Listed] (1996)
José Vicente Rueda-Almonacid / CBFTT / Isla Palma, Valle del Cauca, Colombia
117
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Rhinoclemmys pulcherrima pulcherrima (Gray 1856b)
Guerrero Wood Turtle
Rhinoclemmys pulcherrima manni (Dunn 1930)
Central American Wood Turtle
John B. Iverson / nr. Canas, Guanacaste Prov., Costa Rica
John L. Carr / Mexico [captivity]
Mexico (Guerrero, Oaxaca)
Emys pulcherrimus Gray 1856b:25, Clemmys pulcherrima,
Callichelys pulcherrima, Emys pulcherrima, Rhinoclemmys pulcherrima, Chelopus pulcherrima, Pseudemys
pulcherrima, Chelopus pulcherrimus, Nicoria punctularia
pulcherrima, Geoemyda punctularia pulcherrima, Geoemyda pulcherrima pulcherrima, Rhinoclemmys pulcherrima pulcherrima, Callopsis pulcherrima pulcherrima
Costa Rica, Nicaragua
Geoemyda manni Dunn 1930:33, Geoemyda pulcherrima
manni, Geoemyda punctularia manni, Callopsis pulcherrima manni, Rhinoclemmys pulcherrima manni
Type locality: “San Jose, Costa Rica.”
Rhinoclemmys pulcherrima rogerbarbouri (Ernst 1978)
Western Mexican Wood Turtle
Type locality: “Mexico.” Restricted to “Presidio de Mazatlán, Sinaloa,
Mexico” by Smith and Taylor (1950a:343, 1950b:30); and to “vicinity of San Marcos, Guerrero, Mexico” by Ernst (1978:125).
Rhinoclemmys pulcherrima incisa (Bocourt 1868)
Incised Wood Turtle
John B. Iverson / nr. Colima, Colima, Mexico
Mexico (Colima, Jalisco, Nayarit, Sinaloa, Sonora)
Callopsis pulcherrima rogerbarbouri Ernst 1978:127, Rhinoclemmys pulcherrima rogerbarbouri
Type locality: “Guirocoba, Sonora, Mexico.”
John B. Iverson / Honduras
Rhinoclemmys punctularia (Daudin 1801) (08:12, 09:28)
Spot-legged Turtle
El Salvador, Guatemala, Honduras, Nicaragua, Mexico (Chiapas, Oaxaca)
Emys incisa Bocourt 1868:121, Chelopus incisa, Chelopus
incisus, Glyptemys incisa, Nicoria punctularia incisa,
Clemmys incisa, Chrysemys incisa, Geoemyda punctularia
incisa, Rhinoclemmys incisa, Geoemyda pulcherrima incisa, Rhinoclemmys pulcherrima incisa, Callopsis pulcherrima incisa
Type locality: “l’Union, un des ports du Salvador” [El Salvador].
Restricted to “la Union, un des ports du Salvador sur le Pacifique;
elle a été trouvée sur la montagne de Conchavoua” [El Salvador] by
Duméril and Bocourt (1870:12).
Rhinoclemmys frontalis Gray 1873a:144
Type locality: “Tropical America.”
Rhinoclemmys bocourti Gray 1873f:111
Type locality: “Central America.”
(subspecies: punctularia = red, flammigera = purple)
Brazil (Amapá, Amazonas, Bahia, Maranhão, Pará, Piauí,
Roraima, Tocantins), French Guiana, Guyana, Suriname,
Trinidad and Tobago, Venezuela (Amazonas, Bolívar,
Delta Amacuro, Monagas)
Introduced: Brazil (Rio de Janeiro)
IUCN Red List: Least Concern [Not Listed] (1996)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
TFTSG Draft Red List: Least Concern (2011)
Rhinoclemmys punctularia punctularia (Daudin 1801) (08:12, 09:28)
Eastern Spot-legged Turtle
119
Type locality: Not designated. Restricted to “Dutch Guiana” [Suriname] by Boulenger (1889:124).
Rhinoclemmys ventricosa Gray 1873a:145, Rhinoclemys
ventricosa
Type locality: “Tropical America.” Restricted to “Dutch Guiana”
[Suriname] by Boulenger (1889:124).
Rhinoclemmys punctularia flammigera Paolillo 1985 (08:25, 09:28)
Upper Orinoco Spot-legged Turtle
Venezuela (Amazonas)
Rhinoclemmys punctularia flammigera Paolillo 1985:294,
Rhinoclemmys flammigera
Type locality: “Caño Maica, 10 km SE of Carmelitas, Territorio
Federal Amazonas, Venezuela (4º4' N, 66º31' W).”
Rhinoclemmys rubida (Cope 1870a)
Mexican Spotted Wood Turtle
Jérôme Maran / French Guiana
Brazil (Amapá, Amazonas, Maranhão, Pará, Piauí, Roraima,
Tocantins), French Guiana, Guyana, Suriname, Trinidad and Tobago, Venezuela (Bolívar, Delta Amacuro,
Monagas)
Introduced: Brazil (Rio de Janeiro)
Testudo scabra Linnaeus 1758:198 (nomen oblitum and senior homonym, not = Testudo scabra Retzius in Schoepff
1792 or Testudo scabra Latreille in Sonnini and Latreille
1801) (08:12), Emys scabra, Rhinoclemmys scabra , Rhinoclemys scabra
Type locality: “Indiis” [West Indies]. Restricted to “India orientali,
Carolina” [in error] by Linnaeus (1766:351); and to “Cayenne,
French Guiana” by Rhodin and Carr (2009:11).
Testudo verrucosa Walbaum 1782:116 (unavailable name)
Type locality: Not designated.
Testudo verrucosa Suckow 1798:40 (junior homonym, not
= Testudo (ferox) verrucosa Schoepff 1795), Chemelys
verrucosa
Type locality: Not known.
Testudo punctularia Daudin 1801:249[“349”] (nomen
conservandum, ICZN 1963), Emys punctularia, Chersine
punctularia, Clemmys (Clemmys) punctularia, Clemmys
punctularia, Chelopus punctularius, Nicoria punctularia,
Geoemyda punctularia, Geoemyda punctularia punctularia, Rhinoclemmys punctularia, Rhinoclemmys punctularia
punctularia, Callopsis punctularia punctularia
(subspecies: rubida = red, perixantha = purple)
Mexico (Chiapas, Colima, Guerrero, Jalisco, Michoacán,
Oaxaca)
IUCN Red List: Near Threatened (2007); Previously: Vulnerable (1996)
Rhinoclemmys rubida rubida (Cope 1870a)
Oaxaca Wood Turtle
Type locality: “Amérique méridionale...sur-tout...la Guiane..[&]..
Cayenne” [French Guiana].
Testudo dorsata Schoepff 1801:136 (nomen suppressum,
ICZN 1963), Emys dorsata, Clemmys dorsata
Type locality: Not known.
Testudo alacacca Stedman in Schweigger 1812:429 (nomen
nudum)
Type locality: “Surinam.”
Emys dorsualis Spix 1824:11, Emys dorsalis
Type locality: “juxta flumen Solimoens” [Amazonas, Brazil].
Rhinoclemys bellii Gray 1863c:183, Rhinoclemmys bellii,
Rhinoclemmys scabra bellii
Type locality: “Tropical America.”
Geoclemys callocephalus Gray 1863h:254, Clemmys
callocephala, Geoclemmys callocephala, Geoclemmys
callocephalus, Rhinoclemmys callocephala, Rhinoclemys
callocephala
Type locality: Not known. Restricted to “S. America” by Boulenger
(1889:124).
Rhinoclemmys lunata Gray 1873a:144, Geoemyda punctularia lunata, Callopsis punctularia lunata, Rhinoclemys
lunata, Rhinoclemmys punctularia lunata
Michael Redmer / Mexico [captivity]
Mexico (Chiapas, Oaxaca)
Chelopus rubidus Cope 1870a:148, Geoclemmys rubida,
Emys rubida, Nicoria rubida, Clemmys rubida, Geoemyda
rubida, Geoemyda rubida rubida, Rhinoclemmys rubida,
Rhinoclemmys rubida rubida, Callopsis rubida, Callopsis
rubida rubida, Chelopus rubidus rubidus
Type locality: “Tuchitan Tehuantepec, Mexico” [Juchitán, Oaxaca].
Rhinoclemmys mexicana Gray 1870b:659, Chelopus mexicana, Chelopus mexicanus, Emys mexicana
Type locality: “Mexico; San Juan del Rio” [Oaxaca].
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Rhinoclemmys rubida perixantha (Mosimann and Rabb 1953)
Colima Wood Turtle
Aldabrachelys gigantea (Schweigger 1812) (07:54, 07:55, 08:13, 09:29, 09:30, 11:10,
12:28, 14:32) (56)
Aldabra Giant Tortoise
John B. Iverson / nr. Colima, Colima, Mexico
Mexico (Colima, Guerrero, Jalisco, Michoacán)
Geoemyda rubida perixantha Mosimann and Rabb 1953:1,
Rhinoclemmys rubida perixantha, Callopsis rubida perixantha, Chelopus rubidus perixanthus
Type locality: “8 kilometers south of Tecoman, Colima, Mexico.”
TesTudinidae Batsch 1788
Testudines Batsch 1788:437
Testudia Rafinesque 1814:66
Tortudines Schmid 1819:10
Testudinidae Gray 1825:210
Tylopodae Wagler 1828:861
Dysmydae Ritgen 1828:270
Tylopodes Burmeister 1837:732
Baenodactyli Mayer 1849:198
Testudinina Agassiz 1857a:356
Testudinidi Portis 1890:12
(subspecies: gigantea = yellow dot, daudinii = red dot,
arnoldi and hololissa = orange dots [probable])
Seychelles (Aldabra, Granitic Islands)
IUCN Red List: Vulnerable D2 (1996)
CITES: Appendix II, as Testudinidae spp.
Aldabrachelys gigantea gigantea (Schweigger 1812) (07:54, 08:13, 09:29,
09:30, 11:10) (57)
Aldabra Giant Tortoise
Aldabrachelys Loveridge and Williams 1957 (07:52, 07:53, 08:13,
09:29, 09:30, 12:28, 14:32) (55)
Testudo (Megalochelys) Fitzinger 1843:29 (junior homonym,
not = Megalochelys Falconer and Cautley 1837 [= Testudinidae †])
Type species: Testudo (Megalochelys) gigantea [= Testudo gigantea
Schweigger 1812], by original designation.
Geochelone (Aldabrachelys) Loveridge and Williams 1957:225
Type species: Geochelone (Aldabrachelys) gigantea [= Testudo gigantea Schweigger 1812], by original designation.
Dipsochelys Bour 1982a:117
Type species: Dipsochelys elephantina [= Testudo elephantina Duméril
and Bibron 1835 = subjective synonym of Testudo gigantea Schweigger 1812], by original designation.
Bernard Devaux / Aldabra, Seychelles
Seychelles (Aldabra)
Introduced: Mauritius (Aigrettes, Rodrigues, Round), Seychelles (Assomption, Alphonse, Astove, Cerf, Cosmolédo,
Cousin, Cousine, Curieuse, D’Arros, Desroches, Farquhar,
Frégate, Grande Soeur, Moyenne, North, Rémire, Silhouette), Tanzania (Zanzibar [Changuu])
Testudo gigantea Schweigger 1812:327 (08:13, 09:29, 14:32) (partim, misidentified type) (nomen conservandum, ICZN
2013b), Geochelone (Chelonoidis) gigantea, Geochelone
gigantea, Testudo gigantea gigantea, Geochelone gigantea
gigantea, Aldabrachelys gigantea, Megalochelys gigantea,
Megalochelys gigantea gigantea, Dipsochelys gigantea,
Aldabrachelys gigantea gigantea, Dipsochelys giganteus
Type locality: “Brasilia” [Brazil]. Restricted to “Dune Patates, South
Island, Aldabra Atoll, Republic of Seychelles” by neotype designation by Frazier (2006:278).
Testudo dussumieri Schlegel in Gray 1830e:3 (nomen nudum)
Testudo dussumieri Gray 1831d:9 (09:29, 14:32) (nomen suppressum, ICZN 2013b), Dipsochelys dussumieri, Geochelone
dussumieri, Aldabrachelys dussumieri, Dipsochelys
dussumieri dussumieri
Type locality: “Insula Mauritiana, Insula Aldebra” [Mauritius and
Aldabra, Seychelles]. Restricted to “Aldabra (N.W. of Madagascar)”
by Hubrecht (1881:44); and to “Insula Aldebra” [Aldabra, Seychelles]
by Bour (1985:291).
Testudo elephantina Duméril and Bibron 1835:110, Testudo
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
gigantea elephantina, Geochelone elephantina, Geochelone gigantea elephantina, Dipsochelys elephantina,
Aldabrachelys elephantina, Dipsochelys elephantina
elephantina
121
Aldabrachelys gigantea daudinii (Duméril and Bibron 1835) (09:30)
(Extinct, ca. 1850)
Daudin’s Giant Tortoise
Type locality: “la plupart des îles qui sont situées dans le Canal
de Mosambique, telle que Anjouan, Aldebra, les Comores, d’où
on l’apporte fréquemment à Bourbon et à Maurice.” Restricted
to “Aldabra” [Seychelles] by Boulenger (1889:168); to “North
Aldabra” by Rothschild (1915:418); and to “Ile Malabar, Aldabra
Atoll, Republic of Seychelles” by Bour (1985:291).
Testudo ponderosa Günther 1877:35, Aldabrachelys
ponderosa
Type locality: Not known. Restricted to “Aldabra” [Seychelles] by
Bour (1985:292).
Testudo sumeirei Sauzier 1892:398, Geochelone sumeirei,
Dipsochelys sumeirei, Megalochelys sumeirei, Dipsochelys
elephantina sumeirei, Aldabrachelys sumeirei, Dipsochelys
dussumieri sumeirei
Type locality: “probable...de Maurice..[&]..la Réunion..[&]..des
Séchelles.” Restricted to “Seychelles” by Rothschild (1899:360);
to “Seychelles Islands?” by Auffenberg (1974:144), and to “central
Seychelles” by Bour (1985:292).
Testudo goufféi Rothschild 1906:753, Testudo gouffei,
Geochelone gouffei, Geochelone (Aldabrachelys) gouffei,
Geochelone (Aldabrachelys) gigantea gouffei, Geochelone
gigantea gouffei, Megalochelys gouffei, Aldabrachelys
gouffei
Roger Bour / Seychelles [MNHN, Paris]
Seychelles (Mahé? [extinct])
Testudo daudinii Duméril and Bibron 1835:123, Testudo
gigantea daudinii, Geochelone gigantea daudinii, Dipsochelys daudinii, Geochelone daudinii, Aldabrachelys
daudinii, Aldabrachelys gigantea daudinii, Dipsochelys
dussumieri daudinii
Type locality: “Indes orientales.” Restricted to “les îles Seychelles
granitiques” by Bour (1985:58).
Aldabrachelys gigantea hololissa (Günther 1877) (09:30, 11:10)
Seychelles Giant Tortoise
Type locality: “Therese Island, St. Anne’s Channel, Seychelles
Islands.” Emended to “more likely...from Juan de Novo or Farquhar
Island” [Seychelles] by Rothschild (1915:427).
Aldabrachelys gigantea arnoldi (Bour 1982a) (07:54, 09:30, 11:10)
Arnold’s Giant Tortoise
Justin Gerlach / CBFTT / Seychelles
John Pemberton / CBFTT / Seychelles
Seychelles (Mahé?, North? [all extirpated])
Introduced: Seychelles (North, Cousine, Frégate, Silhouette)
CBFTT Account: Gerlach (2009)
Dipsochelys arnoldi Bour 1982a:121, Testudo arnoldi,
Aldabrachelys arnoldi, Aldabrachelys gigantea arnoldi,
Dipsochelys dussumieri arnoldi
Type locality: “Seychelles...probablement...les îles granitiques.”
Seychelles (Cerf?, Cousine?, Frégate?, Mahé?, Praslin?,
Round?, Silhouette? [all extirpated])
Introduced: Seychelles (Cerf, Cousine, Round)
CBFTT Account: Gerlach (2011)
Testudo hololissa Günther 1877:39, Dipsochelys hololissa,
Geochelone hololissa, Aldabrachelys hololissa, Aldabrachelys gigantea hololissa, Dipsochelys dussumieri
hololissa
Type locality: “Seychelle Islands...[probably] originally imported
from the Aldabra group.” Restricted to “des îles Seychelles” by
Sauzier (1899:142).
Dipsochelys resurrecta Gerlach and Canning 1996:133 (nomen nudum)
Astrochelys Gray 1873j (07:52)
Testudo (Astrochelys) Gray 1873j:4
Type species: Testudo (Astrochelys) radiata [= Testudo radiata Shaw
1802], by original monotypy.
Asterochelys Gray 1874:724 (nomen novum)
Angonoka Le, Raxworthy, McCord, and Mertz 2006:528 (09:31)
Type species: Angonoka yniphora [= Testudo yniphora Vaillant 1885a],
by original designation.
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Astrochelys radiata (Shaw 1802)
Radiated Tortoise, Sokake
Astrochelys yniphora (Vaillant 1885a) (07:56, 09:31)
Ploughshare Tortoise, Angonoka
Anders G.J. Rhodin / TCC / CRM 6 / Cap Sainte Marie Special Reserve, Madagascar
Anders G.J. Rhodin / CCB / Baly Bay National Park, Madagascar
Madagascar
Introduced: Mauritius (Rodrigues, Round), Réunion
IUCN Red List: Critically Endangered A4d, E (2008); Previously: Vulnerable (1996)
CITES: Appendix I
Testudo coui Daudin 1801:271 (nomen oblitum)
Type locality: Not known.
Testudo radiata Shaw 1802:22, Psammobates radiatus,
Asterochelys radiata, Testudo radiata radiata, Geochelone
radiata, Astrochelys radiata
Type locality: “Madagascar.” Restricted to “Soalara (Baie de SaintAugustin), sud-ouest de Madagascar” by Bour (1979:152).
Testudo madagascariensis Schweigger 1812:457 (nomen
nudum)
Type locality: “Madagascar.”
Testudo desertorum Grandidier 1869:257
Type locality: “Madagascar.”
Testudo hypselonota Bourret 1941b:9
Type locality: “provenant d’un Chinois de Cholon qui l’avait...
achetée au marché...il n’est pas certain qu’elle ait été trouvée en
Cochinchine” [Vietnam] [in error, trade specimen]. Restricted to
“Cholon?...Indochina” by Wermuth and Mertens (1961:213) [in error]. Shown to be identical to Testudo radiata from Madagascar by
Auffenberg (1963:465); type locality restricted to “durch den Tierhandel...von Madagascar nach Indochina” by Wermuth (1965:285).
(orange dot = trade)
Madagascar
Introduced: Mauritius (Rodrigues)
IUCN Red List: Critically Endangered A4ad, B2ab(v), C1, E
(2008); Previously: Endangered (1996)
CITES: Appendix I
Testudo yniphora Vaillant 1885a:441, Testudo radiata
yniphora, Asterochelys yniphora, Geochelone yniphora,
Geochelone (Asterochelys) yniphora, Astrochelys yniphora, Angonoka yniphora
Type locality: “un îlot situé au nord-nord-est de...grande
Comore...d’une localité située vers Aldabra” [in error]. Restricted to
“cap d’Amparafaka (Baie de Baly), nord-ouest de Madagascar” by
Bour (1979:152).
Testudo hyniphora Vaillant in Vaillant and Grandidier
1910:40 (nomen novum)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Centrochelys Gray 1872c (07:52)
Peltastes (Centrochelys) Gray 1872c:5
Type species: Peltastes (Centrochelys) sulcatus [= Testudo sulcata
Miller 1779], by original monotypy.
Centrochelys sulcata (Miller 1779) (12:29)
African Spurred Tortoise
1854b = unidentified taxon of Chelonoidis nigra species complex],
by original monotypy.
Testudo (Pampatestudo) Lindholm 1929:285 (nomen novum)
Type species: Testudo (Pampatestudo) chilensis [= Testudo (Gopher)
chilensis Gray 1870a], by original monotypy.
Testudo (Monachelys) Williams 1952:547
Type species: Testudo (Monachelys) monensis † Williams 1952, by
original designation.
Darwintestudo Antenbrink-Vetter and Vetter 1998:4
Type species: Darwintestudo hoodensis [= Testudo hoodensis Van
Denburgh 1907], by original designation.
Chelonoidis carbonarius (Spix 1824) (10:19, 10:20, 14:33) (58)
Red-footed Tortoise
Luca Luiselli and Fabio Petrozzi / Arly, Burkina Faso
Cassiano Zaparoli / Fazenda Caiman, Mato Grosso do Sul, Brazil
(orange dots = probable trade)
Benin, Burkina Faso, Cameroon, Central African Republic,
Chad, Djibouti (?), Eritrea, Ethiopia, Mali, Mauritania,
Niger, Nigeria, Saudi Arabia (?), Senegal, Somalia (?),
Sudan, Yemen (?)
IUCN Red List: Vulnerable A1cd (1996)
TFTSG Draft Red List: Endangered (2013)
CITES: Appendix II, as Testudinidae spp.
Testudo sulcata Miller 1779:pl.26,p.54, Geochelone (Geochelone) sulcata, Geochelone sulcata, Peltastes sulcatus,
Centrochelys sulcatus, Centrochelys sulcata
Type locality: “India occidentali” [in error]. Restricted to “West
Indies” [in error] by Miller (1796:54).
Testudo calcarata Schneider 1784:317 (nomen novum),
Chersine calcarata
Type locality: “Westindien.” [in error].
Testudo radiata senegalensis Gray 1831d:11, Geochelone
senegalensis, Geochelone sulcata senegalensis
Type locality: “Senegal.”
Geochelone sulcata sudanensis Ballasina, Vandepitte, Mochi,
and Fenwick 2006:111 (nomen nudum)
Chelonoidis Fitzinger 1835 (07:52) (58)
Geochelone (Chelonoidis) Fitzinger 1835:112
Type species: Testudo (Chelonoidis) boiei [= Testudo boiei Wagler
1830a = subjective synonym of Testudo carbonaria Spix 1824], by
subsequent designation by Fitzinger (1843:29).
Testudo (Gopher) Gray 1870a:190
Type species: Testudo (Gopher) chilensis Gray 1870a, by original
monotypy.
Elephantopus Gray 1874:724 (junior homonym, not = Elephantopus Agassiz 1846 [= Siphonophora])
Type species: Elephantopus planiceps [= Testudo planiceps Gray
(orange dots = introduced or trade)
Argentina (Formosa), Bolivia (Beni, Cochabamba, La Paz,
Pando, Santa Cruz), Brazil (Alagoas, Amazonas, Bahia,
Maranhão, Mato Grosso, Mato Grosso do Sul, Pará,
Pernambuco, Piauí, Rio de Janeiro, Rondônia, Roraima,
Sergipe), Colombia (Antioquia, Arauca, Atlántico, Bolívar,
Boyacá, Caldas, Caquetá, Casanare, Cauca, Cesar, Chocó,
Córdoba, Cundinamarca, Guainía, La Guajira, Magdalena,
Meta, Santander, Sucre, Tolima, Vichada), French Guiana,
Guyana, Panama, Paraguay, Suriname, Venezuela (Apure,
Barinas, Bolívar, Carabobo, Cojedes, Falcón, Guárico,
Mérida, Miranda, Portuguesa, Sucre, Yaracuy, Zulia)
Introduced (modern or prehistoric?): Anguilla, Antigua, Barbuda, Barbados, Brazil (Rio de Janeiro), British Virgin
Islands, Colombia (Providencia), Dominica, Grenada,
Guadeloupe, Martinique, Montserrat, Netherlands Antilles,
Nicaragua (Maíz Grande), Saint-Barthélemy, Saint Kitts
and Nevis, Saint Lucia, Saint Martin, Saint Vincent and the
Grenadines, Trinidad, US Virgin Islands, Venezuela (Isla
Margarita, Los Tostigos)
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IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II, as Testudinidae spp.
Testudo carbonaria Spix 1824:22, Testudo tabulata carbonaria, Geochelone carbonaria, Chelonoidis carbonaria,
Geochelone carbonaria carbonaria, Chelonoidis carbonaria carbonaria, Chelonoidis carbonarius
petersi
Type locality: “Kishka, La Banda, Santiago del Estero” [Argentina].
Chelonoidis denticulatus (Linnaeus 1766) (10:19) (58)
Yellow-footed Tortoise
Type locality: “flumen Amazonum” [Pará, Brazil].
Testudo boiei Wagler 1830a:pl.XIII (14:33), Geochelone (Chelonoidis) boiei, Geochelone boiei, Chelonoidis boiei
Type locality: Not known.
Testudo hercules truncata Gray 1830e:3 (10:7, 10:20)
Type locality: “South America.”
Chelonoidis chilensis (Gray 1870a) (07:57, 10:21, 12:30) (59)
Chaco Tortoise, Pampas Tortoise
Carlos Alberto Jimenez / Madre de Dios, Peru
Maurice Rodrigues / TTN / Monte Desert, nr. Neuquén, Neuquén, Argentina
(red dots = possibe extirpated or trade;
orange dots = probable introduced or trade)
Argentina (Buenos Aires, Catamarca, Chaco, Córdoba,
Formosa, La Pampa, La Rioja, Mendoza, Neuquén, Rio
Negro, Salta, San Juan, San Luis, Santa Fe, Santiago del
Estero, Tucumán), Bolivia (Santa Cruz, Tarija), Paraguay
IUCN Red List: Vulnerable A1cd (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II, as Testudinidae spp.
Testudo (Gopher) chilensis Gray 1870a:190, Testudo chilensis, Geochelone chilensis, Geochelone chilensis chilensis,
Chelonoidis chilensis, Chelonoidis chilensis chilensis
Type locality: “Chili” [Chile; in error], see Sclater (1870:470).
Emended to “Chili...N. Patagonia...Mendoza and the Pampas..
[&]..Monte Video and Buenos Ayres” by Gray (1870d:707); and
restricted to “Mendoza” [Argentina] by Boulenger (1889:159).
Testudo argentina Sclater 1870:471 (nomen novum)
Geochelone donosobarrosi Freiberg 1973:83 (12:30), Geochelone chilensis donosobarrosi, Chelonoidis chilensis
donosobarrosi, Chelonoidis donosobarrosi
Type locality: “San Antonio, Rio Negro” [Argentina].
Geochelone petersi Freiberg 1973:86 (12:30), Chelonoidis
chilensis petersi, Chelonoidis petersi, Geochelone chilensis
Bolivia (Beni, La Paz, Pando, Santa Cruz), Brazil (Acre,
Amapá, Amazonas, Bahia [?, extirpated?], Espírito Santo
[?, extirpated?], Maranhão, Mato Grosso, Mato Grosso do
Sul, Pará, Rio de Janeiro [?, extirpated?], Roraima), Colombia (Amazonas, Arauca, Caquetá, Casanare, Guainía,
Guaviare, Meta, Putumayo, Vaupés, Vichada), Ecuador,
French Guiana, Guyana, Peru (Cusco, Loreto, Madre de
Dios, Pasco, Ucayali), Suriname, Trinidad, Venezuela
(Amazonas, Bolívar, Delta Amacuro, Monagas)
Introduced: Guadeloupe
IUCN Red List: Vulnerable A1cd+2cd (1996)
TFTSG Draft Red List: Near Threatened (2011)
CITES: Appendix II, as Testudinidae spp.
Testudo denticulata Linnaeus 1766:352, Chersine denticulata, Geochelone (Geochelone) denticulata, Geochelone denticulata, Chelonoidis denticulata, Chelonoidis
denticulatus
Type locality: “Virginia” [USA] [in error].
Testudo tabulata Walbaum 1782:122 (unavailable name)
Type locality: Not designated.
Testudo tessellata Stobaeus in Schneider 1792:262, Chersine
tessellata
Testudo stobaeana Gmelin in Schoepff 1793:48 (nomen
nudum)
Testudo tabulata Walbaum in Schoepff 1793:56, Chersine
tabulata, Geochelone (Chelonoidis) tabulata, Geochelone
tabulata, Chelonoidis tabulata
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “Africa australi?” [in error].
Testudo gigantea Schweigger 1812:327 (08:13, 09:29) (partim,
misidentified type)
Type locality: “Brasilia” [Brazil]. Restricted to “Dune Patates, South
Island, Aldabra Atoll, Republic of Seychelles” by neotype designation by Frazier (2006:278).
Chelonoidis abingdonii (Günther 1877) (09:34, 12:31)
(Extinct, 2012)
Pinta Giant Tortoise, Abingdon Island Giant Tortoise
Testudo terrestris brasiliensis Seba in Schweigger 1812:445
(nomen nudum), Testudo brasiliensis
Type locality: Not designated. [Brazil].
Testudo terrestris americana Stobaeus in Schweigger
1812:445
Type locality: Not designated.
Testudo tabulata cayennensis Schweigger 1812:445 (nomen
nudum), Testudo terrestris cayennensis
Type locality: Not designated. [French Guiana].
Testudo terrestris surinamensis Stedman in Schweigger
1812:445 (nomen nudum)
Type locality: Not designated. [Suriname].
Testudo hercules Spix 1824:20
Type locality: “sylvis ad flumen Solimoens” [Amazonas, Brazil].
Testudo sculpta Spix 1824:21
Type locality: “sylvis juxta flumen Amazonum” [Pará, Brazil].
Testudo cagado Spix 1824:23
Type locality: “campis et nemoribus campestribus Bahiae” [Bahia,
Brazil].
Anders G.J. Rhodin / TCC / Pinta [Lonesome George, CDRS, Santa Cruz]
Testudo planata Gmelin in Gray 1831d:9 (nomen nudum)
Testudo foveolata Schinz 1833:40 (nomen nudum)
Chelonoidis niger species complex (09:32, 12:31) (58, 60)
Galapagos Giant Tortoises
Ecuador (Galápagos: Española [Hood]; Fernandina [Narborough] [possibly extinct]; Floreana [Charles] [extinct];
Isabela [Albemarle]; Pinta [Abingdon] [extinct]; Pinzón
[Duncan]; San Cristóbal [Chatham]; Santa Cruz [Indefatigable]; Santiago [San Salvador] [James]
Ecuador (Galápagos: Pinta [Abingdon] [extinct])
Introduced: Ecuador (Galápagos: Isabela [Albemarle]) (hybrids
with C. becki)
IUCN Red List: Extinct (2016); Previously: Extinct in the Wild
(1996)
CITES: Appendix I, as Chelonoidis nigra
Testudo ephippium Günther 1874:422 (partim, nomen nudum)
Type locality: “Charles Island” [Floreana, Galápagos, Ecuador] [in
error].
Testudo ephippium Günther 1875a:271 (partim, misidentified
type)
Type locality: “Charles Island” [Floreana, Galápagos, Ecuador] [in
error]. Restricted to “Duncan” [Pinzón, Galápagos, Ecuador] by
Van Denburgh (1914:259) [in error].
Testudo abingdonii Günther 1877:85, Testudo elephantopus
abingdonii, Geochelone abingdonii, Geochelone elephantopus abingdonii, Chelonoidis abingdonii, Chelonoidis
elephantopus abingdonii, Geochelone (Chelonoidis) nigra
abingdonii, Geochelone nigra abingdonii, Chelonoidis
nigra abingdonii, Chelonoidis elephantopus abingdonii
Type locality: “Abingdon Island” [Pinta, Galápagos, Ecuador].
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Chelonoidis becki (Rothschild 1901) (12:31) (60)
Volcán Wolf Giant Tortoise, Wolf Volcano Giant Tortoise
Chelonoidis chathamensis (Van Denburgh 1907) (07:59, 12:31) (61)
San Cristóbal Giant Tortoise, Chatham Island Giant Tortoise
Washington Tapia / San Cristóbal
Paul M. Gibbons / Volcán Wolf, Isabela / non-hybrid
Ecuador (Galápagos: Isabela [Albemarle])
Invasives: Hybrids with Chelonoidis abingdonii and C. niger
IUCN Red List: Vulnerable D1+2 (1996), as Chelonoidis nigra
becki
TFTSG Draft Red List: Vulnerable (2012)
CITES: Appendix I, as Chelonoidis nigra
Testudo becki Rothschild 1901:372, Geochelone becki, Geochelone elephantopus becki, Chelonoidis becki, Chelonoidis elephantopus becki, Geochelone (Chelonoidis) nigra
becki, Geochelone nigra becki, Chelonoidis nigra becki
Type locality: “Cape Berkeley, northern point of Albemarle Island,
Galapagos Archipelago” [Isabela, Galápagos, Ecuador]. Restricted
to “Bank’s Bay, Albemarle” [Isabela, Galápagos, Ecuador] by Van
Denburgh (1914:259).
Ecuador (Galápagos: San Cristóbal [Chatham])
IUCN Red List: Vulnerable D1+2 (1996), as Chelonoidis nigra
chathamensis
TFTSG Draft Red List: Endangered (2017)
CITES: Appendix I, as Chelonoidis nigra
Testudo chathamensis Van Denburgh 1907:4, Testudo
elephantopus chathamensis, Geochelone chathamensis,
Geochelone elephantopus chathamensis, Chelonoidis
chathamensis, Chelonoidis elephantopus chathamensis,
Geochelone (Chelonoidis) nigra chathamensis, Geochelone nigra chathamensis, Chelonoidis nigra chathamensis
Type locality: “Chatham Island, Galapagos Archipelago” [San
Cristóbal, Galápagos, Ecuador].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelonoidis darwini (Van Denburgh 1907) (12:31) (60)
Santiago Giant Tortoise, James Island Giant Tortoise
Chelonoidis donfaustoi Poulakakis, Edwards, and Caccone in Poulakakis, Edwards, Chiari, Garrick, Russello, Benavides,
Watkins-Colwell, Glaberman, Tapia, Gibbs, Cayot, and
Caccone 2015 (61)
Eastern Santa Cruz Giant Tortoise, Cerro Fatal Giant Tortoise,
Don Fausto’s Giant Tortoise
Tui De Roy/Roving Tortoise Photos / Santiago
Peter Paul van Dijk / Cerro Fatal, Santa Cruz
Ecuador (Galápagos: Santiago [San Salvador] [James])
IUCN Red List: Critically Endangered A1bde (2016); Previously: Endangered (1996)
CITES: Appendix I, as Chelonoidis nigra
Testudo schweiggeri Fitzinger 1826:44 (nomen nudum)
Geochelone (Geochelone) schweiggeri Fitzinger 1835:122
(nomen dubium et oblitum; junior homonym, not =
Testudo schweiggeri Gray in Duméril and Bibron 1835 [=
Testudo schweigeri Gray 1830e]), Geochelone schweiggeri
Type locality: Not designated. Restricted to “America: Insel St. Jacob der Gallopagen” [= Santiago, Galápagos, Ecuador] by Fitzinger
(1853:110).
Testudo darwini Van Denburgh 1907:4, Testudo elephantopus
darwini, Geochelone darwini, Geochelone elephantopus
darwini, Chelonoidis darwini, Chelonoidis elephantopus
darwini, Geochelone (Chelonoidis) nigra darwini, Geochelone nigra darwini, Chelonoidis nigra darwini
Type locality: “James Island, Galapagos Archipelago” [Santiago,
Galápagos, Ecuador].
Ecuador (Galápagos: Santa Cruz [Indefatigable])
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Critically Endangered (2015)
CITES: Appendix I, as Chelonoidis nigra
Chelonoidis donfaustoi Poulakakis, Edwards, and Caccone in
Poulakakis, Edwards, Chiari, Garrick, Russello, Benavides,
Watkins-Colwell, Glaberman, Tapia, Gibbs, Cayot, and Caccone 2015:12, Chelonoidis nigra donfaustoi
Type locality: “Cerro Fatal in Santa Cruz” [Galápagos, Ecuador].
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Chelonoidis duncanensis (Pritchard 1996) (07:60, 12:31)
Pinzón Giant Tortoise, Duncan Island Giant Tortoise
Chelonoidis guntheri (Baur 1889) (07:64, 08:14, 09:36, 12:31) (60)
Sierra Negra Giant Tortoise
Tui De Roy/Roving Tortoise Photos / Sierra Negra, Isabela [captivity]
Peter C.H. Pritchard / CRM 1 / Pinzón [Onan]
Ecuador (Galápagos: Isabela [Albemarle])
IUCN Red List: Endangered C2a (1996), as Chelonoidis nigra
guentheri
TFTSG Draft Red List: Endangered (2012)
CITES: Appendix I, as Chelonoidis nigra
Testudo güntheri Baur 1889:1044 (07:64, 09:36) (invalid diacritic requiring emendation), Geochelone elephantopus
güntheri, Geochelone güntheri, Geochelone (Chelonoidis)
nigra güntheri, Geochelone nigra güntheri, Chelonoidis
elephantopus güntheri
Type locality: “Galapagos.” Restricted to “Vilamil, Albemarle”
[Isabela, Galápagos, Ecuador] by Van Denburgh (1914:259).
(orange dot = introduced)
Ecuador (Galápagos: Pinzón [Duncan])
Introduced: Ecuador (Galápagos: Santa Cruz [Indefatigable])
IUCN Red List: Extinct in the Wild (1996), as Chelonoidis
nigra duncanensis
TFTSG Draft Red List: Vulnerable (2012)
CITES: Appendix I, as Chelonoidis nigra
Testudo ephippium Günther 1874:422 (partim, nomen nudum)
Testudo ephippium Günther 1875a:271 (partim, misidentified
type), Testudo elephantopus ephippium, Geochelone elephantopus ephippium, Geochelone ephippium, Chelonoidis ephippium, Chelonoidis nigra ephippium, Chelonoidis
elephantopus ephippium, Geochelone (Chelonoidis) nigra
ephippium, Geochelone nigra ephippium
Type locality: “Charles Island” [Floreana, Galápagos, Ecuador] [in
error]. Restricted to “Duncan” [Pinzón, Galápagos, Ecuador] by
Van Denburgh (1914:259) [in error].
Testudo duncanensis Garman 1917:269 (nomen nudum)
Geochelone nigra duncanensis Pritchard 1996:47, Chelonoidis nigra duncanensis, Chelonoidis duncanensis
Type locality: “Duncan Island” [Pinzón, Galápagos, Ecuador].
Testudo wallacei Rothschild 1902:619 (12:31), Testudo elephantopus wallacei, Geochelone elephantopus wallacei,
Geochelone wallacei, Chelonoidis elephantopus wallacei,
Chelonoidis nigra wallacei, Geochelone nigra wallacei
Type locality: “Chatham Island” [San Cristóbal, Galápagos,
Ecuador] by supposition. Restricted to “Jervis” [Rábida, Galápagos,
Ecuador] by Van Denburgh (1914:259).
Geochelone elephantopus guntheri Baur in Pritchard
1971a:26 (07:64, 09:36) (nomen novum, justified emendation),
Geochelone elephantopus guntheri, Geochelone guntheri,
Geochelone (Chelonoidis) nigra guntheri, Geochelone
nigra guntheri, Chelonoidis elephantopus guntheri
Geochelone elephantopus guentheri Baur in Pritchard
1971b:50 (07:64, 09:36) (nomen novum, unjustified emendation), Testudo guentheri, Geochelone guentheri, Chelonoidis guentheri, Chelonoidis elephantopus guentheri,
Geochelone nigra guentheri, Chelonoidis nigra guentheri
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelonoidis hoodensis (Van Denburgh 1907) (12:31)
Española Giant Tortoise, Hood Island Giant Tortoise
Chelonoidis microphyes (Günther 1875a) (07:64, 08:14, 09:36, 12:31)
Volcán Darwin Giant Tortoise, Darwin Volcano Giant Tortoise,
Tagus Cove Giant Tortoise
Anders G.J. Rhodin / Volcán Darwin, nr. Tagus Cove, Isabela
Peter C.H. Pritchard / TCC / Española [CDRS, Santa Cruz]
(orange dot = introduced)
Ecuador (Galápagos: Española [Hood])
Introduced: Ecuador (Galápagos: Santa Fé [Barrington])
IUCN Red List: Critically Endangered D (1996), as Chelonoidis
nigra hoodensis
TFTSG Draft Red List: Critically Endangered (2015)
CITES: Appendix I, as Chelonoidis nigra
Testudo hoodensis Van Denburgh 1907:3, Testudo elephantopus hoodensis, Geochelone elephantopus hoodensis,
Geochelone hoodensis, Chelonoidis hoodensis, Chelonoidis elephantopus hoodensis, Geochelone (Chelonoidis)
nigra hoodensis, Geochelone nigra hoodensis, Chelonoidis nigra hoodensis, Darwintestudo hoodensis
Type locality: “Hood Island, Galapagos Archipelago” [Española,
Galápagos, Ecuador].
Ecuador (Galápagos: Isabela [Albemarle])
IUCN Red List: Vulnerable D1+2 (1996), as Chelonoidis nigra
microphyes
TFTSG Draft Red List: Endangered (2012)
CITES: Appendix I, as Chelonoidis nigra
Testudo microphyes Günther 1874:422 (nomen nudum)
Type locality: ““Hood’s Island” [Española, Galápagos, Ecuador] [in
error].
Testudo microphyes Günther 1875a:275 (08:14)
Geochelone elephantopus microphyes, Geochelone microphyes, Chelonoidis microphyes, Chelonoidis elephantopus
microphyes, Geochelone (Chelonoidis) nigra microphyes, Geochelone nigra microphyes, Chelonoidis nigra
microphyes
Type locality: “Hood’s Island” [Española, Galápagos, Ecuador]
by supposition [in error]. Emended to “Tagus Cove, about 4 miles
inland...northern Albemarle Island” [Isabela, Galápagos, Ecuador]
by Günther (1877:78).
Testudo macrophyes Garman 1917:273
Type locality: “Santa Isabela Island (Albemarle) near Tagus Cove”
[Isabela, Galápagos, Ecuador].
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Chelonoidis niger (Quoy and Gaimard 1824b) (07:58, 09:33, 12:31) (58, 62)
(Extinct, ca. 1850)
Floreana Giant Tortoise, Charles Island Giant Tortoise
Chelonoidis phantasticus (Van Denburgh 1907) (07:62, 12:31) (58, 63)
Fernandina Giant Tortoise, Narborough Island Giant Tortoise
Peter C.H. Pritchard / CRM 1 / Fernandina [CAS, San Francisco]
Peter C.H. Pritchard / CRM 1 / Floreana [BMNH, London]
Ecuador (Galápagos: Floreana [Charles] [extinct])
Introduced: Ecuador (Galápagos: Isabela [Albemarle]) (hybrids
with C. becki)
IUCN Red List: Extinct (1996), as Chelonoidis nigra nigra
TFTSG Draft Red List: Extinct (2012)
CITES: Appendix I
Testudo californiana Quoy and Gaimard 1824a:90 (62) (nomen
oblitum)
Type locality: “La Californie...donnée vivante aux îles Sandwich”
[California, USA; in error]. Erroneously given with “Sandwich
Islands” [Hawaii, USA] by Wermuth and Mertens (1961, 1977).
Testudo nigra Quoy and Gaimard 1824b:174 (62)
(nomen novum), Chelonoidis nigra, Geochelone nigra,
Geochelone nigra nigra, Chelonoidis nigra nigra, Geochelone elephantopus nigra, Chelonoidis niger
Type locality: “la Californie...donnée vivante, aux îles Sandwich”
[California, USA; in error]. Restricted arbitrarily to “Cerro Azul
d’Isabela” [Galápagos, Ecuador] by Bour in David (1994:83); and
to “Charles Island (Santa María or Floreana)” [Galápagos, Ecuador]
by Pritchard (1996:49).
Testudo galapagoensis Baur 1889:1044, Testudo elephantopus galapagoensis, Geochelone elephantopus galapagoensis, Chelonoidis galapagoensis, Chelonoidis elephantopus
galapagoensis, Geochelone (Chelonoidis) nigra galapagoensis, Geochelone nigra galapagoensis, Chelonoidis
nigra galapagoensis
Type locality: “Charles Island” [Floreana, Galápagos, Ecuador].
Ecuador (Galápagos: Fernandina [Narborough] [possibly
extinct])
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Critically Endangered (Possibly Extinct)
(2016); Previous Draft: Extinct (63)
CITES: Appendix I, as Chelonoidis nigra
Testudo phantasticus Van Denburgh 1907:4, Testudo phantastica, Testudo elephantopus phantastica, Geochelone
elephantopus phantastica, Geochelone phantastica, Chelonoidis phantastica, Geochelone phantasticus, Chelonoidis
elephantopus phantastica, Geochelone (Chelonoidis)
nigra phantastica, Geochelone nigra phantastica, Chelonoidis nigra phantastica, Chelonoidis phantasticus
Type locality: “Narborough Island, Galapagos Archipelago” [Fernandina, Galápagos, Ecuador].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelonoidis porteri (Rothschild 1903) (07:63, 09:35, 12:31) (62)
Western Santa Cruz Giant Tortoise, Indefatigable Island Giant
Tortoise
Anders G.J. Rhodin / Caseta, Santa Cruz
Ecuador (Galápagos: Santa Cruz [Indefatigable])
IUCN Red List: Endangered C2a (1996), as Chelonoidis nigra
porteri
TFTSG Draft Red List: Critically Endangered (2016)
CITES: Appendix I, as Chelonoidis nigra
Testudo nigrita Duméril and Bibron 1835:80 (62)
(nomen dubium), Testudo elephantopus nigrita, Geochelone nigrita, Chelonoidis nigrita, Geochelone elephantopus nigrita, Chelonoidis elephantopus nigrita, Geochelone
(Chelonoidis) nigra nigrita, Geochelone nigra nigrita,
Chelonoidis nigra nigrita
Type locality: Not known. Restricted to “Insel Indefatigable” [Santa
Cruz, Galápagos, Ecuador] by Mertens and Wermuth (1955:376).
Testudo porteri Rothschild 1903:119, Geochelone elephantopus porteri, Geochelone porteri, Geochelone nigra porteri,
Chelonoidis elephantopus porteri, Chelonoidis nigra
porteri, Chelonoidis porteri
Type locality: “Indefatigable Island, Galapagos group” [Santa Cruz,
Galápagos, Ecuador].
Chelonoidis vandenburghi (DeSola 1930) (07:64, 08:14, 09:36, 12:31) (60)
Volcán Alcedo Giant Tortoise, Alcedo Volcano Giant Tortoise
Peter C.H. Pritchard / CRM 1 / TCF / Volcán Alcedo, Isabela
Ecuador (Galápagos: Isabela [Albemarle])
IUCN Red List: Vulnerable D2 (1996), as Chelonoidis nigra
vandenburghi
TFTSG Draft Red List: Vulnerable (2012)
CITES: Appendix I, as Chelonoidis nigra
Testudo vandenburghi DeSola 1930:79, Geochelone
vandenburghi, Geochelone elephantopus vandenburghi,
Chelonoidis vandenburghi, Chelonoidis elephantopus
vandenburghi, Geochelone (Chelonoidis) nigra vandenburghi, Geochelone nigra vandenburghi, Chelonoidis
nigra vandenburghi
Type locality: “forty miles from Villamil...at the coast on the
southern border of Perry Isthmus...mid-Albemarle Island...Cowley
Mountain nearly north...to the south...Villamil Mountain” [Isabela,
Galápagos, Ecuador], emended to “Cowley Mountain...the first
mountain north of Villamil Mountain” [Volcán Alcedo, Isabela,
Galápagos, Ecuador] by Iverson (1992:250).
131
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Chelonoidis vicina (Günther 1875a) (07:64, 08:14, 09:36, 12:31) (60)
Cerro Azul Giant Tortoise, Iguana Cove Giant Tortoise
Type locality: “Galapagos” [Galápagos, Ecuador].
Chersina Gray 1830e (10:7)
Testudo (Chersina) Gray 1830e:5 (10:7)
Type species: Testudo (Chersina) angulata Duméril in Schweigger
1812], by original monotypy.
Testudo (Goniochersus) Lindholm 1929:285
Type species: Testudo (Goniochersus) angulata [= Testudo angulata
Schweigger 1812], by original designation.
Neotestudo Hewitt 1931:504
Type species: Neotestudo angulata [= Testudo angulata Schweigger
1812], by original monotypy.
Vincenzo Ferri / Cerro Azul, Isabela [captivity]
Ecuador (Galápagos: Isabela [Albemarle])
IUCN Red List: Endangered C2a (1996), as Chelonoidis nigra
vicina
TFTSG Draft Red List: Vulnerable (2017)
CITES: Appendix I, as Chelonoidis nigra
Testudo vicina Günther 1874:422 (nomen nudum)
Chersina angulata (Duméril in Schweigger 1812) (09:37)
Angulate Tortoise, South African Bowsprit Tortoise
Margaretha D. Hofmeyr / CBFTT / West Coast National Park, South Africa
Type locality: “Galapagos” [Galápagos, Ecuador] without precise
locality.
Testudo vicina Günther 1875a:277, Geochelone vicina,
Geochelone (Chelonoidis) nigra vicina, Geochelone nigra
vicina, Geochelone elephantopus vicina, Chelonoidis elephantopus vicina, Chelonoidis nigra vicina, Chelonoidis
vicina
Type locality: “Galapagos.” Restricted to “South Albemarle” [Isabela, Galápagos, Ecuador] by Günther (1877:73); and to “Iguana
Cove, Albemarle” [Isabela, Galápagos, Ecuador] by Van Denburgh
(1914:259).
Chelonoidis, sp. indet.
Testudo elephantopus Harlan 1827:284 (09:33)
(nomen dubium), Testudo elephantopus elephantopus,
Geochelone elephantopus, Geochelone elephantopus
elephantopus, Chelonoidis elephantopus, Chelonoidis elephantopus elephantopus, Geochelone nigra elephantopus
Type locality: “Gallapagos Islands” [Galápagos, Ecuador]. Restricted
to “South Albemarle” [Isabela, Galápagos, Ecuador] by Pritchard
(1967:168); restricted to “Charles Island (Floreana, Santa Maria),
Galapagos” [Ecuador] by Olson and Humphrey (2017:114).
Testudo planiceps Gray 1854b:12 (nomen dubium and junior
homonym, not = Testudo planiceps Schneider 1792),
Geochelone planiceps, Chelonoidis planiceps
Type locality: “Galapagos Islands” [Galápagos, Ecuador].
Testudo clivosa Garman 1917:283 (nomen dubium), Geochelone clivosa, Chelonoidis clivosa
Type locality: “Mascarenes” by supposition [in error].
Testudo typica Garman 1917:285 (nomen dubium), Geochelone typica, Chelonoidis typica
(orange dots = probable trade)
Namibia, South Africa
CBFTT Account: Hofmeyr (2009)
IUCN Red List: Least Concern (2017); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix II, as Testudinidae spp.
Testudo angulata Duméril in Schweigger 1812:321, Testudo
(Chersina) angulata, Chersina angulata, Testudo (Goniochersus) angulata, Goniochersus angulata, Goniochersus
angulatus, Neotestudo angulata, Chersine angulata
Type locality: Not known.
Testudo bellii Gray 1828:2
Type locality: “Cape of Good Hope” [South Africa].
Chersina angulata pallida Gray 1831d:69
Type locality: Not known.
Testudo flavofusca Gray 1844:11 (nomen nudum)
Testudo sculpta Brandt in Gray 1856b:12 (nomen nudum)
Chersobius Fitzinger 1835 (07:67, 10:23) (64)
Testudo (Chersobius) Fitzinger 1835:112
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
133
Type species: Testudo (Chersobius) signatus Walbaum [= Testudo
signata Gmelin 1789], by subsequent designation by Lindholm
(1929:284).
Pseudomopus Hewitt 1931:496
Type species: Pseudomopus signatus Walbaum [= Testudo signata
Gmelin 1789], by original designation.
Chersobius boulengeri (Duerden 1906) (64)
Karoo Dwarf Tortoise, Karoo Padloper
South Africa
IUCN Red List: Vulnerable A2acd (2017), as Homopus signatus; Previously: Near Threatened (1996)
CITES: Appendix II, as Testudinidae spp.
Testudo signata Walbaum 1782:120 (unavailable name)
Type locality: Not known.
Victor J.T. Loehr / Western Cape, South Africa
Testudo signata Gmelin 1789:1043, Chersine signata,
Homopus signatus, Pseudomopus signatus, Pseudomopus
signatus signatus, Chersobius signatus, Homopus signata,
Homopus signatus signatus
Type locality: “Virginia” [USA, in error]. Restricted to “vicinity of
Springbok, Cape Province, South Africa” by Bour (1988:3).
Testudo cafra Daudin 1801:291 (10:24), Homopus signatus
cafra, Homopus signatus cafer
Type locality: “Afrique...la Cafrerie” [South Africa]. Restricted to
“Drainage of the Olifants River, Cape Province, South Africa” by
Bour (1988:3).
Testudo juvencella Daudin 1802:380 (10:25)
Type locality: “Afrique.”
South Africa
IUCN Red List: Near Threatened (2017), as Homopus boulengeri; Previously: Least Concern [Not Listed] (1996)
CITES: Appendix II, as Testudinidae spp.
Homopus boulengeri Duerden 1906:406, Pseudomopus
boulengeri, Chersobius signatus
Pseudomopus signatus peersi Hewitt 1935:345, Homopus
signatus peersi, Chersobius peersi
Type locality: “Klaver District, C.P. [Cape Province], near Van
Rhynsdorp” [South Africa].
Chersobius solus Branch 2007 (07:67) (64)
Nama Tortoise, Nama Padloper
Type locality: “South Africa—Districts of Willowmore, Aberdeen,
and Beaufort West.”
Chersobius signatus (Gmelin 1789) (10:24) (64)
Speckled Tortoise, Speckled Padloper
Maurice Rodrigues / TCC / Namibia
Victor J.T. Loehr / Clanwilliam, Western Cape, South Africa
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IUCN Red List: Extinct (1996)
Testudo indica Schneider 1783:355, Chelonura indica, Cylindrapis indica, Megalochelys indica, Geochelone indica,
Cylindraspis indica
Type locality: “Ostindien”. Restricted to “Réunion” by Austin and
Arnold (2001:2517).
Testudo tabulata africana Schweigger 1812:322 (09:38)
Type locality: “Africa” by inference.
Chersine retusa Merrem 1820:29 (nomen novum), Testudo
retusa
Type locality: “India orientali.”
Testudo perraultii Duméril and Bibron 1835:126 (nomen
novum), Geochelone (Cylindraspis) perraultii, Testudo
indica perraultii
Namibia
IUCN Red List: Vulnerable C2a (1996), originally listed as
Homopus bergeri
TFTSG Draft Red List: Endangered (2013)
CITES: Appendix II, as Testudinidae spp.
Homopus bergeri Lindholm 1906:348 (partim)
Type locality: “nach Gibeon in Deutsch-Südwestafrika...möglicherweise weiter im Innern Südafrikas” [Namibia or South Africa].
Homopus solos Devaux 2003:40 (nomen nudum)
Homopus solus Branch 2007:11, Chersobius solus
Type locality: “vicinity of Aus, Luderitz District, Namibia.”
Cylindraspis Fitzinger 1835 (55)
Type locality: “Indes Orientales.”
Testudo graii Duméril and Bibron 1835:135 (nomen novum),
Geochelone graii, Cylindraspis graii
Type locality: “Afrique?”
Chersina grayi Strauch 1865:36 (nomen novum), Geochelone
grayi, Geochelone (Cylindraspis) grayi
Type locality: “Afrika.”
Cylindraspis borbonica Bour 1978:492
Type locality: “Réunion.”
Cylindraspis inepta (Günther 1873)
(Extinct, ca. 1735)
Mauritius Giant Domed Tortoise
Chelonura Rafinesque 1815:74 (nomen nudum)
Chelonura Rafinesque 1832:64 (junior homonym, not = Chelonura Fleming 1822)
Type species: Chelonura indica [= Testudo indica Schneider 1783], by
original designation.
Geochelone (Cylindraspis) Fitzinger 1835:112
Type species: Chersina (Cylindraspis) vosmaeri [= Testudo indica
vosmaeri Suckow 1798], by subsequent designation by Fitzinger
(1843:29).
Cylindraspis indica (Schneider 1783) (07:65)
(Extinct, ca. 1840)
Reunion Giant Tortoise
Roger Bour / Mare aux Songes, Mauritius [BMNH, London]
Roger Bour / Réunion [MNHN, Paris]
Mauritius (Mauritius [extinct])
IUCN Red List: Extinct (1996)
Testudo neraudii Gray 1831d:14 (nomen oblitum)
Type locality: “Isle of France” [Mauritius]. Restricted to “Quatre
Cocos, Flacq, Maurice” [Mauritius] by Bour (1985:37).
Testudo inepta Günther 1873:397, Geochelone inepta, Cylindraspis inepta
Type locality: “Mauritius.” Restricted to “La Mare aux Songes...
near Mahe’bourg...Mauritius” by Günther (1875b:297).
Testudo boutonii Günther 1875b:297
Type locality: “La Mare aux Songes...near Mahe’bourg...
Mauritius.”
Réunion [extinct]
Testudo sauzieri Gadow 1894:315, Geochelone sauzieri
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “Mare aux Songes, in Mauritius.”
Cylindraspis peltastes (Duméril and Bibron 1835)
(Extinct, ca. 1800)
Rodrigues Domed Tortoise
Roger Bour / Rodrigues [MNHN, Paris]
Mauritius (Rodrigues [extinct])
IUCN Red List: Extinct (1996)
Testudo rotunda Latreille in Sonnini and Latreille 1801:107
(partim, nomen dubium), Chersine rotunda, Geochelone
(Geochelone) rotunda
Type locality: Not known. Restricted to “Mascarenes...Rodrigues”
by Bour (2005:25).
Testudo peltastes Duméril and Bibron 1835:138, Geochelone
peltastes, Geochelone (Cylindraspis) peltastes, Cylindraspis peltastes
Type locality: Not known. Restricted to “Rodriguez” [Rodrigues]
by Günther (1877:53).
Cylindraspis triserrata (Günther 1873)
(Extinct, ca. 1735)
Mauritius Giant Flat-shelled Tortoise
Roger Bour / plates from Boulenger (1889:pls.2-3) / Mauritius [BMNH, London]
Mauritius (Mauritius [extinct])
IUCN Red List: Extinct (1996)
Testudo schweigeri Gray 1830e:3 (10:7) (nomen oblitum)
Type locality: Not known. Restricted to “Mauritius” by Austin and
Arnold (2001:2517).
Testudo schweiggeri Gray in Duméril and Bibron 1835:108
(nomen novum et oblitum; senior homonym, not = Geochelone (Geochelone) schweiggeri Fitzinger 1835)
Testudo triserrata Günther 1873:397, Geochelone triserrata,
Cylindraspis triserrata
Type locality: “Mauritius.” Restricted to “La Mare aux Songes...
near Mahe’bourg...Mauritius” by Günther (1875b:297).
Testudo leptocnemis Günther 1875b:297, Geochelone leptocnemis, Cylindraspis leptocnemis
Type locality: “district of Flacq..[&]..La Mare aux Songes...near
Mahe’bourg...Mauritius.” Restricted to “District of Flacq” [Mauritius] by Günther (1877:14).
Testudo microtympanum Boulenger 1891:4, Geochelone
microtympanum
Type locality: “Mauritius” by supposition.
Testudo guentheri Gadow 1894:320 (senior homonym)
Type locality: “Mare aux Songes...Mauritius.”
Testudo güntheri Gadow in Van Denburgh 1914:257 (nomen
novum, invalid name, junior homonym), Testudo guntheri
Testudo gadowi Van Denburgh 1914:257 (nomen novum),
Geochelone (Megalochelys) gadowi, Geochelone (Cylindraspis) gadowi
135
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Cylindraspis vosmaeri (Suckow 1798) (07:66)
(Extinct, ca. 1800)
Rodrigues Giant Saddleback Tortoise
Geochelone Fitzinger 1835 (07:52)
Geochelone Fitzinger 1835:112
Type species: Geochelone stellata [= Testudo stellata Schweigger
1812 = objective synonym of Testudo elegans Schoepff 1795], by
subsequent designation by Fitzinger (1843:29).
Geochelone elegans (Schoepff 1795) (65)
Indian Star Tortoise
Roger Bour / Rodrigues [Ste. Geneviève]
Peter Paul van Dijk / CRM 2 / northeast Tamil Nadu, India
Mauritius (Rodrigues [extinct])
IUCN Red List: Extinct (1996)
Testudo indica vosmaeri Suckow 1798:57, Testudo vosmaeri,
Geochelone (Cylindraspis) vosmaeri, Chersina (Cylindraspis) vosmaeri, Geochelone vosmaeri, Cylindraspis
vosmaeri
Type locality: Not known. Restricted to “Rodriguez” [Rodrigues]
by Günther (1877:53).
Testudo rotunda Latreille in Sonnini and Latreille 1801:107
(partim, nomen dubium), Chersine rotunda, Geochelone
(Geochelone) rotunda
Type locality: Not known. Restricted to “Mascarenes...Rodrigues”
by Bour (2005:25).
Testudo rodericensis Günther 1873:397
Type locality: “Rodriguez” [Rodrigues].
Testudo commersoni Vaillant 1898:138, Geochelone commersoni, Cylindraspis commersoni
Type locality: “Rodrigue” [Rodrigues].
(orange dots = trade)
India (Andhra Pradesh, Gujarat, Karnataka, Kerala, Madhya
Pradesh, Rajasthan, Tamil Nadu), Pakistan, Sri Lanka
IUCN Red List: Vulnerable A4cd (2016); Previously: Least
Concern (2000), Least Concern [Not Listed] (1996)
CITES: Appendix II, as Testudinidae spp.
Testudo elegans Schoepff 1795:111 (65), Chersine elegans,
Peltastes stellatus elegans, Geochelone elegans, Geochelone elegans elegans
Type locality: “India orientali.”
Testudo stellata Schweigger 1812:325 (65) (nomen novum),
Geochelone (Geochelone) stellata, Geochelone stellata,
Peltastes stellatus
Type locality: “India orientali.”
Testudo actinodes Bell 1828a:419
Type locality: “Africâ” [in error].
Testudo actinoides Bell in Gray 1844:7 (nomen novum),
Peltastes stellatus actinoides
Testudo megalopus Blyth 1854:640 (66)
Type locality: Not known. Restricted to “Calcutta” [India] by Das
et al. (1998:127).
Peltastes stellatus maura Gray 1870c:8
Type locality: Not known.
Peltastes stellatus seba Gray 1870c:8
Type locality: Not known.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Geochelone platynota (Blyth 1863)
Burmese Star Tortoise
Kalyar Platt / CBFTT / Lawkanandar Wildlife Sanctury, Bagan, Myanmar
(orange dots = trade)
Myanmar
CBFTT Account: Platt, Thanda Swe, Win Ko Ko, Platt, Khin
Myo Myo, Rainwater, and Emmett (2011)
IUCN Red List: Critically Endangered A1cd+2cd, C2a (2000); Previously: Critically Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix I
Testudo platynotus Blyth 1863:83, Peltastes platynotus,
Testudo platynota, Geochelone platynota, Geochelone
elegans platynota
Type locality: “Lower Pegu..[&]..Valley of the Irawadi” [Myanmar]. Restricted to “Irrawaddy Valley” [Myanmar] by Smith
(1931:140).
Gopherus Rafinesque 1832 (67)
Gopherus Rafinesque 1815:74 (nomen nudum)
Gopherus Rafinesque 1832:64
137
Gopherus agassizii (Cooper 1861) (10:22, 11:11)
Mojave Desert Tortoise, Mohave Desert Tortoise, Agassiz’s
Desert Tortoise
Bev Steveson / CCB / Mojave Desert, California
USA (Arizona, California, Nevada, Utah)
IUCN Red List: Vulnerable A1acde+2cde, E (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Testudinidae spp.
Xerobates agassizii Cooper 1861:120 (10:22), Testudo agassizii,
Gopherus agassizii, Gopherus polyphemus agassizii,
Geochelone agassizii, Scaptochelys agassizii
Type locality: “mountains of California, near Fort Mojave”
[USA].
Xerobates lepidocephalus Ottley and Velázquez Solis
1989:497 (11:11)
Type locality: “western base of the Sierra San Vicente, approximately 1 km N of the Buena Mujer Dam, Baja California Sur, Mexico”
[in error or introduced].
Gopherus berlandieri (Agassiz 1857a) (14:34)
Texas Tortoise, Berlandier’s Tortoise
Type species: Gopherus polyphemus [= Testudo polyphemus Daudin
1801], by original designation.
Xerobates Agassiz 1857a:252,446
Type species: Xerobates berlandieri Agassiz 1857a, by subsequent
designation by Brown (1908:115).
Bysmachelys Johnston 1937:439
Type species: Bysmachelys canyonensis † Johnston 1937 [= subjective
synonym of Testudo pertenuis † Cope 1892 (see TEWG 2015)], by
original monotypy.
Scaptochelys Bramble 1982:852
Type species: Scaptochelys agassizii [= Xerobates agassizii Cooper
1861], by original designation.
Larry Ditto / CCB / Texas
138
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(orange dots = introduced)
Mexico (Coahuila, Nuevo Leon, San Luis Potosi, Tamaulipas),
USA (Texas)
IUCN Red List: Least Concern (1996)
TFTSG Draft Red List: Near Threatened (2011)
CITES: Appendix II, as Testudinidae spp.
Testudo tuberculatu Berlandier 1850:287 (14:34) (nomen
oblitum)
Type locality: “Tamaulipas” [Mexico].
Testudo bicolor Berlandier 1850:287 (14:34) (nomen oblitum)
Type locality: “Tamaulipas” [Mexico].
Xerobates berlandieri Agassiz 1857a:392,447, Testudo
berlandieri, Xerobates gopher berlandieri, Gopherus berlandieri, Gopherus polyphemus berlandieri, Scaptochelys
berlandieri
Mexico (Chihuahua, Sinaloa, Sonora)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Vulnerable (2016)
CITES: Appendix II, as Testudinidae spp.
Gopherus evgoodei Edwards, Karl, Vaughn, Rosen, Meléndez Torres, and Murphy 2016:140
Type locality: “Alamos (approximate location 27°02' N, 108°55' W,
elevation 433 m), Sonora, Mexico.”
Gopherus flavomarginatus Legler 1959 (12:32)
Bolson Tortoise
Type locality: “southern Texas [USA] and Mexico.” Restricted
to “Lower Rio Grande, Texas” [USA] by Stejneger and Barbour
(1917:121); and to “Brownsville, Cameron County, Texas, USA”
by Smith and Taylor (1950a:361).
Testudo tuberculata Berlandier in True 1882:441 (14:34) (nomen
novum et nudum)
Gopherus auffenbergi † Mooser 1972:61 [Late Pleistocene,
Mexico (Aguascalientes)], Xerobates auffenbergi
Type locality: “Cedazo Ravine; 3 1/2 km. NE of Aguascalientes,
Aguascalientes, Mexico.”
Eric V. Goode / TCC / Bolson de Mapimí, Durango, Mexico
Gopherus evgoodei Edwards, Karl, Vaughn, Rosen, Meléndez
Torres, and Murphy 2016 (68)
Goode’s Thornscrub Tortoise, Sinaloan Thornscrub Tortoise
Eric V. Goode / Monte Mojino Reserve, Alamos, Sonora, Mexico
Mexico (Chihuahua, Coahuila, Durango)
Introduced: USA (New Mexico)
IUCN Red List: Vulnerable A1cd (2007); Previously: Vulnerable (1996)
TFTSG Draft Red List: Critically Endangered (2017)
CITES: Appendix I
Gopherus flavomarginatus Legler 1959:337, Gopherus polyphemus flavomarginatus, Gopherus flavomarginata
Type locality: “30 to 40 miles from Lerdo, Durango, Mexico.”
Gopherus huecoensis † Strain 1966:24 [Early Pleistocene,
Blancan, USA (Texas)]
Type locality: “Madden Arroyo...Hudspeth County, Texas” [USA].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Gopherus morafkai Murphy, Berry, Edwards, Leviton, Lathrop,
and Riedle 2011 (11:11)
Sonoran Desert Tortoise, Morafka’s Desert Tortoise
Roy C. Averill-Murray / CCB / Maricopa Co., Arizona
USA (Florida, Georgia, South Carolina, Alabama, Mississippi,
Louisiana)
IUCN Red List: Vulnerable A1acde (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Testudinidae spp.
Testudo polyphaemus Bartram 1791:18 (nomen nudum)
Testudo polyphemus Daudin 1801:256, Emys polyphemus,
Gopherus polyphemus, Xerobates polyphemus, Gopherus
polyphemus polyphemus
Type locality: “l’Amérique septentrionale, principalement sur
les bords de la rivière Savannah et près de l’Alatamaha” [USA].
Restricted to “vicinity of Savannah, Georgia” [USA] by Schmidt
(1953:104).
Testudo depressa Guérin 1829:pl.1,f.1
Type locality: Not designated. Restricted to “vicinity of Savannah,
Georgia” [USA] by Schmidt (1953:105).
(orange dots = introduced or trade)
Mexico (Sinaloa, Sonora), USA (Arizona)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Vulnerable (2011)
Gopherus morafkai Murphy, Berry, Edwards, Leviton,
Lathrop, and Riedle 2011:53 (11:11), Xerobates morafkai
Type locality: “Tucson (approximate location 32°7' N, 110°56' W,
elevation 948 m), Pima County, Arizona, U.S.A.”
Gopherus polyphemus (Daudin 1801)
Gopher Tortoise
(12:33)
Testudo carolina Le Conte 1830:97 (nomen novum and junior
homonym, not = Testudo carolina Linnaeus 1758), Xerobates carolinus, Gopherus carolinus
Type locality: “pine forests of Georgia and Florida, ...never found
north of Savannah river” [USA].
Testudo gopher Bartram in Gray 1844:4, Xerobates gopher
Type locality: Not designated. Restricted to “vicinity of Savannah,
Georgia” [USA] by Schmidt (1953:105).
Testudo atascosae † Hay 1902:383 (nomen dubium) [Pleistocene, USA (Texas)], Gopherus atascosae
Type locality: “Atascosa county, Texas” [USA].
Gopherus praecedens † Hay 1916a:55 [Late Pleistocene,
USA (Florida)]
Type locality: “Florida...Vero, St. Lucie county” [USA].
Gopherus, sp. indet.
Testudo australis Girard 1858:470 (12:34) (nomen dubium et
oblitum), Gopherus australis
Type locality: “Bay of Islands, New Zealand” [in error, based on a
trade specimen].
Kevin Main / CCB / Lake Wales Ridge, Florida
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Homopus Duméril and Bibron 1834 (07:67, 10:23) (64)
Homopus Duméril and Bibron 1834:357
(10:23)
Homopus femoralis Boulenger 1888a (64)
Greater Dwarf Tortoise, Greater Padloper
Type species: Homopus areolatus [Tortue Aréolée Schoepff = Testudo
areolata Thunberg 1787], by subsequent designation by Duméril
and Bibron (1835:7).
Homopus areolatus (Thunberg 1787) (64)
Parrot-beaked Tortoise, Common Padloper
Victor J.T. Loehr / Karoo, Northern Cape, South Africa
Gerald Kuchling / Elandsberg, South Africa
South Africa
IUCN Red List: Least Concern (2017); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix II, as Testudinidae spp.
Homopus femoralis Boulenger 1888a:251, Testudo femoralis
South Africa
IUCN Red List: Least Concern (2017); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix II, as Testudinidae spp.
Testudo areolata Thunberg 1787:180, Chersine areolata,
Homopus areolatus, Homopus areolata
Type locality: “Indien” [India] [in error].
Testudo minuta Thunberg 1788:206 (nomen nudum)
Testudo miniata Lacepède 1788:166, synopsis[table] (09:6) (nomen suppressum, ICZN 2005a)
Type locality: “South Africa...Cradock.”
Indotestudo Lindholm 1929
Testudo (Indotestudo) Lindholm 1929:285
Type species: Testudo (Indotestudo) elongata [= Testudo elongata
Blyth 1854], by original designation.
Indotestudo elongata (Blyth 1854) (66)
Elongated Tortoise, Yellow-headed Tortoise
Type locality: “Cap de Bonne-espérance” [South Africa].
Testudo fasciata Daudin 1801:294 (junior homonym, not =
Testudo fasciata Suckow 1798), Chersine fasciata
Type locality: “l’île de Ceilan” [Sri Lanka] [in error].
Testudo africana Hermann 1804:218
Type locality: “Africa” by inference.
Chersine tetradactyla Merrem 1820:32
Type locality: “India orientali” [in error].
Testudo areolata pallida Gray 1831d:13
Type locality: Not designated. Restricted to “Süd-Afrika” [South
Africa] by Wermuth and Mertens (1961:175).
Flora Ihlow / CBFTT / Doi Phu Nang National Park, Phayao Prov., Thailand
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Testudinidae spp.
Testudo forstenii Schlegel and Müller 1845:30 (69), Peltastes
forstenii, Geochelone forstenii, Indotestudo forstenii,
Indotestudo elongata forstenii, Geochelone elongata
forstenii
Type locality: “Gilolo...Indischen Archipel” [Halmahera, Moluccas,
Indonesia] [probably in error]; restricted here to “near Gorontalo,
Sulawesi, Indonesia” by TTWG (69).
Indotestudo travancorica (Boulenger 1907) (07:68)
Travancore Tortoise
Bangladesh, Bhutan, Cambodia, China (?) (Guangxi, Yunnan),
India (Assam, Bihar, Jharkhand, Meghalaya, Mizoram,
Odisha, Sikkim, Tripura, Uttarakhand, Uttar Pradesh, West
Bengal), Laos, Malaysia (Peninsular), Myanmar, Nepal,
Thailand, Vietnam
CBFTT Account: Ihlow, Dawson, Hartmann, and Som (2016)
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Testudinidae spp.
Testudo elongata Blyth 1854:639 (66), Peltastes elongatus,
Indotestudo elongata, Geochelone elongata, Indotestudo
elongata elongata, Geochelone elongata elongata
V. Deepak / CBFTT / Anamalai Hills, Western Ghats, Tamil Nadu, India
Type locality: “Arakan” [Myanmar].
Testudo parallelus Annandale 1913:76
Type locality: “Chaibassa (Singhbhum) district, Chota Nagpur”
[India].
Indotestudo forstenii (Schlegel and Müller 1845) (69)
Forsten’s Tortoise, Sulawesi Tortoise
India (Karnataka, Kerala, Tamil Nadu)
CBFTT Account: Deepak, Ramesh, Bhupathy, and Vasudevan
(2011)
IUCN Red List: Vulnerable A1cd (2000)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Testudinidae spp.
Testudo travancorica Boulenger 1907:560, Geochelone travancorica, Indotestudo travancorica, Indotestudo elongata
travancorica, Geochelone elongata travancorica
Jérôme Maran / Sulawesi, Indonesia
Type locality: “near Trivandrum..[&]..Travancore hills between 500
and 1,000 feet altitude” [India].
Kinixys Bell 1827 (12:35)
Kinixys Bell 1827:398
Type species: Kinixys homeana Bell 1827, by subsequent designation
by Bell (1828c:514).
Kinyxis Gray 1830e:6 (nomen novum)
Cinixys Wagler 1830b:138 (nomen novum)
Cinixys (Cinothorax) Fitzinger 1835:108
Type species: Cinixys (Cinothorax) bellianus [= Testudo (Kinyxis)
belliana Gray 1830e], by subsequent designation by Fitzinger
(1843:29).
(orange dot = erroneous original type locality)
Indonesia (Sulawesi)
Kinixis Hallowell 1839:161 (nomen novum)
Cinyxis Peters 1866:887 (nomen novum)
Kinothorax Gray 1873j:16 (nomen novum)
Madakinixys Vuillemin 1972b:169
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Type species: Madakinixys domerguei Vuillemin 1972b, by original
monotypy.
Kinixys belliana Gray 1830e (07:69, 08:11, 10:7, 12:35)
Bell’s Hinge-back Tortoise
Vincenzo Ferri / Entebbe, Uganda
Angola, Benin, Cameroon, Central African Republic, Congo
(DRC), Congo (ROC), Equatorial Guinea, Gabon, Ghana,
Guinea, Ivory Coast, Liberia, Nigeria, Sierra Leone, Togo,
Uganda
CBFTT Account: Luiselli and Diagne (2014)
IUCN Red List: Data Deficient (1996)
TFTSG Draft Red List: Endangered (2013)
CITES: Appendix II, as Testudinidae spp.
Testudo erosa Schweigger 1812:321, Kinixys erosa, Cinixys
erosa, Kinixis erosa, Kinixys belliana erosa
Type locality: “America septentrionali” [North America] [in error].
Testudo schoepfii Fitzinger 1826:44 (nomen nudum)
Kinixys castanea Bell 1827:398, Cinixys (Cinixys) castanea,
Cinixys castanea, Kinixis castanea
Type locality: “Africa”.
Kinixis denticulata Hallowell 1839:161, Kinixys denticulata
Type locality: “Liberia...banks of the St. Paul and Mesurado rivers.”
Angola, Burundi, Congo (DRC), Congo (ROC), Eritrea,
Ethiopia, Kenya, Rwanda, Somalia, South Sudan, Sudan,
Tanzania, Uganda
IUCN Red List: Least Concern [Not Listed] (1996)
SARCA Draft: Least Concern (regional) (2010)
TFTSG Draft Red List: Vulnerable (2013)
CITES: Appendix II, as Testudinidae spp.
Testudo (Kinyxis) belliana Gray 1830e:6 (10:7), Kinixys belliana, Cinixys (Cinothorax) belliana, Cinixys belliana,
Cinothorax bellianus, Kinixys belliana belliana
Kinixys homeana Bell 1827 (12:35)
Home’s Hinge-back Tortoise
Type locality: Not known.
Kinixys schoensis Rüppell 1845:226, Kinixys belliana
schoensis
Type locality: “Schoa, südlich von Abyssinien” [Ethiopia].
Kinixys belliana mertensi Laurent 1956:27
Tomas Diagne / CBFTT / Cameroon
Type locality: “Dika, Uele...le nord du Congo” [Democratic
Republic of Congo (DRC)].
Kinixys erosa (Schweigger 1812) (12:35)
Forest Hinge-back Tortoise, Serrated Hinge-back Tortoise
Tomas Diagne / Oyo State, Nigeria
Benin, Cameroon, Central African Republic, Congo (DRC) (?),
Equatorial Guinea, Gabon (?), Ghana, Ivory Coast, Liberia,
Nigeria, Togo
CBFTT Account: Luiselli and Diagne (2013)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
143
IUCN Red List: Vulnerable A2cd (2006); Previously: Data
Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2013)
CITES: Appendix II, as Testudinidae spp.
Kinixys homeana Bell 1827:400, Cinixys homeana, Cinixys
(Cinothorax) homeana, Testudo (Kinyxis) homeana,
Kinixys belliana homeana
Type locality: “Africâ occidentali” [West Africa].
Kinixys lobatsiana Power 1927 (12:35)
Lobatse Hinge-back Tortoise
Mozambique, South Africa, Swaziland
IUCN Red List: Near Threatened (1996)
SARCA Draft: Least Concern (2010)
TFTSG Draft Red List: Vulnerable (2013)
CITES: Appendix II, as Testudinidae spp.
Kinixys natalensis Hewitt 1935:353, Kinixys belliana
natalensis
Type locality: “Jameson Drift, Tugela River, and...Dimane stream,
near Jameson Drift” [Natal] [South Africa].
Flora Ihlow / Lapalala Reserve, Limpopo, South Africa
Kinixys nogueyi (Lataste 1886) (08:11, 09:39, 12:35)
Western Hinge-back Tortoise
Tomas Diagne / Dene, Senegal
(orange dots = introduced)
Botswana, South Africa
IUCN Red List: Least Concern (2017)
TFTSG Draft Red List: Vulnerable (2017)
CITES: Appendix II, as Testudinidae spp.
Cinixys lobatsiana Power 1927:410, Kinixys lobatsiana,
Kinixys belliana lobatsiana
Type locality: “Lobatsi [Bechuanaland Protectorate]” [Botswana].
Kinixys natalensis Hewitt 1935 (12:35)
Natal Hinge-back Tortoise, KwaZulu-Natal Hinge-back Tortoise
Benin, Burkina Faso, Cameroon, Central African Republic,
Chad, Congo (DRC), Gambia, Ghana, Guinea, GuineaBissau, Ivory Coast, Mali, Mauritania (?), Niger, Nigeria,
Senegal, Sierra Leone, Togo
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Vulnerable (2013)
CITES: Appendix II, as Testudinidae spp.
Homopus nogueyi Lataste 1886:286, Cinixys nogueyi, Cinixys belliana nogueyi, Kinixys nogueyi, Kinixys belliana
nogueyi
Type locality: “Médine (Haut-Sénégal)” [Senegal].
Victor J.T. Loehr / KwaZulu-Natal, South Africa
Cinixys dorri Lataste 1888:164
Type locality: “Haut-Sénégal” [Senegal]. Restricted to “Bakel,
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Haut-Sénégal” [Senegal] by Mertens and Wermuth (1955:372).
Kinixys spekii Gray 1863d (12:35)
Speke’s Hinge-back Tortoise
Kinixys zombensis Hewitt 1931 (12:35)
Southeastern Hinge-back Tortoise
Victor J.T. Loehr / nr. Lephalale, Limpopo, South Africa
(subspecies: zombensis = red, domerguei = orange dots)
Kenya, Malawi, Mozambique, South Africa, Tanzania, Madagascar (prehistoric introduction?)
IUCN Red List: Not Evaluated
SARCA Draft: Least Concern (regional) (2010)
TFTSG Draft Red List: Vulnerable (2013)
CITES: Appendix II, as Testudinidae spp.
Angola, Botswana, Burundi, Congo (DRC), Kenya, Malawi,
Mozambique, Namibia (Caprivi), South Africa, Swaziland,
Tanzania, Zambia, Zimbabwe
IUCN Red List: Not Evaluated
SARCA Draft: Least Concern (regional) (2010)
TFTSG Draft Red List: Vulnerable (2013)
CITES: Appendix II, as Testudinidae spp.
Kinixys spekii Gray 1863d:381, Cinixys spekii, Kinixys belliana spekii
Kinixys zombensis zombensis Hewitt 1931 (12:35)
Southeastern Hinge-back Tortoise
Type locality: “Central Africa”. Restricted to “Tanganyika Territory
east of the lakes” [Tanzania] by Loveridge (1936:218).
Homopus darlingi Boulenger 1902b:15, Kinixys darlingi,
Kinixys belliana darlingi
Type locality: “Mashonaland...Rhodesia...district about Salisbury...
at Mazoë and between Umtali and Marandellas”. Restricted to
“Mashonaland” [Zimbabwe] by Mertens and Wermuth (1955:372)
and to “Salisbury District, Mashonaland, Southern Rhodesia”
[Zimbabwe] by Fritz and Havaš (2007:286).
Testudo procterae Loveridge 1923:928, Malacochersus
procterae
Type locality: “Ikikuyu, Dodoma Dist., Tanganyika Territory”
[Tanzania]. Restricted to “Ikikuyu, south of Gulwe, Tanganyika
Territory” [Tanzania] by Fritz and Havaš (2007:286).
Kinixys australis Hewitt 1931:477, Kinixys australis australis, Kinixys belliana australis
Type locality: “White River, Eastern Transvaal” [South Africa].
Kinixys jordani Hewitt 1931:482
Type locality: “Isoka, N. Rhodesia” [Zambia].
Kinixys youngi Hewitt 1931:486
Type locality: “Nyasaland...near Livingstonia, on the shore of the
lake” [Malawi].
Kinixys australis mababiensis FitzSimons 1932:37, Kinixys
belliana mababiensis
Type locality: “Tsotsoroga Pan, Mababe Flats” [South Africa].
Flora Ihlow / Bonamanzi Game Reserve, KwaZulu Natal, South Africa
Malawi, Mozambique, South Africa, Tanzania
Kinixys belliana zombensis Hewitt 1931:469, Kinixys zombensis, Kinixys zombensis zombensis
Type locality: “Zomba, Nyassaland” [Malawi].
Kinixys belliana zuluensis Hewitt 1931:471, Kinixys
zuluensis
Type locality: “Richards Bay, Zululand” [South Africa].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Kinixys zombensis domerguei (Vuillemin 1972b) (12:35)
Madagascan Hinge-back Tortoise
[Tanzania] by Mertens and Wermuth (1955:374).
Testudo loveridgii Boulenger 1920:263, Malacochersus
loveridgii
Type locality: “Afrique orientale à Dodoma, entre Kilossa et
Tabora” [Tanzania]. Restricted to “Dodoma, Ugogo, Tanganyika
Territory” [Tanzania] by Fritz and Havaš (2007:287).
Manouria Gray 1854a
Manouria Gray 1854a:133
Type species: Manouria fusca Gray 1854a [= subjective synonym of
Testudo emys Schlegel and Müller 1840], by original monotypy.
Teleopus Le Conte 1854:187
Type species: Teleopus luxatus Le Conte 1854 [= subjective synonym of Testudo emys Schlegel and Müller 1840], by monotypy.
Thomas E.J. Leuteritz / CRM 6 / Nosy Faly, Madagascar
Madagascar (prehistoric introduction?)
Madakinixys domerguei Vuillemin 1972b:169, Kinixys belliana domerguei, Kinixys zombensis domerguei
Type locality: “Madagascar”. Restricted to “canton
d’Antsakoamanondro, N.W. de Madagascar” by Bour (1985:60).
Scapia Gray 1869a:167
Type species: Testudo (Scapia) falconeri Gray 1869 [= subjective
synonym of Testudo emys Schlegel and Müller 1840 or Testudo
phayrei Blyth 1854], by monotypy.
Manouria emys (Schlegel and Müller 1840)
Asian Giant Tortoise
Malacochersus Lindholm 1929
Testudo (Malacochersus) Lindholm 1929:285
Type species: Testudo (Malacochersus) tornieri [= Testudo tornieri
Siebenrock 1903b], by original designation.
Malacochersus tornieri (Siebenrock 1903b)
Pancake Tortoise
Fabian Schmidt / Ruaha National Park, Tanzania
(subspecies: emys = red, phayrei = purple)
(orange dots = possible trade)
Kenya, Tanzania, Zambia
IUCN Red List: Vulnerable A1bd (1996)
TFTSG Draft Red List: Critically Endangered (2013)
CITES: Appendix II, as Testudinidae spp.
Testudo tornieri Siebenrock 1903b:443, Testudo (Malacochersus) tornieri, Malacochersus tornieri
Type locality: “Bussisia am Viktoria Nyanza” [Tanzania]. Restricted
to “Bussisia, Süden des Victoria-Sees, Tanganyika-Territorium”
Bangladesh, Brunei, India (Assam, Meghalaya, Mizoram, Nagaland), Indonesia (Kalimantan, Sumatra), Malaysia (Peninsular, East), Myanmar, Singapore (extirpated), Thailand
CBFTT Account: Stanford, Wanchai, Schaffer, Schaffer, and
Thirakhupt (2015)
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Testudinidae spp.
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Manouria emys emys (Schlegel and Müller 1840)
Asian Brown Giant Tortoise
Manouria impressa (Günther 1882)
Impressed Tortoise
Chuck Schaffer / CBFTT / Tabin Wildlife Reserve, Sabah, Malaysia (East)
Brunei, Indonesia (Kalimantan, Sumatra), Malaysia (Peninsular, East), Singapore (extirpated), Thailand
Testudo emys Schlegel and Müller 1840:pl.4, Manouria
emys, Manouria emys emys, Geochelone emys, Geochelone emys emys, Testudo emys emys
David Emmett / TL / Cardamom Mts., Cambodia
Type locality: “Sumatra” [Indonesia].
Testudo emydoides Duméril and Bibron in Duméril and
Duméril 1851:4 (nomen novum), Manouria emydoides
Manouria fusca Gray 1854a:134
Type locality: “Singapore.”
Teleopus luxatus Le Conte 1854:187, Manouria luxata
Type locality: “Java” [Indonesia] (in error).
Testudo (Scapia) falconeri Gray 1869a:169 (partim, nomen
dubium), Testudo falconeri, Scapia falconeri
Type locality: “India?”
Manouria emys phayrei (Blyth 1854) (65)
Burmese Black Giant Tortoise
(orange dots = probable trade)
Cambodia, China (Yunnan), Laos, Malaysia (Peninsular),
Myanmar, Thailand, Vietnam
IUCN Red List: Vulnerable A1acd, B1+2acd (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Testudinidae spp.
Geoemyda impressa Günther 1882:343, Testudo impressa,
Geochelone impressa, Manouria impressa
Type locality: “Siam” [Thailand].
Geoemyda latinuchalis Vaillant 1894:68, Testudo latinuchalis
Type locality: “la rivière Noire, Tonkin” [Vietnam].
Testudo pseudemys Boulenger 1903a:144
Craig B. Stanford / CBFTT / Kaeng Krachan National Park, Thailand
Bangladesh, India (Assam, Meghalaya, Mizoram, Nagaland),
Myanmar, Thailand
Testudo phayrei Blyth 1854:639 (65), Scapia phayrei, Manouria emys phayrei
Type locality: “Arakan; Tenasserim Provinces...Burma”
[Myanmar].
Testudo (Scapia) falconeri Gray 1869a:169 (partim, nomen
dubium), Testudo falconeri, Scapia falconeri
Type locality: “India?”
Testudo nutapundi Reimann and Nutaphand in Nutaphand
1979:193, Geochelone nutapundi, Manouria emys nutapundi, Geochelone emys nutapundi, Manouria nutapundi
Type locality: “Northern Thailand (Tak Province) and western Central Region (Kanchanaburi Province); Assam, Burma”. Restricted to
“Nord-Thailand, Tak-Provinz” by Obst (1983:253).
Type locality: “Batang Padang district, South Perak (1,000 feet to
2,000 feet)” [Malaysia].
Psammobates Fitzinger 1835
Psammobates Fitzinger 1835:113
Type species: Psammobates geometricus Fitzinger [= Testudo geometrica Linnaeus 1758], by subsequent designation by Fitzinger
(1843:29).
Peltastes (Chersinella) Gray 1870c:8
Type species: Chersinella geometrica [= Testudo geometrica Linnaeus 1758], by subsequent designation by Hewitt (1933:259).
Lindholm (1929:286) previously designated Testudo graeca Linnaeus 1758 as type species, but it was not originally included in
Chersinella by Gray (1870c), so therefore has no validity as type.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Psammobates geometricus (Linnaeus 1758)
Geometric Tortoise
(orange dots = trade or possible misidentified)
Eric V. Goode / South African Geometric Tortoise Preserve, South Africa
Botswana, Namibia, South Africa, Zimbabwe
IUCN Red List: Least Concern [Not Listed] (1996)
SARCA Draft: Least Concern (2010)
TFTSG Draft Red List: Least Concern (2013)
CITES: Appendix II, as Testudinidae spp.
Testudo oculifera Kuhl 1820:77, Emys oculifera, Clemmys
oculifera, Chersinella oculifera, Psammobates oculifera,
Psammobates oculifer, Psammobates oculiferus
Type locality: “Cap” [Cape of Good Hope, South Africa] [in
error].
Emys occilifera Kuhl in Gray 1830e:9 (nomen novum)
Emys kuhlii Gray 1831d:73 (nomen dubium)
Type locality: Not known.
Testudo semiserrata Smith 1839a:Reptilia, pl.6, Peltastes
semiserratus
South Africa
IUCN Red List: Critically Endangered A2acde+4acde (2015);
Previously: Endangered (1996)
CITES: Appendix I
Testudo geometrica Linnaeus 1758:199, Chersine geometrica, Hydrone geometrica, Psammobates geometricus,
Peltastes geometricus, Peltastes geometrica, Peltastes
(Chersinella) geometrica, Chersinella geometrica, Psammobates geometrica, Geochelone geometrica
Type locality: “districts between Latakoo and the Tropic of Capricorn” [South Africa].
Psammobates tentorius (Bell 1828a) (61)
Tent Tortoise
Type locality: “Asia.” Restricted to “southwestern Cape Province,
South Africa” by Baard (1991:9).
Testudo luteola Daudin 1801:277
Type locality: Not known.
Peltastes geographicus Gray 1869a:173 (nomen novum),
Testudo geographica
Testudo strauchi Lidth de Jeude 1893:312, Chersinella
strauchi
Type locality: “Cape of Good Hope” [South Africa].
Psammobates oculifer (Kuhl 1820)
Serrated Tent Tortoise, Kalahari Tent Tortoise
William R. Branch / Botswana
(subspecies: tentorius = red, trimeni = purple, verroxii = green;
overlap = intergrades; orange dots = possible trade or misidentified)
Namibia, South Africa
IUCN Red List: Least Concern (2017); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix II, as Testudinidae spp.
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Psammobates tentorius tentorius (Bell 1828a)
Southern Tent Tortoise, Common Tent Tortoise
Psammobates tentorius trimeni (Boulenger 1886a)
Western Tent Tortoise
William R. Branch / Groenrivier, South Africa / male
Victor J.T. Loehr / nr. Prince Albert, Western Cape, South Africa
South Africa
Testudo tentoria Bell 1828a:420, Testudo geometrica tentoria, Peltastes tentorius, Peltastes (Chersinella) tentoria,
Chersinella tentoria, Chersinella tentoria tentoria,
Psammobates tentoria, Psammobates tentoria tentoria,
Psammobates tentorius, Psammobates tentorius tentorius,
Testudo tentoria tentoria
Type locality: “Africâ?”
Namibia (?), South Africa
TFTSG Draft Red List: Endangered (2013)
Testudo trimeni Boulenger 1886a:541, Chersinella trimeni,
Psammobates trimeni, Psammobates tentorius trimeni,
Testudo tentoria trimeni, Psammobates tentoria trimeni
Type locality: “Mouth of the Orange River” [Little Namaqualand,
South Africa].
Psammobates tentorius verroxii (Smith 1839)
Northern Tent Tortoise
Testudo geometrica nigriventris Gray 1856b:8
Type locality: “South Africa.”
Chersinella tentoria albanica Hewitt 1933b:266, Psammobates tentoria albanica
Type locality: “neighbourhood of farm Mayfair, Albany District”
[South Africa].
Chersinella tentoria tentorioides Hewitt 1933b:268, Psammobates tentoria tentorioides
Type locality: “Bowden Hall, Middelburg district, C.P.” [Cape
Province, South Africa].
Chersinella tentoria piscatella Hewitt 1933b:269, Psammobates tentoria piscatella
Type locality: “Little Fish River, Somerset East district” [South
Africa].
Chersinella tentoria subsulcata Hewitt 1933b:270
Type locality: “farm “Brighton” near Steytlerville” [South Africa].
Chersinella tentoria karuica Hewitt 1933b:272, Psammobates tentoria karuica
Type locality: “farm “Drogekloof” near Klaarstroom, Prince Albert
district” [South Africa].
Chersinella tentoria duerdeni Hewitt 1933b:279, Psammobates tentoria duerdeni
Type locality: “Graaff Reinet, C.P.” [Cape Province, South Africa].
Chersinella tentoria lativittata Hewitt 1933b:281
Type locality: “Willowmore” [South Africa].
Chersinella tentoria karuella Hewitt 1933b:283
Type locality: “Uniondale, C.P.” [Cape Province, South Africa].
Chersinella tentoria hexensis Hewitt 1933b:286
Type locality: “Hex River, Worcester district” [Cape Province,
South Africa].
Victor J.T. Loehr / nr. Beaufort West, Northern Cape, South Africa
Namibia, South Africa
Testudo verroxii Smith 1839b:Reptilia,pl.8, Peltastes verroxii, Chersinella verroxii, Chersinella verroxii verroxii,
Psammobates tentorius verroxii, Testudo tentoria verroxii,
Psammobates tentoria verroxii, Testudo tentorius verroxii
Type locality: “South Africa, near the sources of the Garriep or
Orange River.” Restricted to “somewhere north of Aliwal North,
between the Orange and Caledon Rivers...roughly 260 miles east of
Niekirk’s Hope” [South Africa] by Power (1932:466).
Peltastes verreauxii Gray 1870b:656 (nomen novum), Testudo verreauxii, Psammobates verreauxii
Testudo fiski Boulenger 1886a:542, Testudo tentoria fiski,
Chersinella fiski, Chersinella fiski fiski, Psammobates fiski,
Psammobates fiski fiski
Type locality: “De Aar, not far from Hopetown” [South Africa].
Testudo smithi Boulenger 1886a:542, Chersinella verroxii
smithi, Testudo smithi smithi, Testudo verroxii smithi
Type locality: “S. Africa” [South Africa].
Testudo seimundi Boulenger 1903b:216, Chersinella fiski
seimundi
Type locality: “3 miles east of Deelfontein” [Richmond District,
Cape Province, South Africa].
Testudo boettgeri Siebenrock 1904a:194 (junior homonym,
not = Testudo graeca boettgeri Mojsisovics 1889), Chersinella verroxii boettgeri, Chersinella boettgeri
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “Groß-Namaland in Südwestafrika” [Great Namaqualand, Namibia].
Homopus bergeri Lindholm 1906:348 (partim), Testudo
bergeri, Chersinella verroxii bergeri, Testudo smithi
bergeri, Testudo verroxii bergeri
149
Pyxis arachnoides arachnoides Bell 1827 (07:70)
Spider Tortoise, Common Spider Tortoise
Type locality: “nach Gibeon in Deutsch-Südwestafrika...möglicherweise weiter im Innern Südafrikas” [Namibia or South Africa].
Testudo oscarboettgeri Lindholm 1929:295 (nomen novum)
Chersinella schonlandi Hewitt 1934:303
Type locality: “Namaqualand, C.P.” [Little Namaqualand, Cape
Province, South Africa].
Chersinella fiski cronwrighti Hewitt 1934:317, Psammobates
fiski cronwrighti
Type locality: “Hanover, C.P.” [Cape Province, South Africa].
Chersinella fiski orangensis Hewitt 1934:319
Type locality: “between Philipstown and Petrusville district” [Cape
Province, South Africa].
Chersinella fiski colesbergensis Hewitt 1934:321, Psammobates fiski colesbergensis
Type locality: “Colesberg” [Cape Province, South Africa].
Chersinella fiski grica Hewitt 1934:323
Type locality: “Marydale, situated about midway between Prieska
and Kenhardt” [Prieska District, Cape Province, South Africa].
Chersinella fiski gricoides Hewitt 1934:326
Type locality: “Niekerks Hope, about midway between Griquatown
and Prieska, C.P.” [Niekerkshoop, Hay District, Cape Province,
South Africa].
Chersinella fiski amasensis Hewitt 1934:333
Type locality: “Ukamas district” [Cape Province, South Africa].
Psammobates depressa FitzSimons 1938:154
Michael Ogle / Anakao, Madagascar
Madagascar
Pyxis arachnoides Bell 1827:395, Testudo (Pyxis) arachnoides, Testudo arachnoides, Pyxis arachnoidea, Bellemys
arachnoides, Pyxis arachnoides arachnoides
Type locality: Not known. Restricted to “Soalara (Baie de SaintAugustin), sud-ouest de Madagascar” by Bour (1979:153).
Testudo (Pyxis) aranoides Gray 1830e:6 (nomen novum),
Pyxis aranoides
Pyxis madagascariensis Lesson 1831a:120
Type locality: “Madagascar.”
Pyxis arachnoides brygooi (Vuillemin and Domergue 1972)
Northern Spider Tortoise
Type locality: “8 miles west of Aus, Great Namaqualand”
[Namibia].
Pyxis Bell 1827
Pyxis Bell 1827:395
Type species: Pyxis arachnoides Bell 1827, by original monotypy.
Acinixys Siebenrock 1902b:12
Type species: Acinixys planicauda [= Testudo planicauda Grandidier
1867], by original monotypy.
Bellemys Williams 1950:512 (nomen novum)
Pyxoides Vuillemin and Domergue 1972:193
Type species: Pyxoides brygooi Vuillemin and Domergue 1972, by
original monotypy.
Pyxis arachnoides Bell 1827
Spider Tortoise
Roger Bour / Salary, nr. Forêt des Mikea, Madagascar
Madagascar
Pyxoides brygooi Vuillemin and Domergue 1972:193, Pyxis
arachnoides brygooi
Type locality: “la côte Sud-Ouest, entre Morombe et Tuléar, aux
alentours du lac Ihotry, dans la forêt des Mikea” [Madagascar].
Restricted to “Ampanonga (N.-W. Lac Ihotry), sud-ouest de Madagascar” by Bour (1979:153).
Pyxis arachnoides oblonga Gray 1869a
Southern Spider Tortoise
(subspecies: arachnoides = red, brygooi = purple, oblonga = orange)
Madagascar
IUCN Red List: Critically Endangered A4cd, E (2008); Previously: Vulnerable (1996)
CITES: Appendix I
Anders G.J. Rhodin / TCC / CRM 6 / Berenty, Madagascar
Madagascar
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Pyxis arachnoidea oblonga Gray 1869a:173, Pyxis arachnoides oblonga
Stigmochelys pardalis (Bell 1828a) (07:71, 10:27)
Leopard Tortoise
Type locality: Not known. Restricted to “Asia” by Gray (1873j:14);
to “Madagascar; Mauritius” by Boulenger (1889:145); and to “Cap
Sainte-Marie, province de Tuléar (Toliara), sud du Madagascar” by
Bour (1982b:30).
Pyxis arachnoides matzi Bour 1979:143
Type locality: “Cap Sainte-Marie (Province de Tuléar), extrême sud
du Madagascar.”
Pyxis planicauda (Grandidier 1867)
Flat-tailed Tortoise, Flat-shelled Spider Tortoise
Victor J.T. Loehr / nr. Beaufort West, Northern Cape, South Africa
Miguel Pedrono / CRM 6 / Kirindy Forest, Madagascar
Madagascar
IUCN Red List: Critically Endangered A4acd (2008); Previously: Endangered (1996)
CITES: Appendix I
Testudo planicauda Grandidier 1867:223, Acinixys planicauda, Pyxis (Acinixys) planicauda, Pyxis planicauda
Type locality: “Mouroundava...cote sud-ouest de Madagascar”
[Morondava].
Testudo morondavaensis Vuillemin 1972a:127
Type locality: “Morondava (côte sud-ouest de Madagascar).”
Stigmochelys Gray 1873j (07:52, 10:26)
Testudo (Stigmochelys) Gray 1873j:5
Type species: Testudo (Stigmochelys) pardalis Bell 1828a, by original
monotypy.
Megachersine Hewitt 1933b:257
Type species: Megachersine pardalis [= Testudo pardalis Bell
1828a], by original designation.
(orange dots = introduced or questionable)
Angola, Botswana, Burundi, Ethiopia, Kenya, Malawi, Mozambique, Namibia, Rwanda, Somalia, South Africa, South
Sudan, Swaziland, Tanzania, Uganda, Zambia, Zimbabwe
IUCN Red List: Least Concern (2015); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix II, as Testudinidae spp.
Testudo pardalis Bell 1828a:420, Geochelone (Geochelone)
pardalis, Geochelone pardalis, Testudo (Stigmochelys)
pardalis, Megachersine pardalis, Testudo pardalis pardalis, Geochelone pardalis pardalis, Stigmochelys pardalis,
Centrochelys pardalis, Centrochelys pardalis pardalis,
Stigmochelys pardalis pardalis, Psammobates pardalis
Type locality: “Promont. Bonae Spei” [Cape of Good Hope,
South Africa].
Testudo biguttata Cuvier 1829:10 (nomen nudum)
Testudo armata Boie in Gray 1830e:4 (nomen nudum)
Testudo bipunctata Gray 1830e:4 (nomen nudum)
Testudo pardalis babcocki Loveridge 1935:4 (10:27), Geochelone pardalis babcocki, Geochelone babcocki, Stigmochelys pardalis babcocki
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “western slopes of Mount Debasien, Karamoja,
Uganda at 5,500 feet.”
Testudo Linnaeus 1758 (07:72) (56, 70)
Testudo Linnaeus 1758:197
Type species: Testudo graeca Linnaeus 1758 [= subjective synonym
of Testudo ibera Pallas 1814], by subsequent designation by
Fitzinger (1843:29) and Lindholm (1929:284); not Testudo graeca
Auct. [= subjective synonym of Testudo hermanni Gmelin 1789]
by earlier designation by Bell (1828c:514).
Chersine Merrem 1820:29 (09:40)
151
kleinmanni Lortet 1883], by original designation.
Protestudo Chkhikvadze 1970:245
Type species: Protestudo bessarabica [= Testudo bessarabica †
Riabinin 1918], by original designation.
Furculachelys Highfield 1990:1
Type species: Furculachelys nabeulensis Highfield 1990 [= Testudo
graeca nabeulensis], by subsequent designation by Fritz and
Havaš (2007:295).
Testudo (Testudo) graeca Linnaeus 1758 (07:73, 11:12, 12:36, 14:35) (71, 72)
Spur-thighed Tortoise, Greek Tortoise, Moorish Tortoise
Type species: “T. graeca »L.« auct. = T. hermanni Gmelin” [=
Testudo hermanni Gmelin 1789], by subsequent designation by
Lindholm (1929:286).
Chersini Merrem in Gray 1825:210 (nomen novum)
Chersus Wagler 1830b:138
Type species: Chersus marginatus [Testudo marginata Schoepff
1793], by monotypy.
Cherseus Gray 1856b:10 (nomen novum)
Peltastes Gray 1869a:167,171 (junior homonym, not = Peltastes
Illiger 1807 [= Hymenoptera] or Peltastes Agassiz 1838
[= Echinodermata] or Peltastes Fischer-Waldheim 1839 [=
Orthoptera])
Type species: Testudo graeca Linnaeus 1758 [= Peltastes graecus
sensu Gray 1869] [= subjective synonym of Testudo ibera Pallas
1814], by subsequent designation by Lindholm (1929:286).
1
Peltonia Gray 1872c:4 (nomen novum)
Testudo (Medaestia) Wussow 1916:170 (09:40)
Type species: Medaestia graeca sensu Wussow 1916 [= subjective
synonym of Testudo hermanni Gmelin 1789] by subsequent
designation by Mertens (1949:232).
Testudo (Pseudotestudo) Loveridge and Williams 1957:166, 276
Type species: Testudo (Pseudotestudo) kleinmanni [= Testudo kleinmanni Lortet 1883], by original designation.
Protestudo Chkhikvadze 1970:245
Type species: Protestudo bessarabica [= Testudo bessarabica †
Riabinin 1918], by original designation.
Furculachelys Highfield 1990:1
Type species: Furculachelys nabeulensis Highfield 1990 [= Testudo
graeca nabeulensis], by subsequent designation by Fritz and
Havaš (2007:295).
2
Eurotestudo Lapparent de Broin, Bour, Parham, and Perälä
2006a:803
Type species: Eurotestudo hermanni [= Testudo hermanni Gmelin
1789], by original designation.
Testudo (Testudo) Linnaeus 1758 (07:72) (70)
Testudo Linnaeus 1758:197
Type species: Testudo graeca Linnaeus 1758 [= subjective synonym
of Testudo ibera Pallas 1814], by subsequent designation by
Fitzinger (1843:29) and Lindholm (1929:284); not Testudo graeca
Auct. [= subjective synonym of Testudo hermanni Gmelin 1789]
by earlier designation by Bell (1828c:514).
Chersus Wagler 1830b:138
Type species: Chersus marginatus [Testudo marginata Schoepff
1793], by monotypy.
Cherseus Gray 1856b:10 (nomen novum)
Peltastes Gray 1869a:167,171 (junior homonym, not = Peltastes
Illiger 1807 [= Hymenoptera] or Peltastes Agassiz 1838
[= Echinodermata] or Peltastes Fischer-Waldheim 1839 [=
Orthoptera])
Type species: Testudo graeca Linnaeus 1758 [= Peltastes graecus
sensu Gray 1869] [= subjective synonym of Testudo ibera Pallas
1814], by subsequent designation by Lindholm (1929:286).
Peltonia Gray 1872c:4 (nomen novum)
Testudo (Pseudotestudo) Loveridge and Williams 1957:166, 276
Type species: Testudo (Pseudotestudo) kleinmanni [= Testudo
3
(subspecies [map #]: graeca = red [1], armeniaca = purple [3],
buxtoni = orange [3], cyrenaica = pink [2], ibera = brown [2,3],
marokkensis = green [1], nabeulensis = gray [1], soussensis = tourmaline [1],
terrestris = olive [2,3], zarudnyi = blue [3]; overlap = intergrades [2,3];
orange dots = uncertain, possible or probable trade, or introduced)
Afghanistan (?), Algeria, Armenia, Azerbaijan, Bulgaria,
Georgia, Greece, Iran (Alborz, Ardabil, Chahar Mahal
Va Bakhtiari, East Azarbaijan, Esfahan, Fars, Gilan,
Hamedan, Hormozgan, Ilam, Kerman, Kermanshah, Khorasan Razavi, Kohgiluyeh Va Boyer Ahmad, Kordestan,
Lorestan, Markazi, Qazvin, Qom, Sistan Va Baluchestan,
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South Khorasan, Tehran, West Azarbaijan, Yazd, Zanjan),
Iraq, Israel, Jordan, Kosovo, Lebanon, Libya, Macedonia,
Moldova, Morocco, Pakistan (?), Palestine (West Bank),
Romania, Russia (Chechnya [?], Dagestan, Krasnodarskiy), Serbia, Spain, Syria, Tunisia, Turkey (Asian,
European), Turkmenistan (?)
Introduced: Egypt, France, Italy (Continental, Sardinia [prehistoric], Sicily), Malta (?), Spain (Continental, Balearic
Islands)
IUCN Red List: Global: Vulnerable A1cd (1996); Regional:
Europe: Vulnerable A2bcde+4bcde (2004)
CITES: Appendix II, as Testudinidae spp.
Testudo (Testudo) graeca armeniaca Chkhikvadze and Bakradze
1991 (11:12)
Araxes Tortoise
Testudo (Testudo) graeca graeca Linnaeus 1758 (09:41)
Mediterranean Spur-thighed Tortoise
Pavel Široký / Kuzeh Rash, West Azarbaijan Prov., Iran
Armenia, Azerbaijan, Georgia, Iran (Ardabil, East Azarbaijan,
West Azarbaijan), Russia (Dagestan), Turkey
Testudo graeca pallasi Chkhikvadze 1989:67 (nomen
nudum)
Testudo graeca armeniaca Chkhikvadze 1989:67 (nomen
nudum)
Testudo graeca armeniaca Chkhikvadze and Bakradze
1991:60, Testudo armeniaca, Testudo terrestris armeniaca
Type locality: “Мегри, ЮВ Армянской ССР” [Megri, SE Armenian SSR] [Armenia].
Testudo graeca pallasi Chkhikvadze and Bakradze 2002:276,
Testudo pallasi, Testudo marginata pallasi
Chris Leone / Morocco [captivity]
Algeria, Morocco, Spain
Introduced: Spain (Balearic Islands)
Testudo graeca Linnaeus 1758:198, Testudo graeca graeca,
Peltastes graecus
Type locality: “Africa.” Restricted to “Santa Cruz in Barbarie” [Algeria] by Leven (1764:56), emended to “Santa-Cruz (alte spanische
Festung bei Oran in der Algérie)” by Strauch (1862:67).
Testudo pusilla Linnaeus 1758:199 (senior homonym, not =
Testudo pusilla † Bergounioux 1936), Chersine pusilla
Type locality: “Дагестан, окрестности села Гиляры-Даг”
[Dagestan, near the village of Gilyary-Dag]. Emended here to
“Gilyar, near Gilyary-Dag mountain, Magaramkent District,
Dagestan, Russia (41.558499 N, 48.257204 E).”
Testudo dagestanica Chkhikvadze, Mazanaeva, and Shammakov 2011:337 (11:12), Testudo graeca dagestanica
Type locality: “Папас (Южный Дагестан)” [Papas (Southern
Dagestan)] [Russia].
Testudo (Testudo) graeca buxtoni Boulenger 1921 (10:28)
Buxton’s Tortoise
Type locality: “India.” Restricted to “Santa-Cruz (alte spanische
Festung bei Oran in der Algérie)” by Strauch (1862:67).
Testudo mauritanica Duméril and Bibron 1835:44, Testudo
graeca mauritanica, Chersus mauritanicus, Peltastes
mauritanicus, Testudo ibera mauritanica
Type locality: “Mauritanie..[&]..les côtes occidentales de la mer
Caspienne..[&]..les environs d’Alger.” Restricted to “Umgebung
von Algier” [Algeria] by Mertens and Wermuth (1955:378).
Testudo whitei Bennett in White 1836:361 (09:41), Peltastes
marginatus whitei, Testudo marginata whitei, Furculachelys whitei, Testudo graeca whitei
Type locality: Not known. Restricted to “Algiers and its environs,
Algeria” by Highfield and Martin (1989a:21).
Pavel Široký / Dehlili, Kermanshah Prov., Iran
Iran (Alborz, Ardabil, Chahar Mahal Va Bakhtiari, East Azarbaijan, Esfahan, Fars, Gilan, Hamedan, Ilam, Kermanshah, Kordestan, Lorestan, Markazi, Qazvin, Tehran, West
Azarbaijan, Zanjan), Turkey
Testudo ecaudata Pallas 1814:19 (10:28) (nomen dubium and
junior homonym, not = Testudo ecaudata Daudin 1801)
Type locality: “nemorosis Persiae mari caspio conterminis”
[forests of Persia along the Caspian Sea] [Iran].
Testudo buxtoni Boulenger 1921:251, Testudo terrestris
buxtoni, Testudo ibera buxtoni, Testudo graeca buxtoni
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “Manjil, between Resht and Kasuin, South Coast of
the Caspian Sea, on a hill-side about 7,000–7,500 feet...northern
Persia” [Iran].
Testudo perses Perälä 2002c:81, Testudo graeca perses,
Testudo ibera perses
Type locality: “vicinity of Lalabad village, some 25 mi NW
Kermānshāh, Kermānshāhān province, W Iran...34°27'N
46°50'E.”
Testudo (Testudo) graeca cyrenaica Pieh and Perälä 2002
Cyrenaican Spur-thighed Tortoise
153
[Taurus Mts., Cilicia, Turkey].
Testudo ibera racovitzai Calinescu 1931:169
Type locality: “Turtucaia (jud. Durostor)” [Romania] [now
Tutrakan, Silistra, Bulgaria].
Testudo graeca nikolskii Chkhikvadze and Tuniyev
1986:618, Testudo ibera nikolskii, Testudo terrestris nikolskii, Testudo nikolskii
Type locality: “Поселок Небуг Туапсинского района
(Краснодарский край)” [Nebug Settlement, Tuapse Co., Krasnodar District] [Russia].
Testudo graeca pontica Khosatzky 1987:58 (nomen nudum)
Testudo (Testudo) graeca marokkensis Pieh and Perälä 2004 (09:42)
Morocco Tortoise
Chris Leone / Libya [captivity]
Libya
Testudo graeca cyrenaica Pieh and Perälä 2002:8, Testudo
cyrenaica
Type locality: “Derna 32°46'N, 22°39'E (= Darnah, Cyrenaika
Ostlibyen” [Libya].
Testudo (Testudo) graeca ibera Pallas 1814 (11:12)
Asia Minor Tortoise
Chris Leone / Fez, Morocco [captivity]
Morocco
Introduced: Spain
Testudo graeca marokkensis Pieh and Perälä 2004:22 (09:42)
Type locality: “Tarmilete (33°23'N 6°04'W)” [Morocco].
Testudo graeca lamberti Pieh and Perälä 2004:33 (09:42)
Type locality: “22 km nördlich von Tetuan (= Tetouan; Tétuan
35°34'N 5°22'W)” [Morocco].
Testudo (Testudo) graeca nabeulensis (Highfield 1990)
Nabeul Tortoise
Alexander A. Inozemtsev / CCB / TCF / nr. Novorossiysk, Krasnodarskiy, Russia
Albania (?), Armenia, Azerbaijan, Bulgaria, Georgia, Greece,
Kosovo, Macedonia, Moldova, Romania, Russia (Krasnodarskiy), Serbia, Turkey
IUCN Red List: The currently synonymized taxon Testudo graeca nikolskii listed as Critically Endangered
A1abcde+2bcde (1996)
Testudo ibera Pallas 1814:18, Chersus iberus, Cherseus
iberus, Medaestia ibera, Testudo graeca ibera, Testudo
ibera ibera, Testudo terrestris ibera
Type locality: “Caucasi meridionalibus..&..convallibus orae
meridionalis montosae Chersonesi tauricae” [southern Caucasus
..&.. steep mountainous southern coast of Crimea]. Restricted to
“Iberia, haud pro cul a Tiflisio” [Tbilisi, Georgia] by Eichwald
(1831:196); to “das Gebiet des mittleren Kura-Tales im Kaukasus” [Georgia] by Mertens (1946:113); and to “Tbilissi (Tiflis),
Géorgie, URSS” [Georgia] by Bour (1987a:112).
Testudo georgicana Güldenstedt in Pallas 1814:18 (nomen
nudum)
Testudo iberia Blyth 1854:642 (64) (nomen novum)
Testudo ibera bicaudalis Venzmer 1920:289
Type locality: “cilicischen Taurus in der kleinasiatischen Türkei”
Norbert Halasz / Nabeul, Tunisia [captivity]
Libya, Tunisia
Introduced: Italy (Sardinia, Sicily)
Testudo flavominimaralis Highfield and Martin 1989b:[9]
(nomen dubium), Testudo graeca flavominimaralis
Type locality: “North Africa.” Restricted to “Libya” by Highfield
(1990:41).
Furculachelys nabeulensis Highfield 1990:32, Testudo
nabeulensis, Testudo graeca nabeulensis
Type locality: “forested area in the region of Nabeul, Tunisia.”
Restricted to “Waldgebiet 7–8 km nordwestlich von Nabeul
Richtung Grombalia (Tunesien)” [Tunisia] by Pieh and Perälä
(2004:42).
Testudo graeca sarda Ballasina 1995:166 (nomen nudum)
Testudo graeca sardinia van der Kuyl, Ballasina, Dekker,
Maas, Willemsen, and Goudsmit 2002:180 (nomen
nudum)
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Testudo (Testudo) graeca soussensis Pieh 2001
Souss Valley Tortoise
Andreas Nöllert / east of Taroudannt, Morocco
Morocco
Testudo graeca soussensis Pieh 2001:211, Testudo soussensis
Type locality: “Umgebung von Agadir 30°28'N, 9°55'W, Sousstal,
Südwest-Marokko” [Morocco].
Testudo (Testudo) graeca terrestris Forskål 1775 (10:29) (72)
Mesopotamian Tortoise
Testudo (Testudo) graeca zarudnyi Nikolsky 1896
Iranian Tortoise
Asghar Mobaraki / Saravan, Sistan Va Baluchestan, Iran [captivity]
Afghanistan (?), Iran (Esfahan, Fars, Hormozgan, Kerman,
Khorasan Razavi, Sistan Va Baluchestan, South Khorasan,
Yazd), Pakistan (?), Turkmenistan (?)
Testudo zarudnyi Nikolsky 1896:369, Testudo graeca zarudnyi, Testudo ibera zarudnyi, Testudo terrestris zarudnyi
Type locality: “Persia orientali” [Iran]. Restricted to “montibus
provinciae Birdschan in Persiae orientali” [Iran] by Nikolsky (1897:308); and to “Birjand in Khorāsan province, NE
Iran (32º53' N 59º03' E)” by lectotype designation by Perälä
(2002c:84).
Testudo (Testudo) kleinmanni Lortet 1883 (07:74) (73)
Egyptian Tortoise
Norbert Halasz / Aleppo, Syria [captivity]
Iraq, Israel, Jordan, Lebanon, Palestine (West Bank), Syria,
Turkey
Testudo terrestris Forskål 1775:viii,12 (junior homonym, not
= Testudo terrestris Garsault 1764 or Testudo terrestris
Fermin 1765; nomen conservandum, ICZN 1963), Testudo
graeca terrestris, Testudo terrestris terrestris, Testudo
ibera terrestris
Eric V. Goode / TCC / Egypt or Libya [captivity]
Type locality: “Lohajae..&..Kahirae..&..Aleppo..&..Libanon” [Al
Luhayyah..&..Cairo..&..Aleppo..&..Mount Lebanon]. Restricted
to “Arabien” [Saudi Arabia] by Wermuth (1956:402); to “Libanon-Gebirge, Israel” by Wermuth (1958:152); and to “Aleppo
(Alep, Halab; 36°12' N, 37°09' E), Syria (Syrian Arab Republic)”
by neotype designation by Perälä and Bour (2004:117).
Testudo zolhafa Forsskål in Gray 1830e:5 (nomen nudum)
Testudo floweri Bodenheimer 1935:197 (72), Testudo graeca
floweri, Testudo terrestris floweri, Testudo ibera floweri
Type locality: “the Negeb...Palestine” [Israel]. Restricted to
“Negev, Palestine (environs de Gaza, Israël)” [Israel] by Bour
(1989:14).
Testudo graeca anamurensis Weissinger 1987:14, Testudo
ibera anamurensis, Testudo terrestris anamurensis, Testudo anamurensis
Type locality: “Strand von Anamurum, 7 km westlich von
Anamur, SW–Küste der Türkei” [Turkey].
Testudo antakyensis Perälä 1996:16, Testudo graeca antakyensis, Testudo terrestris antakyensis, Testudo ibera
antakyensis
Type locality: “mountains to the east of Antakya in southern
Turkey.”
(orange dots = probable trade or extirpated)
Egypt, Israel, Libya, Palestine? (Gaza? [extirpated])
IUCN Red List: Critically Endangered A2abcd+3d (2003); Previously: Endangered (1996)
CITES: Appendix I
Testudo leithii Günther 1869:502 (junior homonym, not =
Testudo leithii † Carter 1852), Peltastes leithii, Medaestia
leithii
Type locality: “Sindh” [Pakistan] [in error].
Testudo kleinmanni Lortet 1883:188, Pseudotestudo kleinmanni, Testudo kleinmanni kleinmanni
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “la basse Égypte, surtout dans les environs
d’Alexandrie” [Egypt].
Testudo werneri Perälä 2001:570 (07:74), Testudo kleinmanni
werneri
Type locality: “Northern Negev desert, Israel (14 km south of
Be’er Sheva).”
Testudo (Testudo) marginata Schoepff 1793 (07:75, 11:12, 11:13)
Marginated Tortoise
Rethymnon” [Greece].
Testudo marginata sarda Mayer 1992:95
Type locality: “Olbia...Sardinia” [Italy].
Testudo weissingeri Bour 1996:30, Testudo marginata
weissingeri
Type locality: “Kardamili, Messénie, Grèce” [Greece].
Testudo (Agrionemys) Khosatzky and Mlynarski 1966
(07:72, 09:40) (70)
Testudinella Gray 1870c:12 (junior homonym, not = Testudinella Bory de Saint-Vincent 1822 [= Rotatoria])
Type species: Testudinella horsfieldii [= Testudo horsfieldii Gray
1844], by original monotypy.
Agrionemys Khosatzky and Mlynarski 1966:123 (nomen novum)
Type species: Agrionemys horsfieldi [= Testudo horsfieldii Gray
1844], by original designation.
Testudo (Agrionemys) horsfieldii Gray 1844 (07:77, 08:15, 09:43, 10:30, 14:38)
Central Asian Tortoise, Steppe Tortoise, Horsfield’s Tortoise
Andreas Nöllert / Marathea, Peloponnese, Greece
Albania, Greece
Introduced: Cyprus, Italy (Continental, Sardinia [prehistoric])
IUCN Red List: Least Concern (2004); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix II, as Testudinidae spp.
Testudo tabulata campanulata Walbaum 1782:124 (unavailable name)
Testudo marginata Schoepff 1793:52, Chersine marginata,
Chersus marginatus, Peltastes marginatus, Peltastes
marginata, Testudo marginata marginata
Type locality: Not known. Restricted to “Morea oggi Pelopponneso, Grecia” [Greece] by Bruno (1986:238), and “Greece, probably
province of Attica, Stereá Eláda” by Bour (1987a:111).
Testudo graja Hermann in Schoepff 1793:52
Type locality: Not designated. Restricted to “Griechenland”
[Greece] by Mertens and Wermuth (1955:380).
Testudo campanulata Walbaum in Gray 1830e:4 (nomen
nudum)
Testudo graji Gray 1830e:4 (nomen novum)
Testudo campanulata Walbaum in Strauch 1862:65
Type locality: “Griechenland” [Greece].
Peltastes marginatus melas Gray 1870c:10
Type locality: “Greece.”
Testudo nemoralis Schreiber 1875:557
Type locality: “Griechenland und Süditalien” [Greece and Italy].
Restricted to “Griechenland” [Greece] by Mertens and Wermuth
(1955:380).
Testudo marginata cretensis † Bachmayer, Brinkerink, and
Symeonidis 1975:111 [Pleistocene, Greece (Crete)]
Type locality: “Nordküste der Insel Kreta in der Nähe von
(subspecies: horsfieldii = red, bogdanovi = purple,
kazachstanica = green, kuznetzovi = brown, rustamovi = blue;
orange dots = uncertain, possible trade, or introduced)
Afghanistan, China (Xinjiang), Iran (Golestan, Khorasan
Razavi, North Khorasan, Semnan, South Khorasan), Kazakhstan, Kyrgyzstan, Pakistan, Tajikistan, Turkmenistan,
Uzbekistan
Introduced: Latvia
IUCN Red List: Vulnerable A2d (1996)
CITES: Appendix II, as Testudinidae spp.
Testudo (Agrionemys) horsfieldii horsfieldii Gray 1844 (10:30)
Central Asian Tortoise, Steppe Tortoise, Horsfield’s Tortoise
Afghanistan, Pakistan
Testudo horsfieldii Gray 1844:7, Testudinella horsfieldii,
Homopus horsfieldii, Testudo (Homopus) horsfieldii,
Medaestia horsfieldii, Agrionemys horsfieldii, Testudo
horsfieldi, Testudo horsfieldii horsfieldii, Agrionemys
horsfieldii horsfieldii
Type locality: “India, Affghanistan [p.7]...Cabul [p.vi].” Restricted
to “Afghanistan” by Günther (1864:7).
Homopus burnesii Blyth 1854:642 (65)
Type locality: “Afghanistan.”
Testudo baluchiorum Annandale 1906:75 (10:28), Agrionemys
horsfieldii baluchiorum
Type locality: “Baluchistan” [Pakistan].
Testudo (Agrionemys) horsfieldii bogdanovi Chkhikvadze in
Chkhikvadze, Brushko, and Kubykin 2008 (10:30)
Fergana Valley Steppe Tortoise
Kyrgyzstan, Tajikistan, Uzbekistan
155
156
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Agrionemys bogdanovi Chkhikvadze in Chkhikvadze, Brushko, and Kubykin 2008:100, Testudo horsfieldii bogdanovi,
Agrionemys horsfieldii bogdanovi
Type locality: “Узбекистан (Окрестности городов Бухара,
Самарканд, Карши), Восточный Туркменистан (окрестности
Чарджоу) и Киргизстан (Чуйская долина и окрестности
города Ош)” [Uzbekistan (vicinity of the cities of Bukhara,
Samarkand, Karshy), East Turkmenistan (near Chardzhou) and
Kyrgyzstan (Chuy valley and vicinity of Osh)]. Restricted to
“Ферганская долина” [Ferghana Valley] [Uzbekistan] by Chkhikvadze et al. (2009:49).
Testudo (Agrionemys) horsfieldii rustamovi Chkhikvadze, Amiranashvili, and Ataev 1990 (09:43, 10:30)
Kopet-Dag Steppe Tortoise
Testudo (Agrionemys) horsfieldii kazachstanica Chkhikvadze
1988 (09:43, 10:30, 14:38)
Kazakhstan Steppe Tortoise
Andreas Nöllert / Kara-Kum Desert, N. of Ashgabat, Turkmenistan
Afghanistan, Iran (Golestan, Khorasan Razavi, North Khorasan, Semnan), Turkmenistan
Agrionemys horsfieldi rustamovi Chkhikvadze, Amiranashvili, and Ataev 1990:73, Agrionemys horsfieldii rustamovi,
Testudo horsfieldii rustamovi, Agrionemys rustamovi
Type locality: “село Мадау (Кизыл-Атрекский район), ЮгоЗападный Туркменистан” [Madau Village, Kizyl Atrek Region,
south-western Turkmenistan].
Shi Haitao / Huocheng Nature Reserve, Xinjiang, China
Afghanistan, China (Xinjiang), Kazakhstan, Kyrgyzstan,
Tajikistan, Turkmenistan, Uzbekistan
Agrionemys horsfieldi kazachstanica Chkhikvadze 1988:110,
Testudo horsfieldii kazachstanica, Agrionemys horsfieldii
kazachstanica, Agrionemys kazachstanica
Type locality: “Южное Прибалхашье, поселок Каратал” [southern Pribalkhashye (= Balkash), Karatal village]” [Kazakhstan].
Agrionemys kazachstanica terbishi Chkhikvadze 2009:60
(10:30, 14:38)
, Testudo horsfieldii terbishi, Agrionemys horsfieldii terbishi
Type locality: “Монголия, г. Кобдо” [Mongolia, g. Kobdo]
[probable trade specimen].
Testudo (Agrionemys) horsfieldii kuznetzovi Chkhikvadze, Ataev,
Shammakov, and Zatoka in Chkhikvadze, Ataev, and
Shammakov 2009 (10:30)
Turkmenistan Steppe Tortoise
Turkmenistan, Uzbekistan
Agrionemys kazachstanica kuznetzovi Chkhikvadze, Ataev,
Shammakov, and Zatoka in Chkhikvadze, Ataev, and
Shammakov 2009:52, Testudo horsfieldii kuznetzovi,
Agrionemys horsfieldii kuznetzovi
Type locality: “впадины Акчакая (Северный Туркменистан, к
западу от г. Дашогуз – бывш. Ташауз)” [Akchakai depression (North Turkmenistan, west of the city of Dashoguz – prev.
Tashauz)].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Testudo (Chersine) Merrem 1820 (07:72, 09:40) (70)
Chersine Merrem 1820:29
(09:40)
157
Testudo (Chersine) hermanni hermanni Gmelin 1789 (07:76, 14:36)
Western Hermann’s Tortoise
Type species: “T. graeca »L.« auct. = T. hermanni Gmelin” [=
Testudo hermanni Gmelin 1789], by subsequent designation by
Lindholm (1929:286).
Chersini Merrem in Gray 1825:210 (nomen novum)
Testudo (Medaestia) Wussow 1916:170 (09:40)
Type species: Medaestia graeca sensu Wussow 1916 [= subjective
synonym of Testudo hermanni Gmelin 1789] by subsequent
designation by Mertens (1949:232).
Eurotestudo Lapparent de Broin, Bour, Parham, and Perälä
2006a:803
Type species: Eurotestudo hermanni [= Testudo hermanni Gmelin
1789], by original designation.
Testudo (Chersine) hermanni Gmelin 1789 (14:36) (74)
Hermann’s Tortoise
Albert Bertolero / CBFTT / Albera, Catalonia, Spain
France (Continental, Corsica), Italy (Continental, Sicily), Spain
(Continental)
Introduced: Italy (Sardinia [prehistoric introduction]), Spain
(Balearic Islands)
IUCN Red List: Endangered B1+2abcde (1996)
Testudo hermanni Gmelin 1789:1041, Testudo hermanni
hermanni, Protestudo hermanni, Agrionemys hermanni,
Eurotestudo hermanni, Chersine hermanni, Chersine
hermanni hermanni
Type locality: Not designated. Restricted to “Collobrières, massif
de Maures, Var, France” by Bour (1987a:116).
Testudo graeca bettai Lataste 1881:396
Type locality: Not known.
Testudo globosa † Portis 1890:3 (14:37) [Early Pleistocene, Villafranchian, Italy (mainland)]
Type locality: “le Ville...Valdarno Superiore...Toscana” [Italy].
Testudo oriens † Portis 1890:9 (14:37) [Early Pleistocene, Villafranchian, Italy (mainland)]
Type locality: “le Mignaie...Valdarno Superiore...Toscana” [Italy].
Testudo seminota † Portis 1890:10 (14:37) [Early Pleistocene,
Villafranchian, Italy (mainland)]
Type locality: “il Tasso...Valdarno Superiore...Toscana” [Italy].
Testudo hermanni robertmertensi Wermuth 1952:162
Type locality: “Gebirge nördlich von Arta, Mallorca, Balearen”
[mountains north of Arta, Mallorca, Balearic Islands] [Spain].
(subspecies: hermanni = red, boettgeri = purple;
orange dots = probable introduced or trade)
Testudo (Chersine) hermanni boettgeri Mojsisovics 1889
Eastern Hermann’s Tortoise
Albania, Bosnia and Herzegovina, Bulgaria, Croatia, France
(Continental, Corsica), Greece, Italy (Continental, Kosovo,
Sardinia [prehistoric introduction], Sicily), Macedonia,
Montenegro, Romania, Serbia, Slovenia, Spain (Continental, Balearic Islands [prehistoric introduction]), Turkey
(European)
Introduced: Malta (?), Spain (Balearic Islands)
CBFTT Account: Bertolero, Cheylan, Hailey, Livoreil, and
Willemsen (2011)
IUCN Red List: Near Threatened (2004); Previously: Near
Threatened (1996)
CITES: Appendix II, as Testudinidae spp.
Adrian Hailey / CBFTT / Greece
Albania, Bosnia and Herzegovina, Bulgaria, Croatia, Greece,
Italy (?) (Continental), Kosovo, Macedonia, Montenegro,
Romania, Serbia, Slovenia, Turkey (European)
Testudo graeca boettgeri Mojsisovics 1889:242 (senior homonym, not = Testudo boettgeri Siebenrock 1904a), Testudo
158
Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
hermanni boettgeri, Testudo boettgeri, Testudo boettgeri
boettgeri, Eurotestudo boettgeri, Chersine hermanni
boettgeri
Type locality: “Orsova...Cernathal...Süd-Ungarn” [Romania].
Restricted to “Orsova, Banat” [Romania] by Boettger (1893:11).
Testudo graeca hercegovinensis Werner 1899:818, Testudo hercegovinensis, Testudo boettgeri hercegovinensis, Testudo hermanni hercegovinensis, Eurotestudo
hercegovinensis
Type locality: “Trebinje...Hercegovina” [Bosnia and
Herzegovina].
Testudo enriquesi Parenzan 1932:1160
Type locality: “Conca di Elbassan, nella vallata dello Skumbi, in
Albania Centrale.”
TRionyChoidea Gray 1825
Trionicidae Gray 1825:212
Trionychoidea Fitzinger 1826:5
Trionychia Hummel 1929:362
CaReTToChelyidae Boulenger 1887a
Carettochelydidae Boulenger 1887a:171
Carettochelydes Baur 1891a:190
Carettochelyidae Baur 1891c:637
Carettochelys Ramsay 1886
Carettochelys Ramsay 1886:158
Type species: Carettochelys insculpta Ramsay 1886, by original
monotypy.
Carettochelys insculpta Ramsay 1886 (07:78, 08:10)
Pig-Nosed Turtle, Fly River Turtle
IUCN Red List: Vulnerable A1bd (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Carettocchelys insculptus Ramsay 1886:158, Carettochelys
insculptus, Carettochelys insculpta, Carettochelys insculpta insculpta
Type locality: “Fly River, New Guinea.” Restricted to “Strickland River...upper right hand branch of the Fly River” by Waite
(1905:110); and to “Turtle Camp, Strickland River [Papua New
Guinea] [6º25'30" S, 142º04'30" W]” by Mackay (2003:39).
Carettochelys insculpta canni Wells 2002a:1 (07:78, 08:10, 10:43) (unavailable name), Carettochelys canni
Type locality: “near Ooloo Crossing, Daly River, Northern Territory [Australia]”
TRionyChidae Gray 1825
Amydae Oppel 1811:9 (partim)
Trionyces Schmid 1819:18
Trionicidae Gray 1825:212
Trionychoidea Fitzinger 1826:5
Trionychidae Bell 1828c:515
Trionycidae Bonaparte 1831:63
Trionychidi Portis 1890:22
CyClanoRbinae Lydekker 1889
Cyclanosteina Gray 1864b:94
Cyclanorbinae Lydekker 1889:x
Cyclanorbis Gray 1854a
Cryptopus Duméril and Bibron 1835:499 (junior homonym, not
= Cryptopus Latreille 1829 [= Crustacea])
Type species: Cryptopus senegalensis Duméril and Bibron 1835, by
subsequent designation by Bour et al. (1995:82).
Cyclanorbis Gray 1854a:135
Type species: Cyclanorbis petersii Gray 1854a [= subjective synonym of Cryptopus senegalensis Duméril and Bibron 1835], by
original monotypy.
Cryptopodus Duméril 1856:374 (nomen novum)
Cyclanosteus Gray 1856a:201
Type species: Cyclanosteus senegalensis [= Cryptopus senegalensis
Duméril and Bibron 1835], by subsequent designation by Günther
(1865:108).
Tetrathyra Gray 1865a:205
John Cann / CBFTT / Daly R., Northern Territory, Australia
Type species: Tetrathyra baikii Gray 1865a [= subjective synonym
of Cryptopus senegalensis Duméril and Bibron 1835], by original
monotypy.
Baikiea Gray 1869a:215
Type species: Baikiea elegans Gray 1869a, by original monotypy.
(orange dot = introduced or trade)
Australia (Northern Territory), Indonesia (Papua), Papua New
Guinea (Southern)
CBFTT Account: Georges, Doody, Eisemberg, Alacs, and
Rose (2008)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Cyclanorbis elegans (Gray 1869a)
Nubian Flapshell Turtle
Maurice Rodrigues / CBFTT / Africa [captivity]
Benin, Burkina Faso, Cameroon, Central African Republic,
Chad, Ethiopia, Gambia, Ghana, Guinea (?), Guinea-Bissau, Ivory Coast, Liberia, Mali, Mauritania (extirpated?),
Niger, Nigeria, Senegal, Sierra Leone, South Sudan,
Sudan, Togo
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Vulnerable (2016)
CITES: Appendix II, as Cyclanorbis spp.
Cryptopus senegalensis Duméril and Bibron 1835:504,
Emyda senegalensis, Cyclanosteus senegalensis, Cyclanorbis senegalensis
Type locality: “Sénégal.”
Cyclanorbis petersii Gray 1854a:135, Cyclanosteus petersii,
Cycloderma petersii
Type locality: “West Africa, River Gambia” [The Gambia].
Benin (?), Cameroon, Central African Republic, Chad, Ethiopia
(?), Ghana, Nigeria, South Sudan, Sudan, Togo
CBFTT Account: Baker, Diagne, and Luiselli (2015)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Critically Endangered (2016)
CITES: Appendix II, as Cyclanorbis spp.
Baikiea elegans Gray 1869a:222, Cyclanorbis elegans,
Cyclanosteus elegans
Type locality: “Africa.” Restricted to “West Africa” by Gray
(1873j:86); and to “Niger River drainage in west Africa” by
lectotype designation by Webb (1975:349).
Cyclanorbis oligotylus Siebenrock 1902c:810
Type locality: “Nubien...oberen Nil” [Nubia...upper Nile] [Sudan].
Cyclanorbis senegalensis (Duméril and Bibron 1835) (75)
Senegal Flapshell Turtle, Sahelian Flapshell Turtle
Cycloderma senegalense Duméril 1861a:168 (justified
emendation)
Tetrathyra baikii Gray 1865a:205
Type locality: “West Africa, River Niger?” Restricted to “Niger
River drainage in west Africa” by lectotype designation by Webb
(1975:350).
Cyclanosteus senegalensis equilifera Gray 1865b:425
Type locality: “the Niger” [Niger River, West Africa].
Cyclanosteus senegalensis normalis Gray 1865b:425
Type locality: “the Niger” [Niger River, West Africa].
Cyclanosteus senegalensis callosa Gray 1865b:425
Type locality: “the Niger” [Niger River, West Africa].
Tetrathyra vaillantii Rochebrune 1884:36
Type locality: “Sénégambie” [Senegal and The Gambia].
Cycloderma Peters 1854
Cycloderma Peters 1854:216
Type species: Cycloderma frenatum Peters 1854, by original
monotypy.
Heptathyra Cope 1860:294
Type species: Heptathyra aubryi [= Cryptopodus aubryi Duméril
1856], by original monotypy.
Aspidochelys Gray 1860a:6
Type species: Aspidochelys livingstonii Gray 1860a [= subjective
synonym of Cycloderma frenatum Peters 1854], by original
monotypy.
William R. Branch / Northern Prov., Sierra Leone
159
160
Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Cycloderma aubryi (Duméril 1856) (10:31)
Aubry’s Flapshell Turtle
Olivier S.G. Pauwels / Gamba, Ogooué-Maritime Prov., Gabon
Malawi, Mozambique, Tanzania, Zambia, Zimbabwe
CBFTT Account: Broadley and Sachsse (2011)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Endangered (2016)
CITES: Appendix II, as Cycloderma spp.
Cycloderma frenatum Peters 1854:216, Cyclanosteus frenatus, Heptathyra frenata
Type locality: “In fluminibus Zambeze et Licuare..[&]..Tette et
Sena..[&]..terra Boror” [Mozambique]. Restricted to “Sambesi
bei Tete” [Mozambique] by lectotype designation by Fritz et al.
(1994:169).
Aspidochelys livingstonii Gray 1860a:6, Heptathyra
livingstonii
Type locality: “Mozambique in tributaries of River Zambesi?”
Angola (Cabinda), Cameroon, Central African Republic, Congo
(DRC), Congo (ROC), Gabon
IUCN Red List: Not Listed [Least Concern 1996]
TFTSG Draft Red List: Vulnerable (2016)
CITES: Appendix II, as Cycloderma spp.
Cryptopodus aubryi Duméril 1856:374 (10:31), Cryptopus
aubryi, Cycloderma aubryi, Heptathyra aubryi
Type locality: “Gabon.”
Cycloderma frenatum Peters 1854
Zambezi Flapshell Turtle
Lissemys Smith 1931 (11:14)
Emyda Gray 1830e:19 (10:7) (junior homonym, not = Emyda
Rafinesque 1815)
Type species: Emyda punctata [= Testudo punctata Lacepède 1788]
[= Testudo punctata Bonnaterre 1789], by original monotypy.
Lissemys Smith 1931:xxviii, 154 (nomen novum)
Type species: Lissemys punctata [= Testudo punctata Lacepède
1788] [= Testudo punctata Bonnaterre 1789], by original
designation.
Lissemys ceylonensis (Gray 1856a) (11:14)
Sri Lankan Flapshell Turtle
Wulf Haacke / CBFTT / Lake Malawi, Malawi
Anslem de Silva / Udawalawe National Park, Sri Lanka
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Lissemys punctata punctata (Bonnaterre 1789) (11:14)
Southern Indian Flapshell Turtle
Indraneil Das / CBFTT / Mamallapuram, Tamil Nadu, India
Sri Lanka
IUCN Red List: Not Evaluated
CITES: Appendix II [as part of Lissemys punctata]
Emyda ceylonensis Gray 1856a:201, Emyda granosa ceylonensis, Lissemys ceylonensis, Lissemys punctata ceylonensis
Type locality: “Ceylon” [Sri Lanka].
Lissemys punctata sinhaleyus † Deraniyagala 1953:5 [Late
Pleistocene, Ratnapura Beds, Sri Lanka]
Type locality: “near Ratnapura...Ceylon” [Sri Lanka].
Lissemys punctata (Bonnaterre 1789) (09:44, 11:14)
Indian Flapshell Turtle
India (Kerala, Tamil Nadu)
Testudo punctata Lacepède 1788:171, synopsis[table] (09:6)
(senior homonym, not = Testudo punctata Schoepff 1792);
nomen suppressum, ICZN 2005a)
Type locality: “les grandes Indes” [India]. Restricted to “Pondicherry, South Arcot (district), Tamil Nadu (state), India” by Webb
(1980a:552).
Testudo punctata Bonnaterre 1789:30 (senior homonym,
not = Testudo punctata Schoepff 1792), Trionyx (Emyda)
punctatus, Trionyx punctatus, Emyda punctata, Trionyx
punctata, Lissemys punctata, Lissemys punctata punctata,
Trionyx punctatus punctatus
Type locality: “les grandes Indes” [India]. Restricted to “Pondicherry, South Arcot (district), Tamil Nadu (state), India” by Webb
(1980a:552).
Testudo sonnerati Meyer 1790:83 (09:8) (nomen novum et
oblitum)
Testudo granulosa Suckow 1798:48 (nomen novum)
Testudo scabra Latreille in Sonnini and Latreille 1801:164
(nomen novum and junior homonym, not = Testudo scabra
Linnaeus 1758)
Testudo granosa Schoepff 1801:127, Trionyx granosus,
Cryptopus granosus, Emyda granosa, Emyda granosa
granosa, Lissemys punctata granosa, Trionyx punctatus
granosus
Type locality: “Coromandeliae” [India].
(subspecies: punctata = red, andersoni = purple, vittata = orange;
overlap = intergrades; orange dot = introduced)
Bangladesh, India (Andhra Pradesh, Bihar, Goa, Gujarat,
Kerala, Madhya Pradesh, Odisha, Punjab, Tamil Nadu,
West Bengal), Myanmar, Nepal, Pakistan
Introduced: India (Andaman Islands)
CBFTT Account: Bhupathy, Webb, and Praschag (2014)
IUCN Red List: Least Concern (2000); Previously: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
CITES: Appendix II
Testudo granulata Daudin 1801:81 (nomen novum)
Trionyx coromandelicus Geoffroy Saint-Hilaire 1809b:16
(nomen novum)
Emyda dura Buchanan-Hamilton in Anderson 1876:514 (nomen nudum)
Lissemys punctata andersoni Webb 1980a (11:14)
Spotted Northern Indian Flapshell Turtle
Shailendra Singh / CBFTT / Ghaghra R., Uttar Pradesh, India
Bangladesh, India (Assam, Bihar, Haryana, Jammu, Madhya
Pradesh, Meghalaya, Rajasthan, Sikkim, Uttar Pradesh,
West Bengal), Myanmar, Nepal, Pakistan
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Introduced: India (Andaman Islands)
Lissemys punctata andersoni Webb 1980a:554, Lissemys
andersoni
Type locality: “Belbari, Terai, southeastern Nepal, elevation 210 m.”
Lissemys punctata vittata (Peters 1854) (11:14)
Central Indian Flapshell Turtle
Lissemys scutata
Type locality: “Pegú” [Myanmar].
Emyda fuscomaculata Gray 1873c:308
Type locality: Not designated. Restricted to “Asia, Pegu?” [Myanmar] by Gray (1873j:89).
TRionyChinae Gray 1825
Amydae Oppel 1811:9 (partim)
Trionyces Schmid 1819:18
Trionicidae Gray 1825:212
Trionychoidea Fitzinger 1826:5
Trionychinae Lydekker 1889:4
Amyda Schweigger in Geoffroy Saint-Hilaire 1809a
Amyda Schweigger in Geoffroy Saint-Hilaire 1809a:365
Shailendra Singh / CBFTT / Mahanadi River, Odisha, India
India (Andhra Pradesh, Chhattisgarh?, Goa, Gujarat, Karnataka, Madhya Pradesh, Maharashtra, Odisha, Rajasthan)
Emyda vittata Peters 1854:216, Emyda granosa vittata, Lissemys punctata vittata
Type locality: “India orientalis, Goa.”
Emyda granosa intermedia Annandale 1912a:171
Type locality: “Near Purulia, Manbhum Dist.” [India].
Lissemys scutata (Peters 1868) (11:14)
Burmese Flapshell Turtle
Type species: Amyda javanica Schweigger in Geoffroy Saint-Hilaire
1809a [= subjective synonym of Testudo cartilaginea Boddaert
1770], by original monotypy.
Amida Duméril and Bibron 1834:416 (nomen novum)
Potamochelys Fitzinger 1843:30
Type species: Aspidonectes (Potamochelys) javanica Wagler [=
Amyda javanica Schweigger in Geoffroy Saint-Hilaire 1809a] [=
subjective synonym of Testudo cartilaginea Boddaert 1770], by
original monotypy.
Aspilus Gray 1864b:83 (junior homonym, not = Aspilus Schaum
1848 [= Coleoptera])
Type species: Aspilus cariniferus [= Trionyx cariniferus Gray 1856b]
[= subjective synonym of Testudo cartilaginea Boddaert 1770],
by original designation.
Ida Gray 1873h:55 (junior homonym, not = Ida Gray 1838 [=
Sauria] or Ida Gray 1840 [= Gastropoda] or Ida RobineauDesvoidy 1863 [= Diptera])
Type species: Ida ornata [ = Trionyx ornatus Gray 1861a] [= subjective synonym of Testudo cartilaginea Boddaert 1770], by original
monotypy.
Amyda cartilaginea (Boddaert 1770) (76)
Malayan Softshell Turtle, Asiatic Softshell Turtle
Gerald Kuchling / CRM 2 / Myanmar [trade: Ruili, Yunnan, China]
(subspecies: cartilaginea = red, maculosa = purple,
unassigned A. cartilaginea sensu lato = gray;
orange dots = probable trade)
(orange dot = trade)
Myanmar
IUCN Red List: Data Deficient (2000); Previously: Data Deficient (1996)
TFTSG Draft Red List: Near Threatened (2011)
CITES: Appendix II
Emyda scutata Peters 1868:449, Emyda granosa scutata,
Lissemys punctata scutata, Trionyx punctatus scutatus,
Brunei, Indonesia (Bali, Java, Kalimantan, Sumatra), Malaysia
(Peninsular, East), Singapore, Thailand (?)
Introduced or Trade: Indonesia (Lesser Sundas, Moluccas,
Sulawesi)
CBFTT Account: Auliya, van Dijk, Moll, and Meylan (2016)
[includes Amyda ornata]
IUCN Red List: Vulnerable A1cd+2cd (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Amyda cartilaginea cartilaginea (Boddaert 1770) (76)
South Sundas Softshell Turtle
Mark Auliya / Java, Indonesia [trade]
Indonesia (Bali, Java, Kalimantan), Malaysia (East)
Introduced or Trade: Indonesia (Lesser Sundas [Lombok]),
Moluccas [Ambon, Seram]), Sulawesi)
Testudo cartilaginea Boddaert 1770:1, Gymnopus cartilaginea, Trionyx cartilagineus, Aspidonectes cartilagineus,
Potamochelys cartilagineus, Amyda cartilaginea, Amyda
cartilaginea cartilaginea
Type locality: Not known. Restricted to “Java” [Indonesia] by
Baur (1893a:220).
Testudo membranacea Blumenbach 1779:257 (nomen
dubium)
Testudo boddaerti Schneider 1787:12 (nomen novum), Trionyx
boddaerti, Testudo boddarti
Testudo striata Suckow 1798:37 (partim, nomen novum)
Amyda javanica Schweigger in Geoffroy Saint-Hilaire
1809a:365 (senior homonym, not = Trionyx javanicus
Gray 1830e), Trionyx javanicus, Trionyx iavanicus,
Tyrse javanica, Aspidonectes javanicus, Trionyx stellatus
javanica, Aspilus javanicus, Potamochelys javanicus
Amyda cartilaginea maculosa Fritz, Gemel, Kehlmaier, Vamberger, and Praschag 2014a (76)
North Sundas Softshell Turtle
Mark Auliya / CBFTT / Anjungan, West Kalimantan, Indonesia
Brunei (?), Indonesia (Kalimantan, Sumatra), Malaysia (East,
Peninsular?), Thailand (?)
Amyda cartilaginea maculosa Fritz, Gemel, Kehlmaier, Vamberger, and Praschag 2014a:240
Type locality: “Nanga Badau, Kalimantan, Indonesia (“Nanga
Bandang, Borneo”).”
Amyda cartilaginea (Boddaert 1770) (76) or
Amyda ornata (Gray 1861a)
Southeast Asian Softshell Turtle
Type locality: “Java et les îles voisines” [Indonesia].
Trionyx stellatus Geoffroy Saint-Hilaire 1809a:365 (nomen
novum), Potamochelys stellatus
Trionyx cariniferus Gray 1856b:67, Aspilus cariniferus,
Trionyx carinifera
Type locality: “Moluccas” [Indonesia].
Aspilus punctulatus Gray 1864b:84
Type locality: “Amboina or Ceram” [Indonesia].
Trionyx jeudi Gray 1869a:217
Type locality: “Java?” [Indonesia].
Trionyx trinilensis † Jaekel 1911:78 [Pleistocene, Pithecanthropus Trinil Beds, Indonesia (Java)]
Type locality: “Pithecanthropus-schichten...Java...Trinil”
[Indonesia].
(subspecies: ornata = red, phayrei = purple;
unassigned A. ornata sensu lato = gray)
Bangladesh, Cambodia, India (Mizoram), Laos, Myanmar,
Thailand, Vietnam
Introduced: Indonesia (Lesser Sundas, Sulawesi)
CBFTT Account: Auliya, van Dijk, Moll, and Meylan (2016)
[as part of Amyda cartilaginea]
IUCN Red List: Vulnerable A1cd+2cd (2000), as part of Amyda
cartilaginea; Previously: Vulnerable (1996), as part of A.
cartilaginea
TFTSG Draft Red List: Vulnerable (2015)
CITES: Appendix II, as part of A. cartilaginea
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Amyda cartilaginea ornata (Gray 1861) (76) or
Amyda ornata ornata (Gray 1861)
Indochinese Softshell Turtle
Mesodeca Rafinesque 1832:64
Type species: Mesodeca bartrami [= Testudo bartrami Daudin 1801]
[= subjective synonym of Testudo ferox Schneider 1783], by
original monotypy.
Trionyx (Platypeltis) Fitzinger 1835:109
Type species: Platypeltis ferox Fitzinger [= Testudo ferox Schneider
1783], by subsequent designation by Fitzinger (1843:30).
Callinia Gray 1869a:221
Type species: Callinia spinifera [= Trionyx spiniferus LeSueur 1827],
by subsequent designation by Stejneger (1907:514).
Euamyda Stejneger 1944:7
Type species: Euamyda mutica [= Trionyx muticus LeSueur 1827],
by original monotypy.
Apalone ferox (Schneider 1783)
Florida Softshell Turtle
David Emmett / CBFTT / Cardamom Mountains region, Cambodia
Cambodia, Laos, Thailand, Vietnam
Trionyx ornatus Gray 1861a:41, Aspilus ornatus, Ida ornata,
Amyda ornata, Amyda ornata ornata, Amyda cartilaginea
ornata
Type locality: “Camboja” [Cambodia].
Amyda cartilaginea phayrei (Theobald 1868b) (76) or
Amyda ornata phayrei (Theobald 1868b)
Burmese Softshell Turtle
Matthew Aresco / CRM 3 / Leon Co., Florida
Gerald Kuchling / Dokhtawady R., Myanmar
Bangladesh (?), India (?) (Mizoram), Malaysia (Peninsular),
Myanmar, Thailand
Trionyx phayrei Theobald 1868b:18, Aspidonectes phayrei,
Amyda ornata phayrei, Amyda cartilaginea ornata
Type locality: “montium Arakanensium, prope Bassein” [Arakan
Hills, Bassein, Myanmar].
Trionyx ephippium Theobald 1875:177
Type locality: “Tenasserim” [Myanmar].
Trionyx phayrii Boulenger 1889:ix (nomen novum), Amyda
phayrii
Trionyx nakornsrithammarajensis Nutaphand 1979:8, Amyda
nakornsrithammarajensis
Type locality: “Nakorn Sri Thammaraj Province... Thailand.”
Trionyx cartilageneus nakorn Nutaphand 1990:[8] (nomen
novum), Trionyx cartilagineus nakorn
Apalone Rafinesque 1832
Aplaxia Rafinesque 1817:166 (nomen oblitum)
Type species: Aplaxia nasica Rafinesque 1817 (nomen nudum) [=
Trionyx nasica Rafinesque 1822 (nomen suppressum] [= subjective synonym of Trionyx spiniferus LeSueur 1827], by original
monotypy.
Apalone Rafinesque 1832:64
Type species: Apalone hudsonica Rafinesque 1832 [= subjective synonym of Trionyx spiniferus LeSueur 1827], by original monotypy.
(orange dot = introduced)
USA (Alabama, Florida, Georgia, South Carolina)
IUCN Red List: Least Concern (2010); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix III (USA)
Testudo ferox Schneider 1783:330, Emydes ferox, Trionyx
ferox, Amyda ferox, Aspidonectes ferox, Platypeltis ferox,
Amyda ferox ferox, Trionyx ferox ferox, Apalone ferox
Type locality: “Savannach und Alatama..&..Ost-Florida” [USA].
Restricted to “Savannah River (at Savannah), Georgia” [USA] by
Schwartz (1956b:8).
Testudo mollis Lacepède 1788:137, synopsis[table] (09:6) (nomen suppressum, ICZN 2005a)
Type locality: “sud de la Caroline, rivières de Savannah &
d’Alatamaha..&..la Floride orientale” [USA].
Testudo mollis Bonnaterre 1789:25
Type locality: “rivières de Savannah, d’Alatamaha, de la Floride,
& de la Caroline méridionale” [USA].
Testudo (ferox) verrucosa Schoepff 1795:90 (senior homonym, not = Testudo verrucosa Suckow 1798)
Type locality: “Floridae Orientalis” [USA]. Restricted to “Halfway Pond...between...Palatka and Gainesville, Fla...somewhere
in southwestern Putnam County” [Florida, USA] by Harper
(1940:717) by restriction of nomen novum replacement name
Testudo bartrami Daudin 1801.
Testudo bartrami Daudin 1801:74 (nomen novum), Chelys
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
bartrami, Trionyx bartrami, Mesodeca bartrami
Type locality: “la Floride orientale” [Florida, USA]. Restricted to
“Halfway Pond...between...Palatka and Gainesville, Fla...somewhere in southwestern Putnam County” [Florida, USA] by Harper
(1940:717).
Trionyx georgianus Geoffroy Saint-Hilaire 1809a:367 (nomen
novum)
Trionyx carinatus Geoffroy Saint-Hilaire 1809b:14, Aspidonectes carinatus
Type locality: Not known. Restricted to “Savannah River, Georgia” [USA] by Schmidt (1953:108).
Trionyx georgicus Geoffroy Saint-Hilaire 1809b:17 (nomen
novum)
Trionyx brongniarti Schweigger 1812:288 (nomen novum),
Testudo brongniarti, Platypeltis brongniarti
Type locality: Not known. Restricted to “Savannah River, Georgia” [USA] by Schmidt (1953:109).
Trionyx harlani Bell in Harlan 1835:159
Type locality: “East Florida” [USA].
Pennsylvania [extirpated], South Dakota, Tennessee,
Texas, West Virginia, Wisconsin)
Trionyx pusilla Rafinesque 1822:3 (nomen dubium et suppressum, ICZN 1984)
Type locality: “United States” [USA].
Trionyx muticus LeSueur 1827:263, Aspidonectes muticus,
Gymnopus muticus, Amyda mutica, Euamyda mutica, Trionyx muticus, Trionyx muticus muticus, Apalone mutica,
Apalone muticus, Apalone mutica mutica
Type locality: “Newharmony, sur le Wabash, à peu de distance de
son embouchure dans l’Ohio...l’Etat de l’Indiana” [USA].
Potamochelys microcephalus Gray 1864b:87, Callinia microcephala, Potamochelys microcephala
Type locality: “Sarawak” [Malaysia] [in error]. Emended to
“___?” [not known] by Boulenger (1889:262); restricted to “New
Harmony, Indiana” [USA] by Schmidt (1953:110).
Apalone mutica calvata (Webb 1959) (77)
Gulf Coast Smooth Softshell Turtle
Apalone mutica (LeSueur 1827)
Smooth Softshell Turtle
Barry Mansell / CRM 3 / Escambia Co., Florida
USA (Alabama, Florida, Louisiana, Mississippi)
Trionyx muticus calvatus Webb 1959:519, Apalone mutica
calvata, Apalone calvata
Type locality: “Pearl River, Roses Bluff, 14 miles northeast Jackson, Rankin County, Mississippi” [USA].
(subspecies: mutica = red, calvata = purple)
USA (Alabama, Arkansas, Florida, Illinois, Indiana, Iowa,
Kansas, Kentucky, Louisiana, Minnesota, Mississippi,
Missouri, Nebraska, New Mexico, North Dakota, Ohio,
Oklahoma, Pennsylvania [extirpated], South Dakota, Tennessee, Texas, West Virginia, Wisconsin)
IUCN Red List: Least Concern (2010); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix III (USA)
Apalone mutica mutica (LeSueur 1827)
Midland Smooth Softshell Turtle
Peter V. Lindeman / Scioto R., Pike Co., Ohio
USA (Alabama, Arkansas, Illinois, Indiana, Iowa, Kansas,
Kentucky, Louisiana, Minnesota, Mississippi, Missouri,
Nebraska, New Mexico, North Dakota, Ohio, Oklahoma,
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Apalone spinifera (LeSueur 1827) (12:37)
Spiny Softshell Turtle
Nebraska, New Mexico, New York, North Carolina, North
Dakota, Ohio, Oklahoma, Pennsylvania, South Dakota,
Tennessee, Vermont, Virginia, West Virginia, Wisconsin,
Wyoming)
Aplaxia nasica Rafinesque 1817:166 (nomen nudum)
Trionyx nasica Rafinesque 1822:3 (nomen suppressum,
ICZN 1984)
Type locality: “western streams...United States” [USA].
Trionyx spiniferus LeSueur 1827:258, Gymnopus spiniferus,
Gymnopodus spiniferus, Aspidonectes spinifer, Gymnopus spinifer, Trionyx spinifer, Callinia spicifera, Callinia
spinifera, Platypeltis spinifer, Tyrse spinifera, Amyda
spinifera, Platypeltis spinifera, Amyda spinifer, Amyda
spinifera spinifera, Trionyx spinifera, Trionyx spinifera
spinifera, Amyda ferox spinifera, Trionyx ferox spinifera,
Trionyx spinifer spinifer, Trionyx spiniferus spiniferus,
Apalone spinifera, Apalone spiniferus, Apalone spinifera
spinifera
Type locality: “Newharmony, sur le Wabash, à peu de distance de
son embouchure dans l’Ohio...l’Etat de l’Indiana” [USA].
Trionyx ocellatus LeSueur 1827:261 (senior homonym, not =
Trionyx ocellatus Gray 1832a)
Type locality: “Newharmony, sur le Wabash, à peu de distance de
son embouchure dans l’Ohio...l’Etat de l’Indiana” [USA].
Apalone hudsonica Rafinesque 1832:64
Type locality: “River Hudson between the falls of Hadley, Glen and
Baker, and further up to the source” [USA]. Restricted to “Hudson
River, near Baker’s Falls, Saratoga County, New York” [USA] by
Webb (1962:491).
(subspecies: spinifera = red, aspera = purple, atra = orange,
emoryi = pink, guadalupensis = brown, pallida = green;
overlap = intergrades; orange dots = introduced)
Canada (Ontario, Québec), Mexico (Chihuahua, Coahuila,
Nuevo Leon, Tamaulipas), USA (Alabama, Arkansas,
Colorado, Florida, Georgia, Illinois, Indiana, Iowa, Kansas,
Kentucky, Louisiana, Maryland, Michigan, Minnesota,
Mississippi, Missouri, Montana, Nebraska, New Mexico,
New York, North Carolina, North Dakota, Ohio, Oklahoma, Pennsylvania, South Carolina, South Dakota, Tennessee, Texas, Vermont, Virginia, West Virginia, Wisconsin,
Wyoming)
Introduced: USA (Arizona, California, Hawaii, Nevada, New
Jersey, Utah, Virginia)
IUCN Red List: Least Concern (2010); Previously: Least Concern [Not Listed] (1996)
CITES: Appendix III (USA)
Apalone spinifera spinifera (LeSueur 1827) (08:22)
Northern Spiny Softshell Turtle
Trionyx annulifer Wied 1839:140 (nomen novum)
Tyrse argus Gray 1844:48, Trionyx argus
Type locality: “West Africa, Sierra Leone?” [in error]. Emended to
“___?” [not known] by Boulenger (1889:260); restricted to “New
Harmony, Indiana” [USA] by Schmidt (1953:110).
Trionyx annulatus Gray 1856b:69 (nomen novum)
Aspidonectes nuchalis Agassiz 1857a:402,406, Platypeltis
nuchalis
Type locality: “Cumberland River..[&]..head waters of the Tennessee River” [USA]. Restricted to “Cumberland River near
Nashville, Tennessee” [USA] by Schmidt (1953:110).
Gymnopus olivaceus Wied 1865:55 (nomen novum)
Amyda spinifera hartwegi Conant and Goin 1948:1 (08:22),
Amyda ferox hartwegi, Trionyx ferox hartwegi, Trionyx
spinifer hartwegi, Trionyx spiniferus hartwegi, Apalone
spinifera hartwegi
Type locality: “Wichita, Sedgwick County, Kansas” [USA].
Apalone spinifera aspera (Agassiz 1857a)
Gulf Coast Spiny Softshell Turtle
Barry Mansell / CRM 3 / Santa Rosa Co., Florida
John B. Iverson / Muscatine, Muscatine Co., Nebraska / female
Canada (Ontario, Québec), USA (Arkansas, Colorado, Illinois,
Indiana, Iowa, Kansas, Kentucky, Louisiana, Maryland,
Michigan, Minnesota, Mississippi, Missouri, Montana,
USA (Alabama, Florida, Georgia, Mississippi, North Carolina,
South Carolina, Tennessee)
Aspidonectes asper Agassiz 1857a:402,405, Platypeltis asper,
Amyda spinifera aspera, Amyda ferox aspera, Trionyx
ferox aspera, Trionyx spinifer asper, Trionyx spiniferus
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
asper, Trionyx spiniferus asperus, Apalone spinifera asper,
Apalone spinifera aspera
spinifera emoryi, Trionyx spiniferus emoryi, Apalone
spinifera emoryi
Type locality: “Mississippi” [USA]. Restricted to “Lake Concordia, La.” [Louisiana, USA] by Baur (1893:220) and Schmidt
(1953:109); and to “Pearl River at Columbus, Marion County,
Mississippi” [USA] by lectotype designation by Webb (1960:7).
Type locality: “lower Rio Grande of Texas, near Brownsville”
[USA].
Platypeltis agassizii Baur 1888c:1122, Trionyx agassizii,
Pelodiscus agassizii, Aspidonectes agassizii, Trionyx spiniferus agassizii, Amyda agassizii, Amyda ferox agassizii,
Trionyx ferox agassizii
Aspidonectes emyda Gray 1870c:95 (nomen novum)
Aspidonectes georgii Gray 1870c:109 (nomen novum)
Apalone spinifera guadalupensis (Webb 1962)
Guadalupe Spiny Softshell Turtle
Type locality: Not designated. Restricted to “Savannah, Savannah
River” [Georgia, USA] by Schwartz (1956b:17).
Apalone spinifera atra (Webb and Legler 1960) (07:79, 08:23, 12:37)
Black Spiny Softshell Turtle, Cuatro Cienegas Softshell
Peter V. Lindeman / Palmetto State Park, San Marcos R., Gonzales Co., Texas
USA (Texas)
Trionyx spinifer guadalupensis Webb 1962:517, Trionyx
spiniferus guadalupensis, Apalone spinifera guadalupensis
Type locality: “15 miles northeast Tilden, McMullen County,
Texas” [USA].
Suzanne E. McGaugh / CBFTT / Tío Cándido, Cuatrociénegas Basin, Coahuila, Mexico
Apalone spinifera pallida (Webb 1962)
Pallid Spiny Softshell Turtle
Mexico (Coahuila)
CBFTT Account: Cerdá-Ardura, Soberón-Mobarak, McGaugh, and Vogt (2008)
IUCN Red List: Critically Endangered A1ace+2c (1996),
originally listed as Apalone ater
CITES: Appendix I
Trionyx ater Webb and Legler 1960:21, Trionyx spinifer ater,
Trionyx spiniferus ater, Apalone spinifera ater, Apalone
ater, Apalone spiniferus ater, Apalone spinifera atra,
Apalone (Apalone) atra, Apalone atra
Type locality: “16 kilometers south of Cuatro Ciénegas, Coahuila...
Mexico.” Emended to “Poza del Tío Cándido, 16 km southwest of
Cuatrociénegas de Carranza, central Coahuila, México” by CerdáArdura et al. (2008:2).
Apalone spinifera emoryi (Agassiz 1857a) (12:37)
Texas Spiny Softshell Turtle
Vincenzo Ferri / Texas [captivity]
Mexico (Chihuahua, Coahuila, Nuevo Leon, Tamaulipas),
USA (Texas)
Aspidonectes emoryi Agassiz 1857a:392,407, Trionyx emoryi,
Platypeltis emoryi, Amyda emoryi, Amyda ferox emoryi,
Trionyx ferox emoryi, Trionyx spinifer emoryi, Trionyx
Stanley E. Trauth / nr. Horatio, Sevier Co., Arkansas
USA (Arkansas, Louisiana, Oklahoma, Texas)
Trionyx spinifer pallidus Webb 1962:522, Trionyx pallidus,
Trionyx spiniferus pallidus, Apalone spinifera pallida,
Apalone spinifera pallidus
Type locality: “Lake Caddo, Caddo Parish, Louisiana” [USA].
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Indonesia (Java)]
Chitra Gray 1844
Chitra Gray 1844:49
Type species: Chitra indica [= Trionyx indicus Gray 1830e], by
original monotypy.
Chitra chitra Nutaphand 1986
Asian Narrow-headed Softshell Turtle
Type locality: “Pithecanthropus-schichten...Java...Trinil”
[Indonesia].
Chitra chitra javanensis McCord and Pritchard 2003:41,
Chitra chitra javanica
Type locality: “Pasuruan River drainage, near Pasuruan, Probolinggo District, East Java, Indonesia.” Emended to “Bengawan (= river)
Solo, between Kalitidu and Padangan, Bojonegoro District, East
Java” [Indonesia] by Iskandar (2004:129).
Chitra indica (Gray 1830e) (10:7)
Indian Narrow-headed Softshell Turtle
Shekar Dattatri / CBFTT / Chambal R., Rajasthan, India
(subspecies: chitra = red, javanensis = purple)
Indonesia (Java, Sumatra), Malaysia (Peninsular), Thailand
IUCN Red List: Critically Endangered A1cd, B1+2c (2000); Previously: Critically Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix I
Chitra chitra chitra Nutaphand 1986
Siamese Narrow-headed Softshell Turtle
Bangladesh, India (Andhra Pradesh, Madhya Pradesh, Maharashtra, Odisha, Punjab, Rajasthan, Tamil Nadu, Uttar
Pradesh, West Bengal), Nepal, Pakistan
CBFTT Account: Das and Singh (2009)
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Chitra spp.
Trionyx indicus Gray 1830e:18 (10:7), Trionyx egyptiacus indicus, Trionyx aegyptiacus indica, Chitra indica, Gymnopus
indicus, Trionyx aegyptianus indicus, Aspidonectes indicus
Chris Tabaka / TCC / Thailand [captivity]
Malaysia (Peninsular), Thailand
Chitra chitra Nutaphand 1986:64 (nomen conservandum,
ICZN 2005b), Chitra chitra chitra
Type locality: “Khwae Noi and Khwae Yai rivers of Kanchanaburi
Province and in the Mae Klong river of Ratburi Province” [Thailand]. Restricted to “[Mae Klong in Ratburi and Khwae River in
Kanchanaburi]” by Nutaphand (1990:113); and to “Kanburi (=
Kanchanaburi), where the Khwae Noi and the Khwae Yai Rivers
join to form the Mae Klong River in Kanchanaburi Province,
Thailand” by McCord and Pritchard (2003:18).
Chitra chitra javanensis McCord and Pritchard 2003
Javanese Narrow-headed Softshell Turtle
Indonesia (Java, Sumatra)
Chitra selenkae † Jaekel 1911:80 (nomen suppressum,
ICZN 2005b) [Pleistocene, Pithecanthropus Trinil Beds,
Type locality: “India.” Restricted to “Fatehgahr, Ganges” [India] by
Smith (1931:162).
Testudo chitra Buchanan-Hamilton in Gray 1831d:47 (nomen
nudum)
Gymnopus lineatus Duméril and Bibron 1835:491, Trionyx
lineatus
Type locality: “le Gange” [India].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chitra vandijki McCord and Pritchard 2003
Burmese Narrow-headed Softshell Turtle
Win Ko Ko / CBFTT / TCC / Chindwin R., Myanmar
Brunei, Indonesia (Java, Kalimantan, Sumatra), Malaysia (Peninsular, East), Myanmar, Philippines (Luzon, Mindanao,
Mindoro, Palawan), Singapore, Thailand
IUCN Red List: Least Concern (2000); Previously: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
CITES: Appendix II
Trionyx subplanus Geoffroy Saint-Hilaire 1809b:11, Gymnopus subplanus, Amyda subplana, Dogania subplana,
Dogania subplanus
Type locality: Not known.
Trionyx frenatus Gray 1856b:67, Potamochelys frenatus,
Sarbieria frenata
Type locality: “Singapore.”
Myanmar
CBFTT Account: Platt, Platt, Win Ko Ko, and Rainwater (2014)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix I
Chitra burmanica Jaruthanin 2002:40 (nomen nudum)
Chitra vandijki McCord and Pritchard 2003:39
Type locality: “Khayansat Kone Village (23°16.30'N; 95°58.99'E)
on the Ayeyarwaddy River...Myanmar.”
Gymnopus javanicus Bibron in Gray 1856b:67 (nomen
nudum)
Dogania guentheri Gray 1862c:265, Trionyx guentheri,
Trionyx güntheri, Dogania güntheri
Type locality: “India.”
Trionyx dillwynii Gray 1873c:306
Type locality: “Borneo” [Malaysia or Indonesia].
Trionyx vertebralis Strauch 1890:113
Type locality: “Flusse Gabon im aequatorialen West-Afrika”
[in error]. Restricted to “Süd-Asien” by Wermuth and Mertens
(1961:255).
Trionyx pecki Bartlett 1895a:30
Dogania Gray 1844
Type locality: “Borneo” [Sarawak, Malaysia].
Dogania Gray 1844:49
Type species: Dogania subplana [= Trionyx subplanus Geoffroy
Saint-Hilaire 1809b], by original monotypy.
Sarbieria Gray 1869a:211
Type species: Sarbieria frenata [= Trionyx frenatus Gray 1856b] [=
subjective synonym of Trionyx subplanus Geoffroy Saint-Hilaire
1809b], by original monotypy.
Dogania subplana (Geoffroy Saint-Hilaire 1809b)
Malayan Softshell Turtle
Nilssonia Gray 1872a (07:80, 11:15) (78)
Nilssonia Gray 1872a:332
Type species: Nilssonia formosa [= Trionyx formosus Gray 1869a],
by original monotopy.
Isola Gray 1873h:51
Type species: Isola peguensis [= Trionyx peguensis Gray 1870c] [=
subjective synonym of Trionyx formosus Gray 1869a], by original
monotypy.
Aspideretes Hay 1904:274
Type species: Aspideretes gangeticus [= Trionyx gangeticus Cuvier
1825], by original designation.
Peter Paul van Dijk / CRM 2 / Khao Luang, Thailand
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Nilssonia formosa (Gray 1869a)
Burmese Peacock Softshell Turtle
Peter Paul van Dijk / TCC / Myanmar [captivity]
Afghanistan, Bangladesh, India (Bihar, Gujarat, Jammu, Madhya Pradesh, Odisha, Punjab, Rajasthan, Uttar Pradesh),
Nepal, Pakistan
IUCN Red List: Vulnerable A1d+2d (2000); Previously: Least
Concern [Not Listed] (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix I, as Aspideretes gangeticus
Trionyx gangeticus Cuvier 1825:186,203, Aspidonectes
gangeticus, Tyrse gangetica, Isola gangetica, Aspideretes
gangeticus, Trionyx gangeticus, Amyda gangetica, Nilssonia gangetica
Type locality: “du Gange” [India].
Trionyx javanicus Gray 1830e:19 (10:7) (partim, junior homonym, not = Amyda javanica Schweigger in Geoffroy
Saint-Hilaire 1809a), Tyrse javanica
Type locality: “Java and India.”
(orange dot = probably introduced)
China (?) (Yunnan [?]), Myanmar
IUCN Red List: Endangered A1cd+2d, B1+2c (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Trionyx formosus Gray 1869a:217, Nilssonia formosa,
Aspidonectes formosus, Isola formosa, Trionyx formosa,
Amyda formosus, Amyda formosa
Gymnopus duvaucelii Duméril and Bibron 1835:487 (partim,
nomen novum)
Trionyx gangeticus mahanaddicus Annandale 1912b:262
Type locality: “Sambalpur and Cuttack, Orissa” [India]. Restricted
to “Cuttack, Orissa” [India] by Smith (1931:167).
Nilssonia hurum (Gray 1830e) (10:7)
Indian Peacock Softshell Turtle
Type locality: “Pegu” [Myanmar].
Trionyx peguensis Gray 1870c:99, Isola peguensis
Type locality: “Pegu” [Myanmar].
Trionyx grayii Theobald 1875:176
Type locality: “neighbourhood of Thayet-myo...Irawadi valley”
[Myanmar].
Nilssonia gangetica (Cuvier 1825) (12:38)
Indian Softshell Turtle
Chittaranjan Baruah / CBFTT / Brahmaputra R., India
B.C. Choudhury / CRM 2 / India
Bangladesh, India (Assam, Bihar, Madhya Pradesh, Odisha,
Rajasthan, Uttar Pradesh, West Bengal), Nepal, Pakistan
CBFTT Account: Das, Basu, and Singh (2010)
IUCN Red List: Vulnerable A1cd+2d (2000); Previously: Least
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Concern [Not Listed] (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix I, as Aspideretes hurum
Trionyx occellatus Gray 1830d:pl.78 (nomen oblitum)
Type locality: “India.” Restricted to “Barrackpore, India” by Webb
(1980b:71).
Trionyx hurum Gray 1830e:18 (10:7), Isola hurum, Aspideretes
hurum, Aspidonectes hurum, Tyrse hurum, Amyda hurum,
Nilssonia hurum
Type locality: “India.” Restricted to “Barrackpore, India” by Webb
(1980b:71).
Testudo chim Buchanan-Hamilton in Gray 1831d:47 (nomen
nudum)
Trionyx ocellatus Gray 1832a:directions (nomen novum and
junior homonym, not = Trionyx ocellatus LeSueur 1827),
Testudo ocellata, Gymnopus ocellatus
Gymnopus duvaucelii Duméril and Bibron 1835:487 (partim,
nomen novum)
Trionyx sewaare Gray 1872a:336
Trionyx javanicus Gray 1830e:19 (10:7) (partim, junior homonym, not = Amyda javanica Schweigger in Geoffroy
Saint-Hilaire 1809a), Tyrse javanica
Type locality: “Java and India.”
Testudo gotaghol Buchanan-Hamilton in Gray 1831d:48 (nomen nudum)
Trionyx leithii Gray 1872a:334, Isola leithii, Aspideretes
leithii, Amyda leithii, Nilssonia leithii
Type locality: “Poonah” [India].
Aspilus gataghol Gray 1872a:339, Trionyx gataghol, Trionyx
gatajhal
Type locality: “India.”
Trionyx sulcifrons Annandale 1915b:341
Type locality: “Nagpur,...Central Provinces of India.”
Nilssonia nigricans (Anderson 1875) (07:81) (78)
Black Softshell Turtle, Bostami Softshell
Type locality: “Bengal” [India or Bangladesh].
Trionyx bellii Gray 1872a:337
Type locality: “Asia.”
Trionyx buchanani Theobald 1874:78
Type locality: “Bengal” [India or Bangladesh].
Trionyx hurum sivalensis † Lydekker 1889:9 [Late Pliocene
to Early Pleistocene, Siwaliks, India (Punjab)], Trionyx
sivalensis
Type locality: “India...Siwalik Hills.”
Nilssonia leithii (Gray 1872a)
Leith’s Softshell Turtle
Chittaranjan Baruah / TCC / Madhab Temple, Hajo, Assam, India
Rahul Naik / CBFTT / Godavari R. nr. Nashik, Maharashtra, India
(orange dots = temple ponds)
Bangladesh, India (Assam, Tripura, West Bengal)
IUCN Red List: Extinct in the Wild (2002); Previously: Critically Endangered (2000), Critically Endangered (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix I, as Aspideretes nigricans
Trionyx nigricans Anderson 1875:284, Amyda nigricans,
Aspideretes nigricans, Nilssonia nigricans
Type locality: “Chittagong, Bengal” [Bangladesh].
Palea Meylan 1987
Palea Meylan 1987:77
India (Andhra Pradesh, Karnataka, Kerala, Madhya Pradesh,
Maharashtra, Odisha, Tamil Nadu)
CBFTT Account: Das, Sirsi, Vasudevan, and Murthy (2014)
IUCN Red List: Vulnerable A1c (2000); Previously: Near
Threatened (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Type species: Palea steindachneri [= Trionyx steindachneri Siebenrock 1906a], by original designation.
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Pelochelys Gray 1864b
Palea steindachneri (Siebenrock 1906a)
Wattle-necked Softshell Turtle
Pelochelys Gray 1864b:89
Type species: Pelochelys cantorii Gray 1864b, by subsequent designation by Günther (1865:108).
Pelochelys (Ferepelochelys) Hoser 2014a:62 (unavailable name
pending ICZN decision; Rhodin et al. 2015) (4, 79)
Pelochelys bibroni (Owen 1853)
New Guinea Giant Softshell Turtle
Torsten Blanck / Hong Kong, China
Anders G.J. Rhodin / TCF / CCB / Lake Murray, Western Prov., Papua New Guinea
(orange dots = trade or introduced)
China (Guangdong, Guizhou, Yunnan), Vietnam
Introduced: China (Hong Kong), Mauritius, USA (Hawaii
[Kauai])
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Near Threatened (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II
Aspidonectes californiana Rivers 1889:233 (nomen suppressum, ICZN 1982), Pelodiscus californianus
Type locality: “Sacramento River, near the city of Sacramento...
California” [USA] [in error, trade specimen].
Trionyx steindachneri Siebenrock 1906a:579 (nomen
conservandum, ICZN 1982), Amyda steindachneri, Palea
steindachneri, Pelodiscus steindachneri
Type locality: “Kau-Kongriver...Insel Hainan” [China].
Indonesia (Papua), Papua New Guinea (Southern)
IUCN Red List: Vulnerable A1cd+2cd (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II, as Pelochelys spp.
Trionyx (Gymnopus) bibroni Owen 1853:185, Trionyx
bibroni, Pelochelys bibroni, Pelochelys (Ferepelochelys)
bibroni, Ferepelochelys bibroni
Type locality: “Australian.” Restricted to “Laloki River, Astrolabe
Range, 40 miles [ca. 65 km] from its entry into Redscar Bay
(9°20'S 147°14'E), Central District, Papua New Guinea” by
neotype designation by Webb (1995a:302).
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Pelochelys cantorii Gray 1864b (79)
Asian Giant Softshell Turtle, Cantor’s Giant Softshell Turtle
Pelochelys signifera Webb 2003
Northern New Guinea Softshell Turtle
Anders G.J. Rhodin / CCB / Maprik, East Sepik Prov., Papua New Guinea
Annette Olsson / CBFTT / TCC / nr. Kratie, Mekong R., Cambodia
Bangladesh, Brunei (?), Cambodia, China (Fujian, Guangdong,
Guangxi, Hainan, Zhejiang), India (Kerala, Odisha, Tamil
Nadu, West Bengal), Indonesia (Kalimantan, Sumatra),
Laos, Malaysia (Peninsular, East), Myanmar, Philippines
(Luzon, Mindanao, Palawan), Singapore (extirpated),
Thailand, Vietnam
CBFTT Account: Das (2008)
IUCN Red List: Endangered A1cd+2cd (2000); Previously:
Vulnerable (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Pelochelys spp.
Pelochelys cantorii Gray 1864b:90
Type locality: “Malacca” [Malaysia].
Pelochelys cumingii Gray 1864b:90, Chitra indica cumingii
Type locality: “Philippines.”
Pelochelys cantoris Boulenger 1889:ix (nomen novum)
Pelochelys poljakowii Strauch 1890:118
Type locality: “Fu-tschau” [China]. Emended to “Fu-tschau
[Fuzhou Shi], Fujian Province, China” by Fritz and Havaš
(2008:317).
Indonesia (Papua), Papua New Guinea (Northern)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II, as Pelochelys spp.
Pelochelys signifera Webb 2003:100, Pelochelys (Ferepelochelys) signifera, Ferepelochelys signifera
Type locality: “Wanggar River, Weyland Range, Geelvinck Bay,
N. New Guinea (Papua Province, Indonesia).”
Pelodiscus Fitzinger 1835 (07:82, 10:32, 11:16)
Trionyx (Pelodiscus) Fitzinger 1835:110
Type species: Aspidonectes (Pelodiscus) sinensis [= Trionyx (Aspidonectes) sinensis Wiegmann 1834], by subsequent designation by
Fitzinger (1843:30).
Landemania Gray 1869a:211
Type species: Landemania irrorata Gray 1869a [= subjective
synonym of Trionyx (Aspidonectes) sinensis Wiegmann 1834], by
original designation.
Psilognathus Heude 1880:24
Type species: Psilognathus laevis Heude 1880 [= subjective synonym of Trionyx (Aspidonectes) sinensis Wiegmann 1834], by
original monotypy.
Chitra minor † Jaekel 1911:80 [Pleistocene, Pithecanthropus
Trinil Beds, Indonesia (Java)]
Temnognathus Heude 1880:25
Pelochelys clivepalmeri Hoser 2014a:62 (unavailable name
pending ICZN decision; Rhodin et al. 2015) (4, 79)
Pelochelys telstraorum Hoser 2014a:62 (unavailable name
pending ICZN decision; Rhodin et al. 2015) (4, 79)
Gomphopelta Heude 1880:27
Type locality: “Pithecanthropus-schichten...Java...Trinil”
[Indonesia].
Type species: Temnognathus mordax Heude 1880 [= subjective
synonym of Trionyx (Aspidonectes) sinensis Wiegmann 1834], by
original monotypy.
Type species: Gomphopelta officinae Heude 1880 [= subjective
synonym of Trionyx (Aspidonectes) sinensis Wiegmann 1834], by
original monotypy.
Coelognathus Heude 1880 :29
Type species: Coelognathus novemcostatus Heude 1880 [= subjective synonym of Trionyx (Aspidonectes) sinensis Wiegmann
1834], by original monotypy.
Tortisternum Heude 1880:31
Type species: Tortisternum novemcostatum Heude 1880 [= subjective
synonym of Trionyx (Aspidonectes) sinensis Wiegmann 1834], by
original monotypy.
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Ceramopelta Heude 1880:33
Type species: Ceramopelta latirostris Heude 1880 [= subjective
synonym of Trionyx (Aspidonectes) sinensis Wiegmann 1834], by
original monotypy.
Coptopelta Heude 1880:34
Type species: Coptopelta septemcostata Heude 1880 [= subjective
synonym of Trionyx (Aspidonectes) sinensis Wiegmann 1834], by
original monotypy.
Cinctisternum Heude 1880:36
Type species: Cinctisternum bicinctum Heude 1880 [= subjective
synonym of Trionyx (Aspidonectes) sinensis Wiegmann 1834], by
original monotypy.
Pelodiscus axenaria (Zhou, Zhang, and Fang 1991) (10:32, 11:16)
Hunan Softshell Turtle
China (Guangxi, Hunan)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II
Trionyx axenaria Zhou, Zhang, and Fang 1991:382, Pelodiscus axenaria, Pelodiscus axenarius
(orange dots = possibly native or historic introduction)
China (Heilongjiang, Jilin, Liaoning, Nei Monggu), North
Korea, Russia (Amurskaya, Khabarovskiy, Primorskiy,
Yevreyskaya), Japan (historic introduction?), South Korea
Introduced: USA (Hawaii [Oahu])
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II
Trionyx maackii Brandt 1857:110, Amyda maackii, Pelodiscus maackii
Type locality: “in fluminibus australioribus lateralibus Amuris,
nominatim in fluviis Sungari et Ussuri, nec non in ipsius fluminis
Amuris parte inter fluvios modo dictos obvia” [in the more
southern rivers next to the Amur, namely the Sungari and Ussuri
rivers, and also in the part of the Amur river between these rivers]
[Russia]. Restricted to “Southern Amur R. and Ussuri R.” [Russia]
by Stejneger (1907:529).
Pelodiscus parviformis Tang 1997 (10:32, 11:16)
Vietnamese Softshell Turtle
Type locality: “Taoyuan, Pingjiang, Rucheng, Lingling, Shaoyang
Counties, Hunan Province, China” [in Chinese].
Pelodiscus maackii (Brandt 1857) (10:32, 11:16) (80)
Northern Chinese Softshell Turtle
Peter Paul van Dijk / Vietnam [trade; Cuc Phuong Turtle Conservation Center]
Shi Haitao / nr. Nagoya, Japan [captivity]
China (Guangxi, Hunan), Vietnam
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
CITES: Appendix II
Pelodiscus parviformis Tang 1997:13
Type locality: “Quanzhou, Xing’an, Guanyang, Ziyuan, Lingchuan counties of Guangxi Auto. Region; Dong’an, Qiyang,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Daoxian counties of Hunan Province” [China].
Pelodiscus sinensis (Wiegmann 1834) (10:32, 11:16, 14:30) (80, 81)
Chinese Softshell Turtle
Trionyx tuberculatus Cantor 1842a:16 (82) (nomen nudum)
Trionyx tuberculatus Cantor 1842b:482 (82), Potamochelys
tuberculatus, Amyda tuberculata, Trionyx sinensis tuberculatus, Amyda sinensis tuberculata, Pelodiscus sinensis
tuberculatus
Type locality: “Chusan...Island” [Zhoushan Island, Zhejiang,
China].
Tyrse perocellata Gray 1844:48, Trionyx perocellatus,
Potamochelys perocellatus, Landemania perocellata,
Gymnopus perocellatus
Type locality: “Canton, China” [Guangzhou Shi, Guandong,
China].
Trionyx schlegelii Brandt 1857:111, Amyda schlegelii
Type locality: “China borealis...Pekingo” [Beijing, China].
Landemania irrorata Gray 1869a:216
Type locality: “Shanghai” [China].
Tien-Hsi Chen / Keelung R., nr. Taipei, Taiwan
Trionyx peroculatus Günther in Gray 1869a:216 (nomen
nudum)
Gymnopus simonii David 1875:214 (nomen nudum)
Psilognathus laevis Heude 1880:24
Type locality: “la région montagneuse au sud de la ville de Ningkouo fou” [Ningguo Co., Anhui, China].
Temnognathus mordax Heude 1880:26
Type locality: “environs de Chang-hai” [Shanghai, China].
Gomphopelta officinae Heude 1880:27
Type locality: “la Houai, vers l’endroit où cette rivière sort de la
province du Ho-nan” [Huai River, Henan, China].
Coelognathus novemcostatus Heude 1880:29
Type locality: “l’extrémité orientale du lac Tch’ao” [Lake Chao,
Anhui, China].
Tortisternum novemcostatum Heude 1880:31
China (Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou,
Hainan, Hebei, Henan, Hong Kong, Hubei, Hunan,
Jiangsu, Jiangxi, Nei Mongol, Shaanxi, Shandong, Shanxi,
Sichuan, Yunnan, Zhejiang), Taiwan
Introduced: Brazil (Pará), Guam, Indonesia (Kalimantan, Sumatra, Timor), Iran, Japan (Bonin Islands, Honshu, Kyoshu,
Ryukyu Archipelago, Shikoku), Laos, Malaysia (Peninsular, East), Northern Mariana Islands, Philippines (Bohol,
Cebu, Latvia, Leyte, Luzon, Mindanao, Mindoro, Panay),
Singapore, South Korea, Spain, Thailand, Timor-Leste,
USA (Hawaii [Oahu]), Vietnam
IUCN Red List: Vulnerable A1d+2d (2000); Previously: Least
Concern [Not Listed] (1996)
TFTSG Draft Red List: Vulnerable or Endangered (2011)
Testudo rostrata Thunberg 1787:179 (nomen suppressum,
ICZN 1991), Emydes rostrata
Type locality: Not known.
Testudo striata Suckow 1798:37 (partim, nomen novum)
Testudo semimembranacea Hermann 1804:219 (nomen suppressum, ICZN 1963)
Type locality: “Japania” [Japan].
Trionyx stellatus var. Japon Temminck and Schlegel
1834:pls.5,7 (10:18, 14:30) (invalid vernacular name)
Trionyx (Aspidonectes) sinensis Wiegmann 1834:189 (81) (nomen conservandum, ICZN 1963, 1991), Trionyx sinensis,
Pelodiscus sinensis, Tyrse sinensis, Amyda sinensis, Trionyx sinensis sinensis, Amyda sinensis sinensis, Pelodiscus
sinensis sinensis
Type locality: “kleinen Insel im Tigerflusse, dicht bei Macao”
[China].
Trionyx japonica Temminck and Schlegel 1838:139 (14:30), Amyda
japonica, Pelodiscus sinensis japonicus
Type locality: “Japon” [Japan].
Type locality: “lac Tch’ao, département de Lu-tcheou” [Lake
Chao, Anhui, China].
Ceramopelta latirostris Heude 1880:33
Type locality: “environs de Ngan-k’ing fou” [Anqing Shi, Anhui,
China].
Coptopelta septemcostata Heude 1880:35
Type locality: “lacs de Tong-lieou (Ngan-houé)” [Dongliu, Anhui,
China].
Cinctisternum bicinctum Heude 1880:37
Type locality: “marais de Ngan-K’ing” [Anqing Shi, Anhui,
China].
Trionyx cartilagineus newtoni Ferreira 1897:114, Trionyx
sinensis newtoni
Type locality: “Timor” [Timor-Leste] [in error, trade specimen]
Amyda schlegelii haseri Pavlov 1932:27
Type locality: ”Tzu ya ho, the river of Sien hien (central Chili)”
[Ziya He, Xian Co., Hebei, China].
Amyda schlegelii licenti Pavlov 1932:28
Type locality: “Tientsin” [Tianjin, China].
Amyda schlegelii laoshanica Pavlov 1933:3
Type locality: “Chantong, Laoshan, near Tsingtao” [Qingdao Shi,
Shandong, China].
Rafetus Gray 1864b
Rafetus Gray 1864b:81
Type species: Rafetus euphraticus [= Testudo euphratica Olivier in
Daudin 1801], by original monotypy.
Oscaria Gray 1873g:157
Type species: Oscaria swinhoei Gray 1873g, by original monotypy.
Yuen Heude 1880:18
Type species: Yuen leprosus Heude 1880 [= subjective synonym of
Oscaria swinhoei Gray 1873g], by subsequent designation by
Stejneger (1907:514).
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Rafetus euphraticus (Olivier in Daudin 1801) (83, 84)
Euphrates Softshell Turtle
Hanyeh Ghaffari / CBFTT / Balarood R., Khuzestan Prov., Iran
China (Anhui? [extirpated], Jiangsu? [extirpated], Yunnan,
Zhejiang? [extirpated]), Vietnam
IUCN Red List: Critically Endangered A1cd+2cd (2000)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Oscaria swinhoei Gray 1873g:157, Pelodiscus swinhoei,
Trionyx swinhoei, Amyda swinhoei, Rafetus swinhoei
Type locality: “neighbourhood of Shanghai” [China].
Yuen leprosus Heude 1880:20
Type locality: “le Houang-p’ou, à Chang-hai” [Huangpu Jiang,
Shanghai, China].
Yuen maculatus Heude 1880:22, Pelochelys maculatus
Iran (Ilam, Khuzestan, Lorestan), Iraq, Syria, Turkey
CBFTT Account: Taskavak, Atatür, Ghaffari, and Meylan (2016)
IUCN Red List: Endangered A1ac+2c (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Rafetus spp.
Testudo euphratica Olivier in Daudin 1801:305 (83), Trionyx
euphraticus, Gymnopus euphraticus, Rafetus euphraticus, Pelodiscus euphraticus, Amyda euphratica, Tyrse
euphratica
Type locality: “le Houang-p’ou” [Huangpu Jiang, Shanghai,
China].
Yuen elegans Heude 1880:23
Type locality: “Houang-p’ou” [Huangpu Jiang, Shanghai, China].
Yuen viridis Heude 1880:23
Type locality: “Grand Lac, à Sou-tcheou” [Tai Hu, Suzhou Shi,
Jiangsu, China].
Yuen pallens Heude 1880:23
Type locality: “Grand Lac, près de Sou-tcheou” [Tai Hu, Suzhou
Shi, Jiangsu, China].
Type locality: “le Tigre et l’Euphrate” [Tigris and Euphrates
rivers, Iraq]. Restricted to “Euphrates (Al Firat), vicinity of Anah,
Al-Anbar, Iraq” by Bour et al. (1995:85).
Trionyx swinhonis Boulenger 1889:ix (nomen novum)
Pelochelys taihuensis † Zhang 1984:71 [Holocene, Neolithic,
subfossil, China (Zhejiang)]
Type locality: “le Tigre..[&]..l’Euphrate” [Tigris and Euphrates
rivers, Iraq].
Trionyx liupani † Tao 1986:23 [Late Pleistocene, Taiwan]
Testudo rafcht Olivier 1807:328
Testudo rascht Gray 1830e:19 (nomen novum)
Tyrse rafeht Gray 1844:49 (nomen novum), Trionyx rafeht,
Testudo rafeht
Rafetus swinhoei (Gray 1873g) (07:83, 10:33, 11:17) (85)
Red River Giant Softshell Turtle, Yangtze Giant Softshell Turtle,
Swinhoe’s Softshell Turtle
Type locality: “Luojiajiao Relics, Tongxiang County, Zhejiang”
[China].
Type locality: “Penghu Channel in the Taiwan Strait” [Taiwan,
China].
Rafetus hoankiemensis Ha 1995:4 (nomen nudum)
Rafetus hoguomensis Ha 1995:4 (nomen nudum)
Rafetus leloii Ha 2000:104 (11:17)
Type locality: “Hoan Kiem Lake, Ha Noi” [Vietnam].
Rafetus vietnamensis Le, Le, Tran, Phan, Phan, Tran, Pham,
Nguyen, Nong, Phan, Dinh, Truong, and Ha 2010:950 (10:33, 11:17)
Type locality: Not known; holotype from “Hung Ky Pagoda,
Hanoi” [Vietnam].
Trionyx Geoffroy Saint-Hilaire 1809a
Trionyx Geoffroy Saint-Hilaire 1809a:363
Type species: Trionyx “tortue du Nil” [= Trionyx egyptiacus Geoffroy
Saint-Hilaire 1809a] [= subjective synonym of Testudo triunguis
Forskål 1775], by subsequent designation by Bory de SaintVincent (1828:77).
Aspidonectes Wagler 1830b:134
Timothy E.M. McCormack / TCC / nr. Hanoi, Vietnam
Type species: Aspidonectes aegyptiacus [= Trionyx aegyptiacus
Geoffroy Saint-Hilaire 1809b] [= subjective synonym of Testudo
triunguis Forsskål 1775), by subsequent designation by Fitzinger
(1843:30).
Gymnopus Duméril and Bibron 1835:472 (nomen novum)
Tyrse Gray 1844:47
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type species: Tyrse aegyptiacus [= Trionyx aegyptiacus Geoffroy
Saint-Hilaire 1809b] [= subjective synonym of Testudo triunguis
Forskål 1775], by original designation.
Fordia Gray 1869a:219
Type species: Fordia africana Gray 1869a [= subjective synonym of
Testudo triunguis Forskål 1775], by original monotypy.
Trionyx triunguis (Forskål 1775) (10:29, 11:18)
African Softshell Turtle, Nile Softshell Turtle
Subpopulations: Mediterranean: Critically Endangered
C2a (1996)
TFTSG Draft Red List: Vulnerable (2013)
CITES: Appendix II, as Trionyx spp.
Testudo triunguis Forskål 1775:ix (10:29), Amyda triunguis,
Trionyx triunguis, Pelodiscus triunguis, Aspidonectes
triunguis, Tyrse triunguis, Amyda triunguis triunguis
Type locality: “Nilo” [Nile River, Egypt].
Testudo striata Suckow 1798:37 (partim, nomen novum)
Trionyx egyptiacus Geoffroy Saint-Hilaire 1809a:366
Type locality: “l’Egypte” [Egypt]. Restricted to “the lower Nile
River drainage” [Egypt] by Bour et al. (1995:77).
Trionyx aegyptiacus Geoffroy Saint-Hilaire 1809b:12 (nomen novum), Aspidonectes aegyptiacus, Gymnopodus
aegyptiacus, Gymnopus aegyptiacus
Trionyx niloticus Gray 1831d:46 (nomen novum), Tyrse
nilotica, Aspidonectes niloticus
Trionyx labiatus Bell 1835:unnumbered, Trionyx (Pelodiscus) labiatus
Type locality: “Africâ occidentali...Sierra Leone.”
Trionyx mortoni Hallowell 1844:120
Oguz Türkozan / Dalaman, Turkey
Type locality: “Africa.”
Aspidonectes aspilus Cope 1860:295, Gymnopus aspilus
Type locality: “Rembo and Ovenga rivers, tributaries of the
Fernando Vas, Equatorial West Africa” [Gabon].
Fordia africana Gray 1869a:219
Type locality: “Upper Nile, Chartoum” [Sudan].
Trionyx triunguis rudolfianus Deraniyagala 1948:30, Amyda
triunguis rudolfianus
Type locality: “Ferguson’s Gulf, Lake Rudolf, Africa” [Kenya].
pleuRodiRa Cope 1864
(08:20)
Pleurodères Duméril and Bibron 1834:354
Pleurodera Lichtenstein 1856:2 (08:20)
Pleurodera Cope 1864:181
Pleurodira Cope 1865:186
Chelidae Gray 1825 (12:39) (86)
(orange dots = introduced, trade, or uncertain)
Angola, Benin, Cameroon, Central African Republic, Chad,
Congo (DRC), Congo (ROC), Egypt, Equatorial Guinea,
Eritrea, Ethiopia, Gabon, Gambia, Ghana, Greece, Guinea,
Guinea-Bissau, Israel, Ivory Coast, Kenya, Lebanon, Liberia, Mauritania, Namibia, Niger, Nigeria, Senegal, Sierra
Leone, Somalia, South Sudan, Sudan, Syria, Togo, Turkey,
Uganda
Introduced: Israel (inland)
Mediterranean subpopulation: Egypt (extirpated), Greece, Israel,
Lebanon, Syria, Turkey
IUCN Red List: Global: Least Concern [Not Listed] (1996);
Chelides Cuvier 1816:14
Chelydes Schmid 1819:17
Chelidina Gray 1825:211
Chelydoidea Fitzinger 1826:7
Chelydae Gray 1831d:7
Chelydidae Gray 1831d:37
Hydraspidina Bonaparte 1836:3 (partim)
Chelina Bonaparte 1836:4
Hydraspididae Agassiz 1857a:249
Chelydidi Portis 1890:17
Chelyidae Baur 1893a:211
Chelodinidae Baur 1893a:211
Hydromedusidae Baur 1893a:211
Rhinemydidae Baur 1893a:212
Chelidae Lindholm 1929:289
Cheluidae Storr 1978:303 (86)
Chelinae Gray 1825 (12:39)
Chelides Cuvier 1816:14
Chelydes Schmid 1819:17
Chelidina Gray 1825:211
Chelina Bonaparte 1831:63
Chelidinae Georges, Birrell, Saint, McCord, and Donnellan 1998:235
Chelinae Turtle Taxonomy Working Group 2012:289
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Acanthochelys Gray 1873c (14:39)
Acanthochelys Gray 1873c:305
Type species: Acanthochelys spixii [= Platemys spixii Duméril and
Bibron 1835], by original monotypy.
Acanthochelys macrocephala (Rhodin, Mittermeier, and McMorris
1984) (07:84)
Pantanal Swamp Turtle, Big-headed Pantanal Swamp Turtle
(orange dot = introduced)
Thomas and Sabine Vinke / CBFTT / nr. Filadelfia, Chaco, Paraguay
Argentina (Chaco, Formosa, Salta, Santa Fe), Bolivia (Tarija),
Paraguay
Introduced: Argentina (Mendoza)
CBFTT Account: Vinke, Vinke, Richard, Cabrera, Paszko,
Marano, and Métrailler (2011)
IUCN Red List: Endangered A2cde+3cde+4cde; C1+2a(i)
(2016); Previously: Vulnerable (1996)
Platemys pallidipectoris Freiberg 1945:19, Acanthochelys
pallidipectoris
Type locality: “Pcia. [Presidencia] Roque Sáenz Peña, Chaco”
[Argentina].
Acanthochelys radiolata (Mikan 1820) (14:39) (87)
Brazilian Radiolated Swamp Turtle
(orange dot = probable trade)
Bolivia (Santa Cruz), Brazil (Mato Grosso, Mato Grosso do
Sul), Paraguay
CBFTT Account: Rhodin, Métrailler, Vinke, Vinke, Artner,
and Mittermeier (2009)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Near Threatened or Least Concern
(2012)
Phrynops schoepffii Fitzinger in Siebenrock 1904b:27 (nomen nudum, partim)
Platemys macrocephala Rhodin, Mittermeier, and McMorris
1984a:38, Acanthochelys macrocephala
Russell A. Mittermeier / Alagoas, Brazil
Type locality: “Caiçara, Rio Paraguai, Mato Grosso, Brazil
(16°03' S 57°43' W).”
Phrynops chacoensis Fritz and Pauler 1992:299 (07:84), Acanthochelys chacoensis, Mesoclemmys chacoensis
Type locality: “Paraguayanischer Chaco, 22°30'3" S, 59°44'30"
W” [Paraguay].
Acanthochelys pallidipectoris (Freiberg 1945)
Chaco Side-necked Turtle
(orange dots = misidentified or uncertain)
Brazil (Alagoas, Bahia, Espírito Santo, Minas Gerais, Rio de
Janeiro, Sergipe)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Data Deficient (2011)
Emys radiolata Mikan 1820:[unpaginated], Chelodina radiolata, Rhinemys radiolata, Chelys (Hydraspis) radiolata,
Chelys radiolata, Hydraspis radiolata, Platemys radiolata,
Platemys radiolata radiolata, Acanthochelys radiolata
Thomas and Sabine Vinke / CBFTT / Boquerón, Paraguay
Type locality: “Sebastianopoli...Brasilia” [Rio de Janeiro, Brazil].
Platemys gaudichaudii Duméril and Bibron 1835:427,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Hydraspis gaudichaudii
Type locality: “Brésil” [Brazil].
Platemys werneri Schnee 1900:463
Type locality: “Umgebung von São Paulo” [Brazil].
Chelus fimbriata (Schneider 1783) (88)
Matamata Turtle
Platemys radiolata quadrisquamosa Luederwaldt 1926:437,
Platemys quadrisquamosa
Type locality: “Rio Doce (Est. do Espirito Santo)..[&] Belmonte
(Bahia)” [Brazil].
Acanthochelys spixii (Duméril and Bibron 1835)
Black Spiny-necked Turtle, Spix’s Sideneck Turtle
Rafael Bernhard / Cachoeira do Caldeirão, Rio Madeira, Amazonas, Brazil
Russell A. Mittermeier / São Paulo, Brazil
(orange dots = probable introduced or trade)
Brazil (Bahia, Goiás, Minas Gerais, Paraná, Rio Grande do Sul,
Santa Catarina, São Paulo), Uruguay
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Near Threatened (2011)
Emys depressa Spix 1824:4 (junior homonym, not = Emys
depressa Merrem 1820)
Type locality: “provinciarum Rio de Janeiro et fluminis Sti
Francisci” [Brazil]. Restricted to “Rio São Francisco, near Rio dos
Pandeiros, Minas Gerais, Brazil” by Rhodin et al. (1984b:783)
by restriction of nomen novum replacement name Platemys spixii
Duméril and Bibron 1835.
Emys aspera Cuvier in Gray 1830e:16 (10:7) (nomen oblitum)
Type locality: “America.” Restricted to “Brésil” [Brazil] by Bour
and Pauler (1987:14).
Platemys spixii Duméril and Bibron 1835:409 (nomen
novum), Hydraspis spixii, Acanthochelys spixii, Platemys
radiolata spixii
Type locality: “Brésil” [Brazil]. Restricted to “Rio São Francisco,
near Rio dos Pandeiros, Minas Gerais, Brazil” by Rhodin et al.
(1984b:783).
Chelus Duméril 1805 (10:11)
Chelus Duméril 1805:76 (10:11)
Type species: Chelus fimbriata [= Testudo fimbriata Schneider 1783],
by original monotypy.
Chelys Oppel 1811:6 (nomen novum)
Chelyda Rafinesque 1815:75 (nomen novum)
Matamata Merrem 1820:21 (nomen novum)
Bolivia (Beni, Pando, Santa Cruz), Brazil (Amapá, Amazonas,
Goiás, Mato Grosso, Pará, Rondônia, Roraima, Tocantins),
Colombia (Amazonas, Arauca, Caquetá, Casanare, Guainía,
Meta, Putumayo, Vaupés, Vichada), Ecuador, French Guiana, Guyana, Peru (Loreto, Ucayali), Suriname, Trinidad,
Venezuela (Amazonas, Anzoátegui, Apure, Barinas, Bolívar,
Cojedes, Delta Amacuro, Guárico, Monagas, Sucre, Zulia)
CBFTT Account: Pritchard (2008)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Testudo terrestris Fermin 1765:51 (junior homonym, not
= Testudo terrestris Garsault 1764 or Testudo terrestris
Forskål 1775; nomen suppressum, ICZN 1963)
Type locality: “Surinam.”
Testudo fimbriata Schneider 1783:349 (nomen conservandum, ICZN 1963), Chelus fimbriata, Chelys fimbriata,
Matamata fimbriata, Chelus fimbriatus
Type locality: “Surinam...von Aprouague..[&]..Remire.”
Testudo fimbria Gmelin 1789:1043 (nomen novum), Chelys
fimbria
Testudo matamata Bruguière 1792:257, Emydes matamata,
Chelus matamata, Chelys matamata
Type locality: “Cayenne” [French Guiana].
Testudo bispinosa Ruiz de Xelva in Daudin 1801:94, Chelys
bispinosa, Matamata bispinosa
Type locality: “Brésil” [Brazil].
Testudo rapara Gray 1831d:44 (nomen nudum)
Testudo raparara Gray 1844:44 (nomen nudum)
Chelys boulengerii Baur 1890b:968
Type locality: “Orinoco” [Venezuela].
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Mesoclemmys Gray 1873c (07:100)
Mesoclemmys Gray 1873c:305
Mesoclemmys gibba (Schweigger 1812)
Gibba Turtle
Type species: Mesoclemmys gibba [= Emys gibba Schweigger 1812],
by original monotypy.
Batrachemys Stejneger 1909:126
Type species: Batrachemys nasuta [= Emys nasuta Schweigger
1812], by original monotypy.
Bufocephala McCord, Joseph-Ouni, and Lamar 2001:732
Type species: Bufocephala vanderhaegei [= Phrynops tuberculatus
vanderhaegei Bour 1973], by original designation.
Ranacephala McCord, Joseph-Ouni, and Lamar 2001:732
Type species: Ranacephala hogei [= Phrynops hogei Mertens 1967],
by original designation.
Mesoclemmys dahli (Zangerl and Medem 1958) (12:40)
Dahl’s Toad-headed Turtle
Anders G.J. Rhodin / Maipuco, Rio Maranon, Peru
German Forero-Medina / CBFTT / Colombia
Colombia (Atlántico, Bolívar, Cesar, Córdoba, Magdalena,
Sucre)
CBFTT Account: Forero-Medina, Castaño-Mora, CárdenasArevalo, and Medina-Rangel (2013)
IUCN Red List: Critically Endangered B1+2c (1996)
TFTSG Draft Red List: Endangered (2011)
Phrynops (Batrachemys) dahli Zangerl and Medem
1958:376, Phrynops dahli, Batrachemys dahli, Phrynops
nasutus dahli, Mesoclemmys dahli
Type locality: “Vicinity of Sincelejo, Bolivar, Colombia.”
Bolivia, Brazil (Acre, Amapá, Amazonas, Mato Grosso, Pará,
Roraima), Colombia (Amazonas, Arauca, Caquetá,
Casanare, Guainía, Guaviare, Meta, Putumayo, Vaupés,
Vichada), Ecuador, French Guiana, Guyana, Peru (Amazonas, Loreto, Madre de Dios, Puno, Ucayali), Suriname,
Trinidad, Venezuela (Amazonas, Bolívar, Delta Amacuro,
Monagas)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Emys gibba Schweigger 1812:299, Rhinemys gibba, Hydraspis cayennensis gibba, Platemys gibba, Hydraspis
(Podocnemis) gibba, Hydraspis gibba, Phrynops gibbus,
Mesoclemmys gibba, Phrynops (Mesoclemmys) gibba,
Mesoclemmys gibbus
Type locality: Not known. Restricted to “Amérique méridional”
by Duméril and Duméril (1851:20); and to “environs de Cayenne,
Guyane française” [French Guiana] by Bour and Pauler (1987:7).
Emys stenops Spix 1824:12, Hydraspis stenops
Type locality: “fluminis Solimoëns” [Rio Solimões, Amazonas,
Brazil]. Restricted to “l’Amazone entre Tefé et Tabatinga au Brésil
(Amazonas)” [Brazil] by Bour and Pauler (1987:7).
Platemys miliusii Duméril and Bibron 1835:431, Phrynops
miliusii, Hydraspis miliusii
Type locality: “Cayenne” [French Guiana].
Hydraspis gordoni Gray 1868:563
Type locality: “Trinidad, near the mountain of Tamana.”
Hydraspis bicolor Gray 1873c:304
Type locality: “Demerara Falls” [Guyana].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
181
Mesoclemmys heliostemma (McCord, Joseph-Ouni, and Lamar
2001) (07:101, 12:41)
Yellow-crowned Toad-headed Turtle
(orange dot = probable erroneous type locality)
Jiří Moravec / nr. Anguilla, Rio Nanay, Loreto, Peru
Brazil (Espírito Santo, Minas Gerais, Rio de Janeiro, São
Paulo?)
IUCN Red List: Critically Endangered A2bcd+4bcd (2016);
Previously: Endangered (1996)
Phrynops hogei Mertens 1967:73, Mesoclemmys hogei,
Ranacephala hogei
Type locality: “Rio Pequena, südwestlich von São Paulo, Brasilien” [Brazil] [in error]. Emended to “Rio Pequeno, southeast of
São Paulo” [Brazil] [in error?] by Rhodin et al. (1982:179).
Mesoclemmys nasuta (Schweigger 1812)
Guyanan Toad-headed Turtle
(orange dot = trade)
Brazil (Acre, Amazonas, Mato Grosso, Pará, Rondônia, Roraima), Colombia (Amazonas, Caquetá, Guainía, Putumayo,
Vaupés), Ecuador, Peru (Loreto), Venezuela (Amazonas)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
Batrachemys heliostemma McCord, Joseph-Ouni, and Lamar
2001:734, Mesoclemmys heliostemma
Type locality: “base of Pico da Neblina (situated on the Venezuela/
Brazil border) on the left bank of Río Baria (= Río Mawarinuma)
[4º95'N, 66º10'W], a tributary of the Rio Negro, Amazonas,
Venezuela.” GPS coordinates incorrect, emended here to 0º50'N,
66º10'W.
Jérôme Maran / French Guiana
Mesoclemmys hogei (Mertens 1967)
Hoge’s Side-necked Turtle
Russell A. Mittermeier / TCF / TCC / Espírito Santo, Brazil
Brazil (Amapá), French Guiana, Suriname
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Data Deficient (2011)
Emys nasuta Schweigger 1812:298, Rhinemys nasuta,
Hydraspis (Rhinemys) nasuta, Hydraspis nasuta, Platemys
nasuta, Batrachemys nasuta, Phrynops (Batrachemys)
nasuta, Phrynops nasuta, Phrynops nasutus, Phrynops
nasutus nasutus, Phrynops nasuta nasuta, Batrachemys
nasutus, Mesoclemmys nasuta
Type locality: Not known. Restricted to “Amérique méridionale”
by Duméril and Bibron (1835:437); to “Guyanes et au nord-est de
l’Amazonie” by Lescure and Fretey (1976:1318); and to “rivières
Ouaqui et Inini, bassin du Maroni en amont de Maripasoula,
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Guyane française” [French Guiana] by Bour and Pauler (1987:6).
Emys barbatula Gravenhorst 1829:15, Hydraspis barbatula
Type locality: Not known.
Platemys schweiggerii Duméril and Bibron 1835:435 (nomen
novum)
Phrynops walbaumi Fitzinger in Siebenrock 1904b:20 (nomen nudum)
Mesoclemmys perplexa Bour and Zaher 2005
Cerrado Side-necked Turtle
Vinicius T. de Carvalho / Piauí, Brazil
Bolivia, Brazil (Acre, Amazonas, Rondônia, Roraima), Colombia (Amazonas, Caquetá, Guainía, Putumayo, Vaupés),
Ecuador, Peru (Loreto, Madre de Dios, Pasco, Ucayali)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Data Deficient (2011)
Hydraspis raniceps Gray 1856b:55, Platemys raniceps,
Phrynops raniceps, Batrachemys raniceps, Batrachemys
raniceps raniceps, Mesoclemmys raniceps
Type locality: “Brazils; Para” [Pará, Brazil].
Hydraspis maculata Gray 1873c:305 (88)
Type locality: “Tropical America.” Restricted to “S. America” by
Gray (1873j:65); and to “Venezuela” by Boulenger (1889:219) and
Pritchard and Trebbau (1984:127) [in error, see Rivas et al. (2015)].
Phrynops wermuthi Mertens 1969b:132, Phrynops tuberculatus wermuthi, Phrynops (Batrachemys) nasutus wermuthi,
Phrynops nasutus wermuthi, Phrynops nasuta wermuthi,
Batrachemys raniceps wermuthi
Type locality: “Peru...zweifellos der amazonische Teil des Landes.” Restricted to “Iquitos (3°50' S 73°15' W), Loreto, Peru” by
Bour and Pauler (1987:8).
Brazil (Ceará, Goiás, Piauí)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
Mesoclemmys perplexa Bour and Zaher 2005:298
Mesoclemmys tuberculata (Luederwaldt 1926)
Tuberculate Toad-headed Turtle
Type locality: “the regions called “Olho d’Água da Santa” and
“Baixão do Fausto”, southern part of the Parque Nacional da Serra
das Confusões, State of Piauí, Brasil” [Brazil].
Mesoclemmys raniceps (Gray 1856b) (12:41) (89)
Amazon Toad-headed Turtle
Daniel O. Santana / CBFTT / Areia Branca, Sergipe, Brazil
Richard C. Vogt / Iquitos, Peru [captivity]
Brazil (Alagoas, Bahia, Ceará, Maranhão, Minas Gerais, Paraíba, Pernambuco, Piauí, Rio Grande do Norte, Sergipe)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
CBFTT Account: Santana, Marques, Vieira, Moura, Faria, and
Mesquita (2016)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Data Deficient (2011)
Rhinemys tuberculata Luederwaldt 1926:428, Batrachemys
tuberculata, Phrynops (Batrachemys) tuberculata, Phrynops tuberculata, Phrynops (Batrachemys) tuberculatus,
Phrynops tuberculatus, Phrynops tuberculatus tuberculatus, Mesoclemmys tuberculata
Mesoclemmys zuliae (Pritchard and Trebbau 1984) (12:40)
Zulia Toad-headed Turtle
Type locality: “Brasil: Estado da Bahia e Pará” [Brazil]. Restricted
to “Villa Nova, Bahia” by Mertens and Wermuth (1955:400); and
to “Fortaleza, Ceará” [Brazil] by lectotype designation by Bour
and Pauler (1987:9).
Mesoclemmys vanderhaegei (Bour 1973)
Vanderhaege’s Toad-headed Turtle
Peter C.H. Pritchard / nr. Maracaibo, Zulia, Venezuela
Elizângela S. Brito / CBFTT / Cáceres, Mato Grosso, Brazil
Colombia (?) (Norte de Santander), Venezuela (Zulia)
IUCN Red List: Vulnerable B1+2c (1996)
TFTSG Draft Red List: Vulnerable (2011)
Phrynops (Batrachemys) zuliae Pritchard and Trebbau
1984:135, Phrynops zuliae, Batrachemys zuliae, Mesoclemmys zuliae
Type locality: “Caño Madre Vieja near El Guayabo, Distrito
Colón, Edo. Zulia, Venezuela (8°53' N, 72°30' W).”
Phrynops Wagler 1830b (07:100)
Phrynops Wagler 1830b:135
Argentina (Corrientes, Formosa, Misiones), Bolivia, Brazil
(Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais,
Paraná, São Paulo, Tocantins), Paraguay
CBFTT Account: Marques, Böhm, Brito, Cabrera, and Verdade (2014)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Data Deficient or Near Threatened
(2012)
Phrynops schöpfii Fitzinger in Diesing 1839:237 (nomen
nudum)
Phrynops schoepffii Fitzinger in Siebenrock 1904b:22 (nomen
nudum, partim)
Phrynops paraguayensis Vanzolini in Donoso-Barros
1965:13 (nomen nudum)
Phrynops tuberculatus vanderhaegei Bour 1973:175, Phrynops (Batrachemys) vanderhaegei, Phrynops vanderhaegei, Batrachemys vanderhaegei, Bufocephala vanderhaegei, Mesoclemmys vanderhaegei
Type locality: “environs d’Asunción au Paraguay.” Restricted to
“Tobati (25º15' S, 57º04' W), La Cordillera, Paraguay” by Bour
and Pauler (1987:10).
Type species: Phrynops geoffroanus [= Emys geoffroana Schweigger
1812], by original monotypy.
Spatulemys Gray 1872b:463
Type species: Spatulemys lasalae [= subjective synonym of Platemys hilarii Duméril and Bibron 1835], by original monotypy.
Parahydraspis Wieland 1923:2
Type species: Parahydraspis paranaensis † Wieland 1923, by original monotypy.
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Phrynops geoffroanus (Schweigger 1812) (10:44, 14:40) (90)
Geoffroy’s Side-necked Turtle
Platemys waglerii Duméril and Bibron 1835:422, Hydraspis
waglerii, Phrynops waglerii
Type locality: “Brésil” [Brazil].
Platemys neuwiedii Duméril and Bibron 1835:425 (nomen
novum)
Hydraspis boulengeri Bohls 1895:53
Type locality: “nördlichen Theile Paraguays...im Aquidaban,
Tagatiya und anderen linken Nebenflüssen des Paraguaystromes”
[Paraguay]. Restricted to “Departmento Concepción: Río Saladillo, 23ºS” [Paraguay] by Cacciali et al. (2016:45).
Hydraspis lutzi Ihering in Luederwaldt 1926:441, Phrynops
lutzi
Type locality: “Mogy-guassù (Est. de S. Paulo)” [Brazil].
Vinícius T. de Carvalho / Piracicaba, São Paulo, Brazil
Phrynops hilarii (Duméril and Bibron 1835)
Saint-Hilaire’s Side-necked Turtle
Russell A. Mittermeier / Rio Pelotas, Rio Grande do Sul, Brazil
(orange dot = probable trade)
Argentina (Corrientes, Misiones), Bolivia (Beni), Brazil (Acre,
Alagoas, Amazonas, Bahia, Ceará, Espírito Santo, Goiás,
Maranhão, Mato Grosso, Mato Grosso do Sul, Minas
Gerais, Pará, Paraíba, Paraná, Pernambuco, Piauí, Rio
de Janeiro, Rio Grande do Norte, Rio Grande do Sul,
Rondônia, Santa Catarina, São Paulo, Sergipe, Tocantins),
Colombia (Amazonas, Caquetá, Casanare, Guainía, Meta,
Putumayo, Vaupés, Vichada), Ecuador, Paraguay, Peru
(Cusco, Huanuco, Junin, Loreto, Madre de Dios, Pasco),
Venezuela (Amazonas)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Emys geoffroana Schweigger 1812:302, Chelodina geoffroana, Phrynops geoffroanus, Platemys geoffroana,
Hydraspis (Phrynops) geoffroana, Hydraspis geoffroana,
Phrynops geoffroana, Rhinemys geoffroana, Phrynops
geoffroana geoffroana, Phrynops geoffroanus geoffroanus
Type locality: “Brasilia” [Brazil].
Emys depressa Merrem 1820:22 (senior homonym, not =
Emys depressa Spix 1824), Chelys (Hydraspis) depressa,
Chelys depressa, Hydraspis depressa, Platemys depressa
Type locality: “Brasilia” [Brazil].
Emys viridis Spix 1824:3, Chelys (Hydraspis) viridis, Chelys
viridis, Hydraspis viridis
Type locality: “fluminis Carinhanhae, confluentis Sti Francisci”
[Rio Carinhanha, tributary of Rio São Francisco, Brazil].
Emys tritentaculata Saint-Hilaire in Cuvier 1829:11
(nomen
nudum et dubium)
Emys geoffroyana Gray 1830e:16 (nomen novum), Phrynops
geoffroyana, Hydraspis geoffroyana, Platemys geoffroyana
Platemys geoffreana Duméril and Bibron 1835:418 (nomen
novum)
(14:40)
(orange dots = probably introduced)
Argentina (Buenos Aires, Chaco, Córdoba, Corrientes, Entre
Rios, Formosa, La Rioja (?), Mendoza (?), Misiones, Santa
Fe, Santiago del Estero), Brazil (Rio Grande do Sul, Santa
Catarina), Paraguay, Uruguay
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Platemys hilarii Duméril and Bibron 1835:428, Hydraspis
hilarii, Hydraspis geoffroyana hilarii, Phrynops hilarii,
Phrynops geoffroana hilarii, Phrynops geoffroanus hilarii
Type locality: “Brésil” [Brazil].
Hydraspis hilairii Gray 1844:40 (nomen novum), Platemys
hilairii, Phrynops (Phrynops) geoffroanus hilairii, Phrynops geoffroanus hilairii
Spatulemys lasalae Gray 1872b:463
Type locality: “Rio Parana, Corrientes” [Argentina].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Phrynops tuberosus (Peters 1870) (10:44) (90)
Guianan Shield Side-necked Turtle
Argentina (Corrientes, Entre Rios, Misiones), Brazil (Paraná,
Rio Grande do Sul, Santa Catarina), Paraguay, Uruguay
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Vulnerable (2011)
Phrynops williamsi Rhodin and Mittermeier 1983:58
Type locality: “Rio Cadea, Rio Grande do Sul, Brazil.”
Platemys Wagler 1830b (14:39)
Platemys Wagler 1830b:135
Type species: Platemys planiceps [= Testudo planiceps Schneider
1792] [= objective synonym of Testudo platycephala Schneider
1792], by original monotypy.
Peter C.H. Pritchard / Canaima National Park, Rio Caroni, Bolivar, Venezuela
Brazil (Roraima), Guyana, Venezuela (Bolívar)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Least Concern (2011)
Platemys tuberosa Peters 1870:311, Hydraspis tuberosa,
Phrynops tuberosa, Phrynops geoffroana tuberosa,
Phrynops geoffroanus tuberosus, Phrynops (Phrynops)
tuberosus, Phrynops tuberosus
Type locality: “Cotingaflusse am Roraimagebirge in British-Guyana” [Guyana]. Emended to “Amazon drainage, the Río Cotingo...
Brazil...state of Roraima” by Bour (2008:38).
Phrynops williamsi Rhodin and Mittermeier 1983
Williams’ Side-necked Turtle
Platemys platycephala (Schneider 1792) (14:41) (91)
Twist-necked Turtle
(subspecies: platycephala = red, melanonota = purple)
Bolivia (Beni, Cochabamba), Brazil (Acre, Amapá [?], Amazonas, Mato Grosso, Pará, Roraima), Colombia (Amazonas,
Caquetá, Guainía, Guaviare, Meta, Putumayo, Vaupés,
Vichada), Ecuador, French Guiana, Guyana, Peru (Amazonas, Huánuco, Loreto, Madre de Dios, Puno, Ucayali),
Suriname, Venezuela (Amazonas, Bolívar, Delta Amacuro,
Monagas)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Platemys platycephala platycephala (Schneider 1792)
Eastern Twist-necked Turtle, Common Twist-necked Turtle
Vinícius T. de Carvalho / Picada Café, rio Cadeia, Rio Grande do Sul, Brazil
Russell A. Mittermeier / nr. Leticia, Colombia
Bolivia (Beni, Cochabamba), Brazil (Acre, Amapá?, Amazonas, Mato Grosso, Pará, Roraima), Colombia (Amazonas,
Caquetá, Putumayo, Vaupés), French Guiana, Guyana,
Peru (Huánuco, Loreto, Madre de Dios, Puno, Ucayali),
Suriname, Venezuela (Amazonas, Bolívar, Delta Amacuro,
Monagas)
Testudo platycephala Schneider 1792:261, Platemys platycephala, Platemys platycephala platycephala
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Type locality: “Ost-indien.” Restricted to “Cayenne, French
Guiana” by Ernst (1984:350).
Testudo planiceps Schneider 1792:pl.7 (nomen novum and
senior homonym, not = Testudo planiceps Gray 1854b),
Emys planiceps, Hydraspis planiceps, Clemmys planiceps,
Platemys planiceps, Chelys (Hydraspis) planiceps, Chelys
planiceps
Testudo martinella Daudin 1802:377, Platemys martinella
Type locality: “Cayenne et dans l’intérieur de la Guiane” [French
Guiana].
Emys discolor Thunberg in Schweigger 1812:302, Hydraspis
(Phrynops) discolor, Hydraspis discolor
Type locality: Not known. Restricted to “Cayenne, French Guiana” by Ernst (1984:350).
Emys canaliculata Spix 1824:10, Hydraspis canaliculata,
Platemys canaliculata
Type locality: “fluminis Solimoens” [Rio Solimões, Amazonas,
Brazil].
Emys carunculata Saint-Hilaire in Cuvier 1829:11 (14:41) (nomen
nudum et dubium)
Emys constricta Cuvier in Gray 1830e:17 (nomen nudum)
Hydraspis pachyura Boie in Gray 1830e:17 (nomen nudum),
Emys pachyura
Hydraspis constricta Gray 1831d:43
Type locality: Not known. Restricted to “Belem, Brazil” by Ernst
(1984:350).
Platemys platycephala melanonota Ernst 1984
Black-backed Twist-necked Turtle
Ecuador, Peru (Amazonas, Loreto)
Platemys platycephala melanonota Ernst 1984:352
(91)
Type locality: “vicinity of Galilea, on the Río Santiago, Amazonas, Perú (4°1' S, 77°47' W).”
Rhinemys Wagler 1830b (07:100)
Rhinemys Wagler 1830b:134
Type species: Rhinemys rufipes [= Emys rufipes Spix 1824], by subsequent designation by Fitzinger (1843:29).
Rhinemys rufipes (Spix 1824)
Red Side-necked Turtle, Red-footed Sideneck Turtle
(orange dots = trade and uncertain)
Brazil (Amazonas, Pará), Colombia (Amazonas, Guainía, Vaupés), Venezuela (?) (Amazonas)
CBFTT Account: Magnusson and Vogt (2014)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Least Concern (2011)
Emys rufipes Spix 1824:7, Hydraspis rufipes, Rhinemys rufipes, Chelys (Hydraspis) rufipes, Chelys rufipes, Platemys
rufipes, Phrynops rufipes
Type locality: “fluminis Solimoëns” [Rio Solimões, Amazonas,
Brazil].
hydRoMedusinae Baur 1893a (12:39)
Hydromedusidae Baur 1893a:211
Hydromedusinae Georges, Birrell, Saint, McCord, and Donnellan 1998:235
Hydromedusa Wagler 1830b
Hydromedusa Wagler 1830b:135
Type species: Hydromedusa maximiliani [= Emys maximiliani Mikan
1825], by original monotypy.
Hydromedusa (Chelomedusa) Gray 1873c:303
Type species: Hydromedusa (Chelomedusa) depressa [= Hydromedusa depressa Gray 1856b] [= subjective synonym of Emys maximiliani Mikan 1825], by subsequent designation by Lindholm
(1929:289).
Hydromedusa maximiliani (Mikan 1825)
Brazilian Snake-necked Turtle, Maximilian’s Snake-necked
Turtle
William E. Magnusson / CBFTT / Presidente Figuero, Amazonas, Brazil
Franco L. Souza / CBFTT / Parque Estadual Carlos Botelho, São Paulo, Brazil
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Rios, Formosa, Misiones, Santa Fe, Santiago del Estero),
Brazil (Minas Gerais, Paraná, Rio de Janeiro, Rio Grande
do Sul, Santa Catarina, São Paulo), Paraguay, Uruguay
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Hydromedusa tectifera Cope 1870a:147
Type locality: “tributaries of the Parana or Uraguay rivers, either
in the Argentine Confederation or the Banda Oriental” [Argentina
or Uruguay].
Hydromedusa platanensis Gray 1873c:302, Hydromedusa
platensis
Type locality: “Rio de la Plata” [Argentina or Uruguay].
Brazil (Espírito Santo, Minas Gerais, Rio de Janeiro, São Paulo)
CBFTT Account: Souza and Martins (2009)
IUCN Red List: Vulnerable B1+2cd (1996)
TFTSG Draft Red List: Near Threatened (2011)
Emys maximiliani Mikan 1825:[unpaginated], Chelodina
maximiliani, Hydromedusa maximiliani, Hydraspis
maximiliani
Type locality: “Brasiliam...Capitania St. Paulo” [São Paulo,
Brazil].
Platemys antiqua † Ameghino 1882:41 (nomen nudum)
[Pleistocene or Holocene, Argentina (Buenos Aires)]
Platemys fossilis † Ameghino 1882:41 (nomen nudum)
[Pleistocene or Holocene, Argentina (Buenos Aires)]
Platemys laevis † Ameghino 1882:41 (nomen nudum)
[Pleistocene or Holocene, Argentina (Buenos Aires)]
Platemys robusta † Ameghino 1882:41 (nomen nudum)
[Pleistocene or Holocene, Argentina (Buenos Aires)]
Hydromedusa wagleri Günther 1884:423
Type locality: “Buenos Ayres” [Argentina].
Emys maximiliana Gray 1830e:17 (nomen novum), Hydromedusa maximiliana
Chelodina flavilabris Duméril and Bibron 1835:446, Hydromedusa flavilabris, Chelomedusa flavilabris
Chelodininae Baur 1893a (12:39)
Hydromedusa subdepressa Gray 1854a:134
Chelodina Fitzinger 1826 (07:85, 08:2, 10:34)
Type locality: “Brésil” [Brazil].
Type locality: “Brazils” [Brazil].
Hydromedusa depressa Gray 1856b:60 (nomen novum),
Chelomedusa depressa
Hydromedusa bankae Giebel 1866b:19
Type locality: “Insel Banka” [Bangka Island, Sumatra, Indonesia]
[in error].
Hydromedusa tectifera Cope 1870a
South American Snake-necked Turtle
Chelodinidae Baur 1893a:211
Chelodininae Georges, Birrell, Saint, McCord, and Donnellan
1998:235
Chelodina Fitzinger 1826:6
Type species: Chelodina longicollis [= Testudo longicollis Shaw
1794], by original designation.
Hydraspis Bell 1828b:512
Type species: Hydraspis longicollis [= Testudo longicollis Shaw
1794], by original designation.
Chelyodina Agassiz 1846:79 (nomen novum)
Hesperochelodina Wells and Wellington 1985:9 (nomen nudum)
Macrochelodina Wells and Wellington 1985:9
Type species: Macrochelodina oblonga [= Chelodina oblonga Gray
1841], by original designation.
Macrodiremys McCord and Joseph-Ouni 2007b:57
Type species: Macrodiremys oblonga [= Chelodina oblonga Gray
1841], by original designation.
Chelodina (Supremechelys) Hoser 2014b:8 (unavailable name
pending ICZN decision; Rhodin et al. 2015) (4, 92)
Chelodina (Chelodina) Fitzinger 1826 (10:34)
Chelodina Fitzinger 1826:6
Peter C.H. Pritchard / Brazil [captivity]
Type species: Chelodina longicollis [= Testudo longicollis Shaw
1794], by original designation.
Hydraspis Bell 1828b:512
Type species: Hydraspis longicollis [= Testudo longicollis Shaw
1794], by original designation.
Chelyodina Agassiz 1846:79 (nomen novum)
Hesperochelodina Wells and Wellington 1985:9 (nomen nudum)
Argentina (Buenos Aires, Chaco, Córdoba, Corrientes, Entre
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Chelodina (Chelodina) canni McCord and Thomson 2002 (07:86)
Cann’s Snake-necked Turtle
Chelodina (Chelodina) gunaleni
Type locality: “Uta River basin, Mimika District, Central Papua
Province (Irian Jaya), Indonesia.”
Chelodina (Chelodina) longicollis (Shaw 1794) (92)
Eastern Snake-necked Turtle, Common Snake-necked Turtle
John Roe / CBFTT / Booderee National Park, Jervis Bay, New South Wales, Australia
John Cann / Kalala Station, Roper R. drainage, Northern Territory, Australia
Australia (Northern Territory, Queensland)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Near Threatened (2011)
Chelodina rankini Wells and Wellington 1985:8 (nomen
nudum)
Chelodina canni McCord and Thomson 2002:256, Chelodina (Chelodina) canni, Chelodina novaeguineae canni
Type locality: “Malogie Waterhole, near Scarlet Hill on Kalala
Station (16º08' S, 133º36' E), Northern Territory, Australia.”
Chelodina rankini Wells 2007a:2 (07:86, 10:43) (unavailable
name)
Type locality: “north-eastern Australia...Burdekin River,
Queensland.”
Chelodina (Chelodina) gunaleni McCord and Joseph-Ouni 2007a (10:35)
Gunalen’s Snake-necked Turtle
(orange dots = introduced)
Australia (New South Wales, Queensland, South Australia,
Victoria)
Introduced: Australia (Tasmania)
CBFTT Account: Kennett, Roe, Hodges, and Georges (2009)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Testudo longicollis Shaw 1794:19, Emys longicollis, Chelodina longicollis, Hydraspis longicollis, Chelys (Chelodina)
longicollis, Chelys longicollis, Chelodina longicollis
longicollis, Chelodina (Chelodina) longicollis
Type locality: “The vast island or rather continent of Australia,
Australasia, or New Holland.”
Chelodina novaehollandiae Duméril and Bibron 1835:443
(nomen novum)
Chelodina sulcata Gray 1856a:201, Chelodina longicollis
sulcata
Type locality: “Australia.”
Chelodina sulcifera Gray 1856b:59 (nomen novum), Chelodina longicollis sulcifera
Indonesia (Papua)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
Chelodina gunaleni McCord and Joseph-Ouni 2007a:48,
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelodina (Chelodina) mccordi Rhodin 1994b (10:36)
Roti Snake-necked Turtle
(subspecies: mccordi [western Roti] and roteensis [eastern Roti] = red,
timorensis = purple)
Indonesia (Lesser Sundas [Roti]), Timor-Leste
CBFTT Account: Rhodin, Ibarrondo, and Kuchling (2008)
IUCN Red List: Critically Endangered A1d, B1+2e (2000);
Previously: Vulnerable (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II
Chelodina (Chelodina) mccordi mccordi Rhodin 1994b (10:36)
Western Roti Snake-necked Turtle
Chelodina (Chelodina) mccordi timorensis McCord, Joseph-Ouni,
and Hagen 2007a (07:89, 10:36)
Timor Snake-necked Turtle
Bonggi R. Ibarrondo / CCB / Lospalos, Lautém Dist., Timor-Leste
Timor-Leste
Chelodina timorensis McCord, Joseph-Ouni, and Hagen
2007a:54, Chelodina mccordi timorensis, Chelodina (Chelodina) mccordi timorensis
Type locality: “Lake Ira Lalaro, Lautem District (regency),
Tutuala Subdistrict, eastern East Timor (= Timor-Leste; = Timor
Lorosa’E).”
Chelodina mccordi timorlestensis Kuchling, Rhodin, Ibarrondo, and Trainor 2007:213
Type locality: “plain of Lake Iralalaro (= Lagoa Ira Lalaro) (ca.
08º28' S; 127º07' E, elev. ca. 334 m), east of Lospalos, Lautém
District, Timor-Leste.”
Chelodina (Chelodina) novaeguineae Boulenger 1888b
New Guinea Snake-necked Turtle
Anders G.J. Rhodin / TCF / CBFTT / nr. Busalangga, Roti, Indonesia
Indonesia (Lesser Sundas [Roti])
Chelodina rottiensis Brongersma in Rhodin 1994b:3 (nomen
nudum)
Chelodina mccordi Rhodin 1994b:4, Chelodina mccordi
mccordi, Chelodina (Chelodina) mccordi, Chelodina
(Chelodina) mccordi mccordi
Type locality: “Danau Naloek, near Busalangga, ca. 11 km northeast of Tudameda and ca. 8 km southwest of Ba’a, elevation ca.
115 m, southwestem Roti Island (10°48' S, 123°00' E), East Nusa
Tenggara Province, Indonesia.”
Fred Parker / Abam, Western Prov., Papua New Guinea
Chelodina (Chelodina) mccordi roteensis McCord, Joseph-Ouni,
and Hagen 2007b (07:87, 10:36)
Eastern Roti Snake-necked Turtle
Indonesia (Lesser Sundas [Roti])
Chelodina mccordi roteensis McCord, Joseph-Ouni, and Hagen 2007b:59, Chelodina (Chelodina) mccordi roteensis
Type locality: “Lake Enduy, eastern Rote Island, East Nusa Tenggara Province, Indonesia.”
Indonesia (Papua), Papua New Guinea (Southern)
IUCN Red List: Least Concern (2000); Previously: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Chelodina novaeguineae Boulenger 1888b:450, Chelodina
novaeguineae novaeguineae, Chelodina (Chelodina)
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novaeguineae
Type locality: “Katow, S. E. New Guinea” [Papua New Guinea].
Emended to “Mawatta, Binaturi River, Western Province, Papua
New Guinea” by Rhodin (1994a:9).
Chelodina (Chelodina) pritchardi Rhodin 1994a
Pritchard’s Snake-necked Turtle
Anders G.J. Rhodin / CRM 2 / Bore, Kemp Welch R., Central Prov., Papua New Guinea
Indonesia (Papua), Papua New Guinea (Southern)
IUCN Red List: Near Threatened (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Data Deficient (2011)
Chelodina reimanni Philippen and Grossman 1990:95,
Chelodina novaeguineae reimanni, Chelodina (Chelodina)
reimanni
Type locality: “Merauke-River, West-Irian, Neuguinea” [Papua,
Indonesia].
Chelodina (Chelodina) steindachneri Siebenrock 1914 (07:85)
Steindachner’s Snake-necked Turtle
Papua New Guinea (Southern)
IUCN Red List: Endangered B1+2e (2000); Previously: Vulnerable (1996)
TFTSG Draft Red List: Endangered (2011)
Chelodina pritchardi Rhodin 1994a:4, Chelodina (Chelodina) pritchardi
Gerald Kuchling / Borodale R., Western Australia, Australia
Type locality: “Bore, Kemp Welch River, 13 km southeast of
Kwikila, Central Province, Papua New Guinea (9°53' S, 147°46' E).”
Chelodina (Chelodina) reimanni Philippen and Grossmann 1990
Reimann’s Snake-necked Turtle
Australia (Western Australia)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Data Deficient (2011)
Chelodina steindachneri Siebenrock 1914:386, Hesperochelodina steindachneri, Chelodina (Chelodina) steindachneri
Anders G.J. Rhodin / nr. Merauke, Papua, Indonesia [captivity]
Type locality: “Marloo Station am Grey River in Westaustralien”
[Australia].
Chelodina millymillyensis Glauert 1923:54
Type locality: “Milly Milly, Murchison River, W.A.” [Australia].
Emended to “Milly Milly Creek, Milly Milly Station, Murchison
River, W.A.” [Australia] by lectotype designation by Cogger et al.
(1983:62).
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelodina (Macrochelodina) Wells and Wellington 1985
(07:85, 08:2, 10:34)
Macrochelodina Wells and Wellington 1985:9
Type species: Macrochelodina oblonga [= Chelodina oblonga Gray
1841], by original designation.
Chelodina (Supremechelys) Hoser 2014b:8 (unavailable name
pending ICZN decision; Rhodin et al. 2015) (4, 92)
Chelodina (Macrochelodina) burrungandjii Thomson, Kennett, and
Georges 2000
Arnhem Snake-necked Turtle, Sandstone Snake-necked Turtle
Australia (New South Wales, Queensland, South Australia,
Victoria)
CBFTT Account: Bower and Hodges (2014)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Near Threatened (2011)
Chelodina expansa Gray 1857:370, Chelodina oblonga
expansa, Macrochelodina expansa, Chelodina (Macrochelodina) expansa, Chelodina (Supremechelys) expansa
Type locality: “Australia.” Restricted to “nördliches Australien”
[Australia] by Wermuth and Mertens (1977:122).
Rod Kennett / Koolpin Gorge, Kakadu National Park, Northern Territory, Australia
Chelodina (Supremechelys) expansa brisbaneensis Hoser
2014b:9 (unavailable name pending ICZN decision; Rhodin et al. 2015) (4, 93)
Chelodina (Supremechelys) duboisi Hoser 2014b:9 (unavailable name pending ICZN decision; Rhodin et al. 2015) (4, 93)
Chelodina (Macrochelodina) kuchlingi Cann 1997d (07:90, 10:37, 14:42) (94)
Kuchling’s Snake-necked Turtle
Australia (Northern Territory)
CBFTT Account: Thomson, Kennett, Tucker, FitzSimmons,
Featherston, Alacs, and Georges (2011)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Least Concern (2011)
Chelodina burrungandjii Thomson, Kennett, and Georges
2000:676, Macrochelodina burrungandjii, Chelodina
(Macrochelodina) burrungandjii
Gerald Kuchling / Parry Creek, Western Australia, Australia [Western Australia Museum]
Type locality: “Koolpin Gorge, South Alligator River (13º28' S,
132º38' E)... Arnhem Land Plateau, Northern Territory, Australia.”
Chelodina (Macrochelodina) expansa Gray 1857 (93)
Broad-shelled Snake-necked Turtle
Claire Treilibs / CBFTT / Paringa, South Australia, Australia
Australia (Western Australia)
IUCN Red List: Not Evaluated
Chelodina kuchlingi Cann 1997d:41 (07:90, 10:37, 14:42), Macrochelodina kuchlingi, Chelodina (Macrochelodina) kuchlingi
Type locality: “Kalumburu, N. W. Australia, (14°18' S x 126°28' E).”
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Chelodina (Macrochelodina) oblonga Gray 1841(14:43) (94)
[formerly C. (M.) rugosa Ogilby 1890] (07:91, 10:37)
Northern Snake-necked Turtle
Arthur Georges / CBFTT / Morehead R., Western Prov., Papua New Guinea
Australia (Northern Territory, Queensland, Western Australia),
Indonesia (Papua), Papua New Guinea (Southern)
CBFTT Account: Kennett, Fordham, Alacs, Corey, and
Georges (2014)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Near Threatened (2011)
Chelodina oblonga Gray 1841:446 (14:43) (nomen conservandum, ICZN 2013a), Chelodina (Macrochelodina) oblonga
Type locality: “Western Australia.” Emended to “Northern
Territory...possibly...Port Essington” [Australia] by Thomson
(2000:747).
Chelodina rugosa Ogilby 1890:56 (14:43), Chelodina oblonga
rugosa, Macrochelodina rugosa, Macrochelodina rugosa
rugosa, Chelodina (Macrochelodina) rugosa
Chelodina (Macrochelodina) parkeri Rhodin and Mittermeier 1976
Parker’s Snake-necked Turtle
Fred Parker / Balimo, Western Prov., Papua New Guinea
Indonesia (Papua), Papua New Guinea (Southern)
IUCN Red List: Vulnerable D2 (2000); Previously: Vulnerable
(1996)
TFTSG Draft Red List: Near Threatened (2011)
Chelodina parkeri Rhodin and Mittermeier 1976:465, Macrochelodina parkeri, Chelodina (Macrochelodina) parkeri
Type locality: “Mawa, Lake Murray, Western District, Papua New
Guinea.”
Chelodina (Macrochelodina) walloyarrina McCord and JosephOuni 2007b (08:4, 10:38) (94)
Kimberley Snake-necked Turtle
Type locality: “Cape York, Q.” [Queensland, Australia].
Chelodina siebenrocki Werner 1901a:602 (07:91), Chelodina
oblonga siebenrocki, Macrochelodina siebenrocki, Macrochelodina rugosa siebenrocki
Type locality: “Deutsch Neu-Guinea” [northern Papua New Guinea] [in error]. Emended to “New Guinea’s south coast” [southern
Papua New Guinea] by Rhodin and Mittermeier (1976:474).
Chelodina intergularis Fry 1915:88
Type locality: “Australia.”
Macrochelodina billabong Wells and Wellington 1985:9 (nomen nudum), Chelodina billabong
John Cann / TCF / Kimberleys, Western Australia, Australia
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Chelodina (Macrodiremys) oblonga
Type locality: “Western Australia.” Emended to “Northern
Territory...possibly...Port Essington” [Australia] by Thomson
(2000:747).
Chelodina colliei Gray 1856a:200, Macrodiremys colliei,
Chelodina (Macrodiremys) colliei
Type locality: “Swan River” [Western Australia, Australia].
Elseya Gray 1867 (07:92, 14:45) (95)
Elseya Gray 1867:44
Type species: Elseya dentata [= Chelymys dentata Gray 1863a], by
subsequent designation by Lindholm (1929:291).
Pelocomastes De Vis 1897:6
Australia (Northern Territory, Western Australia)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
Macrochelodina walloyarrina McCord and Joseph-Ouni
2007b:59, Chelodina (Macrochelodina) walloyarrina
Type locality: “The Fitzroy River at Fitzroy River Crossing,
Western Australia, Australia, 18°10.834' S and 125°35.849' E.”
Chelodina (Macrodiremys) McCord and Joseph-Ouni 2007b
(10:34, 10:39, 14:44)
Macrodiremys McCord and Joseph-Ouni 2007b:57 (08:3, 10:39)
Type species: Macrodiremys oblonga sensu Burbidge et al. 1974
[= Chelodina oblonga Gray 1841 sensu Burbidge et al. 1974 =
subjective synonym of Chelodina colliei Gray 1856a], by original
designation.
Type species: Pelocomastes ampla † De Vis 1897 [= subjective synonym of Chelymys uberrima † De Vis 1897], by original monotypy.
Elseya (Hanwarachelys) Thomson, Amepou, Anamiato, and
Georges 2015:65
Type species: Elseya (Hanwarachelys) novaeguineae [= Platemys
novaeguineae Meyer 1874], by original designation.
Elseya (Elseya) Gray 1867 (95)
Elseya Gray 1867:44
Type species: Elseya dentata [= Chelymys dentata Gray 1863a], by
subsequent designation by Lindholm (1929:291).
Elseya (Elseya) branderhorsti (Ouwens 1914) (07:93)
White-bellied Snapping Turtle, Branderhorst’s Snapping Turtle
Chelodina (Macrodiremys) colliei Gray 1856a (07:88, 08:3, 10:39, 14:44
[formerly C. (M.) oblonga Gray 1841] (14:43)
Southwestern Snake-necked Turtle
Arthur Georges / Bensbach R., Western Prov., Papua New Guinea
Gerald Kuchling / Moore River Nature Reserve, Western Australia, Australia
Indonesia (Papua), Papua New Guinea (Southern)
IUCN Red List: Vulnerable B1+2e (2000)
TFTSG Draft Red List: Vulnerable (2011)
Emydura branderhorsti Ouwens 1914:31, Elseya branderhorsti, Elseya (Elseya) branderhorsti
Australia (Western Australia)
IUCN Red List: Near Threatened (1996)
TFTSG Draft Red List: Near Threatened (2011)
Chelodina oblonga Gray 1841:446 (14:43) (in error, misidentified type, ICZN 2013a), Macrodiremys oblonga,
Type locality: “Sth. Nw. Guinea” [Southern New Guinea = Papua,
Indonesia]. Restricted to “southeastern Papua, Indonesia, between
the Lorentz River and Merauke” by Thomson et al. (2015:65); and
to “Bensbach River of the Trans-Fly region of Papua New Guinea
(8°50'58.6896" S., 141°14'52.944" E.) by neotype designation by
Thomson et al. (2015:63).
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Elseya (Elseya) dentata (Gray 1863a)
Northern Snapping Turtle
Gerald Kuchling / Carson R., Western Australia, Australia
Australia (Northern Territory)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Not Evaluated
Elseya flaviventralis Georges, Doody, Young, and Cann
2000:7 (nomen nudum)
Elseya jukesi Wells 2002b:7 (nomen nudum)
Elseya jukesi Wells 2007b:5 (07:94, 10:43) (unavailable name)
Elseya (Elseya) flaviventralis Thomson and Georges 2016:20
Type locality: “Pine Creek Crossing, South Alligator River Drainage, Kakadu National Park, Northern Territory, Australia, 13°30' S
132°28' E”.
Elseya (Hanwarachelys) Thomson, Amepou, Anamiato,
and Georges 2015 (95)
Australia (Northern Territory, Western Australia)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Chelymys dentata Gray 1863a:98, Podocnemis dentata,
Elseya dentata, Emydura dentata, Elseya dentata dentata,
Elseya (Elseya) dentata
Elseya (Hanwarachelys) Thomson, Amepou, Anamiato, and
Georges 2015:65
Type species: Elseya (Hanwarachelys) novaeguineae [= Platemys
novaeguineae Meyer 1874], by original designation.
Elseya (Hanwarachelys) novaeguineae (Meyer 1874) (10:40, 14:45) (97)
Western New Guinea Stream Turtle, New Guinea Snapping Turtle
Type locality: “N. Australia; Upper Victoria, in Beagle’s Valley.”
[Northern Territory, Australia]. Emended to “Beagles Valley,
Victoria River, Northern Territory...(15°34' S, 130°54' E)” by
lectotype designation by Thomson et al. (2015:62).
Chelymys elseyi Gray 1864d:132 (nomen nudum)
Chelymys elseya Gray 1870c:76 (nomen nudum)
Elseya intermedia Gray 1872c:23
Type locality: “North Australia, upper part of Victoria” [Northern
Territory, Australia].
Elseya (Elseya) flaviventralis Thomson and Georges 2016 (96)
Yellow-bellied Snapping Turtle
Anders G.J. Rhodin / Bintuni, West Papua, Indonesia [captivity]
Arthur Georges / South Alligator R., Northern Territory, Australia
Indonesia (Misool, Papua, Waigeo)
Introduced: Palau (Babeldaob)
IUCN Red List: Least Concern (2000); Previously: Least
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2015)
Platemys novaeguineae Meyer 1874:128, Emydura novaeguineae, Elseya novaeguineae, Elseya latisternum
novaeguineae, Elseya dentata novaeguineae, Elseya
novaeguineae novaeguineae, Myuchelys novaeguineae,
Elseya (Hanwarachelys) novaeguineae
Elseya (Hanwarachelys) schultzei (Vogt 1911) (07:95, 10:40, 14:45)
Northern New Guinea Stream Turtle, Schultze’s Snapping Turtle
Type locality: “Neu-Guinea” [Papua, Indonesia]. Restricted to
“Passim (Nordwest Neu Guinea)” [Papua, Indonesia] by Meyer
(1887:4); to “southwestern shore of Cenderawasih Bay on the
southeastern Vogelkop” [Papua, Indonesia] by Rhodin and
Genorupa (2000:132); and to “Passim, Barbussi River, ca. 3 km N.
Sieb, 1 km S. Tandjung Sjeri (= Syeri), west shore Geelvink Bay
(= Cenderawasih Bay), Papua, Indonesia (1°41' S, 134°05' E)” by
Thomson et al. (2015:67).
Elseya (Hanwarachelys) rhodini Thomson, Amepou, Anamiato, and
Georges 2015 (97)
Southern New Guinea Stream Turtle; Rhodin’s Stream Turtle
John Cann / Josaker, Eilanden R., Papua, Indonesia
Anders G.J. Rhodin / nr. Aiome, Ramu R., Madang Prov., Papua New Guinea
Indonesia (Papua); Papua New Guinea (Northern)
Introduced: Solomon Islands (?) (Malaita)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Least Concern (2015)
Emydura schultzei Vogt 1911:410, Elseya schultzei, Elseya
novaeguineae schultzei, Elseya (Hanwarachelys) schultzei
Type locality: “Fluss westlich der Tamimündung...HolländischNeu-Guinea” [Papua, Indonesia]. Restricted to “near Sae village,
Seko coast, near Skosai, ca 5 km W. mouth of Tami River, Papua,
Indonesia (2°37' S, 140°54' E)” by Thomson et al. (2015:68).
Elseya (Pelocomastes) De Vis 1897 (95)
Pelocomastes De Vis 1897:6
Indonesia (Aru Islands, Papua); Papua New Guinea (Southern)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Least Concern (2015)
Elseya (Hanwarachelys) rhodini Thomson, Amepou, Anamiato, and Georges 2015:69, Elseya rhodini
Type species: Pelocomastes ampla † De Vis 1897 [= subjective synonym of Chelymys uberrima † De Vis 1897], by original monotypy.
Elseya (Pelocomastes) albagula Thomson, Georges, and Limpus 2006
White-throated Snapping Turtle, Southern Snapping Turtle
Type locality: “Rue Creek (tributary of Wau Creek), Gulf Province,
Papua New Guinea (07°11'67.3" S, 144°37'13.8" E)”; GPS coordinates emended here to 07°11.673' S, 144°37.138' E.
John Cann / CCB / Burnett R., Queensland, Australia
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Elseya (Pelocomastes) lavarackorum (White and Archer 1994)
Riversleigh Snapping Turtle, Gulf Snapping Turtle
Australia (Queensland)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Vulnerable (2011); Australia: Critically
Endangered (2014)
Elseya albagula Thomson, Georges, and Limpus 2006:75,
Elseya dentata albagula, Elseya (Pelocomastes) albagula
John Cann / CBFTT / Riversleigh Station, Gregory R., Queensland, Australia
Type locality: “Ned Churchwood Weir, Burnett River,
Queensland, Australia (25º03' S, 152º05' E).”
Elseya (Pelocomastes) irwini Cann 1997c
Irwin’s Snapping Turtle
Australia (Northern Territory, Queensland)
CBFTT Account: Freeman, Thomson, and Cann (2014)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
Emydura lavarackorum † White and Archer 1994:159
[Pleistocene, Australia (Queensland)], Elseya lavarackorum, Elseya dentata lavarackorum, Elseya (Pelocomastes)
lavarackorum
Anders Zimny / Broken R., Queensland, Australia
Type locality: “Gregory River, Riversleigh Station, northwestern
Queensland, approximately 200 km northwest of Mount Isa”
[Australia].
Elusor Cann and Legler 1994
Elusor Cann and Legler 1994:83
Type species: Elusor macrurus Cann and Legler 1994, by original
monotypy.
Elusor macrurus Cann and Legler 1994
Mary River Turtle
Australia (Queensland)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
Elseya stirlingi Wells and Wellington 1985:9 (nomen nudum),
Elseya sterlingi
Elseya irwini Cann 1997c:36, Elseya dentata irwini, Elseya
(Pelocomastes) irwini
Type locality: “Burdekin River, Queensland, 19°42' S, 147°18' E,
approximately 18 km upstream from Ayr” [Australia].
Elseya stirlingi Wells 2007b:4 (07:96, 10:43) (unavailable name)
Marilyn Connell / Mary R., Queensland, Australia / male
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
(subspecies: macquarii = red, emmotti = purple, krefftii = blue,
nigra = green; overlap = intergrades)
Australia (Queensland)
IUCN Red List: Endangered B1+2c (1996)
TFTSG Draft Red List: Endangered (2011)
Elusor macrurus Cann and Legler 1994:83
Type locality: “Mary River. 45.5 km S and 21.0 km W Maryborough, Queensland, Australia, elevation approximately 30 m
(25°58' S, 152°30' E).”
Australia (New South Wales, Queensland, South Australia,
Victoria)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Emydura macquarii macquarii (Gray 1830e) (07:98, 10:7, 10:41, 10:42)
Macquarie River Turtle
Emydura Bonaparte 1836
Emydura Bonaparte 1836:7
Type species: Emydura macquaria [= Emys macquaria Cuvier
1829] [= Chelys (Hydraspis) macquarii Gray 1830e], by original
monotypy.
Chelymys Gray 1844:42
Type species: Chelymys macquaria [= Emys macquaria Cuvier
1829] [= Chelys (Hydraspis) macquarii Gray 1830e], by original
monotypy.
Euchelymys Gray 1871a:118
Type species: Euchelymys sulcifera [= subjective synonym of Chelys
(Hydraspis) macquarii Gray 1830e], by subsequent designation
by Lindholm (1929:290).
Tropicochelymys Wells and Wellington 1985:9
Type species: Tropicochelymys victoriae [= Hydraspis victoriae Gray
1842], by original designation.
Emydura macquarii (Gray 1830e) (10:7) (98)
Eastern Short-necked Turtle, Southern River Turtle
Arthur Georges / Brisbane R., Queensland, Australia
Australia (New South Wales, Queensland, South Australia,
Victoria)
Emys macquaria Cuvier 1829:11 (nomen nudum)
Chelys (Hydraspis) macquarii Gray 1830e:15 (10:7), Chelys
macquarii, Emys macquarii, Emydura macquarii, Chelymys macquarii, Emydura macquarii macquarii
Type locality: “New Holland” [Australia]. Restricted to “Novâ
Hollandiâ, Macquarrie River” [New South Wales, Australia] by
Gray (1831d:40), and to “Upper Macquarie River, in the vicinity
of Bathurst [NSW]” [Australia] by Cann (1998:101).
Hydraspis macquarrii Gray 1831d:40 (nomen novum), Emydura macquarrii, Emydura macquarrii macquarrii
Platemys macquaria Duméril and Bibron 1835:438 (nomen
novum), Hydraspis macquaria, Chelymys macquaria
Hydraspis australis Gray 1841:445 (10:41) (nomen dubium),
Chelymys australis, Emydura australis, Emydura australis australis
Type locality: “Western Australia?” Restricted to “Australia,
Macquarie River” by Gray (1872d:506).
Euchelymys sulcifera Gray 1871a:118
Type locality: “North Australia.”
Emydura macquariae Boulenger 1889:ix (nomen novum)
Emydura signata Ahl 1932:127 (10:42), Emydura macquarrii
signata, Chelymys signata, Emydura macquarii signata
Type locality: “Umgebung von Brisbane, Australien”
[Queensland, Australia].
Emydura canni Worrell 1970:pl.6 (nomen nudum)
Chelymys cooki Wells and Wellington 1985:8 (nomen
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nudum), Emydura cooki
Chelymys johncanni Wells and Wellington 1985:8 (nomen
nudum)
Emydura macquarii binjing Cann 1998:116 (10:42)
Type locality: “Clarence River and its tributaries in eastern New
South Wales...29°45'S, 152°15'E” [Australia].
Emydura macquarii dharra Cann 1998:120 (10:42)
Type locality: “Macleay River and its tributaries in eastern New
South Wales...30°54'S, 152°10'E” [Australia].
Emydura macquarii gunabarra Cann 1998:123
(10:42)
Type locality: “Hunter River and its tributaries in eastern New
South Wales...32°09'S, 150°58'E” [Australia].
Type locality: “Burnett’s River” [Queensland, Australia].
Chelymys victoriae marmorata Gray 1872d:506
Type locality: “east coast of Queensland, Burnett River”
[Australia].
Chelymys victoriae sulcata Gray 1872d:506
Type locality: “east coast of Queensland, Burnett River”
[Australia].
Emydura macquarii nigra McCord, Cann, and Joseph-Ouni 2003 (98)
Fraser Island Short-necked Turtle
Emydura macquarii dharuk Cann 1998:126 (10:42)
Type locality: “Norton’s Basin, Nepean River, 0.5 km upstream
from the junction of the Warragamba and Nepean Rivers at
33°52'S, 150°37'E...Sydney Basin in eastern New South Wales”
[Australia].
Emydura macquarii emmotti Cann, McCord, and Joseph-Ouni in
McCord, Cann, and Joseph-Ouni 2003 (98)
Cooper Creek Turtle
Arthur Georges / Fraser Island, Queensland, Australia
Australia (Queensland)
Tropicochelymys insularis Wells and Wellington 1985:9 (nomen nudum), Emydura insularis
Emydura macquarii nigra McCord, Cann, and Joseph-Ouni
2003:59, Emydura nigra
Type locality: “Lake McKenzie, Fraser Island, Queensland,
Australia (25°27' S, 153°04' E).”
Emydura subglobosa (Krefft 1876)
Red-bellied Short-necked Turtle
Kate Hodges / Stonehenge, Thomson R., Queensland, Australia / megacephalic female
Australia (Queensland, South Australia)
Chelymys windorah Wells and Wellington 1985:8 (nomen
nudum), Emydura windorah
Emydura macquarii emmotti Cann, McCord, and Joseph-Ouni in McCord, Cann, and Joseph-Ouni 2003:60, Emydura
emmotti
Type locality: “Waterloo Station, shearing-shed waterhole, southwestern Queensland, Australia (24°13' S, 143°17' E).”
Emydura macquarii krefftii (Gray 1871b) (98)
Krefft’s River Turtle
(subspecies: subglobosa = red, worrelli = purple;
orange dots = trade)
Australia (Northern Territory, Queensland), Indonesia (Papua),
Papua New Guinea (Southern)
Introduced: Papua New Guinea (New Britain, Northern)
IUCN Red List: Least Concern (2000); Previously: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Arthur Georges / Grays Waterhole, Burnett R., Queensland, Australia
Australia (Queensland)
Chelymys krefftii Gray 1871b:366, Emydura krefftii, Emydura
australis krefftii, Emydura macquarrii krefftii, Tropicochelymys krefftii, Emydura macquarii krefftii
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Emydura subglobosa subglobosa (Krefft 1876)
New Guinea Red-bellied Short-necked Turtle
Arthur Georges / Suki-Aramba Swamps, Fly R., Western Prov., Papua New Guinea
Australia (Queensland), Indonesia (Papua), Papua New
Guinea (Southern)
Introduced: Papua New Guinea (New Britain, Northern)
Euchelymys subglobosa Krefft 1876:390, Emydura subglobosa, Emydura australis subglobosa, Emydura macquarrii subglobosa, Chelymys subglobosa, Tropicochelymys
subglobosa, Emydura subglobosa subglobosa
Emydura tanybaraga Cann 1997b
Northern Yellow-faced Turtle
Jason Schaffer / nr. Coen, Archer River, Queensland, Australia
Type locality: “Amama River S. E. New Guinea” [Papua New
Guinea]. Restricted to “Naiabui S. E. New Guinea” [Papua New
Guinea] by Boulenger (1888b:450).
Emydura albertisii Boulenger 1888b:449, Emydura australis
albertisii
Type locality: “Katow, S. E. New Guinea” [= Mawatta, Binaturi
River (9°08' S, 142°55' E), Papua New Guinea].
Tropicochelymys goodei Wells and Wellington 1985:9 (nomen nudum), Emydura goodei
Emydura subglobosa worrelli (Wells and Wellington 1985) (07:99)
Worrell’s Short-necked Turtle, Diamond-head Turtle
Australia (Northern Territory, Queensland)
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2011)
Emydura tanybaraga Cann 1997b:24, Emydura subglobosa
tanybaraga
Type locality: “near Policeman Crossing, Daly River, Northern
Territory (13º46' S x 130°43' E)” [Australia].
Emydura victoriae (Gray 1842)
Northern Red-faced Turtle
Jason Schaffer / Dugald R., Flinders basin, Queensland, Australia
Australia (Northern Territory, Queensland)
Tropicochelymys leichhardti Wells and Wellington 1985:9
(nomen nudum)
Tropicochelymys worrelli Wells and Wellington 1985:9,
Emydura worrelli, Emydura subglobosa worrelli
Type locality: “Caranbirini Waterhole, ca. 21 km north of MacArthur River, Northern Territory (16º16' S x 136º05' E)” [Australia].
Gerald Kuchling / Barnett R., Western Australia, Australia
199
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TFTSG Draft Red List: Endangered (2011)
Phrynops bellii Gray 1844:41, Hydraspis bellii, Elseya bellii,
Wollumbinia bellii, Wollumbinia bellii bellii, Elseya latisternum bellii, Myuchelys bellii
Type locality: Not known. Restricted to “the upper reaches of
both the Namoi-MacDonald and Gwydir Rivers, above the New
England escarpment, in New South Wales” [Australia] by Cann
(1998:211).
Elseya dorriani Wells 2002b:16 (07:97) (nomen nudum), Wollumbinia bellii dorriani
Myuchelys georgesi (Cann 1997a)
Bellinger River Sawshelled Turtle
Australia (Northern Territory, Western Australia)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Hydraspis victoriae Gray 1842:55, Chelymys victoriae,
Emydura victoriae, Tropicochelymys victoriae
Type locality: “Victoria River, North-west coast of New Holland”
[Northern Territory, Australia].
Myuchelys Thomson and Georges 2009 (09:45, 10:43) (99)
Wollumbinia Wells 2007c:1 (07:97, 10:43) (unavailable name)
Myuchelys Thomson and Georges 2009:33 (09:45)
Type species: Myuchelys latisternum [= Elseya latisternum Gray
1867], by original designation.
John Cann / CBFTT / Bellinger River, New South Wales, Australia
Flaviemys Le, Reid, McCord, Naro-Maciel, Raxworthy, Amato,
and Georges 2013:257 (99)
Type species: Flaviemys purvisi [= Elseya purvisi Wells and Wellington 1985], by original designation.
Myuchelys bellii (Gray 1844) (07:97)
Bell’s Sawshelled Turtle, Western Sawshelled Turtle
Darren Fielder / CBFTT / Bald Rock Creek, Queensland, Australia
Australia (New South Wales)
CBFTT Account: Cann, Spencer, Welsh, and Georges (2015)
IUCN Red List: Data Deficient (1996)
TFTSG Draft Red List: Critically Endangered (2015)
Elseya georgesi Cann 1997a:18, Wollumbinia georgesi,
Elseya latisternum georgesi, Myuchelys georgesi
Type locality: “Bellinger River 30°25' S, 152º46' E” [New South
Wales, Australia].
Myuchelys latisternum (Gray 1867) (09:47)
Sawshelled Turtle, Common Sawshelled Turtle
Australia (New South Wales, Queensland)
CBFTT Account: Fielder, Chessman, and Georges (2015)
IUCN Red List: Endangered B1+2c (1996)
Alastair Freeman / CBFTT / Chillagoe Creek, Queensland, Australia
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type locality: “a river 15 km S., 32.3 km E. of Nowendoc, New
South Wales (31º39' S x 152º04' E. elevation 183 m)” [Australia].
Rheodytes Legler and Cann 1980
Rheodytes Legler and Cann 1980:2
Type species: Rheodytes leukops Legler and Cann 1980, by original
designation.
Rheodytes leukops Legler and Cann 1980
Fitzroy River Turtle
Australia (New South Wales, Northern Territory, Queensland)
CBFTT Account: Freeman and Cann (2014)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2011)
Elseya latisternum Gray 1867:44, Emydura latisternum,
Elseya latisternum latisternum, Wollumbinia latisternum,
Myuchelys latisternum
Type locality: “North Australia.”
Euchelymys spinosa Gray 1871a:118 (09:47)
Type locality: “North Australia.”
Stephen M. Zozaya / Connors R., upper Fitzroy R., Queensland, Australia
Elseya latisternon Gray 1871b:292 (nomen novum)
Wollumbinia dorsii Wells 2009:2 (09:46, 10:43) (unavailable name)
Myuchelys purvisi (Wells and Wellington 1985) (14:46) (99)
Manning River Sawshelled Turtle
Australia (Queensland)
IUCN Red List: Vulnerable A1c+2c, D2 (1996)
TFTSG Draft Red List: Vulnerable (2011)
Rheodytes leukops Legler and Cann 1980:2, Elseya leukops
Arthur Georges / Barnard R., New South Wales, Australia
Australia (New South Wales)
IUCN Red List: Data Deficient (1996)
TFTSG Draft Red List: Near Threatened (2011)
Elseya purvisi Wells and Wellington 1985:8, Wollumbinia
purvisi, Elseya latisternum purvisi, Myuchelys purvisi,
Flaviemys purvisi
Type locality: “Fitzroy River, 63 km N and 25 km E of Duaringa,
Queensland, Australia, elevation 40 m (23°09' S 149°55' E).”
201
202
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pseudeMyduRinae Gaffney 1977 (100)
Pseudemydurinae Gaffney 1977:24
Pseudemydura Siebenrock 1901 (100)
Pseudemydura Siebenrock 1901:248
Type species: Pseudemydura umbrina Siebenrock 1901, by original
monotypy.
Pseudemydura umbrina Siebenrock 1901 (100)
Western Swamp Turtle
peloMedusidae Cope 1868a (11:19)
Hydraspidina Bonaparte 1836:3 (partim)
Pelomedusidae Cope 1868a:119
Pelomedusa Wagler 1830b (11:19) (101)
Pelomedusa Wagler 1830b:136 (nomen conservandum, ICZN
1989)
Type species: Pelomedusa galeata [= Testudo galeata Schoepff
1792], by original monotypy.
Pentonyx Duméril and Bibron 1835:389
Type species: Pentonyx galeata [= Testudo galeata Schoepff 1792],
by original designation.
Pelomedusa subrufa (sensu lato) species complex (101)
African Helmeted Turtles, African Helmeted Terrapins
Gerald Kuchling / TCC / Ellen Brook Nature Reserve, nr. Perth, Western Australia, Australia
(yellow dots and red distributions = assigned Pelomedusa spp. – see below;
gray dots and gray distributions = unassigned Pelomedusa subrufa sensu lato;
orange dots = introduced or trade or questionable)
Australia (Western Australia)
IUCN Red List: Critically Endangered A1c, B1+2c, C1+2b, D
(1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix I
Pseudemydura umbrina Siebenrock 1901:249
Type locality: “Australien” [Australia].
Emydura inspectata Glauert 1954:125
Type locality: “Warbrook, about 24 miles north of Perth...Swan
River District” [Western Australia, Australia].
Angola, Benin, Botswana, Burkina Faso, Burundi, Cameroon,
Central African Republic, Chad, Congo (DRC), Congo
(ROC), Eritrea, Ethiopia, Gambia, Ghana, Guinea, GuineaBissau, Ivory Coast, Kenya, Lesotho, Liberia, Madagascar
(prehistoric introduction?), Malawi, Mali, Mauritania,
Mozambique, Namibia, Niger, Nigeria, Rwanda, Saudi
Arabia, Senegal, Sierra Leone, Somalia, South Africa,
South Sudan, Sudan, Swaziland, Tanzania, Togo, Uganda,
Yemen, Zambia, Zimbabwe
Pelomedusa barbata Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer, Schleicher,
Široký, and Fritz 2014 (101)
Arabian Helmeted Turtle
Johannes Els / Taif, Saudi Arabia
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203
ICZN 1989), Emys galeata, Pelomedusa galeata, Pentonyx galeata, Hydraspis (Pelomedusa) galeata, Hydraspis
galeata, Pelomedusa galeata galeata
Type locality: “India orientali.” Restricted to “near Cape Town”
[South Africa] by Hewitt (1935:326).
Pentonyx capensis Duméril and Bibron 1835:390
Type locality: “au cap de Bonne-Espérance, dans l’Ile de Madagascar.” Restricted to “Kap der Guten Hoffnung” [Cape of Good
Hope, South Africa] by Mertens (1937:139) [invalid designation];
and to “Cape of Good Hope” [South Africa] by lectotype designation by Fritz et al. (2014b:510).
Pelomedusa nigra Gray 1863b:99, Pelomedusa galeata
nigra, Pelomedusa subrufa nigra
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato;
orange dot = introduced or trade)
Saudi Arabia (Asir, Jizan), Yemen
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Endangered (2016)
Pelomedusa barbata Petzold, Vargas-Ramírez, Kehlmaier,
Vamberger, Branch, Du Preez, Hofmeyr, Meyer, Schleicher, Široký, and Fritz 2014:530
Type locality: “Zinjibar, Abyan, Yemen, N13°7.75 E45°22.81.”
Type locality: “Natal” [South Africa]; with lectotype designation
by Fritz et al. (2014b:514).
Pelomedusa galeata orangensis Hewitt 1935:332, Pelomedusa subrufa orangensis
Type locality: “presumably from the Kimberley neighbourhood”
[South Africa].
Pelomedusa galeata devilliersi Hewitt 1935:337
Type locality: “Besondermeid, Steinkopf, Namaqualand, C.P.”
[South Africa].
Pelomedusa gehafie (Rüppell 1835) (101)
Eritrean Helmeted Turtle
Pelomedusa galeata (Schoepff 1792) (101)
South African Helmeted Turtle
Tomáš Mazuch / Ghinda, Eritrea
William R. Branch / Port Elizabeth, South Africa
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato;
orange dot = questionable record)
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato)
Botswana (?), Lesotho, Mozambique, Namibia (?), South Africa,
Swaziland
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
Testudo scabra Retzius in Schoepff 1792:12 (nomen nudum
and junior homonym, not = Testudo scabra Linnaeus
1758)
Testudo galeata Schoepff 1792:12 (nomen conservandum,
Eritrea, Sudan
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
Pentonyx gehafie Rüppell 1835:2, Pelomedusa galeata
gehafie, Pelomedusa subrufa gehafie, Pelomedusa gehafie
Type locality: “östlichen Abhange der abyssinischen Küstengebirge” [eastern slope of Abyssinian coastal mountains] [Eritrea].
Restricted to “Massaua” [Masawa, Eritrea] by lectotype designation by Mertens (1937:140) [invalid designation] see Fritz et al.
2014b:511).
Pelomedusa gehafiae Gray 1844:38 (nomen novum)
Pelomedusa galeata disjuncta Vaillant and Grandidier
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1910:56
Type locality: Not designated. Restricted to “shore of Lake Abaya,
Sidamo, Ethiopia” by Loveridge (1941:480) [invalid designation];
and to “Abyssinia...the eastern slope of the coastal mountains
in present-day Eritrea,” by lectotype designation by Fritz et al.
(2014b:515).
Pelomedusa kobe Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer, Schleicher,
Široký, and Fritz 2014 (101)
Tanzanian Helmeted Turtle
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato)
Ethiopia, Kenya, Tanzania
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
Pelomedusa neumanni Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer,
Schleicher, Široký, and Fritz 2014:537
Type locality: “Kakamega, Kenya, N0°17.04 E34°44.52.”
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato)
Pelomedusa olivacea (Schweigger 1812) (101)
Sahelian Helmeted Turtle
Tanzania
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
Pelomedusa kobe Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer, Schleicher,
Široký, and Fritz 2014:535
Type locality: “Naberera, Manyara, Tanzania, S4°11.66
E36°55.74.”
Pelomedusa neumanni Petzold, Vargas-Ramírez, Kehlmaier,
Vamberger, Branch, Du Preez, Hofmeyr, Meyer,
Schleicher, Široký, and Fritz 2014 (101)
Neumann’s Helmeted Turtle
Tomáš Mazuch / nr. Turmi, Omo Region, Ethiopia
Laurent Chirio / Maradi, Niger
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato)
Benin, Burkina Faso, Gambia, Ghana, Guinea, Guinea-Bissau,
Mali, Mauritania, Niger, Nigeria, Senegal, Sierra Leone, Togo
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
Emys olivacea Schweigger 1812:307 (senior homonym, not
= Emys olivacea Gray 1856b), Hydraspis (Pelomedusa)
olivacea, Hydraspis olivacea, Pelomedusa subrufa olivacea, Pelomedusa olivacea
Type locality: “sabulosis Nigritiae” [Senegal].
Pelomedusa gasconi Rochebrune 1884:25
Type locality: “Dagana, Saidé, lac de N’Guer, marigot des
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Maringouins” [Senegal]. Restricted to “Dagana, Senegal” by Loveridge (1941:480) [invalid designation]; and to “Dakar” [Senegal]
by neotype designation by Fritz et al. (2014b:514).
Pelomedusa schweinfurthi Petzold, Vargas-Ramírez, Kehlmaier,
Vamberger, Branch, Du Preez, Hofmeyr, Meyer,
Schleicher, Široký, and Fritz 2014 (101)
Schweinfurth’s Helmeted Turtle
Ethiopia, Somalia
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
Pelomedusa somalica Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer,
Schleicher, Široký, and Fritz 2014:540
Type locality: “Borama district, Awdal, Somaliland/Somalia,
N9°55 E43°10, 4500 ft.”
Pelomedusa subrufa (sensu stricto) (Bonnaterre 1789) (10:45, 11:19) (101)
Helmeted Turtle, African Helmeted Terrapin
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato)
Central African Republic, Congo (DRC), South Sudan
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
Pelomedusa schweinfurthi Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer,
Schleicher, Široký, and Fritz 2014:539
Anders G.J. Rhodin / CBFTT / Tsihombe, Cap Sainte Marie, Madagascar
Type locality: “Liria, Central Equatoria, South Sudan, N4°38.66
E32°4.83.”
Pelomedusa somalica Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer, Schleicher,
Široký, and Fritz 2014 (101)
Somalian Helmeted Turtle
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato)
Angola, Botswana, Congo (DRC), Kenya, Madagascar (prehistoric
introduction?), Malawi, Mozambique, Namibia, SouthAfrica,
Tanzania (?), Zambia, Zimbabwe
CBFTT Account: Boycott and Bourquin (2008) [Pelomedusa subrufa sensu lato]
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2013)
Testudo planitia Meuschen 1778:11 (nomen dubium and
senior homonym, not = Testudo planitia Gmelin 1789),
Hydraspis (Pelomedusa) planitia, Hydraspis planitia
Tomáš Mazuch / Daarbuduq, Somaliland
Type locality: Not designated.
Testudo subrufa Lacepède 1788:173, synopsis[table] (07:102, 09:6)
(nomen suppressum, ICZN 2005a)
Type locality: “l’Inde” [India] [in error]. Restricted to “Kap der
Guten Hoffnung” [Cape of Good Hope, South Africa] by Mertens
(1937:139); and to “Taolañaro (Fort-Dauphin), République Malagasy (Madagascar)” by Bour (1982c:535).
Testudo subrufa Bonnaterre 1789:28, Emys subrufa, Pelomedusa subrufa, Chelys (Hydraspis) subrufa, Chelys
subrufa, Hydraspis subrufa, Pelomedusa galeata subrufa,
Pelomedusa subrufa subrufa
Type locality: “l’Inde” [India] [in error]. Restricted to “Taolañaro, Madagascar” by Fritz et al. (2014b:507), following Bour
(1982c:535).
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato)
Testudo rubra Meyer 1790:83 (09:8) (nomen novum, dubium et
oblitum)
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Testudo badia Donndorff 1798:34 (nomen novum)
Testudo rubicunda Suckow 1798:49 (nomen novum)
Pentonix americana Cornalia 1849:13 (105) (nomen dubium),
Pentonyx americana
Type locality: “Flum. prope Novaeboracum” [= Novum Eboracum = New York, USA] [in error].
Pelomedusa mossambicensis Peters in Lichtenstein 1856:2
(nomen nudum)
Pelomedusa mozambica Peters in Gray 1856b:53 (nomen
nudum)
Pelomedusa galeata damarensis Hewitt 1935:338, Pelomedusa subrufa damarensis
Type locality: “Quickborn, near Okahandja, South West Africa”
[Namibia]; with lectotype designation by Fritz et al. (2014b:515).
Pelomedusa subrufa wettsteini Mertens 1937:141
Type locality: “Majunga, West-Madagaskar” [Mahajanga,
Madagascar].
Pelomedusa variabilis Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer, Schleicher,
Široký, and Fritz 2014 (101)
West African Helmeted Turtle
Tomas Diagne / Afram Arm, Ghana
(gray dots = unassigned Pelomedusa spp. = P. subrufa sensu lato)
Burkina Faso (?), Ghana, Ivory Coast
CBFTT Account: Boycott and Bourquin (2008) [as part of
Pelomedusa subrufa sensu lato]
IUCN Red List: Not Evaluated
Pelomedusa variabilis Petzold, Vargas-Ramírez, Kehlmaier, Vamberger, Branch, Du Preez, Hofmeyr, Meyer,
Schleicher, Široký, and Fritz 2014:543
Type locality: “Gold Coast, Ghana.”
Pelusios Wagler 1830b (11:19) (102)
Sternothaerus Bell 1825a:305 (partim, nomen suppressum,
ICZN 1989)
Type species: Sternothaerus leachianus Bell 1825a [= subjective synonym of Emys castanea Schweigger 1812 = Pelusios castaneus],
by subsequent designation by Bell (1828c:515); not Sternothaerus
odoratus Bell [= Testudo odorata Latreille in Sonnini and Latreille
1801], by incorrect designation by Fitzinger (1843:290).
Pelusios Wagler 1830b:137 (nomen conservandum, ICZN
1989)
Type species: Pelusios subniger Wagler [= Testudo subnigra
Lacepède 1788 (nomen suppressum) = Testudo subnigra
Bonnaterre 1789], by subsequent designation by Fitzinger
(1843:29).
Sternothaerus (Tanoa) Gray 1863f:193
Type species: Sternothaerus (Tanoa) sinuatus (Smith 1838), by
subsequent designation by Lindholm (1929:288).
Sternothaerus (Notoa) Gray 1863f:195
Type species: Type species (by monotypy): Sternothaerus (Notoa)
subniger (Lacepède 1788) [= Sternothaerus (Notoa) subniger
(Bonnaterre 1789)], by original monotypy.
Sternothaerus (Anota) Gray 1863f:196 (junior homonym, not =
Anota Hallowell 1852 [= Sauria])
Type species: Sternothaerus (Anota) niger (Duméril and Bibron
1835), by original monotypy.
Pelusios adansonii (Schweigger 1812)
Adanson’s Mud Turtle
Roger Bour / CBFTT / Lac de Guiers, Senegal
(orange dot = probable trade)
Benin (?), Cameroon, Central African Republic, Chad, Ethiopia,
Mali, Mauritania, Niger, Nigeria, Senegal, South Sudan,
Sudan
CBFTT Account: Bour (2008)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2013)
Emys adansonii Schweigger 1812:308, Hydraspis adansonii,
Pelomedusa adansonii, Sternotherus adansonii, Sternothaerus adansonii, Pentonyx andansonii, Pentonyx adansonii, Pelusios adansonii, Pelusios adansonii adansonii
Type locality: “Nigritia” [Senegal]. Restricted to “cap Vert” [Senegal] by Duméril and Bibron (1835:395).
Chelys (Hydraspis) adamsonii Gray 1830e:15 (nomen
novum)
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207
Pelusios bechuanicus FitzSimons 1932
Okavango Mud Turtle
William R. Branch / Moremi National Park, Botswana
Ethiopia (?), Kenya
IUCN Red List: Vulnerable D2 (1996)
TFTSG Draft Red List: Endangered (2013)
Pelusios broadleyi Bour 1986:31
Type locality: “Loiengalani [= Loyengalanij] (2°43' N, 36°43' E),
Marsabit district, Kenya.”
Pelusios carinatus Laurent 1956 (102)
African Keeled Mud Turtle
Angola, Botswana, Namibia, Zambia, Zimbabwe
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2013)
Pelusios bechuanicus FitzSimons 1932:37, Pelusios castaneus bechuanicus, Pelusios bechuanicus bechuanicus
Type locality: “Thamalakane River at Maun, Ngamiland”
[Botswana].
Tomas Diagne / Congo (DRC) [captivity]
Pelusios broadleyi Bour 1986
Turkana Mud Turtle
(orange dot = uncertain identification)
Tomas Diagne / Lake Turkana, Kenya
Congo (DRC), Congo (ROC), Gabon
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2013)
Pelusios carinatus Laurent 1956:39
Type locality: “Eala, Equateur” [Democratic Republic of Congo
(DRC)].
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Pelusios castaneus (Schweigger 1812) (14:47) (102)
African Mud Turtle
Leone, Togo
Emys castanea Schweigger 1812:314, Pelusios castaneus,
Chelys (Sternotherus) castaneus, Chelys castaneus,
Sternotherus castaneus, Clemmys (Pelusios) castanea,
Clemmys castanea, Sternothaerus castaneus, Sternothaerus nigricans castaneus, Sternothaerus nigricans castanea,
Pelusios nigricans castaneus, Pelusios subniger castaneus,
Pelusios castaneus, Pelusios castaneus castaneus
Type locality: Not known. Restricted to “vicinity of Koutchatcha (7°20' N, 1°18' E)...close to the Amou River (ca. 30 km
East of Gléï), Ogou Prefecture, Plateaux Region, Togo” by
neotype designation by Bour (2008:37).
Sternothaerus leachianus Bell 1825a:306, Sternotherus
leachianus
Type locality: Not known.
(orange dot = introduced or trade, red dot = extirpated)
Angola, Benin, Burkina Faso, Cameroon, Central African
Republic, Chad, Congo (DRC), Congo (ROC), Equatorial
Guinea (?), Gabon, Ghana, Guinea, Guinea-Bissau, Ivory
Coast, Liberia, Mali (?), Niger, Nigeria, São Tomé and
Príncipe (probable historic introduction), Senegal, Seychelles (extirpated; possible historic introduction), Sierra
Leone, Togo
Introduced: Guadeloupe
CBFTT Account: Bour, Luiselli, Petrozzi, Segniagbeto, and
Chirio (2016)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2013)
Sternotherus derbianus Gray 1844:37, Sternothaerus derbianus, Sternothaerus (Tanoa) derbianus, Pelusios derbianus,
Pelusios castaneus derbianus
Type locality: “W. Africa, Sierra Leone ? Gambia.” Restricted to
“Gambia” by Loveridge (1941:491).
Pelusios castaneus seychellensis (Siebenrock 1906c) (07:103, 14:47) (103)
(Extinct, ca. 1950)
Seychelles Mud Turtle
Pelusios castaneus castaneus (Schweigger 1812) (14:47)
West African Mud Turtle, Swamp Terrapin
Roger Bour / CBFTT / Mahé, Seychelles [NMW, Vienna]
Gabriel H. Segniagbeto / CBFTT / Togoville, Togo
(orange dot = introduced or trade)
Angola, Benin, Burkina Faso, Cameroon, Central African
Republic, Chad, Congo (DRC), Congo (ROC), Equatorial
Guinea (?), Gabon, Ghana, Guinea, Guinea-Bissau, Ivory
Coast, Liberia, Mali (?), Niger, Nigeria, São Tomé and
Príncipe (probable historic introduction), Senegal, Sierra
Seychelles (Mahé [extinct]; possible historic introduction)
CBFTT Accounts: Bour and Gerlach 2008; Bour, Luiselli,
Petrozzi, Segniagbeto, and Chirio 2016
IUCN Red List: Extinct (2003)
Sternothaerus nigricans seychellensis Siebenrock 1906c:38,
Pelusios subniger seychellensis, Sternothaerus castaneus
seychellensis, Pelusios castaneus seychellensis, Pelusios
seychellensis
Type locality: “Seychellen” [Seychelles]. Restricted to “Seychellen, Insel Gloriosa?” [Seychelles] by Siebenrock (1909a:559), and
to “Insel Mahé” [Seychelles] by Siebenrock (1909b:362).
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209
Pelusios castanoides Hewitt 1931 (11:19, 12:42)
Yellow-bellied Mud Turtle
Seychelles (Cerf, Fregate, La Digue, Mahé, Praslin, Silhouette) (prehistoric introduction?)
CBFTT Account: Gerlach (2008a)
IUCN Red List: Critically Endangered A2c, B2ab (2003)
Pelusios castanoides intergularis Bour 1983:355
Kenya, Madagascar (prehistoric introduction?), Malawi, Mozambique, Seychelles (prehistoric introduction?), South
Africa, Tanzania
IUCN Red List: Least Concern (1996)
SARCA Draft: Least Concern (regional) (2010)
TFTSG Draft Red List: Least Concern (2013)
Type locality: “La Digue Island, Seychelles.”
Pelusios chapini Laurent 1965 (11:19) (102)
Central African Mud Turtle
Pelusios castanoides castanoides Hewitt 1931 (11:19, 12:42)
East African Yellow-bellied Mud Turtle
Jérôme Maran / Mandji, Gabon
Gerald Kuchling / Madagascar
Kenya, Madagascar (prehistoric introduction?), Malawi,
Mozambique, South Africa, Tanzania
Pelusios nigricans castanoides Hewitt 1931:463, Pelusios
subniger castanoides, Pelusios castaneus castanoides,
Pelusios castanoides, Pelusios castanoides castanoides
Type locality: “Richards Bay, Zululand” [South Africa]. Restricted
to “Lake St. Lucia estuary, KwaZulu” [South Africa] by Broadley
(1981:673).
Pelusios castaneus kapika Bour 1979:149, Pelusios castanoides kapika
Type locality: “Delta du Sambirano (Province de Diégo-Suarez),
nord de Madagascar.”
Pelusios castanoides intergularis Bour 1983 (11:19, 12:42)
Seychelles Yellow-bellied Mud Turtle
Justin Gerlach / CBFTT / Seychelles
Central African Republic, Congo (DRC), Congo (ROC), Gabon, South Sudan, Uganda
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2013)
Pelusios castaneus chapini Laurent 1965:21, Pelusios
chapini
Type locality: “Kasenyi, Lake Albert, Bunia Terr., Ituri, Congo”
[Democratic Republic of Congo (DRC)]
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IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2013)
Pentonyx gabonensis Duméril 1856:373, Pelomedusa gabonensis, Sternothaerus gabonensis, Pelusios gabonensis
Pelusios cupulatta Bour and Maran 2003
Ivory Coast Mud Turtle
Type locality: “Gabon.”
Pentonyx gaboonensis Gray 1863f:194 (nomen novum)
Pelomedusa gabonica Peters 1864:644 (nomen novum)
Sternothaerus steindachneri Siebenrock 1902a:6
Type locality: “Madagascar” [in error].
Pelusios marani Bour 2000
Gabon Mud Turtle
Jérôme Maran / San Pédro, Ivory Coast
Jérôme Maran / Fouganou, Gabon
Benin, Ghana, Guinea, Ivory Coast, Liberia, Nigeria, Sierra
Leone, Togo
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Least Concern (2013)
Pelusios cupulatta Bour and Maran 2003:28
Type locality: “environs de San Pédro, Côte d’Ivoire, précisément
entre San Pédro (10 km W) et Grand-Bérébi (20 km E) (4°50' N,
6°47' W).” [Ivory Coast].
Pelusios gabonensis (Duméril 1856) (102)
African Forest Turtle
Congo (ROC), Gabon
IUCN Red List: Not Evaluated
TFTSG Draft Red List: Data Deficient (2013)
Pelusios marani Bour 2000:3
Type locality: “Yombi (01º26' S, 10º37' E), province de N’Gounié,
Gabon; environ 30 km SSE de Fougamou, entre Lambaréné et
Mouïla.”
Pelusios nanus Laurent 1956 (102)
African Dwarf Mud Turtle
Jérôme Maran / Gabon
William R. Branch / northwest Zambia
(orange dots = uncertain, trade, or possibly misidentified)
Angola, Cameroon, Central African Republic, Congo (DRC),
Congo (ROC), Equatorial Guinea, Gabon
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Pelusios rhodesianus Hewitt 1927 (11:19) (102)
Variable Mud Turtle, Mashona Hinged Terrapin
Angola, Congo (DRC), Zambia
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Data Deficient (2013)
Pelusios nanus Laurent 1956:31, Pelusios adansonii nanus
Richard C. Boycott / CBFTT / nr. Lake St. Lucia, KwaZulu-Natal, South Africa
Type locality: “Dilolo, Haut Lualaba” [Democratic Republic of
Congo (DRC)].
Pelusios niger (Duméril and Bibron 1835)
West African Black Mud Turtle
Gerald Kuchling / Kribi, Cameroon
Angola, Botswana, Burundi, Congo (DRC), Congo (ROC), Malawi,
Mozambique, Namibia, Rwanda, South Africa, Tanzania,
Uganda, Zambia, Zimbabwe
CBFTT Account: Broadley and Boycott (2008)
IUCN Red List: Least Concern (1996)
SARCA Draft: Vulnerable (regional) (2010)
Pelusios nigricans rhodesianus Hewitt 1927:375, Pelusios
subniger rhodesianus, Pelusios rhodesianus, Pelusios
castaneus rhodesianus, Pelusios rhodesianus rhodesianus
Type locality: “Mpika district, N.E. Rhodesia” [Zambia].
Pelusios sinuatus (Smith 1838) (11:19)
Serrated Hinged Terrapin
Benin, Cameroon, Equatorial Guinea, Gabon, Nigeria, Togo
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Near Threatened (2013)
Sternotherus niger Duméril and Bibron 1835:397, Sternothaerus niger, Sternothaerus (Anota) niger, Pelusios niger
Type locality: “probablement...originaire de l’île de Madagascar”
[in error].
Sternothaerus oxyrhinus Boulenger 1897b:919
Type locality: “unknown...but probably...from some part of Tropical Africa.”
Sternothaerus heinrothi Kanberg 1924:195
Type locality: “Kamerun” [Cameroon].
Richard C. Boycott / CBFTT / Ndumu Game Reserve, KwaZulu-Natal, South Africa
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(prehistoric introduction?), South Africa, Tanzania, Zambia, Zimbabwe
Introduced: British Indian Ocean Territory (Chagos Archipelago), Glorioso Islands, Mauritius
IUCN Red List: Least Concern (1996)
SARCA Draft: Least Concern (regional) (2010)
TFTSG Draft Red List: Least Concern (2013)
Pelusios subniger subniger (Bonnaterre 1789) (12:43)
East African Black Mud Turtle
Botswana, Burundi, Congo (DRC), Ethiopia, Kenya, Malawi,
Mozambique, Rwanda, Somalia, South Africa, Swaziland,
Tanzania, Zambia, Zimbabwe
CBFTT Account: Broadley and Boycott (2009)
IUCN Red List: Least Concern [Not Listed] (1996)
SARCA Draft: Least Concern (regional) (2010)
TFTSG Draft Red List: Least Concern (2013)
Sternotherus sinuatus Smith 1838:Reptilia,pl.1, Sternothaerus (Tanoa) sinuatus, Sternothaerus sinuatus, Pelusios
sinuatus, Pelusios sinuatus sinuatus
Type locality: “rivers to the north of 25° south latitude” [South
Africa]. Restricted to “the Crocodile/Marico Confluence, N.
Transvaal” [South Africa] by Broadley (1981:675).
Sternotherus dentatus Peters 1848:494 (nomen nudum),
Sternothaerus dentatus
Sternothaerus bottegi Boulenger 1895a:9
Type locality: “Bardera...Giuba e dei suoi affluenti” [Italian
Somaliland] [Somalia].
Pelusios sinuatus zuluensis Hewitt 1927:371
Type locality: “near the Umsinene River, Zululand” [South
Africa].
Pelusios sinuatus leptus Hewitt 1933a:45
Type locality: “Isoka, North-East Rhodesia” [Zambia].
Sternothaerus rudolphi † Arambourg 1947:461 [Pleistocene,
Ethiopia (Lake Turkana)], Pelusios rudolphi
Type locality: “basin du Lac Rodolphe et de la basse vallée de
l’Omo” [Ethiopia]. Emended to “Shungura Formation...Omo
River Basin...Ethiopia” by Lapparent de Broin (2000b:59).
Jérôme Maran / Tamatave (Toamasina), Madagascar
Botswana, Burundi, Congo (DRC), Madagascar (prehistoric
introduction?), Malawi, Mozambique, South Africa, Tanzania, Zambia, Zimbabwe
Testudo subnigra Lacepède 1788:175, synopsis[table] (09:6)
(nomen conservandum, ICZN 1989; nomen suppressum,
ICZN 2005a)
Type locality: Not known. Restricted to “Tamatave, est de Madagascar” by Bour (1979:152).
Testudo subnigra Bonnaterre 1789:30, Emys subnigra, Pelusios subniger, Sternotherus subniger, Clemmys (Pelusios)
subnigra, Clemmys subnigra, Sternothaerus subniger,
Sternothaerus (Notoa) subniger, Pelusios subniger, Pelusios subniger subniger
Type locality: Not known.
Testudo nigricans Donndorff 1798:34, Terrapene nigricans,
Kinosternon nigricans, Sternotherus nigricans, Sternothaerus nigricans, Sternothaerus nigricans nigricans,
Pelusios nigricans, Pelusios nigricans nigricans
Type locality: Not known.
Pelusios subniger parietalis Bour 1983 (12:43)
Seychelles Black Mud Turtle
Pelusios subniger (Bonnaterre 1789) (12:43)
East African Black Mud Turtle
Justin Gerlach / CBFTT / Seychelles
(orange dots = introduced)
Botswana, Burundi, Congo (DRC), Madagascar (prehistoric introduction?), Malawi, Mozambique, Seychelles
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Pelusios williamsi Laurent 1965
Williams’ Mud Turtle
Seychelles (Cerf, Cousin [extirpated], Fregate, La Digue,
Mahé, Praslin, Silhouette, St. Anne [extirpated]) (prehistoric
introduction?)
CBFTT Account: Gerlach (2008b)
IUCN Red List: Critically Endangered A2c, B2ab(ii,iii) (2003)
Pelusios subniger parietalis Bour 1983:359
Type locality: “La Digue Island, Seychelles.”
Pelusios upembae Broadley 1981
Upemba Mud Turtle
(subspecies: williamsi = red, laurenti = purple, lutescens = orange;
orange dots = possible trade)
Congo (DRC), Kenya, Tanzania, Uganda
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Least Concern (2013)
Pelusios williamsi williamsi Laurent 1965
Lake Victoria Mud Turtle
Tomas Diagne / Congo (DRC) [captivity]
Jérôme Maran / Lake Victoria, Uganda [captivity]
Kenya, Tanzania, Uganda
Pelusios williamsi Laurent 1965:12, Pelusios williamsi williamsi, Pelusios castaneus williamsi
Type locality: “Kakamega, Kaimosi, Kenya.”
Pelusios williamsi laurenti Bour 1984a
Ukerewe Island Mud Turtle
Tanzania
Pelusios williamsi laurenti Bour 1984a:29
Type locality: “Ukerewe Island (Lake Victoria), Tanzania, altitude
1150 m.”
Congo (DRC)
IUCN Red List: Data Deficient (1996)
TFTSG Draft Red List: Data Deficient (2013)
Pelusios bechuanicus upembae Broadley 1981:667, Pelusios
upembae
Type locality: “Kanonga River, tributary of the right bank of the
Fungwe River (695 m), Upemba National Park, Shaba Province,
Zaire” [Democratic Republic of Congo (DRC)].
Pelusios williamsi lutescens Laurent 1965
Albert Nile Mud Turtle
Congo (DRC), Uganda
Pelusios williamsi lutescens Laurent 1965:16, Pelusios castaneus lutescens
Type locality: “Semliki River, 1 km below the Lake Edward”
[Democratic Republic of Congo (DRC)].
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podoCneMididae Cope 1868b (07:104, 09:48, 11:20)
Hydraspidina Bonaparte 1836:3 (partim)
Podocnemididae Cope 1868b:282
Peltocephalidae Gray 1870f:718
by original monotypy.
Peltocephalus dumerilianus (Schweigger 1812)
Big-headed Sideneck Turtle
Erymnochelys Baur 1888a
Dumerilia Grandidier 1867:232 (junior homonym, not =
Dumerilia Leach 1824 [= Coleoptera] or Dumerilia
Robineau-Desvoidy 1835 [= Diptera] or Dumerilia Bocage 1866 [= Sauria])
Type species: Dumerilia madagascariensis Grandidier 1867, by
original monotypy.
Erymnochelys Baur 1888a:421 (nomen novum)
Type species: Erymnochelys madagascariensis [= Dumerilia madagascariensis Grandidier 1867], by application of ICZN Article 67.8.
Erymnochelys madagascariensis (Grandidier 1867)
Madagascan Big-headed Turtle, Rere
Richard C. Vogt / Rio Negro drainage, Amazonas, Brazil
Anders G.J. Rhodin / TCC / CRM 6 / Madagascar [CCBC, Ampijoroa]
Madagascar
IUCN Red List: Critically Endangered A4d (2008); Previously:
Endangered (1996)
CITES: Appendix II
Dumerilia madagascariensis Grandidier 1867:232, Podocnemis madagascariensis, Erymnochelys madagascariensis,
Dumerilia madagascarensis, Erymnochelys madagascarensis, Podocnemis madagascarensis
Type locality: “Mouroundava Tsidsibonque flumina in occidentali
insulae Madagascar littore” [Morondava and Tsiribihina rivers on
the western coast of Madagascar Island]. Restricted to “Morondava River...Madagascar” by Bour (2006a:37).
Podocnemis madagascariensis bifilaris Boettger 1893:14
Type locality: “Mojanga, West-Madagaskar” [Madagascar].
Peltocephalus Duméril and Bibron 1835
Peltocephalus Duméril and Bibron 1835:377
Type species: Peltocephalus tracaxa [= Emys tracaxa Spix 1824]
[= subjective synonym of Emys dumeriliana Schweigger 1812],
(orange dot = probable trade)
Brazil (Amapá, Amazonas, Pará, Roraima), Colombia (Amazonas, Caquetá, Guainía, Guaviare, Meta, Putumayo,
Vaupés, Vichada), Ecuador, French Guiana, Peru (Loreto),
Venezuela (Amazonas, Apure)
IUCN Red List: Vulnerable A1acd (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II
Emys dumeriliana Schweigger 1812:300, Podocnemis
dumeriliana, Hydraspis dumeriliana, Peltocephalus
dumerilianus, Chelonemys dumeriliana, Peltocephalus
dumeriliana
Type locality: “America meridionali.” Restricted to “French
Guiana” by neotype designation by Bour (2006a:29).
Emys macrocephala Spix 1824:5 (senior homonym, not
= Emys macrocephalus Gray 1844), Peltocephalus
macrocephala
Type locality: “Airon ad ripam fluminis Yau, confluentis Rio
Negro” [Airão, Rio Jáu, Amazonas, Brazil].
Emys tracaxa Spix 1824:6, Hydraspis tracaxa, Podocnemis
tracaxa, Peltocephalus tracaxa, Peltocephalus tracaxus
Type locality: “fluminis Solimoëns” [Rio Solimões, Amazonas,
Brazil].
Chelys (Hydraspis) dumerilliana Gray 1830e:17 (nomen
novum)
Emys icterocephala Spix in Gray 1830e:17 (nomen nudum)
Peltocephalus tracaya Troschel 1848:646 (nomen novum)
Podocnemis Wagler 1830b
Podocnemis Wagler 1830b:135
Type species: Podocnemis expansa [= Emys expansa Schweigger
1812], by subsequent designation by Fitzinger (1843:29).
Chelonemys Gray 1864d:134 (junior homonyn)
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Type species: Chelonemys dumeriliana sensu Gray 1864 (non
Emys dumeriliana Schweigger 1812) [= subjective synonym of
Podocnemis unifilis Troschel 1848], by original monotypy.
Bartlettia Gray 1870f:720 (junior homonym, not = Bartlettia
Adams 1867 [= Mollusca])
Podocnemis expansa (Schweigger 1812)
Giant South American River Turtle, Giant Amazon River Turtle,
Arrau
Type species: Bartlettia pitipii Gray 1870 [= subjective synonym
of Podocnemis sextuberculata Cornalia 1849], by original
monotypy.
Podocnemis erythrocephala (Spix 1824) (10:46) (104)
Red-headed Amazon River Turtle
Richard C. Vogt / Rio Trombetas, Pará, Brazil
Richard C. Vogt / CBFTT / Rio Jufari, Amazonas, Brazil
Brazil (Amazonas, Pará, Roraima), Colombia (Guainía, Guaviare [?], Vaupés, Vichada), Venezuela (Amazonas)
CBFTT Account: Mittermeier, Vogt, Bernhard, and Ferrara
(2015)
IUCN Red List: Vulnerable A1bd (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II, as Podocnemis spp.
Emys cayennensis Schweigger 1812:298 (10:48, 14:50) (partim,
misidentified type, provisional nomen suppressum), Chelys
(Hydraspis) cayennensis, Chelys cayennensis, Hydraspis
cayennensis, Podocnemis cayennensis
Type locality: “Cayenna” [Cayenne, French Guiana].
Emys erythrocephala Spix 1824:9, Podocnemis erythrocephala, Hydraspis expansa erythrocephala, Chelys (Hydraspis) erythrocephala, Chelys erythrocephala
Type locality: “aquis ripariis fluminis Solimoens” [Rio Solimões,
Amazonas, Brazil].
Emys bitentaculata Cuvier in Gray 1830e:17
(nomen
nudum et dubium)
Hydraspis bitentaculata Gray 1831d:42 (14:48) (nomen oblitum
et dubium)
(14:48)
Type locality: “Brasilia” [Brazil].
Podocnemis agassizii Coutinho in Göldi 1886:277 (10:46)
Type locality: “Rio Negro” [Amazonas, Brazil].
Podocnemis coutinhii Göldi 1886:279 (nomen novum)
Bolivia (Beni, Cochabamba, La Paz, Pando, Santa Cruz), Brazil
(Amapá, Amazonas, Goiás, Mato Grosso, Pará, Rondônia, Roraima, Tocantins), Colombia (Amazonas, Arauca,
Caquetá, Casanare, Guainía, Meta, Putumayo, Vaupés,
Vichada), Ecuador, Guyana, Peru (Loreto, Ucayali),
Venezuela (Amazonas, Anzoátegui, Apure, Bolívar, Delta
Amacuro, Guárico, Monagas)
IUCN Red List: Lower Risk/conservation dependent (1996)
TFTSG Draft Red List: Critically Endangered (2011)
CITES: Appendix II, as Podocnemis spp.
Emys expansa Schweigger 1812:299, Podocnemis expansa,
Chelys (Hydraspis) expansa, Chelys expansa, Hydraspis
expansa
Type locality: “America meridionali.” Restricted to “French
Guiana” by lectotype designation by Bour (2006a:35).
Testudo arrau Humboldt 1819a:243 (14:49), Emys arrau
Type locality: “entre le confluent de l’Orénoque avec l’Apure et
les grandes cataractes ou Raudales...entre Cabruta et la mission
d’Aturès...les trois pêches de l’Encaramada ou Boca del Cabullare, de Cucuruparu ou Boca de la Tortuga, et de Pararuma, un peu
au-dessous de Carichana” [Venezuela]. Restricted to “zwischen
dem Zusammenfluss des Apure mit dem Orinoko und den grossen
Wasserfällen” [Venezuela] by Wermuth and Mertens (1961:296).
Emys amazonica Spix 1824:1
Type locality: “fluvio Solimoens et confluentibus Javary, Rio
Branco” [Rio Solimões, Amazonas, Brazil].
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Podocnemis lewyana Duméril 1852 (12:44)
Magdalena River Turtle
Podocnemis sextuberculata Cornalia 1849 (10:47) (105)
Six-tubercled Amazon River Turtle, Pitiu
Richard C. Vogt / Paraná Floresta, Roraima, Brazil
Alejandra Cadavid / CBFTT / TCC / Río Chicagua, Bolivar, Colombia
(orange dots = probable trade)
(orange dot = introduced or trade)
Colombia (Antioquia, Atlántico, Bolívar, Boyacá, Caldas,
Cesar, Córdoba, Cundinamarca, La Guajira, Magdalena,
Santander, Sucre, Tolima)
Introduced: Venezuela (Zulia)
CBFTT Account: Páez, Restrepo, Vargas-Ramirez, and Bock
(2009)
IUCN Red List: Critically Endangered A2acd+4acd (2016);
Previously: Endangered (1996)
CITES: Appendix II, as Podocnemis spp.
Podocnemis lewyana Duméril 1852:242
Type locality: “Santa Fé de Bogota...et...la République de
Venezuella” [Colombia and Venezuela]. Restricted to “Bogotá,
Colombia” by holotype [= lectotype] designation by Williams
(1954:281).
Brazil (Amazonas, Pará, Roraima), Colombia (Amazonas,
Caquetá, Putomayo, Vaupés), Ecuador (?), Peru (Loreto)
IUCN Red List: Vulnerable A1acd (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II, as Podocnemis spp.
Podocnemis sextuberculata Cornalia 1849:13 (105)
Type locality: “Fl. Amazonum” [Rio Amazonas, Brazil].
Podocnemis pitiu Coutinho 1868:150 (10:47)
Type locality: “l’Amazone” [Amazonas, Brazil].
Bartlettia pitipii Gray 1870f:720, Bartlettia pitipiti
Type locality: “Lakes of the Upper Amazons” [Brazil].
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Podocnemis unifilis Troschel 1848 (07:105, 08:16, 09:49, 10:48, 10:49, 14:50)
Yellow-spotted River Turtle, Yellow-spotted Sideneck Turtle,
Tracaja
Richard C. Vogt / Amazonas, Brazil / female
Bolivia (Beni, Cochabamba, La Paz, Pando, Santa Cruz), Brazil
(Acre, Amapá, Amazonas, Goiás, Mato Grosso, Maranhão, Pará, Rondônia, Roraima, Tocantins), Colombia
(Amazonas, Arauca, Caquetá, Casanare, Guainía, Meta,
Putumayo, Vaupés, Vichada), Ecuador, French Guiana,
Guyana, Peru (Huanuco, Loreto, Madre de Dios, Pasco,
Ucayali), Suriname, Venezuela (Amazonas, Anzoátegui, Apure, Barinas, Bolívar, Cojedes, Delta Amacuro,
Guárico, Monagas)
IUCN Red List: Vulnerable A1acd (1996)
TFTSG Draft Red List: Endangered (2011)
CITES: Appendix II, as Podocnemis spp.
Emys cayennensis Schweigger 1812:298 (10:48, 14:50)(partim,
misidentified type, provisional nomen suppressum), Chelys
(Hydraspis) cayennensis, Chelys cayennensis, Hydraspis
cayennensis, Podocnemis cayennensis
Type locality: “Cayenna” [Cayenne, French Guiana].
Testudo terekay Humboldt 1819a:243 (14:49) (nomen oblitum),
Emys terekay
Type locality: “Haut-Orénoque...l’Apure, l’Uritucu, la Guarico
et...les Llanos de Caracas” [= Upper Orinoco, Apure, Uritucu,
Guarico, and the Llanos of Caracas, Venezuela].
Chelys (Hydraspis) lata Bell in Gray 1830e:17 (08:16, 10:7) (nomen oblitum), Chelys lata, Hydraspis lata
Type locality: “Demerara” [Guyana].
Podocnemis unifilis Troschel 1848:647
Type locality: “Britisch-Guiana...Rupununi und Takutu”
[Guyana].
Podocnemis tracaya Coutinho 1868:149 (10:49)
Type locality: “l’Amazone” [Amazonas, Brazil].
Podocnemis vogli Müller 1935
Savanna Sideneck Turtle, Llanos Sideneck Turtle
Anders G.J. Rhodin / Hato El Cedral, Mantecal, Apure, Venezuela
(orange dots = probable introduced or trade)
Colombia (Arauca, Casanare, Guaviare, Meta, Vichada), Venezuela
(Anzoátegui, Apure, Barinas, Bolívar, Cojedes, Delta Amacuro, Guárico, Monagas, Portuguesa)
Introduced: Venezuela (Zulia)
IUCN Red List: Least Concern [Not Listed] (1996)
TFTSG Draft Red List: Vulnerable (2011)
CITES: Appendix II, as Podocnemis spp.
Podocnemis vogli Müller 1935:104
Type locality: “Barinas (Staat Zamora), Venezuela.” [= Barinas
Prov.].
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Regional Species Richness Maps
Distribution shapefiles for all taxa in these maps are each shown in red at 80% transparency and stacked;
lighest red color = one taxon, darkest red color = > 15 taxa. For composite global species richness map, see p. 17.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
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Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
ANNOTATIONS
Comments on taxonomic change or other annotations in
this checklist and previous checklists are indicated by superscripts. New annotations in each new checklist are simple bold
numbers in separate parentheses, e.g.,(35). Earlier annotations
from any of the previous checklists are indicated in subsequent
checklists by two-part non-bold superscripts in separate parentheses that indicate the year of publication and the annotation
number from that year, e.g.,(07:105, 08:25, 14:50). All annotations from
previous checklists are now listed at the end of this section in
order to facilitate access to these commentaries, and all checklists are available as open-access publications online at www.
iucn-tftsg.org/checklist/.
CURRENT CHECKLIST
1. Phylocode Classification: Crawford et al. (2015) performed a genome-scale analysis of turtle phylogeny, sequencing
2381 ultraconserved element loci representing a total of 1,718,154
bp of aligned DNA sequences in 32 turtle taxa representing 14
turtle families. Their recovered phylogeny corresponded to
well-supported clades that they concluded were consistent with
the temporal appearance of clades and paleobiogeography. They
recommended the alternative hierarchical Phylocode classification of turtles presented at the beginning of our Checklist. We
continue to use our ICZN-compliant Linnaean classification as
outlined on the same page.
In addition, Pereira et al. (2017) recently used publicly available databases for nucleotide sequences and composed a dataset
comprising 13 loci for 294 living species of Testudines, accounting for all living genera and 83% of extant species diversity (as
recognized in our checklist); they constructed a Phylocode-based
classification scheme somewhat at variance with both Crawford
et al. (2015) and our checklist.
2. Macrochelys: Previous molecular studies have demonstrated significant diversity within the long-recognized widespread
species Macrochelys temminckii: mtDNA data supported the
distinction of three geographic clades, and microsatellite data
suggested that six genetic clusters were recognizable (Roman et
al. 1999; Echelle et al. 2010). Based on additional mtDNA data,
a morphometric (osteological) analysis, and an examination of
all available fossil material for the genus, Thomas et al. (2014)
demonstrated the existence of three distinct, Recent, geographically separated populations: M. suwanniensis (Suwannee River
drainage), M. apalachicolae (Apalachicola River and nearby lesser
drainages), and a restricted M. temminckii (Yellow-Conecuh to
Mobile Bay to Mississippi to Neches River basins). Independently,
Murray et al. (2014) examined morphometric variation in the skull
of Macrochelys across most of its range. Their results demonstrated
distinct drainage-specific differences in skull morphology, supported the distinction of the Suwannee River turtles, and suggested
that “further splitting may eventually be warranted” among the
basins west of the Suwannee. Subsequently, Folt and Guyer (2015)
reconsidered the published data, critiqued the methods of Thomas
et al. (2014), and concluded that the evidence to date supported
the distinction of M. suwanniensis, but that M. apalachicolae was
not adequately distinguished from M. temminckii (sensu stricto),
and thus recommended their synonymization, with which we
concur.
3. Macrochelys: Two names for Alligator Snapping Turtles
used by Hoser (2013) (Macrochelys temminckii muscati and
Macrochelys maxhoseri) in his attempt to name some of these
lineages, have been noted by Thomas et al. (2014) to be unavailable for nomenclatural purposes due to technical errors in the
descriptions.
4. Names coined by Hoser: Raymond Hoser (2013, 2014a,b,
and several other papers) has circumvented conventional standards
of scientific analysis and peer-review in his broadly sweeping and
extensive new taxonomies and nomenclatures. We regard his actions
as confusing and unwarranted acts of nomenclatural disruption under
the International Code of Zoological Nomenclature (ICZN 1999),
and we do not regard the documents circulated under the name
Australasian Journal of Herpetology as scientific publications nor
as available publications for the purposes of nomenclature (Kaiser
et al. 2013; Kaiser 2014; Rhodin et al. 2015). In collaboration with
a wide leadership group representing the global herpetological and
zoological communities, we have petitioned the International Commission on Zoological Nomenclature to declare and treat Hoser’s
works in his self-produced Australasian Journal of Herpetology as
nomenclaturally unavailable (Rhodin et al. 2015), and have noted
that all new names created therein are nomenclaturally unavailable pending a ruling by the Commission. Hoser’s production has
heretofore focused primarily on snakes, but recently he has proposed names for purportedly distinct new taxa of turtles, including
American Macrochelys, Australasian Pelochelys, and Australian
Chelodina. We consider all these names to be unavailable unless
the ICZN rules them to be available.
5. Caretta caretta: Continued examination of intraspecific
genetic structure within this species by Shamblin et al. (2014)
using mitochondrial gene sequences from samples from 42 nesting
rookeries identified 59 different haplotypes. However, the authors
made no taxonomic recommendations, and we agree.
6. Eretmochelys imbricata: Gaos et al. (2016) examined
mtDNA haplotype diversity among nesting populations of Hawksbills along the eastern Pacific Ocean. Despite the low genetic
diversity across their samples, their phylogenetic analysis suggested that Eastern Pacific Hawksbills are more closely related
to those from the Indo-Pacific rather than to those in the Atlantic,
despite the recent closure of the Panamanian Portal. In addition,
despite the low haplotype diversity overall, differences among
the four major eastern Pacific rookeries (El Salvador, Nicaragua,
Costa Rica, and Ecuador) led the authors to recommend that these
four rookeries should be considered separate management units
for conservation purposes.
7. Lepidochelys olivacea: This species was described by
Eschscholtz in 1829 in two separate publications (Eschscholtz
1829a, 1829b), the earlier of them in general overlooked until
recently (Flores-Villela et al. 2016). Historically, and in our
previous checklists, the Zoologischer Atlas (Eschscholtz 1829b)
has always been credited as the source for the name Chelonia
olivacea, and although we previously listed his other publication from the journal Die Quatember (Eschscholtz 1829a) as the
source for two other new names (Chelonia castanea and Chelonia grisea) (TTWG 2014), we assumed that his Zoologischer
Atlas had been published first and had nomenclatural priority.
However, Flores-Villela et al. (2016) have now shown that the
Quatember article was published earlier (January 1829) than the
Atlas (after May 1829), and is therefore the original source for
the name olivacea. In addition, the relatively overlooked name
Chelonia grisea, previously synonymized by others and us under
Eretmochelys imbricata, has been shown by Flores-Villela et al.
(2016) to more likely represent an anomalous Chelonia mydas,
and we follow their suggestion to reassign it as such.
8. Chelonia mydas: Okamoto and Kamezaki (2014) examined
plastral coloration and head and shell morphometrics for Chelonia
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captured in the Western Pacific off Japan. Their data demonstrated
the presence of two distinct phenotypes (“yellow” and “black”)
and they argued that the black form represented C. agassizii, and
should be recognized as a full species. However, given previous
studies (see annotation 07:4), we continue to consider C. agassizii to be a synonym of C. mydas, unless integrated range-wide
analyses of morphology, coloration, and genetics conclusively
demonstrate significant lineage divergence.
Naro-Maciel et al. (2014) examined 15 microsatellite markers
across 19 green turtle rookeries across the Atlantic Ocean. These
data revealed a strong barrier to dispersal between the northern and
southern Atlantic, but not a degree meriting taxonomic changes.
9. Wagler 1830b and 1830c: Gutsche and McCranie (2016)
stated that Wagler’s 1830 publication Natürliches System der
Amphibien was issued as two separate parts: first the text (Wagler 1830b), and then the plates (Wagler 1830c), although dates
for the publication of each were not provided. We and others
had always treated these two parts as a single publication, but
it appears that they appeared as separate issues. As a result, the
name Dermatochelys was established in Wagler 1830b, but the
species Dermatochelys porcata first appeared in Wagler 1830c.
Similarly, the name Cinosternon hirtipes was only a nomen nudum
in Wagler 1830b, but described by association with the figures in
Wagler 1830c, as we previously noted in our annotation 09:11.
These changes do not affect any taxonomic or nomenclatural
considerations. We also note in passing that the description of
Testudo boiei Wagler 1830a (= Chelonoidis carbonarius) appears
to have preceded these others.
10. Dermochelys coriacea: Using mtDNA sequences and
nuclear microsatellite markers, Molfetti et al. (2013) and Dutton
et al. (2013) demonstrated considerable genetic structure within
Dermochelys coriacea in the Atlantic Ocean basin. Although they
advised the recognition of their units for conservation purposes,
they made no taxonomic recommendations.
11. Kinosternon: Iverson et al. (2013) sequenced three mitochondrial and three nuclear markers for all recognized species of
kinosternid turtles, and their analysis revealed strong support for a
monophyletic clade of Sternotherus, a second monophyletic clade
of primarily Meso-American taxa that they named Cryptochelys,
and a third restricted monophyletic clade of the remaining species
formerly included in Kinosternon. However, their support for the
non-monophyly of the traditional inclusive genus Kinosternon
was weak. Subsequently, Spinks et al. (2014b) sequenced 14 additional nuclear loci for most (but not all) recognized species, and
their analysis of the nuclear data alone supported the reciprocal
monophyly of Sternotherus and the traditional Kinosternon (sensu
lato), but did not support a monophyletic Cryptochelys. Because
two independent data sets have produced different conclusions,
we retain the alternative generic arrangement from our last edition
until new data emerge or new analyses are performed to settle
this complicated issue.
12. Kinosternon steindachneri: Bourque (2016) treated this
taxon as a distinct species based on morphology, and argued that
no morphological or genetic data have been presented to support
its continued recognition as a subspecies of K. subrubrum (Iverson
1998; Bourque 2012a, b; Iverson et al. 2013; Spinks et al. 2014;
Bourque and Schubert 2015). We concur, and therefore now list
K. steindachneri as a distinct species.
13. Kinosternon stejnegeri: Iverson (1979a) synonymized
extant K. flavescens stejnegeri Hartweg 1938 with the Pliocene–
Pleistocene fossil taxon, K. arizonense Gilmore 1923, based on
similar morphology. Considered a subspecies, K. f. arizonense,
by Iverson (1979b), it was subsequently recognized as a distinct
species based on genetic analysis by Serb et al. (2001). Recently,
McCord (2016) examined all available Pliocene material of K.
arizonense and compared it with extant specimens, and concluded
that the fossil material differs significantly from the extant material and is actually more similar morphologically to K. flavescens
(sensu stricto). As a consequence, he restricted the name arizonense
to the Pliocene fossils and resurrected the old name stejnegeri for
the extant species. This conclusion was also tentatively accepted
by Joyce and Bourque (2016), and we follow these recommendations here, accepting the resurrected name K. stejnegeri for the
extant species previously known as K. arizonense.
14. Kinosternon subrubrum hippocrepis: Bourque (2016)
elevated this taxon to a full species based on morphological data
from living and fossil forms; however, without a range-wide
analysis of morphometric and/or molecular data supporting that
change, we continue to recognize hippocrepis as a subspecies of
K. subrubrum.
15. Sternotherus depressus: Scott and Rissler (2015) reported a 32–56% decline in the historical range of S. depressus,
as well as significant unidirectional mtDNA introgression from
S. minor peltifer. This hybridization is changing the morphology
of S. depressus and severely threatens its continued distinction.
16. Sternotherus minor peltifer: Bourque (2016) argued that
this taxon should be recognized as a full species based on previous published phylogenetic analyses using molecular (Iverson et
al. 2013) and morphological data (Bourque and Schubert 2015).
In addition, Guyer et al. (2016), based on color differences and
the mitochondrial DNA study by Walker et al. (1995), also recommended elevating S. m. peltifer to species status. However,
Walker et al. (1995) sampled part of only one mitochondrial gene
and included no samples from the previously hypothesized area
of intergradation along the Gulf Coast (Iverson 1977). Because
animals that appear to be morphological intergrades have been
described (Iverson 1977), we consider these recommendations
premature and continue to recognize peltifer as a subspecies of
S. minor.
17. Emydidae: Seidel and Ernst (2017) provided an extensive
review of the history and taxonomic status of the phylogeographic
relationships of the family and its two subfamilies, but recommended no significant taxonomic changes.
18. Chrysemys picta: Jensen et al. (2014a) examined genetic
variation among populations of C. picta at the northwestern range
limit in British Columbia (C. p. bellii). Although they documented
very little variation in regional mitochondrial sequences, they
found two unique mitochondrial haplotypes when compared to
previously published range-wide data. In contrast, based on nine
microsatellite loci, they identified six distinct local population
clusters. They urged that these six geographic units be managed
separately, but made no taxonomic recommendations; however,
their data did support the continued recognition of C. dorsalis
and C. picta as separate species. In addition, Jensen et al. (2015a)
examined variation in one mitochondrial and one nuclear gene
across the range of the genus. The nuclear gene provided no
resolution, but the mitochondrial data demonstrated the reciprocal monophyly of C. dorsalis versus C. picta (sensu stricto), but
no clear pattern among the subspecies of the latter. Because the
available evidence for the recognition of C. dorsalis as a species
vs. subspecies is primarily mitochondrial (see TTWG annotations 07:11 and 10:16), we continue to list two alternatives for
its classification.
19. Graptemys: Praschag et al. (2017) examined over 3200
bp of mtDNA sequence data and 7800 bp of nuclear DNA (across
12 loci) for 89 specimens of all recognized taxa of the genus
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
Graptemys. The mtDNA data supported the distinctiveness of G.
geographica (sister to all other Graptemys), and the divergence
of the broad-headed clade from the narrow-headed clade. Most
species in the broad-headed clade were resolved as reciprocally
monophyletic, but in the narrow-headed clade only G. caglei
and G. versa were well supported as monophyletic. Principal
component analyses using coded phased nuclear DNA sequences
revealed eight clusters: 1) G. geographica; 2) G. barbouri; 3) G.
caglei; 4) G. versa; 5) G. sabinensis; 6) G. ernsti, G. gibbonsi, G.
pearlensis, and G. pulchra; 7) G. flavimaculata, G. nigrinoda, and
G. oculifera; and 8) G. ouachitensis and G. pseudogeographica.
The authors concluded that species recognitions of G. geographica,
G. barbouri, G. caglei, G. versa, and G. sabinensis were all
well supported, but that the remaining taxa were oversplit. They
suggested 1) that G. flavimaculata and G. nigrinoda should be
relegated to subspecies of G. oculifera (see also Mertens and
Wermuth 1955); 2) that G. ernsti, G. gibbonsi, G. pearlensis, and
G. pulchra are conspecific; and 3) that G. ouachitensis should
be relegated to a subspecies of G. pseudogeographica. Their
results supported the earlier view by Walker and Avise (1998)
that Graptemys is taxonomically oversplit, and they made some
suggestions for possible changes, but they presented no explicit
taxonomic revisions, and we await further clarification before
changing the checklist.
20. Graptemys caglei: Ward et al. (2013) examined microsatellite variation across the range of Graptemys caglei in the
Guadalupe and San Marcos Rivers of Texas. They found weak
but identifiable divergence between populations in the Upper
Guadalupe Rive versus those in the Middle Guadalupe and San
Marcos Rivers. They made no taxonomic recommendations, but
noted that turtles in these two regions also differed in life history,
morphology, and coloration.
21. Graptemys ernsti and G. barbouri: Godwin et al. (2014)
confirmed the previously unrecognized presence of both Graptemys
ernsti and G. barbouri in the Choctawhatchee River basin, and
demonstrated hybridization between those two species in that basin.
Based on the available evidence they concluded that both species
were likely present in that basin prior to human intervention. In
a follow-up study, Ennen et al. (2016) examined morphometric,
colorimetric, microsatellite and mitochondrial DNA variation in
G. ernsti, across the three inhabited basins. They demonstrated no
morphometric differences between turtles in two of the basins (no
data for the Pea River, a tributary of the Choctawhatchee), weak
colorimetric differences between turtles from the Yellow vs. Conecuh/Pea basins, and both microsatellite and mtDNA evidence for
the distinction of turtles from the Yellow River vs. those from the
other basins. They declined to recognize those populations taxonomically, and hesitated to even recommend their recognition as
Evolutionarily Significant Units, and we concur with these conclusions. Although it is now clear that both G. ernsti and G. barbouri
are established in the Choctawhatchee River basin, whether either
of them is native to that system is still an open question (Godwin
et al. 2014; Ennen et al. 2016).
22. Graptemys nigrinoda: Ennen et al. (2014) examined
morphometric and colorimetric characters, as well as sequence
variation in a single mitochondrial gene, across the range of
Graptemys nigrinoda. Morphological variation was primarily
clinal, and mitochondrial haplotypes differed by less than 0.3%
and were not related to geography. They recommended the synonymy of G. n. delticola with the nominate form, and we have
followed their recommendation.
23. Graptemys pseudogeographica: Lindeman et al. (2015)
demonstrated that False Map Turtles from the Calcasieu River
basin in southwestern Louisiana differ from those in all other
basins in having a unique eye color and a variable chin pattern.
They made no taxonomic recommendations, but encouraged further range-wide morphological and genetic study of this species
in light of these findings.
24. Malaclemys terrapin: Hart et al. (2014) examined 12
nuclear microsatellite loci from 21 populations of Malaclemys
terrapin from across the species’ range, and identified four genetic clusters that did not correspond to currently accepted,
morphology-based subspecies descriptions. Despite gaps in their
coastal sampling along the east coast of Florida and Georgia, and
the Gulf Coast of Florida, their recommended management units
were Massachusetts (part of the range of M. t. terrapin), New
York to South Carolina (most of the range of M. t. terrapin, and
part of the range of M. t. centrata), the Florida Keys to Tampa
Bay (the range of M. t. rhizophorarum and part of that of M. t.
macrospilota), and Louisiana and Texas (the combined ranges
of M. t. pileata and M. t. littoralis). In addition, based on 16
microsatellite loci (12 shared by Hart et al. 2014), Drabeck et
al. (2014) also found low diversity and an absence of structure
among populations of M. terrapin along the Gulf Coast of Louisiana and Texas. Both studies questioned the current subspecies
designations, but made no explicit taxonomic recommendations,
and we agree. More studies with better geographic sampling are
clearly needed.
25. Pseudemys concinna: In a generally overlooked publication, Guérin (1829) provided an illustration as indication of a
species that he named Emys concinna, preceding the description
of Testudo concinna by Le Conte (1830). However, the species
figured appears to represent what is now considered Trachemys
scripta elegans, described by Wied (1839). Whereas Emys concinna Guérin 1829 is technically a senior homonym of Testudo
concinna Le Conte 1830 (later placed in the genus Emys) and
precedes the description of Emys elegans Wied 1839, it is also a
name and attribution that has never been used since its appearance and we therefore declare it a nomen oblitum, not requiring
any change in status or usage of the two subsequently published
widely-used and accepted names.
26. Central American Trachemys: The taxonomy of MesoAmerican Trachemys sliders has been a quagmire for decades.
Because of the level of disagreement among recent authors
concerning this group, with no clear resolution in sight, our last
checklist listed as many as three taxonomic options for some
taxa. However, in an attempt to settle the confusion, Parham et al.
(2015) examined variation in one mitochondrial and one nuclear
gene across every named Trachemys taxon from Mexico. Their
analysis revealed that the samples of “ornata” from Acapulco used
by Fritz et al. (2012) clustered with turtles from the Caribbean
versant, and not with confirmed ornata from near its type locality.
Hence, the species name ornata should be restricted to the western
versant of Mexico, and the name venusta should be restricted to
the eastern drainages of Mexico and Central America. Furthermore, they confirmed the finding of Fritz et al. (2012) that grayi
is not closely related to venusta, but the sister taxon of emolli. We
agree with these insights (although we remain equivocal regarding
whether emolli is a distinct species or a subspecies of grayi) and
have modified our checklist accordingly. Additionally, Fritz et al.
(2012) showed that panamensis is very closely related to grayi,
and should be considered a subspecies of that taxon, rather than
of venusta, further corroborating the split between western and
Caribbean versant taxa; we have now made that change in our
checklist. Also, as a consequence of the recognition that ornata
is a western versant taxon, callirostris is now considered either a
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distinct species or a subspecies of venusta, rather than of ornata,
as it was designated in our previous checklist.
27. Trachemys adiutrix or T. dorbigni adiutrix: Using a
niche modelling approach, Rodrigues et al. (2016) supported the
subspecies status of the Maranhao slider. The invasive potential
of the nominotypical subspecies was better explained when the
different climatic niches of both taxa were combined, reflecting the common pattern of lack of niche conservatism between
subspecies.
28. Trachemys grayi: The species Callichelys concinna Gray
1873a, described from San Mateo, Tehuantepec, Oaxaca, Mexico,
has unfortunately been overlooked in all previous checklists,
including our own (Siebenrock 1909a; Wermuth and Mertens
1961, 1977; Fritz and Havaš 2007; TTWG 2014). However, Gray
(1873f) and Boulenger (1889) synonymized it with T. grayi, and
Harfush-Meléndez and Buskirk (2008) noted it in their analysis of
the distribution of T. grayi, and we now include it in our checklist.
29. Clemmys guttata: Davy and Murphy (2014) examined
variation in 11 nuclear microsatellite loci across Canadian populations of Clemmys guttata. They identified significant genetic
structure, with a minimum of six distinct subpopulations, with the
most distinctive population being in Hastings County, Ontario.
They made no taxonomic recommendations, but strongly urged
management and protection of the latter population, which numbers
less than 50. An expansion of this study across the entire range
of C. guttata would be invaluable.
30. Emys orbicularis orbicularis: The identity of the
Pleistocene fossil names Clemmys schlotheimii † Fitzinger 1835
and Trionyx schlotheimii † Fitzinger 1835 has previously been
uncertain, and we did not even include T. schlotheimii in our
previous checklists. However, Karl and Paust (2014) examined
the original fossil material, designated lectotypes of both taxa,
and confirmed that both are synonyms of Emys orbicularis.
31. Emys orbicularis orbicularis: The Pleistocene fossil
taxon Testudo (Emys) canstadiensis † Plieninger 1847 was included under the synonymy of Testudo hermanni in our previous
checklist (TTWG 2014), based on presumed synonymy suggested
by Auffenberg (1974); however, the specimen is actually an Emys
orbicularis, based on synonymization by Karl and Tichy (2002),
and we corrected it in our fossil checklist (TEWG 2015), and
herein.
32. Emys orbicularis occidentalis: Based on both mitochondrial DNA sequences and 15 nuclear microsatellite markers,
Stuckas et al. (2014) identified two distinct genetic units within
Emys orbicularis in North Africa, one from Morocco (identified as
E. o. occidentalis) and the other from eastern Algeria and Tunisia
(undescribed). The former was found to be very similar to Iberian
specimens (identified as E. o. fritzjuergenobsti). The authors
declined to name the new taxon for want of morphological data,
but synonymized E. o. fritzjuergenobsti under E. o. occidentalis,
and we reflect that change here.
33. Emys orbicularis persica: We spell the name of this taxon
as Emys orbicularis persica, following Fritz (1998). However,
Eichwald (1831:196) recognized three varieties (“var.”) of Emys
europaea Schneider 1783: the nominotypical, not named, α by
inference, and β and γ, with names given as “ibericae var. β”
and “minoris var. γ persicae.” One might therefore assume that
the valid names should be Emys europaea persicae and Emys
europaea ibericae. However, the names ibericae and persicae
are adjectives, referring to the countries of Iberia and Persia,
respectively, and as a result, ICZN Article 11.9.2 applies: “An
adjectival species-group name proposed in Latin text but written otherwise than in the nominative singular because of the
requirements of Latin grammar is available provided that it meets
the other requirements of availability, but it is to be corrected to
the nominative singular if necessary.” Therefore, persica and
iberica are the valid names, and these were subsequently used
by Eichwald (1840:47).
34. Emys trinacris: Vamberger et al. (2015) examined mitochondrial and nuclear microsatellite variation (15 markers) in
Emys from southern Italy. Their results revealed negligent gene
flow between E. orbicularis and E. trinacris, with intergradation
evident between E. o. galloitalica and E. o. hellenica. Their data
support the continued recognition of E. trinacris as a species, and
galloitalica and hellenica as subspecies of E. orbicularis.
35. Emys or Actinemys pallida: Spinks et al. (2014a) examined 89 nuclear single nucleotide polymorphisms (SNPs) and a
mitochondrial gene sequence from range-wide samples of Emys
or Actinemys marmorata, and compared those results with earlier work (Spinks et al. 2010) using nuclear gene sequence data.
The mitochondrial data resolved four poorly supported clades,
whereas the previous nuclear sequence data revealed only two
main groups with considerable admixture between them. However,
the SNP analysis demonstrated strong support for two geographic
clusters: a northern group from the southern San Joaquin valley
to Washington, and a southern group from the Central Coastal
Range of California and southern California to Baja California.
These results are remarkably (though not perfectly) consistent
with Seeliger’s (1945) morphological work, who described the
subspecies pallida for the southern populations. Furthermore,
secondary analyses by Spinks et al. (2014a) of the molecular
data within each of these two groups recognized two subgroups
in the southern cluster, one from the coastal range and southern
California, and the other from Baja California. Although populations from Baja were also recognized as morphologically distinct
(though undescribed) by Seeliger (1945), Spinks et al. only noted
that species recognition may emerge from future studies. Hence,
they recommended that the two California lineages be recognized
as separate species (rather than subspecies as defined by Seeliger
1945), and we follow that recommendation here.
36. Emys or Emydoidea blandingii: Based on variation
across 12 microsatellite loci, Davy et al. (2014) identified four
distinct genetic units among populations of Emydoidea blandingii
in Ontario, Canada, and suggested that they should be managed
separately. Similarly, based on eight microsatellites, Sethuraman
et al. (2014) identified moderate but significant differentiation
among Midwestern USA populations, with four or five unique
genetic clusters, but were unable to explain the close genetic relationship between a population in western Nebraska to those in
eastern Illinois, rather than with intervening populations in Iowa.
McCluskey et al. (2016) examined variation in seven microsatellites in this species in New York, and identified two or possibly
three genetic units there. None of these studies made taxonomic
recommendations. Range-wide genetic studies of E. blandingii
are clearly needed, as is careful population management of this
genetically diverse species.
37. Terrapene carolina complex: These North American
box turtles remain the center of considerable taxonomic controversy (see TTWG 2014, annotation 7:27). Based primarily on
mtDNA data, Martin et al. (2013) recommended the recognition
of the western forms (triunguis, mexicana, and yucatana) as a
polytypic species (T. mexicana being the oldest name) separate
from the eastern forms (under T. carolina). However, in a brief
summary of the turtle taxonomy issues, Fritz and Havas (2013)
declined to accept this major change for the genus Terrapene,
primarily because of evidence for introgression between the
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
proposed species. Martin et al. (2014) responded with evidence
of interspecific hybridization between other well-accepted species pairs, and reaffirmed their conclusion that T. mexicana was
a distinct species by the phylogenetic species concept. Fritz and
Havas (2014) replied with three lines of evidence as to why they
considered it to be premature to recognize T. mexicana as distinct
from T. carolina. First, intergradation is common where they are
sympatric, and indeed, Butler et al. (2011) demonstrated panmixia
(rather than occasional hybridization) in those areas. Second,
mexicana was resolved as monophyletic by Martin et al. (2013)
with relatively weak support based on mtDNA, and no support
when based on a single nuclear gene. Third, previous study of
mitochondrial and nuclear DNA sequence data (Spinks et al.
2009; seven nuclear loci) did not resolve triunguis as reciprocally
monophyletic relative to eastern forms of T. carolina. A recent
analysis of 30 nuclear and one mitochondrial loci (Spinks et al.
2016) indicated that while triunguis appears to be monophyletic
and distinct, it is nested within a paraphyletic carolina; given the
phylogenetic uncertainty surrounding much of this complex and
the demonstrated hybridization among taxa, these authors also
recommended that the traditional carolina taxonomy (all taxa as
subspecies of carolina) be retained pending a comprehensive,
genomic-level analysis of all contained taxa. Unfortunately, some
authors (e.g., Guyer et al. 2015; Powell et al. 2016) have chosen
to recognize the western populations as T. triunguis (rather than
T. mexicana), without justification. Since the taxonomic issues
here appear relatively unresolved, we continue to recognize the
competing alternative taxonomies, pending additional sampling
of the nuclear genome.
38. Terrapene carolina carolina: In a nearly range-wide study
of variation across eleven microsatellite loci for the subspecies T.
c. carolina, Kimble et al. (2014) found surprisingly little genetic
structure, except that the Appalachian Mountains represented a
modest barrier to gene flow among populations.
39. Platysternon megacephalum: In the present checklist we
reallocate the placement of Platysternon megacephalum tristernalis Schleich and Gruber 1984 by moving it from the synonymy
of P. m. megacephalum into the synonymy of P. m. peguense.
Ernst and Laemmerzahl (2002) originally placed P. m. tristernalis
in the synonymy of the nominotypical subspecies, which was
followed by Fritz and Havaš (2007) and all previous editions of
our checklist. However, Vetter and van Dijk (2006) identified the
holotype of tristernalis from southwestern Yunnan as representing peguense, the taxon occurring in adjacent northwestern Laos,
and Zheng et al. (2013) identified animals apparently from the
tristernalis-topotypic area of southwestern Yunnan as peguense,
both morphologically and genetically, and we agree with these
assessments. In addition, Ernst and Laemmerzahl (2002) noted
that the populations on Hainan appear to be intergrades between
P. m. peguense and P. m. shiui, as also reflected on our current
map.
Using the duplicate control region sequences of the mitochondrial genome of 20 P. megacephalum representing all three
subspecies, Zheng et al. (2013) found these morphologically defined taxa to also be genetically distinct, with P. m. megacephalum
and P. m. shiui identified as sister taxa. More comprehensive geographic surveys and the application of additional genetic markers
to samples from across the species’ entire range should hopefully
refine current taxonomy and provide a better understanding of
the geographic ranges of the individual taxa.
40. Cuora: Tiedemann et al. (2014) sampled 16 nuclear
microsatellite markers across all members of the Cuora trifasciata
complex (C. aurocapitata, C. pani, C. zhoui, C. trifasciata, and the
controversial Vietnamese C. cyclornata) (see previous annotations
on the status of cyclornata Blanck et al. 2006a: TTWG 2007b,
07:36; TTWG 2009, 09:23; TTWG 2012, 12:22). The analyses
by Tiedemann et al. (2014) confirmed the genetic distinction of
each of these five taxa, and they recommended their recognition
at the species level. In addition, their data corroborated the genetic
distinction of the two morphotypes within cyclornata that were
previously described as C. c. cyclornata and C. c. meieri (Blanck
et al. 2006a). Finally, they also corroborated the genetic distinction
of two morphotypes within the restricted C. trifasciata, one from
the Chinese mainland (C. t. trifasciata according to the authors)
and the other an undescribed subspecies from Hainan Island.
Independently, Li et al. (2015) sequenced the entire mitochondrial genome of nine species of Cuora; their phylogenetic
analysis revealed that C. trifasciata (sensu lato) is polyphyletic,
and that Chinese (trifasciata) and Vietnamese (cyclornata) populations are distinctly different. They also recommended the species
recognition of C. cyclornata.
Given that morphology and mitochondrial and nuclear DNA
data each support the recognition of C. cyclornata as a species
distinct from C. trifasciata, with the former comprising two genetic
lineages, we now follow Blanck et al. (2006a) in recognizing C.
cyclornata with two subspecies, C. c. cyclornata and C. c. meieri.
41. Cuora amboinensis: In a morphological and colorimetric
study of populations of Cuora amboinensis, Ernst et al. (2016)
recommended the synonymy of C. a. lineata with C. a. kamaroma,
and of C. a. couro with C. a. amboinensis (suggesting that C.
a. couro was an intergrade between C. a. kamaroma and C. a.
amboinensis). However, their samples of lineata and couro were
small, their analyses and conclusions were based primarily on
color patterns, and they did not discuss the size of the inguinal
scute (apparently diagnostic for at least lineata), nor did they
present statistical or graphical results of their morphometric
analyses (i.e., separately from color data).
Subsequently, Protiva et al. (2016) analyzed shell shape and
mitochondrial DNA from C. amboinensis from primarily Borneo,
Sumatra, and Seram. They disagreed with the findings of Ernst et
al. (2016), and documented morphologic and genetic differentiation between Bornean vs. Sumatran populations, affirming the
distinctiveness of C. a. couro from Sumatra and C. a. kamaroma
from Borneo. Additionally, they identified a deeply divergent
lineage from Seram that they indicated might require taxonomic
recognition based on further work and improved sampling. Based
on this, we consider the synonymizations by Ernst et al. (2016)
to be premature, and await comprehensive genetic studies of C.
amboinensis before altering the taxonomy.
42. Cuora aurocapitata, C. cyclornata, and C. trifasciata:
Blanck et al. (2017) analyzed variation in these three Cuora
species from across their range, using morphometric principal
components analysis (PCA) and microsatellite data, comparing
their results with previous genetic work by Tiedemann et al.
(2014) and Li et al. (2015). They concluded that all three species are polytypic and described a new subspecies within each:
C. aurocapitata dabieshani, C. cyclornata annamitica, and C.
trifasciata luteocephala. We tentatively accept these named taxa
as new subspecies pending further analysis, but note that the
described lineages are not completely distinctive and that the
mixing of individuals across the clades suggests that this may
be too fine an application of the lineage approach to defining
taxa.
43. Cuora flavomarginata: The map for this species has been
updated and revised extensively by re-evaluating most previously
recorded localities. We have also added many localities provided
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by T. Blanck based on compilations of Chinese survey literature
and field data.
44. Cyclemys atripons and C. oldhamii: Vamberger et al.
(2017b) examined a recently discovered Cyclemys population
from Phnom Kulen National Park in northwestern Cambodia
using external morphology, 17 unlinked microsatellite loci, and
the mitochondrial cytochrome b gene. Morphologically, the
turtles resemble C. oldhamii, but have mitochondrial haplotypes
of C. atripons, while having microsatellite loci distinct from C.
atripons. The authors concluded that this population represents
either a natural hybrid swarm of C. atripons and C. oldhamii or
a distinct undescribed species with introgressed mitochondria
of C. atripons, without drawing taxonomic consequences. This
underscores that genetic differentiation in Cyclemys is complex
and still incompletely understood.
45. Cyclemys pulchristriata: Using a turtle kept in Shanghai
Zoo, Li et al. (2017) published a complete mitochondrial genome
(mt genome) assigned to Cyclemys pulchristriata and calculated a
phylogenetic tree using complete mt genomes of other geoemydid
species from GenBank. The topology of their tree conflicted with
the trees published by Fritz et al. (2008) and Stuart and Fritz
(2008) using the mitochondrial cytochrome b gene. When the
cytochrome b sequences of the mt genomes used by Li et al. (2017)
are compared to the data set of Fritz et al. (2008), it turns out that
the sequences of ‘C. atripons’ and ‘C. pulchristriata’ used by Li
et al. match the C. pulchristriata from the data set of Fritz et al.,
while the sequence of ‘C. dentata’ used by Li et al. match the C.
atripons from the data set of Fritz et al. (Fritz, pers. comm.). The
data of Fritz et al. (2008) are considered taxonomically reliable.
This underlines the pitfalls of sequencing animals with unclear
identification and using uncritical taxonomic identifications of
GenBank data.
46. Malayemys species: Brophy (2004) examined morphometric variation in Malayemys subtrijuga Schlegel and Müller
1845 (sensu lato) across its known range and concluded that turtles
from the Mekong River basin (Laos, Cambodia, and Vietnam)
differed (primarily in color pattern) from those in the Chao Phrya
River basin (Thailand), and resurrected the name macrocephala
(Gray 1859) for Malayemys from the latter basin.
Despite the facts 1) that Schlegel and Müller (1845) reported
the type locality of subtrijuga as “Java”, 2) that at least three
syntypes are known (RMNH 6082, 6084–6085: Hubrecht 1881;
although two others [BMNH 1947.3.4.53 (specimen “m” listed
in Boulenger 1889) and MNHN 7964 may also be among the
original type series]; King and Burke 1989; Iverson 1992), 3)
that field data associated with the RMNH syntypes indicate collection “most probably in the most western province of Bantam
[=Bantan]” (Hubrecht 1881), 4) that numerous specimens of this
taxon from “Java” were available in the 1800s (in at least eight
European Museums; see Specimens Examined in Brophy 2004),
5) that this species has been recorded from at least six localities
on Java (Brophy 2002); and 6) that Brophy’s own morphometric
analysis (2002: Figs. 30–31; 2004: Figs. 6–7) demonstrated that
Javan specimens were distinct from specimens from both the Chao
Phrya and Mekong basins (with the latter two samples actually
overlapping in morphometric hyperspace), Brophy (2004) argued
a) that the type locality of subtrijuga (“Java”) was in error, b)
that Malayemys does not occur nor has it occurred on Java, and
c) that the holotype must have originated from the Mekong basin.
Hence, Brophy (2004) assigned the name subtrijuga to the population of Malayemys inhabiting the Mekong basin (but refrained
from restricting the type locality of subtrijuga, citing uncertainty
about whether the Javan specimens might be native), and the
name macrocephala to the population in the Chao Phrya basin.
Furthermore, Brophy (2004) distinguished these two populations
primarily on color patterns of the head.
More recently, Sumontha et al. (2016) and Ihlow et al (2016)
observed that the head pattern in Malayemys from the Mekong
tributaries of the Khorat Plateau (northeastern Thailand) and adjacent Laos, differed from that of specimens from elsewhere in the
Mekong basin. Based solely on color differences, Sumontha et al.
(2016) described a population from the northern Khorat Plateau
as Malayemys isan. Nearly simultaneously (but see below), Ihlow
et al. (2016) examined color pattern and some morphometrics,
as well as mtDNA and nuclear microsatellite variation across
three Southeast Asian mainland regions of Malayemys distribution (Chao Phraya basin, Mekong basin, and Khorat Plateau),
and concluded that each of these regions has its own distinctive
species, based on each of their data sets, and therefore described
the form that they recorded from three locations on the Khorat
Plateau as M. khoratensis. Two of those locations of M. khoratensis
are effectively sympatric with the M. isan population, and color
patterns in the two taxa are very similar, suggesting that the two
taxa are subjectively synonymous. Although the description of
M. isan by Sumontha et al. appeared online first, their paper did
not comply with ICZN standards for digital publication, whereas
that by Ihlow et al. did (see below); therefore, we conclude that
M. isan is a junior subjective synonym of M. khoratensis.
Unfortunately, neither of these papers mentioned the possibility of taxonomically recognizing the Javan population as a fourth,
distinctive taxon, whose continued occurrence is confirmed by
ongoing exports for commercial trade (UNEP-WCMC 2017). If
further phylogenetic research demonstrates the Javan population
to be taxonomically distinct, the name Emys subtrijuga Schlegel
and Müller 1845 would apply to that population, as would its later
synonyms Cistuda gibbosa Bleeker 1857b, Emys nuchalis Blyth
1863, and Damonia oblonga Gray 1871. Furthermore, Gray 1870c
also described Damonia crassiceps from “China”, the locality
probably erroneous but possibly representing the Mekong River
basin population. Finally, none of these papers have addressed
Schweigger’s 1812 name Emys herrmanni, which is considered by
most authors a nomen dubium, synonymous with M. subtrijuga.
It is problematic that none of these type specimens have
been included in any morphometric or genetic analysis to date.
For example, if a Javan population does or did exist, and it was
found to be distinctive (as Brophy’s morphometric analyses suggested), then the name subtrijuga should be applied there, and
the Mekong population might be identifiable as crassiceps.
In conclusion, until a more comprehensive analysis of coloration and pattern, morphometrics, and both mitochondrial and
nuclear DNA variation among all populations (including Java)
is undertaken, with the inclusion of all of the type specimens
mentioned above, we cannot be certain which name applies precisely to which population of Malayemys. Finally, the TTWG is
not unanimous in its support for the recognition of macrocephala
or khoratensis as full species, and whether or not isan is synonymous with or distinct from khoratensis. However, to minimize
taxonomic changes as we await further data, we here recognize
three species in the genus Malayemys: subtrijuga, macrocephala,
and khoratensis.
47. Malayemys khoratensis: In early 2016 a distinctive
clade of Snail-eating turtles (Malayemys) from the Khorat Plateau of eastern Thailand was described as a new species with
two different names in two separate articles (Ihlow et al. 2016;
Sumontha et al. 2016). The name M. khoratensis Ihlow et al.
2016 was published on 6 April 2016, the date when the electronic
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
version of the article, published in the journal PLoS One, met the
ICZN Code for nomenclatural availability of electronic articles
under article 8.5.3 (ICZN 2012). The name M. isan Sumontha et
al. 2016 was published electronically on 26 March 2016 in the
journal Taprobanica; however, the Zoobank registration of this
article failed to meet the requirements of ICZN article 8.5.3.1,
rendering the electronic version unavailable for the purposes
of nomenclature. It must instead be considered as published
when it first met the requirements of ICZN article 8.4.1, i.e.,
when numerous identical hard copies printed on paper became
available. There is no evidence of the journal Taprobanica 8(1)
having been printed prior to 13 April 2016, when copies were
requested and were simultaneously sent to the archiving libraries
as listed on the journal’s website (Thomson and Lambertz, in
press). Therefore, the date of publication of the journal Taprobanica 8(1) and the contained article by Sumontha et al. (2016)
is to be corrected to 13 April under ICZN article 21.4, the first
date for which there is evidence of its physical existence. As
a result, the name Malayemys khoratensis has nomenclatural
priority, and Malayemys isan becomes a junior subjective synonym (see above annotation).
48. Damonia crassiceps Gray 1870c: This taxon has been
listed under the synonymy of Malayemys subtrijuga by most
authors since Smith (1931). However, Sumontha et al. (2016)
questioned its identification as representing either this species or
genus, based on the description of the drawing of the species as
lacking facial stripes, but did not suggest an alternative taxonomy.
49. Mauremys annamensis and M. mutica: Zhao et al.
(2016a) sequenced the entire mitochondrial genomes of three
specimens of Mauremys mutica without published locality data,
but purported to originate from Taiwan, China, and Vietnam or
Hainan. Phylogenetic analysis revealed that the latter genome
was more similar to that of M. annamensis than to the other more
northerly mutica genomes, rendering M. mutica paraphyletic.
Zhao et al. (2016b) examined variation in the mitochondrial
barcode gene COI among larger sample sizes of M. annamensis
and M. mutica. They identified reciprocally monophyletic northern (including Taiwan) and southern (Vietnam, Hainan and M.
annamensis) clades, but the latter showed no monophyly among
the constitutent populations. They speculated that M. annamensis might be of hybrid origin. Furthermore, Zhou et al. (2015)
compared the entire mitochondrial genome of samples of eight
species of Mauremys, and their results also nested annamensis
within mutica, sister to mutica from southern China (near Vietnam). Unfortunately, they included no material from Vietnam, and
most of their samples were purchased from the pet trade or food
markets. More extensive geographic sampling (including M. m.
kami from the Ryukyus) and nuclear analysis will be necessary
before any taxonomic changes are warranted.
Independently, Somerová et al. (2015) examined variation in a mitochondrial gene and a nuclear intron in European
zoo specimens of M. annamensis, and also resolved M. mutica
as paraphyletic with respect to annamensis. In addition, their
analysis also revealed two distinct, reciprocally monophyletic
mitochondrial clades within annamensis, which they recommended be maintained separately in captive breeding operations.
Unfortunately, the natural geographic provenance of these two
clades was unknown.
50. Mauremys caspica and Mauremys rivulata: Using microsatellite loci and nuclear and mitochondrial DNA sequences,
Vamberger et al. (2017a) showed that the two species hybridize
only rarely along their contact zone in Turkey. However, there
is evidence for introgression between both species. In addition,
they found hybrid evidence on Cyprus and, unexpectedly, terrapin populations in Israel and Jordan turned out to be a hybrid
swarm of the two species, morphologically resembling M. rivulata. Ecological paleomodeling suggested that the two species
formerly had temporary contact across what is now the Syrian
Desert during more humid climatic episodes.
51. Mauremys leprosa: An analysis of variation in two
mitochondrial markers and one nuclear gene across the range
of Mauremys leprosa by Veríssimo et al. (2016) confirmed two
distinct genetic lineages that generally correspond to the two currently recognized subspecies. One is distributed from southern
France through Iberia to Morocco, north of the Atlas Mountains
(M. leprosa leprosa); the other occurs from northern Libya and
Tunisia westward to Morocco, both north and south of the Atlas
mountains (M. leprosa saharica); in northern Morocco there is
secondary contact between these subspecies. In northeastern Iberia
and southern France, some native populations have been genetically
impacted by introduced M. l. saharica (Palacios et al. 2015).
52. Mauremys reevesii: The historic native distribution of
this species has been difficult to determine accurately, as the
species has been traded extensively in China for several thousand years, and has apparently been introduced to Taiwan and
Japan in historic times (see the genetic analysis by Suzuki et al.
2011). The populations in Korea may also have been introduced
prehistorically, but could represent natural distribution during an
interglacial warming period. Our previous distribution map in the
last checklist was based on Iverson (1992) and the EmySystem
database plus input from the CBFTT account by Lovich et al.
(2011). We have now updated and revised this map extensively
by re-evaluating most localities and restricting the distribution to
areas below approximately 600 m elevation. We have also added
more localities provided by T. Blanck (based on compilations of
Chinese survey literature and field data) and D. Gaillard. While
generally a lowland species, some populations (e.g., in Anhui)
appear to occur up to elevations of ca. 500 m in hill regions, but
in more southern regions (e.g., Hunan and Jiangxi) the species
appears to be limited to lowland areas below 300 m (T. Blanck,
unpubl. data). Most northern Chinese localities for M. reevesii
appear to be trade specimens from ports and market centers (as
also concluded by Pope 1935:46). Many southern and eastern
coastal specimens are also trade specimens from coastal ports
and markets or possibly locally invasive from markets. However,
ancient Chinese writings seem to indicate that this species apparently occurred along the south coast, at least in Guangdong (T.
Blanck, unpubl. data). Native wild populations do occur in the
Chengdu basin of the upper Yangtze in Sichuan and all along the
central Yangtze lowland basin, notably in Anhui, Guizhao, Hubei,
Hunan, and Jiangxi. There also appears to be a native population
in the Wei Valley of Shaanxi. Other scattered records north of the
Yangtze may or may not represent native populations. Further
genetic studies of Korean and southern and eastern coastal specimens with comparisons to specimens from the central Yangtze
and Sichuan portions of the range are clearly needed.
Oh et al. (2017) examined geographic variation in mitochondrial cyt b sequences across the range of M. reevesii. Their
results were complicated by the translocation of this species by
humans since prehistoric times. They suggested that the two
main natural populations in China and Korea were weakly but
distinctly divergent genetically; however, introgression as a result of translocations is diluting that difference, precluding any
taxonomic recognition.
53. Mauremys rivulata: Vamberger et al. (2014) examined variation in a mitochondrial gene fragment and across 13
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microsatellite loci for samples from throughout the range of Mauremys rivulata. The mitochondrial sequence data showed no evident
structuring. However, the microsatellite data revealed a distinct
genetic break in southern Turkey, separating eastern and western
populations. This break is likely to be caused by introgressed
alleles from M. caspica in the eastern portion of the distribution
range of M. rivulata (Vamberger et al 2017a). Vamberger et al.
(2014) attributed the lack of clear substructure among the western
populations of M. rivulata to trans-Mediterranean dispersal.
54. Melanochelys trijuga parkeri: This taxon was described
as a large-bodied, low-shelled subspecies endemic to Sri Lanka,
but recent surveys at all historically recorded localities have
encountered mainly the more common and widespread smaller
Sri Lankan subspecies M. t. thermalis (A. de Silva, unpubl. data).
These two taxa need genetic evaluation to determine if and how
they may be distinct, and if distinct, whether they are undergoing
introgression and intergradation.
55. Aldabrachelys and Cylindraspis: Based on contemporary
Indian Ocean currents, and historic fluctuations in sea levels, Wilmé
et al. (2017) proposed that insular Indian Ocean populations of
tortoises were the result of introductions by humans approximately
4000 ybp. However, given that previous work dates the colonization of these islands by tortoises at 9.5–22 mybp, and fossils of
Aldabrachelys have been dated to at least 138,000 ybp (Cheke
and Hume 2008; Gerlach and Paquette 2014; Cheke et al. 2017;
Hansen et al. 2017; Hawlitschek et al. 2017), this hypothesis is
untenable.
56. Aldabrachelys gigantea and Testudo: Besnard et al.
(2016) used shotgun sequencing to elucidate the entire mitochondrial genome of Aldabrachelys gigantea. Phylogenetic
analysis of the aligned sequence of this tortoise with the available
mitochondrial genomes of twelve other tortoise species generally
supported the results of Le et al. (2006), except for the placement
of Malacochersus and Testudo horsfieldii. However, the analysis
by Besnard et al. suggested a paraphyletic genus Testudo, although
with reduced bootstrap support (77–78%). Synthesis of these data
with nuclear markers is needed before any taxonomic changes
are warranted.
57. Aldabrachelys gigantea: Turnbull et al. (2015) examined variation in body size and sexual size dimorphism among
four subpopulations of Aldabra tortoises. They speculated that
these differences might have a genetic basis, and recommended
population genetic studies.
58. Chelonoidis: The gender of the tortoise genus name Chelonoidis Fitzinger (1835) has long been assumed to be feminine
(e.g., Agassiz 1857) because its root is the feminine Greek noun
Chelone (not Chelonos as assumed by Olson and David [2014]),
and the Latinized suffix -oides (from the Greek -eides, meaning
form or shape [or “like” in English]). However, according to the
ICZN (Article 30.1.4.4) genus names ending in -oides must be
considered masculine unless the original author indicated gender
directly or indirectly (i.e., by usage). Because Fitzinger gave no
direct indication or orthography indicating that he considered the
name Chelonoidis to be feminine (or neuter), Olson and David
(2014) argued that it should be considered masculine. However,
Fitzinger did provide indirect evidence that he considered the genus name masculine, by his clear rendition of other reptile genera
he described as masculine (e.g., Dracontoidis and Elapoidis). In
either case, the conclusion is the same—that a strict application
of the Code would render Chelonoidis masculine, and require
the emendation of the suffix of many of the species currently
recognized within that genus. The name vicina is treated as a
noun in apposition (Art. 31.2.2) and thus is unaffected by the
gender of the genus name. While the conversions to carbonarius,
denticulatus, niger, and phantasticus imply disruption to a few
well-established and widely-used names, they should not lead to
any significant confusion, and in the interest of Code compliance,
we adopt these changed endings.
59. Chelonoidis chilensis: Sánchez et al. (2015) examined
variation in the karyotype of Chelonoidis chilensis across its range.
They identified two karyomorphs, one from tortoises in the Dry
Chaco Ecoregion, and one from the Monte of Steppes and Plains
Ecoregion. However, these karyomorphs were independent of the
external morphotypes of donosobarrosi, petersi, or chilensis, and
therefore they followed Fritz et al. (2012a) in recognizing only a
single species (C. chilensis) in this complex.
60. Chelonoidis niger complex: Molecular studies of Galápagos tortoises have surged over the last several years, although
most of the work has been more directed at population genetics
than establishing species boundaries and their taxonomic implications. The general working assumption is that separate island
populations previously recognized as subspecies of Chelonoidis
niger [= C. nigra] (or synonyms thereof) are now accepted as
species (e.g., Caccone et al. 2002; Russello et al. 2005, 2007;
Poulakakis et al. 2008, 2012, 2015; Chiari et al. 2009; TTWG
2009 [annotation 09:32], 2014; Edwards et al. 2014; Garrick et
al. 2014), and we continue to recognize them as such, now also
resurrecting the three previously synonymized southern Isabela
taxa (guntheri, microphyes, and vandenburghi) from their synonymy under C. vicina. However, Loire et al. (2013) examined
the population genomics of five Galápagos tortoise taxa (purportedly representing three named island populations) based on
248 nuclear genes. Their results suggested panmixis across their
samples, with little genetic differentiation, and they questioned
the species-level recognition of the various island taxa. Expansion
of this study to include Galápagos-wide sampling would clearly
be helpful in settling some of the taxonomic issues facing these
tortoises.
Edwards et al. (2014) examined 14 microsatellite loci for
tortoises from southern Isabela from the ranges of C. vicina and
C. guntheri (the latter was synonymized with the former based
on mtDNA data in Poulakakis et al. 2012). Not only did they find
support for the distinction of those two taxa, but they also identified a third, unnamed, geographically intermediate genetic cluster
they referred to as the “aplastados” [flattened] type. Although
they made no taxonomic recommendations, and an expansion of
this study to include other Isabela populations is needed, these
data warrant our removal of C. guntheri from the synonymy of
G. vicina, with full species recognition. Simultaneously, based on
12 microsatellite loci and mtDNA sequence data, Garrick et al.
(2014; see also Emerson and Faria 2014) identified two distinct
but coalescing genetic lineages within C. becki on northern Isabela
that appear to represent two different colonizations from Santiago
(C. darwini). They also found some evidence of introgression of
C. vandenburghi into the genome of C. becki. Hence, they concluded that “species boundaries in the group may be somewhat
porous,” and questioned the species-level status of C. darwini
relative to C. becki.
Clearly, the evolutionary history of colonizations and divergences by tortoises in the Galápagos has been very complex,
and much more reticulate than previously realized. Sorting out
this history is an on-going challenge. However, as we noted
in a previous checklist (TTWG annotation 12:31), genotyping
of the many type specimens of named Galápagos tortoises is
desperately needed, so that the correct names may be applied
to all genetic lineages. Until those data are available, confusion
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
and nomenclatural uncertainty will no doubt continue, such as
some authors’ use of elephantopus instead of niger or nigra (see
TTWG annotation 09:33), and ephippium instead of duncanensis
(e.g., Poulakakis et al. 2012; Jensen et al. 2015b; Hennessy
2015).
61. Chelonoidis donfaustoi: Chiari et al. (2009) and Poulakakis et al. (2015) analyzed mitochondrial DNA and nuclear
microsatellite characters in the two giant tortoise populations
on Santa Cruz (Reserva and Cerro Fatal). Both studies found
the populations to be genetically different and Poulakakis et al.
(2015) demonstrated that the Reserva population represented the
previously described C. porteri, and that the Cerro Fatal population was distinct and most similar to C. chathamensis from San
Cristóbal. Despite minimal morphological differences from C.
porteri, and that the holotype of C. porteri is a hybrid with Reserva
nuclear DNA and Cerro Fatal mtDNA, they described the Cerro
Fatal population as a new species, and we tentatively agree.
62. Chelonoidis niger: Olson (2015) investigated the history
of some early names applied to Galapagos tortoises. He concluded
that the provenance of the single type specimen (MNHNP 9550)
of both Testudo californiana Quoy and Gaimard 1824a and
Testudo niger Quoy and Gaimard 1824b (= Chelonoidis niger)
was “extremely unlikely” to be determined based on historical
information, and hence the names should be considered nomina
dubia and their use abandoned. Olson also concluded that one of
the two syntypes of Testudo nigrita Duméril and Bibron 1835,
designated the lectotype by Günther (1875a:268), was apparently
lost, and since the paralectotype (MNHNP 9313) is a juvenile,
he also recommended treating that name as a nomen dubium.
In addition, Olson and Humphrey (2017) investigated the
origin of the type specimen of Testudo elephantopus Harlan
1827, and concluded that it may have come from Charles Island
(Floreana), and suggested that the name elephantopus therefore
replace the name Testudo niger Quoy and Gaimard 1824b, currently used for that island’s species.
However, genetic analysis of the types currently in progress
will hopefully demonstrate their geographic origins satisfactorily,
and the allegedly lost lectotype of nigrita (BMNH 1949.1.4.37)
actually remains extant at the British Museum (P. Campbell, in
litt., 2017). We therefore consider all these recommendations by
Olson (2015) and Olson and Humphrey (2017) as premature and
unnecessary at this time.
63. Chelonoidis phantasticus: In our previous checklists we
indicated that this species was Extinct, based on that supposition
by Pritchard (1996), but its formal conservation status has not
previously been assessed using IUCN Red List criteria (IUCN
2001). This has now been done by the TFTSG using updated Red
Listing guidelines in regard to determining whether a species
is actually Extinct or not (IUCN 2016). Although only a single
individual of this species has ever been collected (in 1906),
sightings and signs from 1964, 2009, and 2013 suggest that a
few individual tortoises may remain extant in the exceedingly
fragmented and hard-to-access landscape of Fernandina, most
of which is covered by uninhabitable lava flows (A. Rhodin, L.
Cayot, J. Gibbs, R. Kiester, and W. Tapia, in press). As such, the
TFTSG now determines this species to be Critically Endangered
(Possibly Extinct) and has submitted that assessment to the IUCN
Red List for publication.
64. Chersobius, Homopus, and Psammobates: Based on
three mitochondrial and two nuclear genes, Hofmeyr et al. (2016)
resolved a paraphyletic Homopus with respect to Chersina, and
resurrected the genus Chersobius for the five-toed species (signatus, boulengeri, and solus) formerly in the genus Homopus, and
restricted the genus Homopus to the four-toed species (areolatus
and femoralis). We have tentatively accepted those changes here.
The authors also noted strong support (Hofmeyr and Daniels,
in prep.) for deep genetic divergence among the currently recognized subspecies of Psammobates tentorius, suggesting that
they deserved species status. They also indicated the presence of
phylogeographic structure within Chersina angulata and Homopus
signatus, suggesting possible future taxonomic changes in those
taxa. We await these forthcoming publications before making
any further changes to the checklist.
65. Geochelone elegans: Schweigger (1812:325) described
Testudo stellata based on the same specimens and plate previously described and figured by Schoepff (1795:111) as Testudo
elegans. However, Schweigger coined the new name stellata as
a nomen novum because Schoepff had based his description of
elegans partly on a Seba (1734) specimen (pl. 79, fig. 3) that
Schweigger (1812:325) concluded had been incorrectly identified,
and he instead synonymized that figure under Testudo rotunda
Latreille. Schweigger therefore coined a new name for Schoepff’s
species, renaming it stellata and stating: “Habitat in India orientali. (Schoepf. sub falso nomine: test. elegans Seb.)”. However,
the description of T. elegans by Schoepff was nomenclaturally
available, and therefore T. stellata Schweigger has the same type
specimens as T. elegans, according to Article 72(e) of the ICZN
Code, and thus becomes its objective junior synonym.
66. Date of Publication of Blyth 1854 [“1853”]: The date
of publication of this article in issue number 7 of volume 22 of the
Journal of the Asiatic Society of Bengal has historically been listed
as 1853, the printed date on its page-headers. However, a note in
the Proceedings published in the same issue 7 on p. 684 is dated 4
January 1854, indicating that the issue was actually not published
until 1854, consistent also with the imprint date of 1854 on the bound
volume of all 7 issues of volume 22. This change of date affects
the following taxa: Testudo phayrei [= Manouria emys phayrei],
Testudo elongata [= Indotestudo elongata], Testudo megalopus [=
Geochelone elegans], Testudo iberia [= Testudo graeca ibera], and
Homopus burnesii [= Testudo horsfieldii horsfieldii].
67. Gopherus: In the recent book on the biology of North
American tortoises (edited by Rostal et al. 2014), Bramble and
Hutchison (2014) reviewed and augmented the previous morphological data for the five known living species at that time, as
well as the fossil forms, and concluded that the available evidence
(including molecular data) supported the reciprocal monophyly
of Xerobates, including agassizii, berlandieri, and morafkai, and
Gopherus (sensu stricto), including flavomarginatus and polyphemus. They recommended recognition of Xerobates and Gopherus
as sister genera. In the same volume, Murphy (2014) provided a
critical review of the systematics of the broad genus Gopherus
and its members, and also concluded that flavomarginatus and
polyphemus are sister taxa, and that berlandieri is sister to the
agassizii group (including morafkai; see also Reid et al. 2011, not
cited in Rostal et al. 2014). However, he made no recommendation about the use of Xerobates, and included all species in the
broader genus Gopherus, as most authors since Crumly (1994)
have done. Although most of the other authors of chapters in this
book used the genus Gopherus (sensu lato), in his review of fossils, Franz (2014) recognized Xerobates and Gopherus. As noted
by Murphy (2014), broad nuclear sampling is needed to resolve
definitively the relationships within this species group. Until such
time, and considering that the name Gopherus could still apply
to all currently included species even if the two identified clades
are reciprocally monophyletic, we retain Gopherus as the sole
genus name.
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68. Gopherus evgoodei: The genetic distinctiveness of
the southern Sonoran / northern Sinaloan population of Desert
Tortoises has long been recognized (reviews in Edwards et al.
2015a,b, 2016), and has now been formally described as a new
species by Edwards et al. (2016).
69. Indotestudo forstenii: The type specimen of I. forstenii
was recorded by Schlegel and Müller (1845) as having been collected by Forsten on “Gilolo, Indischen Archipel” (now Halmahera,
Indonesia). However, no further specimens of I. forstenii have
ever been recorded or found anywhere on that island since that
time (D. Iskandar, A. Riyanto, pers. comm.), and the species does
not appear to occur on Halmahera or its adjacent islands. The
collector, Eltio Alegondas Forsten, spent a few months in 1841
collecting botanical and zoological specimens on Ternate island
just off the west coast of Halmahera, where his collectors also
obtained some material from mainland Halmahera. However, he
also made extensive collections later in 1841–1842 on northern
Sulawesi, while headquartered at Gorontalo (http://www.nationaalherbarium.nl/FMCollectors/F/ForstenEA.htm), where I.
forstenii does occur. We hypothesize that the type specimen was
therefore probably obtained by Forsten near Gorontalo, Sulawesi,
Indonesia, and that it was then mislabeled as to its exact origin,
or possibly that it was acquired from Ternate or Halmahera as
a regionally traded specimen originating from Sulawesi. We no
longer consider Halmahera to be part of the natural range of I.
forstenii, and hereby formally restrict its type locality to “near
Gorontalo, Sulawesi, Indonesia”.
70. Testudo: Based on a phylogenetic analysis of morphological traits for twelve fossil and the five extant species in the
genus Testudo (sensu lato), Luján et al. (2016) recognized three
monophyletic subgenera: Chersine for hermanni, Agrionemys for
horsfieldii, and Testudo for graeca, marginata, and kleinmanni. Additionally, Vasilyev et al. (2014) studied variation in the 12S rRNA
mitochondrial gene and three RAPD markers among individuals
of a number of species of Testudo (sensu lato). Although graeca,
marginata, kleinmanni (including synonymous werneri), and
horsfieldii were resolved as monophyletic in the mtDNA analysis,
hermanni was not; however, the RAPD analysis resolved each of
those taxa as monophyletic. Support for the recognition of three
monophyletic clades (corresponding to Chersine, Agrionemys,
and Testudo [sensu stricto]) was ambiguous, with no support from
the mtDNA, and only weak support from the RAPD data. Based
on the combination of morphology of extant and fossil species
and acknowledging the results of molecular genetics studies of
Testudo by others, Luján et al. (2016) recommended that Chersine
and Agrionemys (as well as the fossil Paleotestudo) be recognized as valid subgenera of Testudo (sensu lato). We now adopt
that recommendation here, discontinuing our previous listing of
alternative generic designations.
71. Testudo (Testudo) graeca: In a paper previously not
recorded in our checklist, Türkozan et al. (2010) analyzed morphometric variation in a large sampling of Testudo graeca from
throughout Turkey. They determined that the putative subspecies
anamurensis and antakyensis are not distinct from each other or
from terrestris (and therefore synonymized under terrestris),
and that armeniaca and perses are distinct. Further, they noted
that the populations from northern and southern Turkey, possibly corresponding to the ibera and terrestris mtDNA clades,
respectively, also appeared to be morphometrically distinct. They
made no taxonomic recommendations other than maintaining a
conservative approach pending additional morphologic and genetic assessment of more material from both within and outside
Turkey. Our checklist has reflected these conclusions for many
years, although we have listed perses as a synonym under the
distinct taxon buxtoni since 2007, a placement reconfirmed by
Parham et al. (2012) (see TTWG annotations 07:73 and 12:36).
Our current updated map with its subspecies delineations
has benefitted from review and input by Oguz Türkozan and
Peter Mikulíček. Notably, unpublished mtDNA genetic work in
progress by Türkozan and colleagues now appears to confirm the
earlier impression that ibera and terrestris represent well-separated
northern and southern populations of T. graeca. Their distribution patterns reflect clear separation by the so-called Anatolian
Diagonal, with only a few areas of intergradation in southwestern
Turkey. Additionally, buxtoni appears to intergrade slightly with
terrestris in the southeastern corner of Turkey.
Using the cyt b gene, Javanbakht et al. (2017) examined the
phylogeography of the subspecies of T. graeca in Iran and Transcaucasia and refined the knowledge of their distribution. Based
on species distribution models, they showed that the distribution
ranges changed little since the Last Glacial Maximum.
Using three mitochondrial DNA fragments (cyt b, 12S, ND4
plus adjacent DNA coding for tRNAs), Graciá et al. (2017a)
studied differentiation of the Testudo graeca complex. According
to fossil-calibrated molecular clock calculations, they inferred
a dual diversification burst. The eastern subspecies, including
the last common ancestor of the North African taxa, radiated in
the Mio-Pliocene, whereas a second radiation in North Africa
took place during the Pleistocene. Based on a Libyan tortoise
of unknown exact provenance, a new North African lineage was
discovered. The recent introduction of most Western European
populations was confirmed, with the exception of populations in
southeastern Spain, which are older.
72. Testudo (Testudo) graeca and Testudo floweri: Meiri
et al. (2011) examined body size variation in Testudo graeca
across the Levant, and demonstrated that these tortoises follow
Bergmann’s Rule, with the smallest tortoises found at the southern
end of the range in the Negev Desert. The small Negev tortoises
had earlier been decribed as T. floweri by Bodenheimer (1935),
and these authors agreed that this taxon is synonymous with T.
graeca. Werner et al. (2016) then expanded that analysis to include
all T. graeca (sensu lato; see TTWG 2014) from Morocco to Iran.
They also found a general correlation of body size with latitude.
However, their analyses of sexual size dimorphism demonstrated
that it seems to scale differently among tortoises in Anatolia versus
the Levant, and concluded that this might suggest that those two
populations might deserve consideration as “separate entities”
(presumably with taxonomic consequences).
73. Testudo (Testudo) kleinmanni: Based on morphological
differences between Testudo kleinmanni on opposites sides of the
Nile River, Perälä (2001) described the eastern population as T.
werneri. Subsequent studies (see TTWG 2007, annotation 07:74)
did not support this distinction and werneri was synonymized
with kleinmanni. However, Werner (2016) has re-emphasized the
apparent morphological differences between the populations and
has claimed that some tortoises from west of the Nile have been
translocated to the east to northern Sinai, potentially confounding
the recent studies. Therefore, he suggested that werneri should
be recognized as a distinct subspecies. Until further genetic work
tests this scenario, we continue to retain werneri in synonymy.
74. Testudo (Chersine) hermanni: Based on an analysis of
17 nuclear microsatellite loci and broad geographic sampling,
Zenboudjii et al. (2016) identified three major genetic clusters
within Testudo hermanni hermanni: one continental (Spain,
France and Italy), one insular (Corsica, Sardinia, and Sicily),
and one an island supporting two clusters (Menorca). There is
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
still debate as to which populations are autochthonous and which
are the result of human introductions (or reintroductions). Perez
et al. (2014; see also TTWG 2014 comment 07:36), based on
mtDNA and microsatellites, concluded that the Spanish, Corsican, and Sardinian populations were likely introduced from
Sicily by humans. However, although Zenboudji et al. (2016)
agreed that the contemporary population on Sardinia is the result
of introductions from Sicily following an earlier extirpation on
Sardinia, they argued that the Corsican population is natural and
genetically distinct. They also concluded that the population in
northeastern Spain (Albera) is also a relict, natural population.
Menorca supports two distinct genetic populations. The one in
the west is closely related to mainland populations, and hence is
likely an introduction from a now extirpated mainland source.
The eastern population is more distinct, but more closely related
to the other insular populations, but whether this represents an
ancient autochthonous divergence or an ancient human introduction of a lineage of unknown provenance could not be determined.
Zenboudji et al. (2016) also noted a number of examples of mismatched genotypes in several populations, representing introduced
individuals (and their offspring) from other populations. Finally,
they recommended that six populations be considered genetic
management units for conservation purposes: 1) Albera (Spain) +
western Menorca; 2) France (Var); 3) Italy; 4) Sardinia + Sicily;
5) Corsica; and 6) eastern Menorca.
75. Cyclanorbis senegalensis: Based on mitochondrial DNA
sequence data, Mazuch et al. (2016) reported the presence of
C. senegalensis in western Ethiopia for the first time, and also
found significant genetic divergence between western (Togo
and Benin) and eastern (Ethiopia) populations. They called
for further phylogeographic study, but made no taxonomic
recommendations.
76. Amyda cartilaginea or A. ornata: Fritz et al. (2104a)
examined variation in mitochondrial and nuclear DNA and color
patterns across the range of Amyda cartilaginea (sensu lato). Their
mtDNA tree was not highly resolved; however, because of the concordance of the identified clades with color pattern and geography
(paleodrainages), the authors recognized two named species with
four named subspecies within what was previously recognized as
A. cartilaginea. They restricted A. cartilaginea (sensu stricto) to the
southern portion of the Greater Sundas, with the nominate subspecies
in southern Borneo and Java, and a new subspecies, maculosa, in
the northern portion of the Greater Sundas (northwestern Borneo
and southern Sumatra). Populations of A. cartilaginea (sensu lato)
in northern Sumatra and Peninsular Malaysia were not assessed or
assigned to subspecies. They also identified, but did not describe, a
potential candidate species from northeastern Borneo. They resurrected the older name ornata (Gray 1861) for the Southeast Asian
mainland species, with the nominate subspecies confined to the
Mekong basin, and resurrected the name phayrei (Theobald 1868)
for the subspecies in Thailand and Myanmar. They also identified,
but did not describe, a third possible subspecies from Bangladesh.
Regrettably, no specimens from southern Myanmar, southern peninsular Thailand, or Peninsular Malaysia were included. In view
of the relatively weak resolution of their named mtDNA clades,
but reasonable concordance of those clades with paleogeographic
drainage basins and color patterns, we list their proposed taxonomic
changes provisionally, pending additional geographic sampling and
further genetic and morphologic work.
77. Apalone mutica calvata: In a guide to Alabama turtles,
Guyer et al. (2015) elevated Apalone mutica calvata to a full
species based on its distinctive color pattern, its geographic
isolation, and the mtDNA data (part of only one gene) presented
by Weisrock and Janzen (2000). Powell et al. (2016) followed
their recommendation. However, because of the small sample
size and small DNA fragment available to Weisrock and Janzen
(2000) and incomplete sampling in the presumed area of overlap
in southeastern Louisiana, we consider this change premature,
and continue to recognize calvata as a subspecies of A. mutica.
78. Nilssonia and N. nigricans: Based on mitochondrial Cytochrome C Oxidase Subunit I (mtCOI) barcode sequences, Kundu
et al. (2016) provided a phylogenetic analysis of the Asian softshell
turtles allied with the genus Nilssonia. Their results confirmed the
presence of N. nigricans in the wild in Assam, India, as well as
provided additional support for the synonymy of Aspideretes with
Nilssonia (see also TTWG 2011, annotation 11:15).
79. Pelochelys cantorii: Hoser (2014a) coined a new subgeneric and two new species names for two purported varieties
of Pelochelys cantorii. Since the ICZN has been petitioned by
Rhodin et al. (2015) (see Annotation 4) to declare these and
other Hoser names unavailable under the Code (Article 82.1),
we maintain prevailing usage and do not recognize these names.
80. Pelodiscus sinensis and P. maackii: Suzuki and Hikida
(2014) examined mitochondrial cytochrome b sequence data from
specimens of Pelodiscus across Japan and compared them with
previously published data from Fritz et al. (2010b) from across
the wider species range. They identified two lineages from Japan, corresponding to the previously recognized P. sinensis and
P. maackii. The latter was widely distributed in Japan, whereas
the former had only a sporadic distribution, leading the authors
to conclude that maackii was native to Japan and sinensis was
introduced. They also noted that if future nuclear DNA data
supported this scenario, the older name japonicus (Temminck
and Schlegel 1838) could be available for the lineage currently
recognized as maackii (Brandt 1857).
81. Pelodiscus sinensis: The publication date of F.J.F. Meyen’s
Reise um die Erde, in which Trionyx (Aspidonectes) sinensis (=
Pelodiscus sinensis) was first described by A.F.A. Wiegmann, has
been a source of controversy. Although frequently cited as appearing in 1835, Bauer and Adler (2001) in a previously overlooked
publication determined that the description of sinensis first appeared
in print in 1834, and we have now corrected that date here.
82. Pelodiscus sinensis: As clarified by Adler (2016), the
name Trionyx tuberculatus was first used by Cantor (1842a), but
without a description or indication, and thus a nomen nudum;
then subsequently formally and validly described a few months
later (Cantor 1842b).
83. Rafetus euphraticus: We note that in the original description of Testudo euphratica published by Daudin (1801) that he
specifically credited Olivier with providing him with the description of the new species. We therefore amend the authorship of
the original name to Testudo euphratica Olivier in Daudin 1801.
84. Rafetus euphraticus: Ihlow et al. (2014) examined variation
in two mitochondrial genome fragments across nearly the entire
range of the species, and found no significant genetic variation.
85. Rafetus swinhoei: Le et al. (2014) analyzed sequence
data from two mitochondrial loci and one nuclear gene from all
known populations of Rafetus swinhoei in China and Vietnam.
Their results demonstrated minimal divergence among populations,
and warranted no taxonomic changes. Despite the long history in
China of human use and transport of turtles, the authors cautioned
against speculation that the Chinese distribution is unnatural.
86. Chelidae: A few recent authors have followed Storr (1978)
in using the family name Cheluidae (Michael and Lindermeyer
2010; Wilson and Swann 2013), a grammatically correct derivation from the Greek type genus Chelus, genitive singular Cheluos.
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However, the spelling Chelidae has been in prevailing usage globally
since Lindholm (1929) and most authors continue to use it. Under
ICZN Article 29.5, the original spelling of a family-level name is
to be maintained when it is in prevailing usage, whether or not its
derivation from the name of the type genus is in accordance with
grammatical procedures. We therefore regard the name Cheluidae
as an unjustified emendation, given the long-standing prevalence of
Chelidae in the literature, and we strongly recommend and concur
with the continued usage of Chelidae.
87. Acanthochelys radiolata: Garbin et al. (2016) examined
morphological variation in Acanthochelys radiolata across its
range. Although they described significant variation in this species
(especially in color and shape), they could identify no geographic
pattern to this variation and concluded that A. radiolata is a single,
highly variable species. In addition, they considered records from
Mato Grosso and Sao Paulo States to be in error.
88. Chelus fimbriata: Zug (1977) clarified that the genus
name for the matamata is Chelus, not Chelys, as originally
proposed by Dumeril (1805:76). Most authors since then have
treated the generic name as feminine, rendering the species name
as Chelus fimbriata. However, Ferreira et al. (2016), following
the recommendation of Vlachos (cited as a pers. comm., but also
discussed in Vlachos 2015), argued that the genus name Chelus
is actually masculine (and the species name therefore Chelus
fimbriatus) because it was supposedly based on Latinization of
the classical Greek feminine word χέλυς, and according to ICZN
(1999) Art. 30.1.3, a Latinized suffix should take the gender of the
Latin suffix. However, according to Appendix B of the 1961 and
1985 ICZN Codes, as well as Art. 30.1.2 and 30.1.3 of the 1999
Code, there is a distinction between transcription or transliteration vs. Latinization of a Greek word. Specifically, for the Greek
letter upsilon (υ), its Latin equivalent (i.e., transliteration) is ‘u’,
whereas the Latinized version is ‘y’ (ICZN 1985, App. B). Hence,
the name Chelus is considered a transliteration, not a Latinization,
of χέλυς, whereas Chelys is considered a Latinization. Unfortunately, Vlachos (2015) transposed the terms “transliteration”
and “Latinization” in his Table 1—he rightly used “Latinized
form” to name his column L (where he gives “y” as the Latinized
form of upsilon), but used “transliteration” for it in the caption,
causing considerable confusion. Chelus and Chelys should not
be considered homonyms (1999: Art. 56.2; 1985: Art. 56.b), and
in accordance with ICZN 1999 Art. 30.1.2 (1985: Art. 30a), the
gender of both Chelus and Chelys is feminine. The correct species name is therefore Chelus fimbriata, as it has been rendered
for several decades, and we continue to use it as such.
89. Mesoclemmys raniceps: Rivas et al. (2015) discussed the
alleged distribution of this species in Venezuela, concluding that
of the three specimens recorded from there, two were misidentified, and one (the type specimen of Hydraspis maculata), was
misidentified and incorrectly restricted to Venezuela. Hence, we
delete Venezuela from the range of M. raniceps, and synonymize
H. maculata under M. raniceps instead of under M. nasutus where
it was previously placed.
90. Phrynops geoffroanus and P. tuberosus: In a previous
checklist (TTWG 2010, Annotation 10:44), we noted that the
distribution and taxonomy of these two taxa remained highly
problematic and subject to differing opinions as to their extent
and inter-relationships, and in our last checklist (TTWG 2014)
we provided coarse and uncritical distribution maps for them. In
this checklist we now present new revised distributional maps
that document most known localities of these taxa, sourced from
a combination of the EmySystem database (with several corrections), recently published literature, and previous research on
many collected specimens by Rhodin and Mittermeier during their
work on the P. geoffroanus complex (Rhodin and Mittermeier
1983; unpubl. data). Morphological analysis at that time indicated
the presence of several apparently allopatric and differentiated
populations of P. geoffroanus corresponding to various level 3
and 4 hydroshed basins, with several areas of close parapatry
and possible intergradation. We now depict these distributional
patterns in our revised maps, showing the distributional limits of
the two taxa to delineate the hydroshed-restricted extent of their
mostly separate populations, and showing the various subpopulations of P. geoffroanus. Many of these differentiated populations
have now also been preliminarily substantiated as genetically
distinct lineages by Carvalho et al. (2016). We anticipate that
the P. geoffroanus complex will eventually be recognized as a
polytypic species complex with several distinct lineages, notably
the southeastern Brazilian coastal, the Rio São Francisco basin,
the Rio Paraná basin, the lower Amazon basin, the Colombian,
and the Peruvian-Bolivian populations.
91. Platemys platycephala melanonota: Mendes-Pinto et
al. (2011) reported the collection of a Platemys platycephala in
southwestern Pará, Brazil, that was diagnosable as the subspecies P. p. melanonota. This record lies nearly 2000 km from the
known range of melanonota, well within the known range of the
nominate subspecies, and calls into question the validity of the
subspecies melanonota which was described based primarily
on color patterns. A re-evaluation of geographic variation and
genetics in this species is needed.
92. Chelodina (Chelodina) longicollis: Hodges et al. (2015)
studied mitochondrial phylogeography of Chelodina longicollis,
a highly terrestrially mobile freshwater species, to determine if
its population genetic structure would correspond to hydrological
boundaries or not. They found two ancient haplogroups broadly
with east-west partitioning across the Great Dividing Range, but
made no taxonomic recommendations. Each haplogroup was
characterized by complex genetic structure, demographically
stable subpopulations, and signals of isolation by distance; but
the patterns were also overlaid with signatures of introgression
and recent gene flow, likely facilitated by late Pleistocene and
ongoing anthropogenic landscape changes.
93. Chelodina (Macrochelodina) expansa: Hodges et
al. (2014) carried out a phylogeographic study of Chelodina
expansa based on mitochondrial gene variation to identify two
major clades of mitochondrial haplotypes. The first comprised
populations from the inland Murray-Darling Basin and the Mary
River in southeast Queensland; the second comprised populations
from coastal catchments north of the Mary River. They did not
regard it as appropriate to provide taxonomic recognition for
these populations east and west of the Great Dividing Range (as
previously proposed for other similarly distributed taxa by Cann
1998), because the mitochondrial analysis demonstrated that the
morphological variation observed was not concordant with the
spatial population structure defined by the molecular data (views
subsequently discussed further by Spinks et al. 2015). Nevertheless, Hoser (2014b) inappropriately applied novel names to two
of the mitochondrial clades identified by Hodges et al. (2014).
However, since the ICZN has been petitioned by Rhodin et al.
(2015) (see Annotation 4 on Hoser) to declare these and other
Hoser names unavailable under the Code (Article 82.1), we
maintain prevailing usage and do not use his names.
94. Chelodina (Macrochelodina) kuchlingi and C. walloyarrina: Both of these purported taxa remain enigmatic and
poorly known, with their apparent distinctiveness incompletely
evaluated, and we remain uncertain whether they are indeed valid
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
and recognizable taxa (as either species or subspecies), or poorly
differentiated lineages of C. oblonga (formerly C. rugosa) and C.
burrungandjii, respectively, or possibly their hybrid intergrades.
Ellis and Georges (2015), in their catalogue of turtle type specimens held at the Western Australian Museum, synonymized C.
kuchlingi under C. oblonga and C. walloyarrina under C. burrungandjii, following the earlier recommendation by Georges
and Thomson (2010), but their action was not based on new
data or analysis. However, we note that prior to the description
of Chelodina walloyarrina from the Kimberleys by McCord and
Joseph-Ouni (2007b), and not addressed in our earlier comment
(TTWG 2010, annotation 10:38), is that Thomson et al. (2000)
noted morphological differences between Kimberley specimens (=
C. walloyarrina) and C. burrungandjii from Arnhem Land, while
Georges et al. (2002) compared the still undescribed Kimberley
form and C. burrungandjii from Arnhem Land species using
allozyme data (45 independent nuclear loci), which showed no
fixed differences between these populations. As we noted in our
prior checklist (TTWG 2014, annotation 14:42), further analysis
of the phylogeographic relationships among all these lineages is
still in progress by Kuchling, Georges, and others, and based on
our precautionary principles in regard to data-driven analysis in
the recognition and protection of biodiversity, we remain reluctant
to formally synonymize these taxa until more conclusive genetic
and morphologic data emerge.
95. Elseya: Thomson et al. (2015) divided the broad genus
Elseya into three subgenera based on genetics and skeletal morphology: Elseya (Elseya), Elseya (Hanwarachelys), and Elseya
(Pelocomastes).
96. Elseya flaviventralis: Allopatric populations of the
Elseya dentata group from the Arnhem Land region of Northern
Australia were first suggested to be unique by Legler (1981) based
on morphology. Subsequent allozyme electrophoretic analyses by
Georges and Adams (1992, 1996) and morphological work by
several authors (reviewed in Thomson and Georges 2016) supported the distinction of these populations at the species level,
and Thomson and Georges (2016) formally described this species
as Elseya (Elseya) flaviventralis, which we accept.
97. Elseya rhodini: As indicated in our last checklist (Annotation 14:45), the population of Elseya (formerly included under E.
novaeguineae) from the southern versant of New Guinea, previously noted by Rhodin and Genorupa (2000) and by Georges et al.
(2014) to be genetically and morphologically differentiated from
other New Guinean Elseya, has since been formally described as
Elseya (Hanwarachelys) rhodini by Thomson et al. (2015). We
accept this new species as distinct.
98. Emydura macquarii krefftii: Todd et al. (2014) examined
variation in mitochondrial and nuclear microsatellite markers
across the widespread range of Emydura macquarii krefftii. The
mitochondrial data revealed 1) a distinct divergence between
northern (Burdekin River northward) and southern (Fitzroy River
and southward) populations of E. m. krefftii, 2) that E. m. emmotti
was most closely related to but highly divergent from the northern
krefftii clade, 3) that E. m. nigra and E. m. macquarii were nested
within the southern krefftii clade, and 4) that populations in the
upper Burnett River were highly divergent from other southern
clade populations. Their examination of twelve microsatellite
loci within only krefftii also demonstrated support for the same
north-south divergence as for the mtDNA data, but did not support the distinction of the upper Burnett population (possibly a
human translocation, according to the authors), or the taxonomic
distinction among the subspecies nigra, macquarii, and southern
krefftii. They made no explicit taxonomic recommendations, but
did note that the north-south divergence was within the range
exhibited for other accepted chelonian species. Further analysis
is clearly indicated.
99. Myuchelys purvisi: Legler (1981) foreshadowed splitting
the genus Elseya into two major clades, one containing Elseya
dentata and related species, the other containing Elseya latisternum
and its close relatives, many of which were undescribed at that
time. A subsequent study based on 54 allozymes loci (Georges and
Adams 1994) established that Legler’s “latisternum group” was
indeed a clade (monophyletic), and this was the foundation for the
description of the new genus Myuchelys (Thomson and Georges
2009) with four contained species: M. purvisi, M. georgesi, M.
bellii, and M. latisternum as type. Unfortunately, morphological
characters that diagnose that genus are symplesiomorphies.
Two subsequent analyses based on a limited set of nuclear
and mitochondrial sequence data were equivocal on the monophyly of Myuchelys. Georges et al. (1998) recovered Myuchelys
as paraphyletic with respect to Elseya based on analyses of two
mitochondrial genes (12S rRNA and 16S rRNA), but without
statistical support. More recently, analyses of a single nuclear
locus (c-mos) provided moderate support (83% bootstrap support
values) for grouping Emydura macquarii, Myuchelys latisternum,
and M. georgesi as a clade to the exclusion of M. purvisi, a result
confirmed by analysis of mtDNA (Fielder et al. 2012). However,
Georges and Adams (1994), Georges et al. (1998), and Fielder
et al. (2012) all maintained that the uncertainty surrounding
incongruence among these analyses should preclude taxonomic
revisions and therefore did not propose revisions to correct the
potential paraphyly of Myuchelys with respect to Emydura.
Subsequently, Le et al. (2013) generated phylogenies for the
chelid genera Elseya, Emydura, Myuchelys, Elusor, and Rheodytes
using two mtDNA markers and a single nuDNA marker. Their
phylogeny also recovered Myuchelys as paraphyletic, again owing
to the position of M. purvisi, and they assigned purvisi to a new
genus, Flaviemys, to maintain monophyly of Myuchelys. A more
comprehensive analysis (Spinks et al. 2015) using 13 independent
nuclear DNA markers recovered Myuchelys, including M. purvisi,
as a well-supported clade, in agreement with the previous allozyme
data. Thus, based on the weight of evidence, the taxonomic revision
of Le et al. (2013), using their more limited sequence information,
was considered by Spinks et al. (2015) to be premature.
On the basis of conflicting evidence from the mitochondrial
and nuclear DNA evidence, and because M. purvisi and M. georgesi
are so morphologically similar as to have long been regarded as
a cryptic species pair (but see Thomson and Georges 1996), we
have retained Myuchelys based on the weight of evidence suggesting that it is a well-supported monophyletic group, and placed
Flaviemys into its synonymy, following the recommendation of
Spinks et al. (2015).
100. Pseudemydurinae, Pseudemydura, P. umbrina: The
affinities of Pseudemydura among the Chelidae are not well
established. Early work using serological comparisons revealed
that Pseudemydura, Emydura-Elseya, and Chelodina formed
an unresolved trichotomy (Burbidge et al. 1974). Many of the
defining morphological characters of Pseudemydura have been
regarded as autapomorphies, and so not useful for phylogenetic
analysis. Nevertheless, Gaffney (1977) placed Pseudemydura
as sister to all the remaining extant chelids of both Australasia
and South America (as the new subfamily Pseudemydurinae),
while admitting that his case, based on a single retained primitive character, was weak. Subsequent DNA sequence studies
were unable to resolve the conundrum. 12S mitochondrial rRNA
data were unable to conclusively establish the relationships of
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Pseudemydura, but tentatively resolved it as sister to the other
Australasian shortnecked genera—Emydura, Elseya, Myuchelys,
Rheodytes and Elusor (Seddon et al. 1997). Additional sequence
analysis from mt 16S rRNA and nuclear c-mos supported this arrangement, but bootstrap support remained poor (Georges et al.
1998). A more recent analysis, using the same data drawn from
Genbank, resolved Pseudemydura as sister to Chelodina (Guillon et al. 2012). Subsequently, in a broad analysis of previously
published sequence data representing 13 mitochondrial and nuclear
DNA loci and including 83% of all extant turtle species (as per our
checklist), Pereira et al. (2017) resolved Pseudemydura as sister
to all short-necked Australasian chelids, with those two clades
sister to Chelodina. Most recently, Zhang et al. (2017) analyzed
the whole mitogenome of Pseudemydura and demonstrated it to
be sister to all Australasian chelid turtles (subfamily Chelodininae), with strong bootstrap support; consequently, they proposed
resurrection of the subfamily Pseudemydurinae Gaffney 1977,
which we adopt here.
101. Pelomedusa: Vargas-Ramírez et al. (2010) and Wong et
al. (2010) reported deeply divergent mitochondrial clades within
what was then recognized as the monotypic species Pelomedusa
subrufa, with less, but concordant, variation in nuclear DNA
markers. The extent of mitochondrial divergences of up to 20%
of the cytochrome b gene exceeds pairwise divergences between
any other congeneric chelonian species. Based on mitochondrial
DNA variation, Petzold et al. (2014) and Nagy et al. (2015) identified at least thirteen terminal clades (see also Fritz et al. 2014b).
Petzold et al. (2014) recommended full species status for some of
these clades. They restricted P. subrufa (sensu stricto) to southern
Angola, Botswana, southeastern Democratic Republic of the
Congo, Madagascar (presumably introduced), Malawi, Namibia,
South Africa, and Tanzania. Older names were resurrected from
the synonymy of P. subrufa for three of the clades: P. galeata
(Schoepff 1792; South Africa), P. gehafie (Rüppell 1835; Eritrea
and possibly Sudan), and P. olivacea (Schweigger 1812; Benin,
Burkina Faso, Niger, Nigeria, and Senegal). Six other species were
newly described: P. barbata (Saudi Arabia and Yemen), P. kobe
(Tanzania), P. neumanni (Kenya and Tanzania), P. schweinfurthi
(Central African Republic and South Sudan), P. somalica (Somalia
and Ethiopia; see also Fritz et al. 2015b), and P. variabilis (Ghana
and Ivory Coast). The authors also identified two clades that they
considered candidate species but did not describe: one from Cameroon, and the other from Sudan. Further candidate species could
correspond to distinct clades within P. galeata and P. somalica
(Fritz et al. 2015b), and Nagy et al. (2015) added another possible
candidate species from the southeastern Democratic Republic of
the Congo. All these studies, however, relied primarily on mitochondrial data and preliminary geographic sampling, but also
included some morphologic work and sequencing and allocation of
type specimens (see also Fritz et al. 2015a). We remain uncertain
whether all taxa described and/or identified are valid and at what
systematic level (species vs. subspecies vs. ESUs), pending corroborating data from nuclear genes. However, based on the data and
extensive analyses presented, we tentatively accept the proposed
taxonomic arrangement while urging and awaiting further work,
notably analysis of nuclear loci, more detailed morphologic work,
and further geographic sampling, especially from areas between
assigned species. We document these unsampled and unassigned
areas as Pelomedusa subrufa (sensu lato) species complex.
102. Pelusios: Kindler et al. (2016) examined sequence variation in three mitochondrial and three nuclear genes in Pelusios,
and found no phylogenetic structure in P. chapini, P. gabonensis,
or P. nanus; however, they identified significant structure within P.
rhodesianus (with two deeply divergent clades), P. carinatus, and
P. castaneus. Both their mtDNA and nDNA data also suggested
that rhodesianus was paraphyletic with respect to carinatus. But
because their geographic sampling was incomplete and there was
discordance between their mtDNA and nDNA data, the authors
made no new taxonomic decisions regarding these six taxa.
103. Pelusios castaneus seychellensis: Kindler et al. (2016),
in their wider phylogeographic study of Pelusios, found that their
mtDNA data nested the lectotype of P. seychellensis deep within a
variable P. castaneus, and sister to specimens from the Republic of
Congo, agreeing with Stuckas et al. (2013), leading them to treat
seychellensis as a junior synonym of castaneus. We addressed
the status of seychellensis previously in Annotation 14:47 and
interpreted it at that time as a subspecies of P. castaneus. We
maintain that interpretation at this time; however, we acknowledge
the possibility that the types of seychellensis could conceivably
have been mislabeled or based on transported specimens, in which
case seychellensis should indeed be synonymized with castaneus.
104. Podocnemis erythrocephala: Santos et al. (2016) examined variation in a single mitochondrial gene across the range
of Podocnemis erythrocephala. They identified considerable
genetic structure among populations, with four distinct genotypes
that they deemed “management units,” but made no taxonomic
recommendations.
105. Podocnemis sextuberculata: Podocnemis sextuberculata and Pentonix americana (nomen dubium) (= Pelomedusa
subrufa) were first described on p. 13 of Cornalia (1849), but the
descriptions (with minor changes) were reprinted on p. 312 in
the Appendix of Osculati (1850), who collected the holotype of
sextuberculata described by Cornalia. The two publications have
created some confusion about dates and pagination for these two
names, and we correct our previously cited paginations here.
PREVIOUS CHECKLIST ANNOTATIONS
2007 Checklist Annotations
TTWG 2007b (CRM 4:173–199) (07:1-105)
07:1. Both IUCN (The World Conservation Union, http://www.iucnredlist.
org) and CREO (Committee on Recently Extinct Organisms, http://creo.amnh.
org) have designated 1500 AD as their official cutoff date for determining what
constitutes a recently extinct species, and we follow their criteria in our checklist.
07:2. Chelydra: Phillips et al. (1996) elevated acutirostris and rossignoni
to full species status and retained the subspecies osceola. See Shaffer et al. (in
press) for a complete review.
07:3. Macrochelys [formerly Macroclemys]:Although Macroclemys has
been the most commonly used name, Webb (1995b) showed that Macrochelys
is the oldest available name.
07:4. Chelonia mydas: Bowen et al. (1992) showed that recognition of
the taxon agassizii Bocourt 1868 renders mydas paraphyletic, and agassizii is
no longer generally recognized as either a distinct species or subspecies. See
Parham and Zug (1996) and Karl and Bowen (1999) for a complete review.
07:5. Eretmochelys imbricata: Fritz and Havas (2006, 2007) did not
list bissa as a valid taxon, but no argumentation for this opinion was given.
Genetic data (Okayama et al., 1999) have suggested significant separation of
Atlantic from Pacific stocks.
07:6. Kinosternon species: Serb et al. (2001) elevated two former
subspecies of flavescens (arizonense and durangoense) to full species status.
07:7. Kinosternon chimalhuaca: This new species name appeared prematurely and erroneously first in the hobbyist literature, with the full original
description published a few months later (Berry et al., 1996, 1997).
07:8. Kinosternon scorpioides scorpioides: Includes the previously
recognized subspecies seriei Freiberg 1936 and carajasensis Cunha 1970 in
synonymy (Cabrera and Colantonio, 1997).
07:9. Sternotherus: This genus was included as a junior synonym of
Kinosternon by Iverson (1992) and David (1994) based on work by Seidel et
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
al. (1986) and Iverson (1991). However, this view was never widely accepted,
and Iverson (1998) showed that the species referred to either Sternotherus or
Kinosternon formed reciprocally monophyletic clades and recommended that
both genera be used.
07:10. Sternotherus depressus: Whereas some earlier authors had placed
this taxon as a subspecies of minor, Walker et al. (1998) showed that depressus
was genetically distinct from minor.
07:11. Chrysemys picta dorsalis: This subspecies of Chrysemys picta
was elevated to full species status by Starkey et al. (2003), who recognized two
distinct genetic lineages: C. dorsalis and C. picta. They did not find genetic
support for the other subspecies of C. picta (belli, marginata) but did not recommend that they be abandoned. Fritz and Havas (2006, 2007) argued that full
specific status of dorsalis was not fully demonstrated and retained it and the
other two taxa as subspecies of C. picta, agreeing also with Ernst et al. (2006).
07:12. Graptemys ouachitensis sabinensis: Based on molecular and
morphologic data, Stephens and Wiens (2003) suggested that sabinensis may
not be closely related to ouachitensis. However, statistical support for this was
weak, and they did not discuss or recommend a taxonomic change. Further
study of this complex may warrant the elevation of the sympatric taxon sabinensis to full species status.
07:13. Pseudemys concinna concinna: Includes the previously recognized subspecies hieroglyphica Holbrook 1836, mobilensis Holbrook 1838,
and metteri Ward 1984 in synonymy (Seidel, 1994).
07:14. Pseudemys concinna floridana: This taxon was previously
considered a separate species, but was designated a subspecies of concinna
by Seidel (1994). Jackson (1995) argued for the retention of floridana as a full
species, but Seidel (1995) rejected this argument.
07:15. Pseudemys concinna suwanniensis: Previously considered a
subspecies of concinna, Seidel (1994) argued for the elevation of this taxon
to full species status, but Jackson (1995) argued for its subspecific status.
07:16. Pseudemys gorzugi: This taxon was previously considered a
subspecies of concinna, but was elevated to species status by Ernst (1990)
without argumentation, but then supported through analysis by Seidel (1994).
07:17. Pseudemys peninsularis: This taxon was previously considered
a subspecies of floridana, but was elevated to species status by Seidel (1994).
Jackson (1995) argued for the retention of peninsularis as a subspecies of
floridana, but Seidel (1995) reaffirmed his recognition.
07:18. Trachemys species: Seidel (2002) recommended elevating nine
Mesoamerican taxa, previously recognized as subspecies of Trachemys scripta,
to species rank.
07:19. Trachemys subspecies: Seidel (2002) also recommended reassigning five taxa, previously subspecies of scripta, to subspecies of his various
elevated Trachemys species.
07:20. Trachemys dorbigni: Includes the previously recognized subspecies brasiliensis Freiberg 1969 in synonymy, based on morphologic work (del
Barco and Larriera, 1993).
07:21. Emydoidea and the turtles formerly known as Clemmys: The
four traditional species of Clemmys (guttata [type], insculpta, muhlenbergii,
and marmorata) do not form a monophyletic group with respect to the two
monotypic genera Emys orbicularis and Emydoidea blandingii in phylogenies
based on DNA data (Bickham et al., 1996; Burke et al., 1996; Lenk et al. 1999;
Feldman and Parham, 2002). While there is a general agreement that insculpta
and muhlenbergii are sister-species and should be placed in the genus Glyptemys
(Holman and Fritz, 2001; Parham and Feldman, 2002), there are two schemes
presented for marmorata and blandingii. Holman and Fritz (2001) recommended
that marmorata be placed in the monotypic genus Actinemys, retaining both
Emys orbicularis and Emydoidea blandingii as additional monotypic genera.
Other authors (Bickham et al., 1996; Feldman and Parham, 2002; Parham and
Feldman, 2002) recommended that marmorata and blandingii be placed into
an expanded Emys, a scheme favored in the most recent analysis of variation
in marmorata (Spinks and Shaffer, 2005).
07:22. Emys or Actinemys marmorata: Previously, two subspecies were
distinguished, including pallida Seeliger 1945, but genetic analysis by Spinks
and Shaffer (2005) demonstrated that the typical and previously recognized
subspecies pallida were within the same phylogeographic clade and so pallida
should not be considered valid.
07:23. Emys orbicularis iberica: Includes the recently described subspecies kurae Fritz 1994 in synonymy (Fritz, 1998).
07:24. Emys orbicularis persica: Includes the recently described subspecies orientalis Fritz 1994 in synonymy (Fritz, 1998).
07:25. Mexican Terrapene carolina: Stephens and Wiens (2003) suggested that Mexican subspecies of T. carolina may warrant full species status.
While this convention has also been adopted previously (Smith et al., 1996),
almost all other workers recognize these as subspecies.
07:26. Platysternidae: Krenz et al. (2005) confirmed that nuDNA placed
Platysternon solidly within the Testudinoidea, and Parham et al. (2006a) supported this finding with mtDNA.
07:27. Platysternon megacephalum: Ernst and Laemmerzahl (2002)
synonymized two subspecies of megacephalum (vogeli Wermuth 1969 and
tristernalis Schleich and Gruber 1984) with the nominate subspecies.
07:28. Testudinoidae or Testuguria: Shaffer et al. (1997) coined the
name ‘Testudinoidae’ for the clade that united Testudinidae with Bataguridae/
Geoemydidae. Joyce et al. (2004) listed Testudinoidae as an undesirable derivative of Testudo being to similar to both ‘Testudinidae’ and ‘Testudinoidea.’
In that same paper, the authors coined the new clade name ‘Testuguria’ for
that same clade (while neglecting to list Testudinoidae as an objective senior
synonym). Parham et al. (2006a) explicitly argued for the use of Testuguria
over Testudinoidae.
07:29. Bataguridae or Geoemydidae: Both names are being used to
refer to this group of predominantly Asian testudinoids. McDowell (1964) used
the name Batagurinae for this group (as a subfamily) which was changed to
Bataguridae (as a family) by Gaffney and Meylan (1988). Bour and Dubois
(1986) showed that Geoemydidae has priority, and David (1994), Spinks et
al. (2004) and others have embraced this view. However, this approach was
questioned by Joyce et al. (2004) who, working in a rank-free phylogenetic
taxonomy framework, recommended the continued use of Bataguridae. In the
interest of reconciling phylogenetic nomenclature with traditional Linnaean
rules of priority, Parham et al. (2006a) endorsed a phylogenetic codification
of Geoemydidae.
07:30. Batagur: Praschag et al. (2007b) and Le et al. (2007) demonstrated
that species of Kachuga were genetically paraphyletic with respect to those
referred to Batagur and Callagur and recommended that only one genus be
recognized, and the name Batagur has priority.
07:31. Batagur baska: The subspecies ranongensis Nutaphand 1979 is
not well differentiated and has been synonymized under baska by Fritz and
Havas (2006, 2007), but no specific morphologic or genetic analysis has yet
been performed to formally evaluate the status of this taxon.
07:32. Cuora: Phylogenies based on DNA data (Honda et al., 2002a;
Stuart and Parham, 2004; Parham et al., 2004; Spinks et al., 2004) have shown
that continued recognition of the genus Pyxidea for mouhotii would render
Cuora paraphyletic. All of these studies recommended expanding Cuora to
include mouhotii. Other schemes for Cuora have not been published in the
recent scientific literature, though there has been some use of Cistoclemmys
for flavomarginata and galbinifrons (e.g., Zhao et al., 1997; Zhao, 1997;
Yasukawa and Ota, 1999).
07:33. Hybrid species: The validity of six taxa of Cuora, Mauremys
[including Ocadia], and Sacalia recently described from pet trade specimens
has been refuted by genetic studies that have shown them to be based on hybrids
(Parham et al., 2001; Wink et al., 2001; Spinks et al., 2004; Stuart and Parham,
2004, 2007). The taxa shown to be hybrids are: Cuora galbinifrons serrata
Iverson and McCord 1992b, Mauremys iversoni Pritchard and McCord 1991,
Mauremys pritchardi McCord 1997, Ocadia glyphistoma McCord and Iverson
1994, Ocadia philippeni McCord and Iverson 1992, and Sacalia pseudocellata
Iverson and McCord 1992a.
07:34. Cuora flavomarginata sinensis: Some authors recognize this
taxon as a valid subspecies (McCord and Iverson, 1991; Fong et al., 2002)
while others synonymize it with flavomarginata (Yasukawa and Ota, 1999;
Fritz and Havas, 2006, 2007).
07:35. Cuora galbinifrons: The taxa bourreti and picturata, originally
described as subspecies of Cuora galbinifrons, were elevated to species rank
by Stuart and Parham (2004) based on concordance of morphological with
molecular differentiation. Fritz et al. (2006c) returned bourreti to subspecies
rank based on osteological characters shown by market specimens, and suggested that picturata warrants the same ranking; Fritz and Havas (2006, 2007)
subsequently listed picturata at subspecies rank based on morphologically
intermediate pet trade specimens. Includes the previously recognized hainanensis
Li 1958 in synonymy (Zong and Pan, 1989; Iverson and McCord, 1992b).
07:36. Cuora trifasciata: Blanck et al. (2006a) recommended that
Cuora trifasciata be split into two species (including their newly named species cyclornata and its new subspecies meieri) based on paraphyletic mtDNA
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haplotypes and morphological differences. Spinks and Shaffer (2007) showed
that trifasciata as traditionally recognized is monophyletic based on nuDNA
and therefore recommended that cyclornata should not be recognized, pending additional study.
07:37. Cuora yunnanensis: This species has been listed as extinct by the
IUCN since 2000 (www.iucnredlist.org), based on several decades of not finding
any surviving animals despite intensive searches. Recently, a pair of animals
representing this species were found in markets (Zhou and Zhao, 2004; Zhou,
2005), with subsequent confirmation through genetic analysis (He et al., 2007).
07:38. Cyclemys: Iverson (1992) recognized two taxa of Cyclemys (dentata and tcheponensis). Later, atripons and pulchristriata were described and
oldhamii was resurrected (Iverson and McCord, 1997; Fritz et al., 1997). Genetic
analysis by Guicking et al. (2002) also supported the validity of shanensis.
07: 39. Geoemyda: Yasukawa et al. (1992) elevated japonica to species
status (previously considered a subspecies of spengleri).
07:40. Hardella thurjii: Praschag et al. (2007b) found no genetic or
morphologic evidence for continued recognition of the subspecies indi Gray
1870b, and synonymized it under thurjii.
07:41. Heosemys annandalii [formerly in Hieremys]: Spinks et al. (2004)
showed that annandalii was nested among species of Heosemys. Diesmos et
al. (2005) formally moved annandalii into Heosemys.
07:42. Leucocephalon yuwonoi [formerly in Geoemyda or Heosemys]:
Originally described as a species of Geoemyda (McCord et al., 1995), Fritz
and Obst (1996) placed yuwonoi in Heosemys. McCord et al. (2000) showed
that yuwonoi was not closely related to the type species of Geoemyda or Heosemys, but instead sister to Notochelys platynota, and erected a new genus,
Leucocephalon, for yuwonoi.
07:43. Malayemys macrocephala: Brophy (2004) proposed the recognition of this species as distinct from subtrijuga based on morphological grounds.
07:44. Mauremys [including species formerly in Annamemys, Chinemys, or Ocadia]: Iverson and McCord (1994) included annamensis under an
expanded Mauremys. Subsequent phylogenies based on DNA data (Honda
et al., 2002b; Barth et al., 2004; Feldman and Parham, 2004; Spinks et al.,
2004) showed that the genera Ocadia and Chinemys rendered Mauremys
paraphyletic. Based on these results, some authors (Feldman and Parham,
2004; Spinks et al., 2004) recommended synonymizing Ocadia and Chinemys
under Mauremys. Barth et al. (2004) presented this same scheme as well as one
that would retain Chinemys and Ocadia and further divide Mauremys into the
genera Cathaiemys and Emmenia. Barth et al. (2004) did not favor one scheme
over the other and a competing scheme for Mauremys has not been formally
proposed in the scientific literature.
07:45. Mauremys leprosa: Fritz et al. (2006a) explicitly synonymized
several subspecies of leprosa recently described by Schleich (1996a) (atlantica,
erhardi, marokkensis, wernerkaestlei, and zizi) plus vanmeerhaeghei Bour and
Maran 1998 [1999], and only recognized leprosa and saharica.
07:46. Mauremys reevesii: Iverson et al. (1989) and Barth et al. (2003,
2004) refuted the validity of the terminal taxon megalocephala Fang 1934, but
it has continued to be recognized by Chinese researchers (Guo et al., 1997;
Zhao, 1997; Zhang et al., 1998), and Fritz and Havas (2006, 2007) listed it as
a separate taxon with speculation about its relationships.
07:47. Melanochelys trijuga edeniana: The subspecies wiroti Reimann
1979 was recognized by Iverson (1992), but David (1994) suggested that it
was synonymous with edeniana, and Fritz and Havas (2006, 2007) followed
this arrangement.
07:48. Pangshura [formerly in Kachuga]: Das (2001) and Schleich and
Kästle (2002) used the name Pangshura to refer to small-bodied Kachuga. A
phylogeny based on DNA data (Spinks et al., 2004) showed that Kachuga was
paraphyletic and so removed flaviventer, smithii, sylhetensis, tecta, and tentoria
into the genus Pangshura. Praschag et al. (2007b) using mtDNA confirmed
the well-supported monophyly of Pangshura.
07:49. Pangshura tentoria flaviventer: Schleich and Kästle (2002)
elevated flaviventer to full species status based on sympatry with circumdata,
but Praschag et al. (2007b) performed a phylogeographic analysis and retained
flaviventer as a subspecies of tentoria.
07:50. Siebenrockiella leytensis [formerly in Heosemys]: Diesmos et al.
(2005) placed leytensis into the genus Siebenrockiella based on strong genetic
evidence for its sister relationship to S. crassicollis.
07:51. Vijayachelys silvatica [formerly in Geoemyda]: This species
was originally named as a species of Geoemyda. However, a molecular study
by Praschag et al. (2006) suggested a distant relationship with that genus and
they recommended that it be placed in the new monotypic genus Vijayachelys.
07:52. The Geochelone complex: This generic complex includes the
genera Geochelone, Aldabrachelys, Astrochelys, Angonoka, Centrochelys,
Chelonoidis, Dipsochelys, and Stigmochelys. Lapparent de Broin (2000b),
Gerlach (2001, 2004), Le et al. (2006), and Fritz and Bininda-Emonds (2007)
recommended dividing the Geochelone complex into several genera, although
their schemes differ somewhat. A general consensus on a generic-level revision
for some members of the group is lacking while in other areas (e.g., Astrochelys
radiata, Chelonoidis) there is agreement.
07:53. Aldabrachelys or Dipsochelys: Bour (1982a) originally recommended that Aldabran tortoises (dussumieri or gigantea) be placed in the genus
Dipsochelys instead of Aldabrachelys. However, Aldabrachelys is still widely
used, including sometimes by Bour (Austin et al., 2003), though Dipsochelys
is favored by others (Palkovacs et al., 2002, 2003; Gerlach, 2004). There is
recent disagreement regarding the type specimen of Testudo gigantea, the type
species of Aldabrachelys, that was presumed lost. Frazier (2006) designated a
neotype for T. gigantea, an act that would seemingly validate the use of both
Aldabrachelys and the terminal taxon gigantea. Around the same time, Bour
(2006) rediscovered the original lost type specimen, which is actually an individual of the South American tortoise Chelonoidis denticulata. If this claim is
correct, then the names Aldabrachelys or gigantea might not be applicable to
Aldabran tortoises. Whether Frazier’s neotype designation or Bour’s specimen
rediscovery prevails nomenclaturally remains a matter of ongoing debate, but
since Bour (2006) was the most recently published authority we use the name
dussumieri rather than gigantea in our list.
07:54. Aldabrachelys or Dipsochelys species: Gerlach and Canning
(1998) recognized six species of tortoises in Aldabra, Madagascar, and the
Seychelles (three of which were extinct: abrupta, daudinii, and grandidieri). The
two species from Madagascar became extinct prior to modern times (abrupta
Grandidier 1868 in ca. 1250 AD and grandidieri Vaillant 1885b in ca. 950
AD) so we do not include them in our list of modern taxa. Palkovacs et al.
(2002, 2003) questioned the validity of multiple extant species based on their
analysis of genetic data, recognizing only a single living taxon (Dipsochelys
dussumieri). Gerlach and Bour (2003) reemphasized the validity of the extant
species based on the observation that the hatchlings are diagnostic. Fritz and
Havas (2006, 2007) recognized only one extant species of Indian Ocean giant
tortoise which they referred to Aldabrachelys gigantea, but did not address
the findings of Gerlach and Bour (2003) or Bour (2006). As we consider the
issues surrounding the validity of these species as remaining unresolved, we
list all these species as potentially valid.
07:55. Aldabrachelys or Dipsochelys dussumieri: Iverson (1992) listed
this species as Geochelone gigantea Schweigger 1812. Many authors now use
dussumieri for the Aldabra tortoise (see above), but others persist in using the
older name gigantea (e.g., Fritz and Havas, 2006, 2007), and others have used
the name elephantina Duméril and Bibron 1835 (David, 1994; Devaux, 2007).
07:56. Astrochelys or Angonoka yniphora: Le et al. (2006) named
Angonoka for yniphora because of its uncertain phylogenetic position. Fritz
and Bininda-Emonds (2007) recovered a weak sister relationship between
yniphora and Astrochelys radiata under some algorithms and recommended
that yniphora be placed in Astrochelys.
07:57. Chelonoidis petersi: According to Cabrera (1998), citing morphologic and osteologic work by Fernández (1988), Chelonoidis chilensis
should be divided into two species, chilensis and petersi Freiberg 1973, but
he considered the taxon donosobarrosi Freiberg 1973 to be synonymous with
chilensis. Fritz and Havas (2006, 2007) speculated that petersi may not be
valid and synonymized it under chilensis, citing phenotypic plasticity in other
tortoise species as a reason for not accepting the reported differences between
petersi and chilensis.
07:58. Chelonoidis nigra: Most recent authors have considered the various
taxa of Galapagos tortoises as subspecies of nigra (e.g., Pritchard, 1996; Caccone et al., 1999; Fritz and Havas, 2006, 2007), but Caccone et al. (2002) and
Russello et al. (2005, 2007) treated them as distinct species. The nomenclatural
and survival status of these taxa were discussed in detail by Pritchard (1996).
07:59. Chelonoidis nigra chathamensis: This taxon described from
western Chatham Island (San Cristóbal) appears to have been extirpated from its
original range, but a population of tortoises persists on eastern Chatham Island
that was considered a possible separate subspecies by Pritchard (1996). Pending
genetic analysis and resolution of this issue we continue to list chathamensis as
the extant taxon from Chatham, whereas Fritz and Havas (2006, 2007) listed
it as extinct, but made no mention of the extant population.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
07:60. Chelonoidis nigra duncanensis: This taxon from Duncan Island
(Pinzón) was historically usually referred to ephippium Günther 1875a, but
Pritchard (1996) demonstrated that ephippium was a synonym of abingdonii
and therefore resurrected the old nomen nudum duncanensis Garman 1917.
07:61. Chelonoidis nigra nigra: The nominotypical subspecies nigra
from Charles Island (Santa Maria or Floreana) is considered to be extinct and
is therefore included separately on this list.
07:62. Chelonoidis nigra phantastica: This taxon was listed by Fritz and
Havas (2006, 2007) as extant, but Pritchard (1996) considered it probably extinct.
07:63. Chelonoidis nigra porteri: This taxon from Indefatigable Island
(Santa Cruz) has often been referred to nigrita Duméril and Bibron 1835, but
most recent authors, including Pritchard (1996) and Fritz and Havas (2006,
2007) have used porteri.
07:64. Chelonoidis nigra vicina: This widespread taxon from Albemarle
Island (Isabela) was previously recognized as one of several valid taxa on that
island, including becki Rothschild 1901, microphyes Günther 1875a, guentheri
Baur 1889, and vandenburghi De Sola 1930. Pritchard (1996) synonymized
microphyes, guentheri, and vandenburghi under vicina, and recognized only
vicina and becki from Albemarle.
07:65. Cylindraspis indica: Includes the recently described borbonica
Bour 1978 in synonymy, based on genetic work by Austin and Arnold (2001).
07:66. Cylindraspis vosmaeri: Fritz and Havas (2006) credited Fitzinger
1826 with authorship of this name, but corrected it to Suckow 1798 in their
2007 checklist.
07:67. Homopus: A separate taxon of Homopus was referred to H. bergeri
Lindholm 1906 by Branch (1989). However, that name was a junior synonym
of Psammobates tentorius verroxii Smith 1839 (Branch, 1992; Boycott and
Bourquin, 2000), and the new taxon was recently described as H. solus by
Branch (2007).
07:68. Indotestudo travancorica: This taxon was previously considered
a subspecies of forstenii (Hoogmoed and Crumly, 1984; Iverson, 1992), but
was resurrected to species status by Pritchard (2000) based on morphology, a
conclusion supported by mtDNA analysis by Iverson et al. (2001c).
07:69. Kinixys belliana: Fritz and Havas (2006, 2007) recognized only
belliana and nogueyi, following Broadley (1993) uncritically, but others (Iverson, 1992; David, 1994; Iverson et al., 2001a) also recognized domerguei and
zombensis. As the phylogeography of this broadly distributed species complex
has not been analyzed, we list the four most widely recognized subspecies.
07:70. Pyxis arachnoides: The three recognized subspecies have recently
been confirmed as genetically distinct lineages (Chiari et al., 2005).
07:71. Stigmochelys or Psammobates pardalis: Based on genetic analysis, Le et al. (2006) recommended that this taxon be included in an expanded
genus Psammobates. Fritz and Bininda-Emonds (2007) argued for the retention of a monophyletic Psammobates exclusive of pardalis. Le at al. (2006)
also found a high level of mitochondrial divergence between two specimens
assigned to the two subspecies pardalis and babcocki. In conjunction with
morphological distinctions between these two taxa (Loveridge and Williams,
1957; Broadley, 1989), the preliminary genetic data suggest that they may be
different at the species level.
07:72. Testudo or Agrionemys: The species horsfieldii and hermanni
have been alternatively placed in the genera Testudo or Agrionemys (Khosatzky
and Mlynarski, 1966; Gmira 1993, 1995) and hermanni also recently in Eurotestudo. Lapparent de Broin (2000a,b) and Parham et al. (2006b) supported
the placement of horsfieldii in the genus Agrionemys, but suggested that a new
genus name was needed for hermanni. Later Lapparent de Broin et al. (2006a)
created the name Eurotestudo for hermanni, but Fritz and Bininda-Emonds
(2007) demonstrated that older genus names (Chersine and Medaestia) are
available for that species. Fritz and Bininda-Emonds (2007) recovered a
weakly monophyletic clade that included horsfieldii, hermanni, and the three
core species of Testudo (graeca, kleinmanni, and marginata). Based on this
phylogeny they recommended that all of these species be placed in the genus
Testudo. The genetic support for some nodes within this clade is not strong and
the decision to lump or split is subjective (e.g., whether Agrionemys should be
used for horsfieldii is open to debate), therefore the taxonomy of this group
may remain in flux for some time.
07:73. Testudo graeca: This species complex has been the subject of
massive taxonomic revisions at the species and subspecies level. These revisions have resulted in the naming and elevation of numerous taxa (e.g., Perälä,
2002a,b,c). Several studies (van der Kuyl et al., 2002, 2005; Harris et al., 2003;
Carretero et al., 2005; Parham et al., 2006b,c; Fritz et al., 2007) have explicitly
refuted the validity of many of these taxonomic acts. Fritz et al. (2007) proposed
a taxonomic scheme that recognized five mitochondrial clades in the eastern
part of the range of T. graeca as subspecies, but did not address the status of
several North African subspecies. Since this is the most recent taxonomic
suggestion, it is listed here. However, in their recent checklist, Fritz and Havas
(2006, 2007) included not only the eleven taxa we list, but also anamurensis
Weissinger 1987, antakyensis Perälä 1996, floweri Bodenheimer 1935, nikolskii
Chkhikvadze and Tuniyev 1986, pallasi Chkhikvadze and Bakradze 2002, and
perses Perälä 2002c. The relationships within this species complex remain
uncertain and we expect its taxonomy to continue fluctuating.
07:74. Testudo kleinmanni: Baha el Din (2006), Široký and Fritz (2007),
and Attum et al. (2007) explicitly refuted the validity of werneri Perälä 2001
as a species distinct from kleinmanni.
07:75. Testudo marginata: Fritz et al. (2005b) explicitly refuted the
validity of weissingeri Bour 1996 as a subspecies of marginata.
07:76. Testudo hermanni: Fritz et al. (2006b) explicitly refuted the validity of hercegovinensis Werner 1899 (previously resurrected by Perälä, 2002b)
and recommended that boettgeri be considered a subspecies of hermanni.
07:77. Testudo horsfieldii: In a conference proceedings, Perälä (2002a)
elevated two subspecies of horsfieldii (kazachstanica and rustamovi) to full
species status. This was accepted by Lapparent de Broin et al. (2006b), but
warrants reconsideration, especially considering the evidence for unjustified
taxonomic inflation in related tortoises in the same work (van der Kuyl et al.,
2002, 2005; Fritz et al., 2005b, 2006b; Parham et al., 2006b,c).
07:78. Carettochelys insculpta canni: This subspecies from northern
Australia described by Wells (2002a) was only weakly defined as different
from the nominotypical subspecies from New Guinea. We list it tentatively
pending further analysis, as did Fritz and Havas (2006), although they excluded
it from their 2007 checklist.
07:79. Apalone spinifera atra: This taxon has usually been designated
a subspecies of spinifera (usually with the original spelling ater), but others
(e.g., Flores-Villela, 1993; David, 1994) have listed it as a full species, though
usually without specific argumentation.
07:80. Aspideretes or Nilssonia: Engstrom et al. (2004) found Aspideretes
to be paraphyletic with respect to Nilssonia formosa based on morphologic
and genetic criteria. Praschag et al. (2007a) formally synonymized Aspideretes
into an expanded concept of Nilssonia based on their analysis of mtDNA of
all five included taxa.
07:81. Aspideretes or Nilssonia nigricans: Recent morphologic and
genetic work on this species previously known only from a single captive
population has demonstrated that it also occurs in the wild (Praschag and
Gemel, 2002; Praschag et al., 2007a).
07:82. Pelodiscus: The genus has recently been recognized as including
up to four separate species by some authorities (David, 1994; Zhao, 1997; Chen
et al., 2005, 2006; Fritz and Havas, 2006, 2007). Relationships within the genus
are far from resolved and also complicated by translocation and mixing of huge
numbers of farm-raised individuals from many parts of the range.
07:83. Rafetus swinhoei: Includes the recently described Pelochelys
taihuensis Zhang 1984 (Farkas, 1992) and Rafetus leloii Duc 2000 in synonymy
(Farkas and Webb, 2003).
07:84. Acanthochelys macrocephala: Includes the recently described
Phrynops chacoensis Fritz and Pauler 1992 in synonymy (Fritz and Pauler, 1999).
07:85. Chelodina: This genus was split into three genera by Wells and
Wellington (1985), using Chelodina for the narrower-headed shorter-necked
species (longicollis, novaeguineae), and establishing Macrochelodina for the
broader-headed longer-necked species (oblonga, expansa, rugosa, siebenrocki),
and Hesperochelodina for steindachneri. Iverson et al. (2001b) refuted the
availability of the name Hesperochelodina, but validated Macrochelodina.
Georges et al. (2002) retained Chelodina for the entire genus, but identified
three phylogenetic clades within the genus and recommended recognition of
three subgenera (but did not name them). Fritz and Havas (2006, 2007) accepted two of these clades (Chelodina and Macrochelodina) as separate genera.
07:86. Chelodina canni: This taxon is the same as the previously described rankini Wells and Wellington 1985, but that name was declared invalid
as a nomen nudum by Iverson et al. (2001b). Wells (2007a) recently disputed
this interpretation and redescribed rankini, but canni McCord and Thomson
2002 retains nomenclatural precedence and rankini Wells 2007a is therefore
a junior synonym of canni.
07:87. Chelodina mccordi roteensis: This recently named subspecies
described in the hobbyist literature needs genetic confirmation of its distinctive-
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238
Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
ness, but we recognize it pending further analysis.
07:88. Chelodina oblonga: Thomson (2000) showed that the holotype of
oblonga Gray 1841 is a specimen of what is currently regarded as Chelodina
rugosa Ogilby 1890. An application is before the International Commission
for Zoological Nomenclature (ICZN) to conserve current usage of the name
C. rugosa Ogilby 1890 for the northern snake-necked turtle and to apply the
earlier available name Chelodina collei Gray 1856a to the long-necked species
of southwestern Australia, while retaining the nomenclatural availability of the
name oblonga for potential future designation of distinct populations of rugosa
(Thomson, 2006). Though no decision has yet been rendered by the ICZN, Fritz
and Havas (2006, 2007) used the name colliei for this southwestern population.
Georges et al. (2002) found support that this taxon represents a third subgenus
under Chelodina, but did not formally establish it under a generic-level name.
07:89. Chelodina timorensis: This species recently described in the
hobbyist literature by McCord et al. (2007b) was also described a few months
later as a new subspecies of mccordi (‘timorlestensis’) by Kuchling et al.
(2007), but the McCord et al. description has chronologic precedence. Concerns
surrounding the history and methodology of the description of timorensis by
McCord et al. are discussed by Kuchling et al. (2007) and serve to emphasize
our recommendations (made in our other chapter in this volume) to follow
certain procedural guidelines for descriptions of new taxa (Turtle Taxonomy
Working Group, 2007a).
07:90. Chelodina kuchlingi: This species was described from a single
specimen, leading to doubts about its validity (Georges and Thomson, 2006;
Fritz and Havas, 2006, 2007), but it remains listed pending further exploration
of its remote area of provenance.
07:91. Chelodina rugosa: The species siebenrocki Werner 1901 was
considered valid by Rhodin and Mittermeier (1976) and Rhodin and Genorupa
(2000), but synonymized under rugosa by Georges et al. (2002) based on weakly
differentiated allozymes within the broader rugosa complex.
07:92. Elseya: This genus has been recognized as consisting of two
separate lineages (Georges and Rose, 1996; Georges and Thomson, 2006).
It was subsequently split into two genera, Elseya and Wollumbinia, by Wells
(2007c), with latisternum designated genotype of Wollumbinia. Papers by Wells
(2002a,b; 2007a,b,c) and Wells and Wellington (1985) have been self-published
without any peer review and also highlight our recommendations to follow
certain procedural guidelines for descriptions of new taxa (Turtle Taxonomy
Working Group, 2007a).
07:93. Elseya branderhorsti: This species was considered valid by Rhodin
and Genorupa (2000), Thomson et al. (2006), and Georges and Thomson (2006).
07:94. Elseya jukesi: The name jukesi Wells 2002b was a nomen nudum
since no type specimen was designated, but the species was recently redescribed
by Wells (2007b).
07:95. Elseya schultzei: This species was listed by Thomson et al. (2006)
and Georges and Thomson (2006), but neither morphologic nor genetic data
have been analyzed from the type population and its status remains unclear.
07:96. Elseya stirlingi: The previously named taxon stirlingi Wells and
Wellington 1985 was declared invalid as a nomen nudum by Iverson et al.
(2001b) (though spelled erroneously as sterlingi), but was recently redescribed
as a valid species by Wells (2007b).
07:97. Elseya or Wollumbinia bellii: The taxon dorriani Wells 2002b
is a nomen nudum without a type designation, but was recently considered a
valid subspecies of bellii by Wells (2007c).
07:98. Emydura macquarii: The taxonomy of E. macquarii was previously
reviewed by Georges and Adams (1996). Later, Cann et al. (2003) and McCord
et al. (2003) described two new subspecies, but taxa previously described by
Cann in 1998 (binjing, dharra, dharuk, and gunabarra), plus signata Ahl 1932
were not specifically evaluated by those authors. However, these taxa were all
recognized as subspecies of macquarii by Fritz and Havas (2006, 2007), and
since phylogeographic variation in the macquarii species complex has not yet
been fully resolved with adequate genetic work, we tentatively list all these
subspecies as valid, pending further analysis.
07:99. Emydura subglobosa worrelli: Originally described as Tropicochelymys worrelli, this taxon was synonymized under Emydura victoriae
Gray 1842 by Iverson (1992) and the nomenclatural validity of the species
name confirmed by Iverson et al. (2001b). Cann (1998) considered it a distinct
species, but Georges and Thomson (2006), partially based on electrophoretic
work by Georges and Rose (1996), concluded that it was best referred to as
a subspecies of subglobosa Krefft 1876. Fritz and Havas (2006, 2007) also
listed it as a subspecies of subglobosa, but Georges et al. (2006) referred to it
as a species, though without providing data or argument.
07:100. Phrynops: Wermuth and Mertens (1977) divided this genus into
three subgenera: Phrynops, Batrachemys, and Mesoclemmys. Cabrera (1998)
and Georges et al. (1998) elevated these subgenera to generic level. McCord
et al. (2001) further divided the remaining monophyletic Phrynops into a total
of four genera (Bufocephala, Phrynops, Ranacephala, and Rhinemys). Joyce et
al. (2004) did not accept the taxonomic acts of McCord et al. (2001). Bour and
Zaher (2005) synonymized Bufocephala and Ranacephala with Mesoclemmys,
but recognized Rhinemys as distinct.
07:101. Mesoclemmys heliostemma: Rueda-Almonacid et al. (2007)
questioned the validity of this taxon which is completely sympatric with
raniceps, suggesting that it may simply represent a juvenile color morph of
that taxon, and recommended genetic analysis.
07:102. Pelomedusa subrufa: Gasperetti et al. (1993) recommended
that the two previously recognized subspecies (nigra Gray 1863b and olivacea
Schweigger 1812) be abandoned.
07:103. Pelusios seychellensis: The taxonomic status of this species is
unclear. Gerlach and Canning (2001) concluded that it is extinct.
07:104. Podocnemididae or Podocnemidae: Cope (1868) used the name
Podocnemididae to refer to this clade. Baur (1893b) later referred to this group
as Podocnemidae. Joyce et al. (2004) phylogenetically defined Baur’s name
(Podocnemidae) to refer to this clade.
07:105. Podocnemis unifilis: This long-recognized species was briefly
referred to as P. cayennensis Schweigger 1812 by David (1994), but that name
was previously often used for what is now recognized as P. erythrocephala
(Mittermeier and Wilson, 1974), and most authors have continued to use unifilis.
2008 Checklist Annotations
Rhodin et al. 2008 (000.1-38.checklist.v.1) (08:2-25)
08:2. Chelodina or Macrodiremys: The southwestern long-necked
turtle of Australia (Chelodina oblonga or colliei, see annotation below) represents one of three lineages that were considered unnamed subgenera of Chelodina by Georges et al. (2002). McCord and Joseph-Ouni (2007b) created
the name Macrodiremys for oblonga/colliei, designating Chelodina oblonga
Gray 1841, as type species by original designation and monotypy,and elevated this to a full monotypic genus. Whether Chelodina sensu stricto will be a
subgenus of Chelodina sensu lato along with Macrodiremys and Macrochelodina or if all three will be used as full genera is subjective and not yet stable.
08:3. Chelodina or Macrodiremys oblonga: Within Chelodina, the
specific epithet oblonga has long been applied to a long-necked species in
southwestern Australia. Thomson (2000) showed that the holotype of oblonga Gray 1841 is a specimen of what is currently regarded as Chelodina
rugosa Ogilby 1890 from northern Australia. An application (Thomson 2006,
2007) is before the International Commission of Zoological Nomenclature
(ICZN) to conserve current usage of the name C. rugosa Ogilby 1890 for the
northern snake-necked turtle and to apply the next available name, Chelodina
colliei Gray 1856a, to the long-necked species of southwestern Australia.
Separately, McCord and Joseph-Ouni (2007b) designated the holotype of
colliei as the neotype of oblonga which would render colliei a junior synonym of oblonga which would be incompatible with an identification of the
holotype and name oblonga as pertaining to the northern Australian taxon.
We list the southwestern long-necked species as oblonga because McCord
and Joseph-Ouni (2007b) is the latest published action but note that, given
the differing taxonomic acts and opinions, this name may remain unstable in
the coming years.
08:4. Macrochelodina or Chelodina walloyarrina: McCord and Joesph-Ouni (2007b) described the new species Macrochelodina walloyarrina
based on morphological criteria.
08:5. Chelydra serpentina: Shaffer et al. (2008) recommended synonymization of Chelydra serpentina osceola Stejneger 1918 into Chelydra
serpentina (Linnaeus 1766) based on range-wide patterns in variability of
mtDNA.
08:6. Cyclemys: Fritz et al. (2008b) performed a revision of the genus
based on molecular and morphological data that included the description of
three new species (enigmatica, fusca, and gemeli).
08:7. Cyclemys dentata: Stuart and Fritz (2008) analyzed mtDNA from
type specimens of Cyclemys belli Gray 1863e, Cyclemys orbiculata Bell 1834
and Cyclemys ovata Gray 1863e, and confirmed their previous morphologybased synonymizations with Cyclemys dentata (Gray 1831d) as accurate.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
08:8. Cyclemys oldhamii: Stuart and Fritz (2008) placed the names
shanensis Annandale 1918, and tcheponensis Bourret 1939a, into the synonymy of oldhamii, based on the absence of significant genetic variation between the type specimen of oldhamii, topotypes of shanensis, and samples of
tcheponensis from near the type locality.
08:9. Batagur: Praschag et al. (2008) examined mtDNA variation within Batagur baska sensu lato and recommended that the southern populations
should be elevated to full species status and for which the available name
affinis Cantor 1847 should be used.
08:10. Carettochelys insculpta: Fritz and Havas (2007) and Georges et
al. (2008) indicated that Carettochelys insculpta canni Wells 2002a is not an
available name because it had not been published in accordance with criteria
established by the International Code of Zoological Nomenclature.
08:11. Kinixys nogueyi: This taxon was treated as a full species by McCord et al. (2005) with minimal argumentation for the change from traditional
recognition as a subspecies of K. belliana.
08:12. Rhinoclemmys punctularia: Testudo scabra Linnaeus 1758 has
previously been referred (as a nomen dubium) to the synonymies of both
Melanochelys trijuga (Schweigger 1812) and Rhinoclemmys punctularia
(Daudin 1801) (see Fritz and Havas 2007). Examination of the holotype specimen still extant in the Uppsala Linnaean collection indicates that it appears to
be a Rhinoclemmys punctularia (Rhodin and Carr, in press) [2009]. However,
since the name Testudo scabra has not been used as the name for a recognized
taxon since the early 1800s, it remains a nomen oblitum and does not replace
the name punctularia Daudin 1801, recognized and used as valid since its description, and also protected by the ICZN (1963) as a nomen conservandum.
08:13. Aldabrachelys or Dipsochelys: The generic and specific names
of the Aldabra tortoise are still being debated (reviewed in TTWG 2007). The
original type specimen of Testudo gigantea Schweigger 1812 is a Chelonoidis denticulata from Brazil (Bour 2006), but since the name gigantea had
been associated with tortoises from Aldabra for a long time, Frazier (2006)
designated a neotype from Aldabra, leading to some confusion. The matter
is currently being petitioned to the International Commission on Zoological
Nomenclature (Frazier 2008).
08:14. Chelonoidis nigra vicina: The type locality for Testudo microphyes Günther 1875a was given as follows: “I suppose that the specimen…
has come from Hood’s Island,” and Pritchard (1996) concluded that the name
microphyes was a nomen dubium since its type specimen was not identifiable
as either a Hood Island tortoise or any other recognizable taxon. Fritz and
Havas (2007) indicated that Günther (1877) had subsequently designated the
type locality for microphyes as “Tagus Cove, northern Albemarle Island” and
they placed microphyes in the synonymy of Testudo vicina Günther 1875a,
but this is not necessarily correct. Günther (1877) simply referred latercollected specimens from Tagus Cove to his type-based concept of microphyes—the name microphyes therefore remains a nomen dubium until such
time as the original type specimen can perhaps be identified as to its exact
provenance using genetic analysis.
08:15. Agrionemys or Testudo horsfieldii: Vasilyev et al. (2008) demonstrated minimal mitochondrial variation between populations of horsfieldii
Gray 1844 and kazachstanica Chkhikvadze, Amiranashvili, and Ataev 1990
and so recommended that these taxa be considered subspecies of Agrionemys
horsfieldii. Elsewhere in the paper they referred to A. h. rustamovi as a third
subspecies, but made no definitive comment on taxonomic status or validity.
08:16. Podocnemis unifilis: The terminal taxon lata Bell in Gray 1830e
has previously been included under the synonymy of Peltocephalus dumerilianus (Schweigger 1812) by many previous authors, including Fritz and
Havas (2007), but Bell (in Gray 1830e) described Chelys (Hydraspis) lata
as having a depressed black shell and orange-spotted head more typical of
Podocnemis unifilis or P. expansa. Later, Gray (1870f) placed Hydraspis lata
in the synonymy of his concept of Chelonemys dumeriliana (= Podocnemis
unifilis) and added P. unifilis, P. cayennensis, and P. erythrocephala to the
same synonymy, while differentiating Peltocephalus tracaxa (= Peltocephalus dumerilianus) as a distinct taxon. Though Chelys (Hydraspis) lata Bell in
Gray 1830e is an older name than Podocnemis unifilis Troschel 1848, it is a
nomen oblitum not used for a valid taxon since its description, and therefore
does not replace unifilis as the valid name for the species.
08:17. Trachemys decussata: Authorship of this taxon was actually
first by Bell (in Griffith and Pidgeon 1830) with a plate. Seidel (1988a) listed
the author as Gray 1831:28 (= Gray 1831d), and Fritz and Havas (2007) listed
the author as Gray 1831:11 (= Gray 1830e). Griffith and Pidgeon 1830 was
published in September 1830, whereas Gray 1830e was published in December 1830, and Gray 1831d was published in May 1831.
08:18. Graptemys geographica: The name lesueuri Gray 1830d (=
lesueurii Gray 1830e) was shown by Bour and Dubois (1983) to be a junior synonym of geographica LeSueur 1827 rather than a senior synonym
of pseudogeographica Gray 1831d as recorded by Fritz and Havas (2007).
08:19. Graptemys pseudogeographica: The name pseudogeographica
Gray 1831d was originally published only as a nomen nudum of a LeSueur
manuscript name in junior synonymy under geographica LeSueur 1827, but
gradually achieved wide usage by many authors. Holbrook (1842) was the
first to actually describe the taxon under the name pseudogeographica, and
arguably his name should perhaps be associated with it, but Stejneger and
Barbour (1917) established the name as pseudogeographica Gray 1831d as
pointed out by Bour and Dubois (1983), who agreed that Gray should be
listed as the author.
08:20. Cryptodira and Pleurodira: These subordinal names were
based on the French vernacular names, Cryptodères and Pleurodères, originally used by Duméril and Bibron 1834. Cope (1864, 1865, 1868b) has generally been credited with authorship of these names, and he was the first to use
the exact names Cryptodira in 1868 and Pleurodira in 1865, but previously
used the name Pleurodera in 1864. Cope was preceded by Lichtenstein (1856)
who used Cryptodera and Pleurodera as subordinal names, but in a printed
catalogue distributed to zoological colleagues and museums, and not apparently sold in bookstores, so therefore perhaps not nomenclaturally available.
08:21. Cuora evelynae: In the previous checklist (TTWG 2007), Cuora
flavomarginata (Gray 1863e) had three subspecies. Ernst et al. (2008) recommended returning Cuora evelynae Ernst and Lovich 1990 to full species
status, based on new morphological and previously published molecular data.
They also argued that the mainland populations assigned to taxon sinensis
Hsü 1930 are synonymous with nominotypical flavomarginata.
08:22. Apalone spinifera: In the previous checklist (TTWG 2007), seven subspecies of spinifera LeSueur 1827 were listed. McGaugh et al. (2008)
performed a rangewide phylogeographic study that uncovered patterns of discordant molecular and morphological variation. These authors conservatively
refrained from making sweeping nomenclatural changes, but noted that there
was “little utility” in recognizing the taxon hartwegi Conant and Goin 1948
and synonymized it under spinifera.
08:23. Apalone spinifera atra: In the previous checklist (TTWG 2007),
atra Webb and Legler 1960 was referred to as a subspecies of spinifera
LeSueur 1827 as it is here. This taxon is sometimes considered a full species
(e.g., argumentation cited in Flores-Villela 1993). Several recent studies on
atra (McGaugh 2008, McGaugh and Janzen 2008, McGaugh et al. 2008)
have argued in support of subspecies status based on low levels of genetic
distinctiveness and habitat-driven color variation. The subspecific status of
atra was followed by Cerdá-Ardura et al. (2008).
08:24. Sacalia quadriocellata: Shi et al. (2008) performed a mitochondrial survey of S. quadriocellata based on known-locality and trade specimens. They found that populations on Hainan are genetically distinct and can
also be diagnosed by morphological characters. They did not elevate these
populations to species status, but noted that eventual study might validate this
conclusion, in which case the name Sacalia insulensis (Adler 1962) would be
available. They also noted that samples of S. quadriocellata from northern
Vietnam are genetically distinct from those from Laos and the type region of
central Vietnam, but there appear to be no obvious morphological differences
between these populations.
08:25. Rhinoclemmys flammigera: Barrio-Amorós and Narbaiza
(2008) elevated Rhinoclemmys punctularia flammigera Paolillo 1985 to species status based on distinct head coloration pattern and allopatric isolated
distribution.
2009 Checklist Annotations
TTWG 2009 (000.39-84.checklist.v.2) (09:3-49)
09:3. Chelydridae: Chandler and Janzen (2009) analyzed the phylogenetic
position of the Chelydridae based on nucleotide sequence data, and found
weak support for a sister group relationship with either the Kinosternoidea
(Kinosternidae + Dermatemydidae) or Chelonioidea (Cheloniidae +
Dermochelyidae). In a more extensive analysis, Barley et al. (in press) show
that Chelydridae is sister to Kinosternoidea.
09:4. Macrochelys temminckii: Roman et al. (1999) showed that M.
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temminckii could be divided into three major mitochondrial clades which they
treated as Evolutionarily Significant Units (ESUs). They noted that samples
from the Suwannee drainage in Florida showed high divergence from the rest
of the range. Echelle et al. (2009) performed a microsatellite study and further
subdivided M. temminckii into six ESUs. They also noted that the Suwannee
population was the most distinct and concluded that it might eventually be
recognized as a distinct taxonomic unit.
09:5. Cheloniidae: Bowen and Karl (2007) reviewed population genetics
and phylogeography of marine turtles and while they noted mtDNA divergence
between Indo-Pacific andAtlantic Chelonia mydas and Eretmochelys imbricata,
they recognized no taxa below the species level.
09:6. Lacépède 1788 and Bonnaterre 1789: The International Commission on Zoological Nomenclature (ICZN) previously rejected the names created
by Lacépède in his 1788 Histoire Naturelle des Serpens and subsequent editions
since they were published in non-binomial works (ICZN 1987). However, all
names published in his earlier 1788 volume, Histoire Naturelle de Quadrupèdes Ovipares (which contained all his turtle descriptions), continued to be
nomenclaturally available until recently, when they were also rejected as being
published in a non-binomial work (ICZN 2005). A few of these turtle names
from the 1788 Histoire Naturelle de Quadrupèdes Ovipares volume had already
been individually suppressed by the ICZN (1963). Therefore, Bonnaterre (1789),
who republished Lacépède’s descriptions with proper binomials, becomes the
authorship source for these rejected Lacépède turtle names.
09:7. Testudo nasicornis: Testudo nasicornis Lacépède 1788 was described as distinct from Testudo caouana Lacépède 1788 (= Caretta caretta)
based on the possession of a soft nasal projection and on being fit for human
consumption like Testudo marina (= Chelonia mydas). The species was included in the synonymy of various other marine turtle taxa until Loveridge and
Williams (1957) placed it alongside T. caouana in the synonymy of Caretta
caretta (Linnaeus, 1758). Bonnaterre (1789) provided an amplified description
and drawing of Testudo nasicornis, but his concept of the taxon differed from
that of Lacépède, illustrating a specimen (pl. 3, fig. 3; hereby designated as the
holotype) with 13 large carapacial scutes, costal tubercles forming an interrupted
lateral keel, a strongly serrated carapacial margin, and apparently two pairs of
prefrontal scales. The 13 carapacial scutes are inconsistent with Caretta but
correspond to Chelonia or Eretmochelys, while the serrated carapacial margin
and apparently split prefrontals indicate that the figured specimen is an Eretmochelys imbricata (Linnaeus, 1766). As long as the name Testudo nasicornis was
nomenclaturally occupied by Lacépède’s 1788 description, Bonnaterre’s 1789
description could simply be dismissed as an incorrect subsequent attribution.
However, now that ICZN Opinion 2104 (ICZN 2005) has rejected availability
of all of Lacépède’s turtle names, Bonnaterre’s becomes the first available usage
of the name Testudo nasicornis. Bonnaterre’s description is clearly attributable
to Eretmochelys imbricata, and thus we consider Testudo nasicornis Bonnaterre
1789 to be a subjective junior synonym of that taxon, while retaining Testudo
nasicornis Lacépède 1788 as a nomen rejectum attributable to Caretta caretta.
09:8. Meyer 1790 species names: Meyer (1790), in a long-overlooked
review article, provided short diagnoses and replacement names (nomina nova)
for four species described by Lacépède (1788). These names have never appeared in any subsequent synonymies and are clearly nomina oblita. The Meyer
names are Testudo bomarii for Testudo viridisquamosa (= Lepidochelys kempii
or Chelonia mydas), Testudo lauanna for Testudo caouana (= Caretta caretta),
Testudo sonnerati for Testudo punctata (= Lissemys punctata punctata), and
Testudo rubra for Testudo subrufa (= Pelomedusa subrufa).
09:9. Eretmochelys imbricata: This checklist (TTWG 2007b; Rhodin
et al. 2008) has previously treated bissa as a valid subspecies of imbricata
in the absence of a definitive, data-based synonymization. Whereas genetic
results have demonstrated distinct genetic lineages in the Atlantic and IndoPacific Oceans, no genetic studies or reviews (Okayama et al. 1999; Bowen
and Karl 2007) have argued for continued recognition of the subspecies bissa.
The phylogenetic structure within Eretmochelys is comparable to that within
Chelonia, for which only a single monotypic species is currently recognized,
and thus we now treat bissa as a synonym of imbricata.
09:10. Kinosternon arizonense: The authorship of this name was given as
arizonense Gilmore 1922 in our previous checklist as well as TTWG (2007b)
and Fritz and Havas (2007). However, the article appeared in February 1923
and the year is therefore corrected.
09:11. Kinosternon hirtipes: The authorship of this name was given
as Wagler 1833 in our previous checklist as well as in TTWG (2007b) and
Fritz and Havas (2007), whereas previous authors (e.g., Iverson 1992) have
recognized hirtipes Wagler 1830, a name sometimes interpreted as a nomen
nudum. However, the ICZN (1999) rules for availability of names published
prior to 1931 (Article 12) state that species names must be accompanied by a
“description or definition” or by an “indication”. The 1830 citation for hirtipes
is not accompanied by a description or definition, but is accompanied by an
indication—the associated illustration of the holotype of the taxon being named
(see Article 12.2.7). Wagler 1830 is therefore the correct authorship designation.
09:12. Deirochelyinae: Spinks et al. (2009b) performed a phylogenetic
analysis of the Emydidae based on mitochondrial and nuclear DNA. Their
mitochondrial phylogeny did not recover a monophyletic Deirochelyinae,
instead it placed Deirochelys as the sister to the rest of Emydidae. In contrast,
their nuclear data recovered a monophyletic Deirochelyinae.
09:13. Pseudemys: Spinks et al. (2009b) performed a phylogenetic
analysis of Emydidae based on mitochondrial and nuclear DNA. Although
their sampling within Pseudemys was limited and uneven, their samples of
concinna and floridana did not yield a monophyletic P. concinna with respect to
peninsularis (mtDNAand nuclear DNA) or nelsoni (nuclear DNAonly). Further
genetic and morphological study of known locality samples will be necessary
in order to resolve the taxonomic status of the terminal taxa within Pseudemys.
09:14. Trachemys: Seidel (2002) made several recommendations about
the specific and subspecific taxonomy of Trachemys, as reflected in our earlier
checklist. Jackson et al. (2008) performed a mitochondrial survey of the genus
and supported those taxonomic revisions.
09:15. Trachemys scripta: Rhodin and Carr (2009) demonstrated that the
original authorship of the taxon name scripta should be attributed to Thunberg
in Schoepff (1792), rather than just Schoepff.
09:16. Emys or Actinemys and Emys or Emydoidea: Using nuclear
markers, Spinks and Shaffer (2009) re-emphasized a close phylogenetic
relationship among marmorata, orbicularis/trinacris, and blandingii as was
previously shown from mitochondrial DNA(see Feldman and Parham 2002 and
case summary in annotation 21 of TTWG 2007b). Spinks and Shaffer (2009)
also showed that those species share a complex evolutionary history including
prehistoric hybridization, and that blandingii and orbicularis/trinacris are sister
taxa. In light of this evidence they strongly recommended that all these species
be included in the genus Emys rather than continued recognition of the genera
Actinemys and Emydoidea. Other authors (Iverson et al. 2008) argue for the
continued recognition of all three genera in this clade.
09:17. Emys orbicularis orbicularis: Fritz et al. (2009b) demonstrated
that the mitochondrial DNA differentiation of the two previously recognized
subspecies colchica and luteofusca were insufficient to continue to recognize
them as distinct and therefore synonymized both under orbicularis.
09:18. Emys orbicularis fritzjuergenobsti: Velo-Antón et al. (2008)
performed a genetic analysis of multiple populations of Emys orbicularis on
the Iberian peninsula and found no significant genetic divergence between
the two previously-defined subspecies hispanica and fritzjuergenobsti, and
therefore synonymized the former under the latter.
09:19. Emys orbicularis persica: Fritz et al. (2009b) demonstrated
that the mitochondrial DNA differentiation of the previously recognized
subspecies iberica was insufficient to continue to recognize it as distinct and
therefore synonymized it under persica, thereby also bringing kurae under
the synonymy of persica.
09:20. Geoemydidae and Rhinoclemmys: Le and McCord (2008) evaluated the molecular phylogeny of Rhinoclemmys and other geoemydid genera
and affirmed the monophyly of the Geoemydidae, but recommended that
Rhinoclemmys be afforded subfamilial recognition as the Rhinoclemminae, a
grouping concept first proposed by Gray (1873j) as the Tribe Rhinoclemmyina.
09:21. Batagur affinis edwardmolli: Praschag et al (2009) assessed the
taxonomic status of B. affinis using mitochondrial and nuclear genetic analysis,
and described the populations of Cambodia and the eastern coast of Peninsular
Malaysia as the new subspecies edwardmolli, with the populations of western
Peninsular Malaysia and Sumatra (Indonesia) retained as the nominotypical
subspecies affinis.
09:22. Cuora galbinifrons complex: In our previous checklists we
listed the three taxa galbinifrons, bourreti, and picturata as subspecies of
galbinifrons. However, the preponderance of well-documented evidence now
supports the elevation of these three taxa to recognition as three closely related
full species, based on both morphology and genetics (Stuart and Parham 2004,
Spinks et al. 2009a).
09:23. Cuora trifasciata: Spinks et al. (2009a) assessed the validity
of the recently described Cuora cyclornata Blanck, McCord, and Le 2006a
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
using a combination of mitochondrial and nuclear genetic markers. Their genetic evidence elucidates a complex history of introgression involving Cuora
trifasciata and the Cuora pani complex. They also provide a critique of the
morphological analysis of Blanck et al. (2006), concluding that the available
evidence is not sufficient to diagnose C. cyclornata. Following their previous
analysis of this group (Spinks et al. 2006), they continue to recommend that
C. cyclornata be considered a junior synonym of C. trifasciata.
09:24. Cyclemys species: Praschag et al. (2009b) analyzed mitochondrial
and nuclear genes in these species, and found that gemeli and fusca were distinct
but closely related, and that atripons, dentata, and pulchristriata were also
well-differentiated and formed a well-supported clade. The taxonomy of the
genus Cyclemys has been subject to intense debate over the past several years
and will likely continue for some time before it is stabilized.
09:25. Geoemyda spengleri: Gong et al. (2009) demonstrated phylogeographic structure in mitochondrial DNA within this taxon.
09:26. Mauremys: Hirayama et al. (2007) recommended splitting the
genus Mauremys (sensu Feldman and Parham 2004) into five genera (Mauremys, Cathaiemys, Chinemys, Ocadia, and an unnamed new genus) based on
the morphology of the palate. The relative utility of single character typological
taxonomies and monotypic genera versus restricting familiar names to welldefined evolutionary clades has been discussed elsewhere for Mauremys and
turtles in general (Parham and Feldman 2002; Feldman and Parham 2004;
Spinks et al. 2004; Turtle Taxonomy Working Group 2007a; Spinks et al. 2009).
Following the philosophy outlined in these papers we retain the larger aggregate
Mauremys and recommend consideration of subgenera for phenetically distinctive subclades (e.g., Parham and Feldman 2002; Smith and Chizsar 2006).
09:27. Mauremys caspica: Fritz et al. (2008a) performed a rangewide
genetic survey of Mauremys caspica. Their study revealed discordant patterns
of morphological and genetic differentiation in this species. They did not
recommend abandoning the current subspecies, but highlighted the need for
future taxonomic revision.
09:28. Rhinoclemmys punctularia flammigera: Barrio-Amorós and
Narbaiza (2008) elevated this taxon to species status based on a brief statement about head coloration and allopatric distribution, a change we reflected
in our previous checklist; however, based on the relative lack of supportive
data, we treat it again as a subspecies pending further analysis.
09:29. Aldabrachelys or Dipsochelys: The nomenclatural validity of the
generic and specific names of the Aldabra tortoise (Aldabrachelys gigantea or
Dipsochelys dussumieri) is currently being debated (reviewed in TTWG 2007b
and Rhodin et al. 2008). There is recent disagreement regarding the type specimen of Testudo gigantea, the type species of Aldabrachelys, that was presumed
lost. Frazier (2006) designated an Aldabran neotype for T. gigantea, an act that
would validate the use of both Aldabrachelys and the terminal taxon name
gigantea. Around the same time, Bour (2006) reported to have rediscovered
the original lost type specimen, which is actually an individual of the South
American tortoise Chelonoidis denticulata, making gigantea and Aldabrachelys
junior synonyms of denticulata and Chelonoidis, respectively, which would
thereby result in the use of Dipsochelys dussumieri as the valid name for the
Aldabra tortoise. Whether Frazier’s neotype designation or Bour’s specimen
rediscovery prevails nomenclaturally remains a matter of major ongoing debate.
The matter has been petitioned to the International Commission on Zoological
Nomenclature (Frazier 2009), with multiple commentaries on both sides of the
issue (Zug et al. 2009; Bour et al. 2009; Takahashi et al. 2009), but no decision
has yet been made by the ICZN.
09:30. Aldabrachelys or Dipsochelys species: Gerlach and Canning
(1998) recognized six species of tortoises in Aldabra, Madagascar, and the
Seychelles (three extant: gigantea or dussumieri, arnoldi, and hololissa; and
three extinct: abrupta, daudinii, and grandidieri). The two species from Madagascar became extinct prior to modern times (Testudo abrupta Grandidier 1868
in ca. 1250 AD and Testudo grandidieri Vaillant 1885b in ca. 950 AD) so we
do not include them in our list of modern taxa. Palkovacs et al. (2002, 2003)
rejected the validity of multiple extant species inhabiting the Indian Ocean
Islands based on their analysis of genetic data, recognizing only a single living
taxon (gigantea or dussumieri). Gerlach and Bour (2003) re-emphasized the
validity of their recognized species based on the observation that the hatchlings
are diagnostic. Further morphologic and genetic research is clearly needed to
determine whether more than one taxon of giant tortoise persists on the Indian
Ocean Islands. Based on their lack of demonstrable genetic differences, we
suggest that these morphologically-defined taxa be listed as subspecies of
gigantea/dussumieri pending further analysis.
09:31. Astrochelys yniphora: Le et al. (2006) proposed the genus name
Angonoka for the single taxon yniphora, but Fritz and Bininda-Emonds (2007)
showed that this species is closely related to radiata and so placed both species in the genus Astrochelys Gray 1873j. The genus name Angonoka has not
been adopted by other authors and we recommend the use of the genus name
Astrochelys for both radiata and yniphora.
09:32. Chelonoidis nigra species complex: Many recent authors and our
previous two checklists have considered the various taxa of Galápagos tortoises
as subspecies (e.g., Pritchard 1996; Caccone et al. 1999; Beheregaray et al.
2003; Fritz and Havas 2007; TTWG 2007b; Rhodin et al. 2008). However,
previous authors have considered them as full species based on morphology (Bour 1980; Fritts 1983; Ernst and Barbour 1989) and recently several
researchers (Caccone et al. 2002; Russello et al. 2005, 2007; Poulakakis et al.
2008; Chiari et al. 2009) have re-elevated them to species based on congruent
patterns of mitochondrial and nuclear variation. Given the allopatric distribution
of Gálapagos taxa, combined with the concordant patterns of mitochondrial,
nuclear, and morphological variation, we support their recognition as distinct
species. In raising these taxa to species, we prefer to highlight their close monophyletic relationship (as distinct from mainland South American Chelonoidis)
by listing them as a species complex.
09:33. Chelonoidis nigra: The correct epithet for the extinct Floreana
tortoise is nigra Quoy and Gaimard 1824. Poulakakis et al. (2008) used the
epithet elephantopus Harlan 1827, but this is in error because that name is
younger and the now-lost holotype of elephantopus cannot be assigned to
any island based on descriptions (Pritchard 1996). Extinct on Floreana since
the 1850s, hybrid descendants of this species were recently discovered on Isla
Isabela (Poulakakis et al. 2008; Parham 2008) indicating that the lineage persists
but has interbred with becki Rothschild 1901. Captive cross-breeding of these
becki x nigra hybrids could be used to partially reconstitute the nigra lineage.
09:34. Chelonoidis abingdonii: Hybrid descendants of C. abingdonii
have recently been found on Volcan Wolf on Isla Isabela (Russello et al. 2007),
and since only a single male (Lonesome George) of this species survives, the
lineage could be partially reconstituted by captive cross-breeding.
09:35. Chelonoidis porteri: Chiari et al. (2009) performed an extensive
analysis of morphological, mitochondrial, and nuclear genetic variation in the
two separate populations of tortoises on Santa Cruz presently referred to the
taxon porteri. They demonstrated that the Cerro Fatal population is genetically
and morphologically distinct from the La Caseta population and warrants a
formal new taxon description, currently in preparation.
09:36. Chelonoidis vicina: Pritchard (1996) previously synonymized
guentheri Baur 1898 under this taxon (see TTWG 2007b) based on lack of
morphological distinctiveness. Recent genetic work by Ciofi et al. (2006) has
confirmed a lack of significant genetic distinctiveness between these previously
recognized taxa on southern Isabela Island.
09:37. Chersina angulata: Daniels et al. (2007) have demonstrated
that this taxon includes two parapatric mitochondrial lineages. These lineages
are morphometrically distinct and also show ecological and behavioral differences. Taken together, these data suggest the existence of more than one
taxon within C. angulata and the matter is under further study (Daniels et al.
2007; Hofmeyr 2009).
09:38. Cylindraspis indica: In our previous checklist (Rhodin et al.
2008) we followed Fritz and Havas (2007) in synonymizing Testudo tabulata
africana Schweigger 1812 under Chersina angulata (Schweigger 1812).
However, Bour (1985) previously identified the type specimen of africana
as being a Cylindraspis graii, and Bour (2008) reaffirmed it as a synonym of
Cylindraspis indica.
09:39. Kinixys belliana nogueyi: This taxon was treated as a full species
by McCord et al. (2005) with minimal argumentation for the change from
traditional recognition as a subspecies of K. belliana, and we followed that
usage in our previous checklist. However, we now agree with Branch (2008)
and traditional usage, and therefore restore nogueyi to a subspecies of belliana.
09:40. Testudo or Chersine or Agrionemys: The type species and synonymizations of the genera Chersine Merrem 1820 and Medaestia Wussow
1916 have recently come under discussion. Bour and Ohler (2008) argued
that Testudo graeca Linnaeus 1758 is the type of Medaestia and that Testudo
hermanni Gmelin 1789 is the type of Chersine, whereas Fritz and Kraus (2008)
concluded that hermanni is the type species for both. In either case, the oldest
available generic name for hermanni or the clade (hermanni + horsfieldii) is
Chersine Merrem, 1820, of which Eurotestudo and perhaps Medaestia are
objective junior synonyms. Agrionemys is a subjective junior synonym if
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horsfieldii is considered congeneric with hermanni outside the genus Testudo,
but remains available for a monotypic genus containing horsfieldii.
09:41. Testudo graeca graeca: Fritz et al. (2009c) demonstrated that
the mitochondrial haplotype of topotypic T. g. whitei is identical to samples
of T. g. graeca, a taxon with overall low genetic variation. They therefore
reconfirmed placement of whitei Bennett in White 1836 in the synonymy of
graeca Linnaeus 1758.
09:42. Testudo graeca marokkensis: Fritz et al. (2009c) demonstrated that
T. g. lamberti and T. g. marokkensis share the same mitochondrial haplotype.
They also questioned the morphometric analyses and proposed geographical
separation of lamberti and marokkensis that were used to justify these taxa, and
recommended combining them into a single subspecies. Since both lamberti and
marokkensis were proposed in the same publication (Pieh and Perälä 2004), they
invoked the principle of first reviser and chose marokkensis as the valid name.
09:43. Testudo or Agrionemys horsfieldii: A recent study by Hitschfeld et
al. (2008) showed that carpal osteological characters used previously to elevate
the subspecies kazachstanica and rustamovi to species level are ontogenetically
variable. In addition, Fritz et al. (2009a) have demonstrated the presence of
three major mitochondrial haplotype clades that do not correspond well with the
presently understood geographic distribution of the three currently recognized
morphologically-defined subspecies. Whether or not to continue to recognize
kazachstanica and rustamovi as distinct subspecies remains uncertain, but we
retain them on the list pending further analysis and resolution.
09:44. Lissemys punctata: Rohilla et al. (2009) demonstrated some
geographic differentiation in allozymes in this wide-ranging taxon.
09:45. Wollumbinia or Myuchelys: Thomson and Georges (2009) described the new genus Myuchelys for these taxa (but not including dorsii Wells
2009), choosing not to recognize the previous description of Wollumbinia Wells
2007c as nomenclaturally available. Whether Wells’ work, distributed online
without adequate hardcopy dissemination, is nomenclaturally available needs
to be decided by the International Commission on Zoological Nomenclature,
and we therefore list both names.
09:46. Wollumbinia or Myuchelys dorsii: Wells (2009) described the new
species Wollumbinia dorsii, but whether the name is nomenclaturally available
is open to question, as the description was distributed online without adequate
hardcopy dissemination. As Australian chelid taxonomy is in a state of flux and
the validity of Wells’ multiple papers in his Australian Biodiversity Record is
under question, we list the name here. However, we make no determination
as to its validity either nomenclaturally or taxonomically.
09:47. Wollumbinia or Myuchelys latisternum: Wells (2009) resurrected
the taxon Wollumbinia spinosa (originally Euchelymys spinosa Gray 1871a)
as a separate valid species, based on the supposed distinctiveness of the single
holotype without known locality data collected in 1866 (as noted by Cann
1998). Whether this resurrection will be accepted as valid is open to question,
as no further specimens of the taxon have been identified and its distribution
is unknown. We therefore retain spinosa as a junior synonym of latisternum
pending further data.
09:48. Podocnemididae: Vargas-Ramirez et al. (2008) performed
mitochondrial and nuclear genetic analysis of all eight extant species and
demonstrated strong support for the Madagascan genus Erymnochelys being
sister to a strongly monophyletic South American Podocnemis, and the South
American Peltocephalus being sister to Erymnochelys + Podocnemis. This
phylogenetic analysis renders the occasionally used subfamilial clade name
Podocnemidinae Broin 1988 (for Podocnemis and Peltocephalus) paraphyletic.
09:49. Podocnemis unifilis: As we noted in our previous two checklists
(TTWG 2007b; Rhodin et al. 2008), most authors since Troschel 1848 have
used the name unifilis for this species (the yellow-spotted river turtle), though
some early authors erroneously used the epithet dumeriliana Schweigger 1812.
Recently, the name unifilis was referred to the synonymy of cayennensis Schweigger 1812 by David (1994), but the latter name has historically been used
for what is now recognized as P. erythrocephala (the red-headed Amazon River
turtle) (Mittermeier and Wilson, 1974). Bour (2006a) then redescribed what
he concluded to be one of the original three specimens used by Schweigger in
his concept of cayennensis and designated it as lectotype, but noted that this
specimen was actually a representative of the taxon currently known as unifilis,
and he recommended that the name cayennensis therefore be used instead of
unifilis. However, since the measurements of the lectotype provided by Bour
do not correspond exactly with those originally provided by Schweigger, and
since Schweigger evidently had examined three specimens for his description,
it remains unclear whether the lectotype has been correctly identified. In view
of the long history of stable usage of the epithet unifilis for the yellow-spotted
river turtle, we recommend its continued usage; suppression of cayennensis by
petition to the ICZN may be needed for nomenclatural stabilization.
2010 Checklist Annotations
TTWG 2010 (000.85-164.checklist.v.3) (10:4-49)
10:4. Testudines: In a paper published too close to our manuscript
deadline to fully analyze its implications, Dubois and Bour (2010b) discuss
the distinction between nomenclature at family-series and class-series rank,
and its application to the widely used Order group name Testudines Batsch
1788. Arguing that group names established at a family-series level cannot be
applied at a class-series level, and that the family-group name based on the
genus Testudo is already validly applied at the Family level (as “Testudinidae
Batsch 1788”), they conclude that the name Testudines Batsch 1788 cannot
also be applied to the Order of turtles, but do not suggest an available name
for the Order. However, the International Code of Zoological Nomenclature
does not regulate use of names above the superfamily level, and there remain
a variety of uncertainties and possible alternative interpretations on the validity, format, use, and authorship attribution of these names. Additionally,
some modern authors continue to use and defend the use of the original name
Testudines Linnaeus 1758 to designate all modern turtles, even though the
name was used primarily in a vernacular fashion in the original publication.
Therefore, we do not make any changes at this time, but continue to refer to
all turtles as the Order Testudines Batsch 1788, and expect to revisit this issue
in more detail in a future checklist.
Additionally, in recent years, the rank level of turtles has been recommended by some to be elevated from its traditional rank of Order within the
Class Reptilia to full Class rank on its own (e.g., Collins et al. 2010). Under
this scheme, Reptilia would be the Class containing only squamates and tuataras (traditionally known as the Lepidosauria), Class Eusuchia would contain
the crocodiles, Mammalia the mammals, and Aves the birds. This possible
class-level rank for turtles is to some extent supported by studies indicating
the paraphyly of the traditional Class Reptilia with regard to birds. However,
other studies indicate a sister-group relationship between turtles and diapsid
reptiles, or placement of turtles within diapsids, and hence, uncertainty about
the phylogenetic relationship of turtles to other groups abounds (e.g., Laurin
and Reisz 1995, deBraga and Rieppel 1997, Kirsch and Mayer 1998, Modesto
and Anderson 2004, Bhullar and Bever 2009). Considering that the monophyly
of turtles has never been challenged, and that ranking of turtles at class-level
provides no improved resolution of the group’s phylogenetic position, but
simply shifts its distinctiveness to a different rank, we continue to treat turtles
as an Order, with no implied judgment of its placement among other living
and fossil tetrapod groups.
10:5. Caretta caretta: Dubois and Bour (2010a) noted that Garsault (1764)
depicted and named a marine turtle as Testudo marina, which they considered
a junior synonym of Caretta caretta (Linnaeus 1758) based on morphology
and geography. Testudo marina Garsault 1764 is also a senior homonym of
Testudo marina Wilhelm 1794, a junior synonym of Dermochelys coriacea
(Vandelli 1761).
10:6. Chrysemys dorsalis or picta dorsalis: Phylogeography of the entire
Chrysemys picta complex was studied by Starkey et al. (2003), who demonstrated
two distinct mtDNA genetic lineages: dorsalis and picta. They recommended
elevating dorsalis to species status, but did not find genetic support for the
other traditional subspecies (bellii, marginata), although they recommended
that they not be abandoned. Ernst et al. (2006) documented morphologic intergradation between dorsalis and marginata in Missouri, but did not express
an opinion as to the validity of the proposed elevation of dorsalis by Starkey
et al., although they referred to their work. Fritz and Havas (2007) suggested
that full specific status of dorsalis was not fully demonstrated by Starkey et
al.’s data and retained it and the other two taxa as subspecies of picta. Iverson
et al. (2008) agreed with Starkey et al.’s analysis and listed dorsalis as a full
species, as have other recent authors and database managers (e.g., McAllister
et al. 2007, NatureServe), while others retain dorsalis at subspecies rank (e.g.,
Ernst and Lovich 2009, ITIS). The sequencing of the entire Chrysemys picta
genome is currently in draft form and should help resolve this problem. We
now choose to list this taxon provisionally at species rank, recognizing the
validity of arguments on either side of the issue, which remains unresolved.
10:7. Gray 1830e and Gray 1831d: The date of publication of Gray 1830e
(A Synopsis of the Species of the Class Reptilia) is cited by most sources as
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
1831, since the title page of Griffith and Pidgeon, Volume 9 (Reptilia), in which
the Synopsis appears as a Supplement, is dated 1831. However, Gray’s Synopsis
is dated on its first page as having been written in October 1830, and Volume 9
of Griffith and Pidgeon was actually published in three separate sections from
1830 to 1831 (see Cowan 1969). The first section, Part 25 (pp. 1–192), which
included Griffith and Pidgeon’s own text on Chelonia (plus part of Sauria),
appeared in September 1830 (this part also includes three new Bell and Gray
names that we cite as Bell 1830a and Gray 1830c). The second section, Part
26 (part of Sauria plus Ophidia and probably including Gray’s Supplement)
was apparenty published in December 1830; this date was interpolated by
Cowan (1969) as the planned three-month time interval between the publication
dates of Parts 25 and 27. The last section, Part 27 (Batrachia plus 18 plates)
was published in March 1831. Cowan (1969) did not indicate when Gray’s
Supplement was published, nor with which Part it appeared. It was certainly
not published in Part 25 in September 1830, when only pp. 1–192 of the main
text were published, and prior to the October date recorded by Gray on his
Synopsis, nor was it published with Part 27 in March 1831 with the Batrachia
and plates. The page header for the first few pages of Gray’s supplement has
“Order Ophidia” printed at the top, and was therefore printed at the same time
as the Ophidia section published in Part 26.
In his later publication, Gray 1831d (Synopsis Reptilium; or Short Descriptions of the Species of Reptiles), dated on p. viii as having been written in
January 1831, on p. 77 Gray referred to the exact pagination for the citation for
Hydraspis lata in the earlier published Griffith version (p. 17 in Gray 1830e).
Therefore, Gray 1830e was available for page-citation in January 1831, and
was therefore published with Part 26, probably in December 1830. Also, in
Gray’s own listing of his publications (Gray 1873k) he recorded 1830 as the
date for publication of his Synopsis in Griffith’s Vol. 9. In addition, Cogger et
al. (1983) recognized 1830 as the date of publication for this work, as do we.
The date of publication of Gray 1831d has always been considered to
be 1831, but the exact date has not been recorded. As noted, it is dated on p.
viii as having been written in January 1831. The back cover of the publication
lists other books by Gray already available for sale at the same time. Listed as
already published are Gray’s Illustrations of Indian Zoology, Parts 1 through
6, with a statement that a total of 20 parts were to be completed, one published
every three months. Sawyer (1953) recorded that Part 1 of Indian Zoology
was published on 6 January 1830, Part 2 on 30 March 1830, Part 3 on 15 July
1830, Part 4 on 6 October 1830, Part 5 on 25 January 1831, Part 6 on 7 April
1831, and Part 7 on 27 July 1831. Gray 1831d was therefore published between
Parts 6 and 7, i.e., between April and July 1831, and we have chosen May as
the probable month of publication.
10:8. Graptemys gibbonsi and G. pearlensis: Ennen et al. (2010) analyzed
morphological and genetic variation in Graptemys gibbonsi throughout its
range and concluded that the western population inhabiting the Pearl River
system of eastern Louisiana and western Mississippi is sufficiently distinct to
warrant description as a full species, Graptemys pearlensis, with Graptemys
gibbonsi restricted to the Pascagoula River system of eastern Mississippi only.
10:9. Pseudemys concinna, floridana, and peninsularis: These three
taxa remain difficult to resolve morphologically and genetically, and their taxonomic relationships have vacillated among various views held by Seidel (1994,
1995), Jackson (1995, 2006), and Thomas and Jansen (2006). Our checklist
has historically listed floridana as a subspecies of concinna, and peninsularis
as a separate species, based primarily on Seidel’s work, and we continue to do
so. However, recent ongoing fieldwork (e.g., Jensen et al. 2008; M. Aresco and
D. Jackson, in litt.) potentially supports recognition of floridana and concinna
as separate species, based on marked differences in their habitat preferences
across wide areas of sympatry, with peninsularis apparently a subspecies of
floridana. In view of the long history of taxonomic uncertainty surrounding
these taxa and the unresolved nature of the data, we now note these conflicting
views by providing alternative listings of floridana as either a subspecies of
concinna or a possible separate species, and peninsularis as either a species or
possible subspecies of floridana. However, we make no actual change in the
taxonomic status of these turtles at this time, but await further field data and
genetic analyses of this difficult species complex, both of which are ongoing.
10:10. Trachemys venusta: McCord et al. (2010) described three new
subspecies of Trachemys venusta based on patterns of head and neck stripes,
carapace and plastron patterns and coloration, plastral scute formulae, maximum
sizes, and distribution. We provisionally list these subspecies pending genetic
analysis.
10:11. Duméril 1805: This reference has historically been given as 1806,
as that date is printed on its frontispiece, but recent work by Gregory (2010),
brought to our attention by R. Bour, has shown that it was actually published
in November 1805.
10:12. Emys, Actinemys, and Emydoidea: Wiens et al (2010) analyzed
multiple mitochondrial and nuclear loci for many emydid terminal taxa. They
considered that their results did not provide phylogenetic support for the placement of Actinemys and Emydoidea in Emys (as recommended most recently
by Spinks and Shaffer 2009); instead, Wiens et al. recommended recognizing
Actinemys and Emydoidea as monotypic genera, with Actinemys apparently
more closely related to Clemmys. We note the discordance among various
published data sets regarding the relationships and analyses of the Emys +
Emydoidea + Actinemys group, and hence, we retain our previous listings
pending greater resolution.
10:13. Emys orbicularis orbicularis: Dubois and Bour (2010a) noted
that Garsault (1764) depicted and named a freshwater turtle as Testudo terrestris, which they identified as an Emys orbicularis (Linnaeus 1758), and
which they considered a subjective junior synonym of E. o. orbicularis based
on geography. Testudo terrestris Garsault 1764 is also a senior homonym of
Testudo terrestris Forskål 1775; however, the latter name has extensive usage
over the past half century, and is a nomen conservandum (ICZN 1963), thus
safeguarding its continuing usage.
10:14. Emys orbicularis galloitalica: Pedall et al. (2011) investigated
genetic differentiation of populations of Italian, Corsican, Sardinian, and Sicilian
Emys orbicularis populations, based on mtDNAand polymorphic microsatellite
loci. They found no significant divergence of Corsican and Sardinian populations from populations of the southwestern Italian mainland, supporting the
view that the subspecies capolongoi (Sardinia) and lanzai (Corsica) described
from these islands are invalid. Their results also suggested that Sardinian and
Corsican populations represented reintroduced populations following earlier
extirpations of native taxa. While Pedall et al. (2011) did not explicitly synonymize capolongoi and lanzai into galloitalica, they indicated synonymization
to be warranted and we consider their results adequate justification to do so.
This leaves the status of the subspecies ingauna, restricted to a small isolated
area in Liguria in the middle of the range of galloitalica, unresolved. Fritz and
Havas (2007:184) noted that ingauna could be synonymous with galloitalica,
but no data-supported analyses of the status of ingauna have apparently been
published since its original description; in the absence of evidence to the contrary, we continue to recognize ingauna as a valid subspecies.
10:15. Emys or Actinemys marmorata: An extensive mitochondrial
and nuclear gene study by Spinks et al. (2010) indicated that southern and
northern lineages, with a zone of contact somewhere in the central Coast Range
of California, exist and may well be diagnosable. However, given the lack of
concordance between the traditionally defined ranges of pallida in the south
and marmorata in the north (as previously recognized subspecies), and with
two nuclear and four mitochondrial lineages identified, we follow Spinks et
al. in waiting for any formal recognition of this variation pending publication
of a much larger, ongoing genetic analysis, and we do not re-elevate pallida
from synonymy at this time.
10:16. Glyptemys muhlenbergii: The name Emys biguttata Say was
previously recorded as being published in 1824; however, though the paper
was read in 1824, it was not published until 1825, in Volume 4, Part 2 of the
Journal of the Academy of Natural Sciences.
10:17. Terrapene: The genus name Cistudo was previously attributed
to Say 1825 as a nomen novum, as also recorded by Fritz and Havas (2007);
however, an Errata sheet published with Say’s article in Volume 4, Part 2 of
the Journal of the Academy of Natural Sciences noted that the name should
have been Cistuda (as originally published by Fleming 1822). However, many
subsequent authors used Cistudo as a valid generic name, often citing Say as
the original author. The first authors to do so appear to have been Duméril and
Bibron (1835), but many others, including Gray (1856b), Agassiz (1857a), and
Boulenger (1889) followed suit.
10:18. Mauremys japonica: Hoogmoed et al. (2010) indicated that this
species name was actually published in 1834 rather than 1835 as previously
recorded by most other authors.
10:19. Chelonoidis carbonaria and C. denticulata: Vargas-Ramirez et
al. (2010a) investigated phylogeographic differentiation in carbonaria and
denticulata, and found distinct mitochondrial clades in carbonaria but only
weak differentiation in denticulata. They recommended further investigation,
but proposed no taxonomic changes at this time.
10:20. Chelonoidis carbonaria: Nowak-Kemp and Fritz (2010) examined
243
244
Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
the holotype of Testudo hercules truncata Gray 1830e in the Bell collection of
the Oxford University Museum and determined that it is a C. carbonaria, rather
than a C. denticulata, where it was previously synonymized as a nomen dubium.
10:21. Chelonoidis chilensis and C. petersi: The validity of the taxa petersi
and donosobarrosi remains subject to debate; the latest contribution to the case
was made by Vinke et al. (2008), who considered petersi synonymous with
chilensis based on syntopic occurrence and lack of consistent morphological
differentiation of populations attributed to petersi vs. chilensis. We consider the
situation unresolved and tentatively retain our recognition of distinct southern
and northern taxa (chilensis and petersi) within the chilensis group, pending
further analysis of range-wide patterns of morphological and molecular variation.
10:22. Gopherus agassizii: Cooper’s description appeared in print in
a section of the Proceedings of the California Academy of Sciences that was
printed and distributed in 1861, not 1863 when the completed volume (including wrappers dated 1863) was issued (R.B. Murphy, pers. comm.). The two
bird species described by Cooper in the same paper (Whitney’s Owl, Athene
whitneyi, now Micrathene whitneyi, and Lucy’s Warbler, Helminthophaga
luciæ, now Vermivora luciae) are consistently attributed to Cooper 1861 in
the ornithological literature.
10:23. Homopus: The date of authorship of the genus Homopus has
traditionally been given as 1835, referring to Tome 2 of Duméril and Bibron
(1835:145). However, Roger Bour (pers. comm.) has drawn our attention to the
fact that the name was actually created and diagnosed in Tome 1 of Duméril
and Bibron (1834:357).
10:24. Homopus signatus: Daniels et al. (2010) investigated systematics
and phylogeography of Homopus signatus using mitochondrial and nuclear
DNA, neither of which supported the recognition of the two traditional subspecies, signatus and cafer. As a result they recommended abandoning subspecies
designations for H. signatus, and we concur.
10:25. Homopus signatus: Our previous checklists and earlier checklists
by other authors have consistently listed Testudo juvencella Gray 1831d:14 as
a junior synonym of this taxon. However, Gray attributed the name to Daudin,
and in fact, it was formally described by Daudin in 1802; hence we correct
this oversight.
10:26. Stigmochelys: Le et al. (2006) proposed placing pardalis in the
genus Psammobates, while Fritz and Bininda-Emonds (2007) argued for
placement in the monotypic genus Stigmochelys. Since then, at least three peerreviewed publications have consciously (i.e. citing Le et al. 2006, among their
references) chosen to use Stigmochelys, as did two ecological papers (which
did not refer to Le et al. 2006) and the CITES Nomenclature Specialist (CITES
2010); in contrast, only a single peer-reviewed paper (Spinks et al. 2009) was
published after 2006 using the combination Psammobates pardalis, and this
was to name the species as an outgroup. Noting the emerging consensus, we
no longer accept Psammobates as an alternative name for Stigmochelys.
10:27. Stigmochelys pardalis: Fritz et al. (2010a) analyzed phylogeographic patterns in mitochondrial DNA and found that seven, largely parapatric,
mtDNA lineages could be identified; these clades did not correspond to the
traditional subspecies (pardalis vs. babcocki) nor to the pronounced geographic
size variation. They concluded that there is no rationale for recognizing subspecies within S. pardalis.
10:28. Testudo graeca buxtoni: The taxon Testudo ecaudata Pallas 1814
has historically been listed in the synonymy of Testudo graeca, and Fritz and
Havas (2007) and our earlier checklists synonymized it under T. g. buxtoni as a
nomen dubium. Wermuth and Mertens (1961) synonymized it under T. g. ibera,
and in their later checklist (Wermuth and Mertens 1977) again under T. g. ibera,
but with a question mark. However, in an overlooked reference, Darevsky and
Mertens (1973) examined the unpublished plate from Pallas (1814) depicting
the type specimen of ecaudata, and determined it to be a variety of the South
African Psammobates tentorius verroxii (Smith 1839), despite the fact that the
specimen was allegedly obtained in the forests of northern Persia along the
Caspian Sea. Despite being a co-author, Mertens was evidently not completely
convinced of the synonymization, and only added a question mark to the name
in his 1977 checklist, while retaining it under the synonymy of T. g. ibera.
Because of the uncertainty of the identification of ecaudata by Darevsky and
Mertens, we solicited input from several specialists regarding the identity of
the figured specimen. Examination by Roger Bour, Ernst Baard, Brian Henen,
Oguz Türkozan, and Jim Buskirk confirmed that it was not a Psammobates, but
a juvenile specimen of a Testudo, most likely referable to T. graeca.
10:29. Forskål or Forsskål: The spelling of the family name of Pehr
Forsskål has varied through the years, with widespread usage of both single
and double ‘s’ spelling (as in our previous checklists). Dubois and Bour (2010a)
declared the “Forsskål” spelling to be a mistake, but whether to spell his name
with a single or double ‘s’ depends on its usage. In the original paper describing
Testudo terrestris and T. triunguis his name is given on the title page in Latin
as Petrus Forskål with one ‘s’, but in his own Swedish vernacular he always
spelled his name as either Petter or Pehr Forsskål, with a double ‘s’. Current
references to his name are reasonably evenly split between the two spellings
(as determined by a search on Google), with the scientific literature preferring
Forskål and the sociohistorical literature preferring Forsskål. We therefore now
spell his name as rendered on the original publication, Forskål, just as we render
the name Carolus Linnaeus in the original published Latin form rather than
the name he was known by in his own Swedish vernacular, Carl von Linné.
10:30. Testudo or Agrionemys horsfieldii: Three new taxa from this
species complex have recently been described by Chkhikvadze and colleagues:
Agrionemys bogdanovi, A. kazachstanica kuznetzovi, and A. kazachstanica
terbishi. In addition, they have elevated two other taxa, A. horsfieldii rustamovi
and Testudo baluchiorum, to species status. The taxonomy of Central Asian
steppe tortoises remains in a state of flux; in contrast to the deeply-dissected
arrangement of several species and subspecies in Agrionemys as proposed by
Chkhikvadze and colleagues based on morphological characters, Fritz et al.
(2009) found phylogeographic structure as evidenced by mtDNA to be in weak
agreement with morphologically-defined taxa, suggested synonymy of rustamovi
with horsfieldii, and recognized only a single species, placed in Testudo, with
implicit recognition of subspecies kazachstanica and an unnamed ESU in
the Fergana valley. To minimize nomenclatural changes in a highly dynamic
situation, and to attempt a middle road between the views of Chkhikvadze et
al. and Fritz et al., we retain the recent arrangement of a single species with
several subspecies, with the newly-described taxa bogdanovi, kuznetzovi, and
terbishi provisionally listed as subspecies of horsfieldii, keeping rustamovi as a
subspecies, and not elevating baluchiorum from synonymy. We anticipate that
further changes will occur in the near future. Additionally, we drop Chersine as
an alternative generic name for the horsfieldii species complex, as all authors
use either Testudo or Agrionemys for this group. This issue was previously
discussed in TTWG 2009.
10:31. Cycloderma aubryi: Duméril (1856) described this species
under the name Cryptopodus aubryi, sp. nov., in his text, but labeled the plate
Cryptopus aubryi. Cryptopodus is considered an ex errore name for Cryptopus
Duméril and Bibron 1835.
10:32. Pelodiscus: Fritz et al. (2010b) carried out preliminary genetic
analyses of softshells of the Pelodiscus sinensis group, and demonstrated the
taxonomic validity and species status of P. maackii; further taxonomic research
was called for to elucidate the status and proper name of the lineages currently
recognized by some as P. axenaria and P. parviformis, as we continue to do
in this checklist.
10:33. Rafetus swinhoei: Le et al. (2010) described a purported new
species of giant softshell, Rafetus vietnamensis, with the type specimen a
complete mounted skeleton located in the Hung Ky pagoda in Hanoi, but
without locality data, while at the same time indicating that R. leloii Ha 2000
was probably not a taxonomically valid description. The study analyzed mostly
the same materials as Le and Pritchard (2009), who reached the opposite
conclusion, that Vietnamese records of Rafetus all pertain to R. swinhoei. A
critical re-assessment of the description of R. vietnamensis is in progress (M.
Le, B. Farkas, pers. comm.). Based on the conclusions of Le and Pritchard
(2009), we provisionally list R. vietnamensis in the synonymy of R. swinhoei.
10:34. Chelodina subgenera: Georges and Thomson (2010) summarized
the history and rationale of grouping species of Australasian snake-necked
turtles and naming these groups, and concluded that the preferred nomenclatural arrangement would be to place all species in the genus Chelodina, while
recognizing three subgenera within Chelodina: subgenus Chelodina for the
narrow-headed species traditionally assigned to ‘Group A’ related to C. longicollis, subgenus Macrodiremys for the single species C. colliei (= C. oblonga
of many authors), and subgenus Macrochelodina for the broad-headed species
of ‘Group B’ related to C. expansa. However, as explained in annotation 39,
the name Macrodiremys may be invalid.
10:35. Chelodina (Chelodina) gunaleni: Georges and Thomson (2010)
considered gunaleni McCord and Joseph-Ouni 2007a as synonymous with
novaeguineae Boulenger 1888b, but provided no data supporting their conclusion; we retain gunaleni as distinct pending further analysis.
10:36. Chelodina (Chelodina) mccordi: Georges and Thomson (2010)
synonymized Chelodina timorensis (McCord et al. 2007a) under C. mccordi
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
(Rhodin 1994b), without recognizing any subspecies of mccordi, instead
referring to the taxa roteensis and timorensis as Diagnosable Terminal Taxa
or ESU’s (Evolutionarily Significant Units). Kuchling et al. (2007) interpreted
these taxa as subspecies of mccordi, and we provisionally agree until further
work clarifies their taxonomic status. The CITES Nomenclature Specialist
(CITES 2010) also synonymized timorensis under mccordi.
10:37. Chelodina (Macrochelodina) rugosa: Georges and Thomson
(2010) reiterated their earlier (Georges and Thomson 2006) synonymy of C.
kuchlingi Cann 1997d into C. rugosa Ogilby 1890, and we now synonymize
it as well.
10:38. Chelodina (Macrochelodina) walloyarrina: Georges and Thomson
(2010) treated walloyarrina McCord and Joseph-Ouni 2007b as synonymous
under burrungandjii Thomson et al. 2000, referring to morphological and molecular information available in the literature, and their own research indicating
hybridization and introgression of burrungandjii with rugosa. However, we
provisionally retain walloyarrina as distinct until published molecular data
resolves the issue.
10:39. Chelodina (subgenus name undetermined) oblonga or colliei:
Thomson (2006), Georges and Thomson (2010) and Kuchling (2010) summarized the convoluted nomenclatural history of the name Chelodina oblonga
Gray 1841, and attributed it to the northern populations (currently named
C. rugosa Ogilby 1890, a name provisionally retained by them) as a senior
synonym, pending the outcome of ICZN Case 3351 (Thomson 2006). With
the name oblonga thus potentially unavailable for the snake-necked turtle of
southwestern Western Australia, they referred to this turtle either by its next
available name, Chelodina colliei Gray 1856a [Georges and Thomson], or C.
oblonga (= C. colliei) [Kuchling] to maintain prevailing usage. In an attempt
to retain traditional usage of the name oblonga for the southwestern snakeneck,
McCord and Joseph-Ouni (2007a) had designated a neotype for oblonga Gray
1841, being the lectotype of C. colliei. However, as a type specimen already
exists for oblonga, their neotype designation is invalid (Kuchling 2010).
There is general agreement that the southwestern snakeneck represents
a distinct lineage within the Chelodina group (Burbidge 1967, Goode 1967,
Kuchling 1988, Georges and Adams 1992, Georges et al. 1998, 2002), warranting recognition at subgenus level alongside Macrochelodina and Chelodina
sensu stricto. The genus name Macrodiremys was created by McCord and
Joseph-Ouni (2007a) to recognize this lineage. They designated oblonga as
its type species, in the mistaken belief that their neotype designation had fixed
the name oblonga to the southwestern taxon. However, with oblonga being
applicable to the northern taxon (rugosa) by virtue of its original valid holotype,
Macrodiremys becomes a junior synonym of Macrochelodina, in contrast to
the intent of McCord and Joseph-Ouni. Kuchling (2010), as first reviewer,
listed possibilities under which articles of the Code the name Macrodiremys
could potentially be fixed to the southwestern longneck, but stated his conviction that it was reasonable to await an opinion on case 3351 of the ICZN
plenum to be published before such nomenclatural acts should be attempted,
and expressed his hope that in their ruling on case 3351, the ICZN plenum
would take suitable action to solve these problems. Independently, Georges
and Thomson (2010) made an explicit attempt to correct the error and follow
the intent of McCord and Joseph-Ouni 2007a under Article A 67.13.1 of the
Code by correcting the type species of Macrodiremys to Chelodina oblonga
McCord and Joseph-Ouni 2007a = C. colliei Gray 1856a, making it potentially
available as a genus-group name for the southwestern snakeneck. However,
Kuchling (2010) had already pointed out the possibility that a nomenclatural
act such as the attempted correction by Georges and Thomson (2010) could
be invalid under the ICZN Code.
Further nomenclatural acts in this group of animals must await the outcome of the anticipated ICZN plenary decision to avoid further complicating
this issue. For that reason, we list Macrodiremys as a synonym of Chelodina
sensu lato and we do not employ a subgenus designation for the southwestern
species in this year’s checklist, recognizing well that the southwestern snakeneck warrants its own subgenus and that a name is potentially available for it.
10:40. Elseya novaeguineae: Georges and Thomson (2010) returned
E. schultzei Vogt 1911 to the synonymy of E. novaeguineae Meyer 1874, and
also tentatively placed E. novaeguineae in the genus Myuchelys. However, we
retain the species in Elseya pending further genetic analysis.
10:41. Emydura m. macquarii: Georges and Thomson (2010) reviewed
the complicated history of the name Hydraspis australis Gray 1841, and supported the conclusion of Cogger et al. (1983) that the name is a junior synonym
of E. m. macquarii. The name australis had previously been inconsistently
used for a variety of Emydura populations in northern Australia (e.g., Cann
1998); Georges and Thomson (2010) instead used the name victoriae for some
of these populations. We interpret the name australis as a nomen dubium and
agree with its synonymization under macquarii.
10:42. Emydura m. macquarii: Georges and Thomson (2010) synonymized the previously recognized subspecies E. m. binjing, E. m. dharra, E. m.
dharuk, E. m. gunabarra, and E. signata, into typical E. m. macquarii, based
on lack of distinction based on allozyme electrophoresis, and they attributed
variation in shell shape and body size to phenotypic plasticity. They considered that the Bellinger River population of Emydura macquarii is genetically
unremarkable and that it was almost certainly established by introduction of
animals from adjacent drainages.
10:43. Myuchelys and Wells taxa: Thomson and Georges (2009)
described the new genus Myuchelys and noted that the name Wollumbinia
Wells 2007c, used for the same group of species, but proposed online in an
unpublished web-based document without adequate hardcopy dissemination,
is nomenclaturally unavailable. They also considered all web-based names
proposed by Wells in his Australian Biodiversity Record (Wells 2002, 2007a,b,c,
2009), as unpublished and nomenclaturally unavailable. In previous editions
of this checklist we included Wollumbinia as an alternative genus name and
listed some of the species described in the other documents; however, based on
arguments against availability of all these names proposed by Wells (Fritz and
Havas 2007, Thomson and Georges 2009, Georges and Thomson 2010), we now
concur in considering these names unavailable under the ICZN Code and treat
each of them as an unavailable name (nomen illegitimum) placed in synonymy.
10:44. Phrynops geoffroanus and tuberosus: The taxonomic status and
distribution of these two taxa remains problematic, without clear consensus of
taxon limits or range delimitations. Pritchard and Trebbau (1984) documented
that tuberosus is isolated from the broad range of geoffroanus and restricted
to a small upland area in eastern Venezuela, extreme western Guyana, and
northern Roraima in Brazil. Métrailler and Le Gratiet (1996) documented
that neither tuberosus nor geoffroanus occurs in French Guiana, and neither
taxon has ever been recorded in Suriname or lowland Guyana. However, McCord et al. (2001), based on an examination of three specimens of what they
identified as tuberosus from eastern Venezuela, the Brazil–Guyana border, and
Piauí, northeastern Brazil, depicted large apparently well-defined allopatric
ranges for both taxa, with tuberosus supposedly distributed throughout the
Guyanan lowlands, the lower Brazilian Amazon, and northeastern Brazil.
Rueda-Almonacid et al. (2007), while acknowledging that the systematics of
the Phrynops geoffroanus complex (Rhodin and Mittermeier 1983) remained
controversial and unresolved, nevertheless reproduced the vastly different
ranges of tuberosus and geoffroanus depicted by McCord et al. We choose
at this time to instead recognize the earlier documented restricted view of the
distribution of tuberosus, since it is based on more extensive fieldwork, while
also noting that the P. geoffroanus complex remains in need of a thorough
range-wide phylogeographic study.
10:45. Pelomedusa subrufa: Phylogeographic patterns of mitochondrial
and nuclear DNA variation in Pelomedusa subrufa were analyzed by VargasRamírez et al. (2010b) and Wong et al. (2010). Both studies reached the same
conclusion that the species is structured into three major clades: northwestern,
eastern, and southern lineages, with evidence that Madagascan populations
may have been introduced there in prehistoric times. Vargas-Ramírez et al.
(2010b) suggested that valid taxonomic units may currently be included in
the synonymy of Pelomedusa subrufa, and recommended further study to
clarify the taxonomy.
10:46. Podocnemis erythrocephala: We previously listed the name
Podocnemis agassizi Coutinho 1868 as a synonym, as did other earlier
checklists. However, although Coutinho described a new species of turtle in
1868 that he proposed to name in honor of “M. Agassiz”, no formal name was
created at that time. Later, Göldi (1886) translated and reprinted Coutinho’s
earlier writings and created the binomen Podocnemis agassizii, attributing it to
Coutinho, but recommending that it be changed to Podocnemis coutinhii since
the name agassizii had already been used for another turtle, Chelonia agassizii.
10:47. Podocnemis sextuberculata: Coutinho (1868) provided an
excellent description of this species that he named Podocnemis pitiu, a name
overlooked in our previous checklists as well as other earlier checklists.
10:48. Podocnemis unifilis: We continue to use this name for the Yellowspotted River Turtle, as it has a long and continuous usage, although the name
Emys cayennensis Schweigger 1812 has recently been shown to have priority
based on the available type material (Bour 2006). However, the name cayennensis
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was for a long time erroneously applied to Podocnemis erythrocephala and it
would introduce unacceptable confusion to use that name for what has nearly
always been referred to as unifilis. We therefore maintain prevailing usage of
unifilis and retain the name cayennensis in synonymy as a provisional nomen
rejectum pending petition to the ICZN (R.C. Vogt et al., in prep.).
10:49. Podocnemis unifilis: Coutinho (1868) provided an excellent description of this species that he named Podocnemis tracaya, a name overlooked
in our previous checklists as well as other earlier checklists.
2011 Checklist Annotations
TTWG 2011 (000.165-242.checklist.v.4) (11:5-20)
11:5. Malaclemys terrapin: Parham et al. (2008) demonstrated, based
on radiometric dating of a fossil found in a cave on Bermuda, that the small
population of diamondback terrapins present there was the result of a natural
dispersal event dating from before human presence on the island. As a result, we
now consider the Bermuda population as native rather than introduced. Genetic
comparison of Bermuda samples with U.S. populations also demonstrated
closest similarity to samples from the Carolinas, so we provisionally list the
Bermuda population under the subspecies M. t. centrata pending further studies.
11:6. Genus Trachemys: Fritz et al. (2011c) assessed the phylogeny and
taxon boundaries of Trachemys from Central and South America based on
mitochondrial and nuclear DNA. Their nuclear data were largely uninformative, but based on their mitochondrial and combined analyses, they concluded
that the Pacific Coast taxa grayi, emolli, and panamensis form a distinct clade
that warrants recognition at the species level, as Trachemys grayi, with three
subspecies. Fritz and co-authors also recommended combining the taxa cataspila, ornata, venusta, callirostris, and chichiriviche as subspecies under a
single species, Trachemys ornata. They further proposed placing adiutrix at
subspecies level in dorbigni, and considered the taxon uhrigi to be synonymous with typical venusta. Their analysis did not include samples of the taxon
iversoni, whose range on the Yucatan Peninsula is embedded in the range of
the redefined ornata group, nor did they sample most Trachemys taxa from
the arid western region of Mexico.
While Fritz et al. (2011c) provide a useful set of hypotheses for future
testing, we recognize that taxonomy of Trachemys will remain dynamic and
expect further changes when additional nuclear data, and deeper geographic
sampling of field-verified specimens, are brought to bear on the problem. We
therefore provide the alternative taxonomies of Seidel (2002) and Fritz et al.
(2011) in this year’s checklist.
11:7. Emys/Emydoidea/Actinemys: Fritz et al. (2011b) reviewed competing generic concepts for Blanding’s, Western Pond, and European Pond turtles,
and reiterated their support for recognition of Emydoidea and Actinemys as
separate genera distinct from Emys.
The TTWG members continue to be deeply divided in their perspectives
on whether to recognize three genera, or a single genus, for the four species
concerned (blandingii, marmorata, orbicularis, and trinacris). The challenge
is that available primary research findings result in different relationship trees
among the four, based on mitochondrial and nuclear DNA and on morphology
(Spinks and Shaffer 2009; Wiens et al. 2010; Fritz et al. 2011b). Overall, the
majority of data indicate that blandingii, marmorata, orbicularis, and trinacris
collectively are each others’ closest relatives. Whether to recognize this by
combining the four species in an expanded genus Emys, or to emphasize the
morphological distinctiveness of blandingii and marmorata from (orbicularis
+ trinacris) by recognizing Actinemys and Emydoidea as monotypic genera,
is largely a subjective matter. Correspondingly, the herpetological taxonomic
community and recent scientific literature have not come to a clear consensus
or prevailing usage. With further research in progress, we agree that the situation remains undecided, and continue to present alternative taxonomies in
this year’s checklist, with no implication that this represents the preferred or
supported arrangement of individual TTWG members.
11:8. Terrapene carolina: Butler et al. (2011) conducted morphological and molecular analyses to address the status of lineages within Terrapene
carolina. They found that box turtles phenotypically corresponding to T. c.
carolina, T. c. bauri, and T. c. triunguis all occur within the range of T. c. major,
and that the latter does not demonstrate a diagnosable morphology. They also
found that carolina, bauri, and triunguis possess divergent mtDNA haplotypes,
which are present alongside a fourth, distinct, haplotype in the range of T. c.
major. Butler et al. interpreted these findings as the introgressed genetic signal
of the extinct Pleistocene T. c. putnami perpetuating in a morphologically mixed
population, and advocated equating the taxon major with putnami; they argued
that major, which has precedence over putnami, should only be used to refer
to the large extinct form.
Their genetic analysis also placed bauri as sister to ornata, and triunguis as
sister taxon to the [ornata + bauri] clade. Butler et al. thus suggested that bauri
should be elevated to species status, although this would leave the remaining
taxa in T. carolina paraphyletic by continued inclusion of triunguis. Butler et
al. did not address the status of mexicana and yucatana, which have variously
been considered subspecies of carolina or full species each. Furthermore,
their analysis did not support the previously recognized species groups (e.g.,
Milstead 1969; Spinks et al. 2009).
We are reluctant to change the widely-recognized taxonomic arrangement for a species complex of notable conservation and legislative significance
based on a single study that relied heavily on a short mtDNA segment. Until
the alternative taxonomy presented by Butler et al. is independently corroborated by further research, and potentially finds widespread acceptance in the
herpetological and taxonomic communities, we prefer to retain the traditional
arrangement and defer possible adoption of their hypothesis until a later version of this checklist.
11:9. Cuora flavomarginata evelynae: In previous checklists we had
overlooked that Ota et al. (2009) had transferred evelynae back to subspecies
status under Cuora flavomarginata, but we now follow them here.
11:10. Aldabrachelys gigantea or Dipsochelys dussumieri: Gerlach
(2011a) studied morphological development of juveniles of the three forms of
Indian ocean giant tortoises reared under identical captive conditions, and concluded that animals consistently develop into the morphotypes characterized by
their parents. Gerlach thus concluded that the morphotypes cannot be explained
by environment alone, and may have a genetic basis, supporting the recognition
of arnoldi and hololissa as taxonomically distinct from gigantea/dussumieri.
11:11. Gopherus agassizii and G. morafkai: Murphy et al. (2011) investigated taxonomic problems affecting the Desert Tortoise. They designated
the sole remaining of three syntypes of Gopherus agassizii as lectotype and
genetically confirmed that it originated from California. They also determined
that the holotype of G. lepidocephalus originated from the Mojave desert
population, at least based on mitochondrial DNA, reconfirming lepidocephalus
as a junior synonym of agassizii. A suite of morphological, molecular and
ecological differences between the Mojave and Sonoran Desert populations
led Murphy and co-authors to describe the Sonoran form, long recognized as
an Evolutionarily Significant Unit, as a new and distinct species, Gopherus
morafkai. Further research will be necessary to determine if the southernmost
populations of G. morafkai in the Sinaloan thornscrub ecosystem also deserve
taxonomic recognition (Lamb et al. 1989; Murphy et al. 2011).
11:12. Testudo graeca and T. marginata: Chkhikvadze et al. (2011)
described Testudo dagestanica from Lake Papas, Dagestan, and considered
the taxa anamurensis, pallasi, terrestris, weissingeri, and zarudnyi (variously
considered subspecies or synonyms of T. graeca or T. marginata in earlier
versions of this checklist) as valid subspecies of T. marginata. However,
Parham et al. (2006) and Fritz et al. (2007) placed the Dagestan population
of Testudo firmly in T. graeca based on a combination of morphological and
genetic characters, although those studies differed in their assignment of this
population to T. g. ibera and T. g. armeniaca, respectively. Until further data
become available supporting the radical changes proposed by Chkhikvadze
et al., we take a conservative approach and make no changes to the taxonomy
of T. graeca or T. marginata.
11:13. Testudo marginata: Perez et al. (2012) studied the effects of
landscape features and demographic history on the genetic structure of Testudo
marginata using microsatellites. They found that their samples from Sardinia
clustered with samples from northern Greece, suggesting that the Sardinian
population may have originated from a small founder population approximately 200 generations ago, while the source population from which those
founders originated was estimated to be very large. In contrast, the samples
from the ‘dwarfed’ marginata population of the southwestern Peloponnesus
demonstrated a low but significant differentiation from all other marginata
populations. While Perez et al. did not recommend taxonomic recognition of
the dwarf population, they did emphasize the conservation significance of what
in effect is an Evolutionarily Significant Unit (ESU).
11:14. Genus Lissemys: Praschag et al. (2011) analyzed variability across
the range of the genus Lissemys, based on 2286 bp of mitochondrial DNA
sequences, with additional morphological and biogeographical considerations.
They concluded that scutata is a distinct divergent lineage and reaffirmed Webb’s
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
(1982) conclusion to recognize scutata as a full species. They found that the Sri
Lankan population shows similar divergence, and recognized this population as
a full species, Lissemys ceylonensis (Gray 1856a). The remaining populations
of Lissemys fell into three lineages, one broadly corresponding to the spotted
northern taxon andersoni (though with some intergradation with unspotted
animals in Orissa) inhabiting the Indus-Ganges-Brahmaputra systems, and
the other two lineages comprised of unspotted populations in peninsular India.
Praschag et al. recommended recognition of the two peninsular lineages by the
names punctata for the southernmost Indian lineage, and vittata for the central
Indian lineage. Because the three mainland Indian lineages are more closely
related to each other than to either scutata or ceylonensis, and intergradation
between at least andersoni and vittata is known, Praschag et al. proposed recognition of vittata and andersoni at subspecies rank under punctata. They also
concluded that Emyda granosa intermediaAnnandale 1912, traditionally placed
in synonymy of the southern, unspotted form punctata (see Webb 1980a:554),
is based on intergrades between vittata and andersoni and cannot be used as
the valid name for any Lissemys taxon. As intermedia is clearly not a valid
synonym of andersoni, and its type locality (Purulia, western West Bengal) is
far outside the redefined range of L. p. punctata, we take this opportunity to
place intermedia into the synonymy of the central unspotted taxon L. p. vittata.
11:15. Genus Nilssonia: In earlier versions of this checklist, we used
the provisional designation ‘Nilssonia or Aspideretes’ as a transitional phase
between the widespread usage of Aspideretes since Meylan’s (1987) morphological analysis, and the more recent findings that Aspideretes is paraphyletic
with regard to Nilssonia and consequent recommendation to synonymize
Aspideretes into Nilssonia (Engstrom et al. 2004; Praschag et al. 2007; Fritz
and Havas 2007). We note that the species gangetica, hurum, leithii, and nigricans are now widely accepted as belonging in the genus Nilssonia, making
Aspideretes a synonym.
11:16. Genus Pelodiscus: Yang et al. (2011) evaluated the validity of P.
parviformis and inferred it to be a distinct species based on the results of their
morphological and molecular analyses. Stuckas and Fritz (2011) sequenced
DNA from the lectotype of P. sinensis and analyzed its placement in relation
to other Pelodiscus. They found it distinct from the lineages identified as P.
axenaria and P. parviformis, and concluded that P. sinensis is not a senior
synonym of either of these two names. Consequently, Stuckas and Fritz
proposed recognition of axenaria, maackii, parviformis, and sinensis as valid
species; they recognized that older names may be available for some of these,
but recommended use of these four names for the time being. The respective
distribution ranges of the different forms, and possible areas of co-occurrence,
remain unclear; the reported occurrence of both axenaria and parviformis in
Guangxi and Hunan warrants further research.
11:17. Rafetus swinhoei: Farkas et al. (2011) reviewed the description
of Rafetus vietnamensis Le et al. 2010. They declared vietnamensis an objective synonym of R. leloii, and reasserted their view that R. leloii (and thus R.
vietnamensis) is a subjective synonym of R. swinhoei.
11:18. Trionyx triunguis: Gidis et al. (2011) sequenced up to seven genes
of 20 known-locality samples and reported shallow divergence among Mediterranean Coast, Nile, and Cameroon samples, in contrast to the results of Guçlu et
al. (2009), who found four different haplotypes for the four unknown-locality
Sub-Saharan specimens that they compared to their Mediterranean samples.
11:19. Pelomedusidae: Fritz et al. (2011a) examined the phylogeny
of Pelusios and Pelomedusa species based on three mitochondrial and three
nuclear DNA fragments. They reported divergent lineages within Pelusios
rhodesianus and P. sinuatus, found no clear differentiation of P. chapini from
P. castaneus, and attributed the nesting of the sole P. williamsi sequence within
P. castaneus to misidentification of the williamsi sample in GenBank. They also
demonstrated very shallow divergences within P. castanoides, suggesting that
populations in Madagascar and the Seychelles were only recently colonized;
these findings agreed with those of Silva et al. (2010), who found limited mtDNA
differentiation of Seychelles from Madagascar specimens of P. castanoides,
but cautioned that their sampling of Madagascar material was limited. Fritz
et al. (2011a) did not propose explicit changes to taxonomy, but indicated that
chapini could be re-instated to subspecies rank under P. castaneus, and that the
recognition of subspecies of P. castanoides could be unwarranted; they also
suggested the existence of cryptic taxa within P. rhodesianus and P. sinuatus,
and reaffirmed the view of Vargas-Ramirez et al. (2010) that Pelomedusa
subrufa represents a species complex.
11:20. Podocnemididae: Gaffney et al. (2011) analyzed the phylogeny
of fossil and living species of Podocnemididae in the context of their earlier
analysis of extinct related groups (Gaffney et al. 2006). They reconfirmed the
monophyly of the family Podocnemididae, and largely agreed with França and
Langer (2006) in not recognizing Erymnochelyinae as a subfamily.
2012 Checklist Annotations
TTWG 2012 (000.243-328.checklist.v.5) (12:6-44)
12:6. Testudines: Crawford et al. (2012) and Lourenço et al. (2012)
analyzed the placement of turtles in a wider context based on molecular phylogeny, and each team concluded that their data provided strong support for
turtles being the sister group to Archosauria (i.e., Crocodilians + Birds, and
extinct related groups), rejecting hypothesized relationships of turtles as sister
group to Lepidosauria (lizards, snakes, and tuataras), or as the most basal branch
of the reptilian (including birds) lineage. Lourenço et al. (2012) estimated the
divergence of turtles and archosaurs as dating back to the late Permian around
255 million years ago (MYA), and dated the divergence between Cryptodira
and Pleurodira at about 157 MYA in the late Jurassic. Guillon et al. (2012)
analyzed mitochondrial and nuclear DNA sequences available in GenBank,
including whole genomes for a few species, and constructed a phylogeny for
the group involving 230 turtle species representing all families and nearly all
genera. Their results reconfirmed the monophyly of Testudines and of Pleurodira
and Cryptodira, added support for the placement of Platysternon among
Testudinoids, placed Dermatemys in the Kinosternoids, placed Trionychoids
(Trionychidae + Carettochelyidae) as sister to all other Cryptodires (a finding
shared with Lourenço et al. [2012] and Wang et al. [2012]), and indicated the
need for additional research to better resolve various groups of chelids and
testudinids at the genus level.
12:7. Cheloniidae: The recognition of higher taxa within the Family
Cheloniidae has been inconsistent, at least since Gray’s (1825) recognition of
an imprecisely defined Carettidae. The inconsistency has been due primarily
to the uncertainty regarding the phylogenetic relationships of Natator and
Eretmochelys, and the relationships of fossil to living cheloniids. However, in
the midst of the uncertainty of most of the relationships within this family, the
close (sister) relationship of Caretta and Lepidochelys has not been disputed at
least since Deraniyagala (1934), who initially argued for the recognition of the
latter as the Carettidae, with the remaining taxa in the Cheloniidae. By 1952,
Deraniyagala had changed his position to subfamilial recognition of these two
groups. Some authors have followed this latter arrangement of recognizing
these groups at subfamily rank (e.g., Mlynarski 1976; Pritchard and Trebbau
1984; Smith and Smith 1979), but most have followed Zangerl and Turnbull
(1955) and Zangerl (1958) in recognizing these groups as two tribes within
the Cheloniidae: Carettini (including Caretta and Lepidochelys) and Chelonini
(including the remaining taxa: Chelonia, Natator, and Eretmochelys).
Impetus for continuing to recognize the Carettini as a tribe (rather
than a subfamily) emerges from 1) the strong support for Caretta as sister to
Lepidochelys (virtually every study since Deraniyagala); 2) the controversy
about the phylogenetic positions of Natator and Eretmochelys (i.e., the possible paraphyly of the living Cheloniinae or Chelonini; review in Bowen et
al., 1993); and 3) the uncertain phylogenetic relationships of numerous fossil
cheloniid taxa, rendering any hierarchy likely to introduce paraphyly (e.g., see
Parham and Fastovsky 1997).
Fortunately, four recent publications, by Naro-Mariel et al. (2008; based
on two mitochondrial and five nuclear genes), Parham and Pyenson (2010; based
on osteology), Duchene et al. (2012; based on the entire mitogenome), and Guillon et al. (2012; based on all available GenBank sequences), seem to reveal the
relationships among the living genera quite definitively. These studies clearly
resolved Natator as sister to Chelonia, and Eretmochelys as sister to (Caretta +
Lepidochelys), with each of these two clades being reciprocally monophyletic.
However, Duchene et al. (2012) explicitly recognized these clades as subfamilies,
whereas Naro-Maciel et al. (2008) explicity referred to them as tribes; Parham
and Pyenson (2010) defined the tribe Carettini for (Caretta + Lepidochelys) only,
but did not use or define group names for other groups below family level, while
Guillon et al. (2012) took no position. In an effort to recognize recent research
consensus, to promote stability, and until the relationships among fossil cheloniid
taxa are better resolved, we here recognize the two living clades as subfamilies
(Carettinae and Cheloniinae), and hope that this will stimulate further research
and discussion of the phylogeny of living and extinct cheloniid sea turtles.
12:8. Cheloniidae: Vilaça et al. (2012) reviewed the occurrence of
natural interspecific hybrids among marine turtle species, and using nuclear
markers demonstrated that hybridization among marine turtle species is very
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common along the Brazilian coast. Most of the hybridization involves male
Eretmochelys and female Caretta, but problematic introgression is occurring
among all four genera Caretta, Eretmochelys, Lepidochelys, and Chelonia.
Vilaça et al. hypothesized that the incidence of this hybridization may have
escalated only about 40 years ago, and may be the result of overhunting and
local warming of the beaches due to coastal deforestation.
12:9. Cheloniidae and Dermochelyidae: Duchene et al. (2012) studied
variation across the entire mitochondrial genome of all seven living marine
turtle species, and demonstrated divergent intraspecific haplotype clades in the
Pacific versus Atlantic and Indian Ocean basins for Eretmochelys imbricata,
Chelonia mydas, and Dermochelys coriacea. However, they made no recommendations regarding intraspecific taxonomy.
12:10. Chelonia mydas: Shamblin et al. (2012) demonstrated distinctive
mitogenomic haplotype frequencies among the nesting populations of Green
Turtles at Buck Island (US Virgin Islands), Aves Island (Venezuela), Suriname,
and Tortuguero (Costa Rica), and recommended that these populations receive
separate management unit status. However, although these populations are genetically distinct, the authors made no recommendations for taxonomic changes.
12:11. Kinosternon subrubrum steindachneri: Bourque (2012a, 2012b)
analyzed the phylogenetic placement of two new fossil Kinosternon taxa, and
in the course of his morphological analysis found that Kinosternon subrubrum
steindachneri was placed as sister taxon to the (K. subrubrum + K. baurii) group;
consequently he suggested (2012a) and then elevated (2012b) steindachneri
to full species rank. Regrettably, he did not specify whether his data for K.
subrubrum were based on characters of K. s. subrubrum, K. s. hippocrepis,
or a combination of these. The extreme morphological similarity of baurii to
hippocrepis was already noted by Iverson (1992), and we provisionally retain
steindachneri at subspecies rank under subrubrum pending further analysis.
12:12. Chrysemys: Gemel and Grillitsch (2008) reported that Wagler
(1821) had nomenclatorally occupied the genus name Hydrochelys for the species Testudo picta (now Chrysemys picta), and explicitly qualified Hydrochelys
Wagler 1821 as a nomen oblitum and invalid, and qualified Chrysemys Gray
1844 as nomen protectum and valid.
12:13. Chrysemys picta: The description of Testudo picta has generally
been attributed to Schneider (1783). However, careful reading of his work
indicates that the description of picta was based on descriptive information
contained in letters from Johann Hermann of Strasbourg, and shows no indication that Schneider had access to the actual specimen, instead adding information from Hermann’s letters for the sake of completeness of his (Schneider’s)
monograph of turtles.
Article 50.1.1 of the Code (ICZN 1999) states “However, if it is clear
from the contents that some person other than an author of the work is alone
responsible both for the name or act and for satisfying the criteria of availability
other than actual publication, then that other person is the author of the name
or act.” In the case of picta, it is not evident how much Schneider was directly
quoting from Hermann’s writings, so we continue to attribute authorship of the
name to Schneider, rather than ‘Hermann in Schneider 1783’.
As an aside, we note that Hermann used different spellings for his
surname over time, involving single or double 'r' and single or double 'n', as
well as using French (Jean) and German (Johann or Johannes) versions of his
given name. We elect to use the spelling 'Hermann' as that matches the spelling
on his death certificate (indicating his preferred spelling later in life), and the
spelling employed by Gmelin six years later for the tortoise named for him,
Testudo hermanni.
12:14. Graptemys: Freedman and Myers (2012) identified and sequenced
a species-specific mitochondrial control region marker and two nuclear markers
in a population of Graptemys pseudogeographica that had hybridized historically
with G. geographica, after which geological events prevented further inbreeding for several generations. Both mitochondrial and nuclear introgression was
documented. They provided additional evidence for hybridization events among
G. pseudogeographica, G. ouachitensis, and G. geographica in other parts of
the range, and combined with literature records of Graptemys hybridization
events in captivity, concluded that reproductive isolation in Graptemys has
evolved incompletely. In the context of conflicting evidence for monophyly
vs. paraphyly of G. pseudogeographica and G. ouachitensis, they suggested
that the two groups may maintain reproductive isolation in parts of their range
while interbreeding in other areas.
12:15. Graptemys ouachitensis: Brown et al. (2012) examined mitochondrial DNA control region sequence variation throughout the range of the
subspecies ouachitensis, and found 18 haplotypes forming two minimally
divergent groups. They also found modest divergence between o. ouachitensis
and o. sabinensis, and slightly greater divergence between ouachitensis and
pseudogeographica. They acknowledged earlier studies suggesting that sabinensis might warrant species status, and noted the need for additional studies
to clarify the relationship among these taxa.
12:16. Pseudemys: Using mitochondrial DNA, Jackson et al. (2012) inferred phylogenetic relationships of the taxa within Pseudemys. While recovering
the genus as a strongly supported monophyletic group, they found no support
for monophyly of the traditionally recognized redbelly (rubiventris) and cooter
(concinna) species groups. They also did not find great support for monophyly
of individual taxa other than gorzugi and texana, suggesting mitochondrial
introgression since the glacial retreat in the Pleistocene, or recent speciation, as
possible explanations. They made no specific recommendations for taxonomic
changes based on their results, and indeed cautioned against making taxonomic
changes within this highly complex group without comprehensive data.
12:17. Trachemys taylori: McGaugh (2012) analyzed population genetic
structure of Trachemys taylori, including a detailed genetic assessment of
potential hybridization, and found significant genetic differentiation among
populations, consistent with population bottlenecks, but detected no genetic
evidence for hybridization of T. taylori with invasive T. scripta elegans.
12:18. Trachemys venusta: Perry (1810) published a description of
Testudo panama, a name overlooked and/or ignored ever since, until Bauer
and Petit (2004) discussed it. They attributed it to the genus Trachemys based
on the illustration and text description, and based on provenance and color
pattern, regarded Testudo panama as referring to Trachemys venusta (Gray
1856b). Invoking ICZN Article 23.9, Bauer and Petit (2004) asserted that
Testudo panama Perry 1810 was to be regarded as a nomen oblitum and invalid,
and Emys venusta Gray 1856b as a nomen protectum whenever the two are
considered as synonyms.
12:19. Emys orbicularis hellenica: Schreiber 1875, in his section on
Cistudo lutaria, listed and defined eight varieties, including var. h), to which
he attributed “Emys Hoffmanni Fitzinger Mus. Vindob.” Our previous inclusion in earlier checklists of Emys orbicularis hoffmani [sic] Schreiber 1875 as
a synonym of E. o. hellenica referred in fact to a subsequent use of Fitzinger’s
(1835) name Emys (Emys) hofmanni (with a minor spelling difference) rather
than to a new taxon description, and as such we now delete it from our listing
of synonymized primary taxa.
12:20. Terrapene ornata: Joyce et al. (2012) described a new fossil box
turtle and carried out a phylogenetic analysis of extant and fossil box turtle
species based on morphological and osteological characters. The fossil taxon
T. longinsulae Hay 1908, from an undefined location in the general region
of Long Island, Kansas (possible age ranging from Miocene to Pleistocene;
see Joyce et al. 2012:185), scored identical for all available characters to the
character states shown by both living T. ornata ornata and T. o. luteola, including two unique shared characters. They therefore placed T. longinsulae in the
synonymy of T. ornata.
12:21. Geoemydidae: The deep divergence of living New World from
Old World geoemydids has been well-established based on mitochondrial and
nuclear DNA data (Spinks et al. 2004; Le and McCord 2008; Guillon et al.
2012), and has recently been dated at 57 mya (Lourenço et al. 2012). Le and
McCord (2008) first proposed the recognition of living New World taxa (genus
Rhinoclemmys) as the subfamily Rhinoclemminae. Given that the divergence
of the latter is as old as, or older than, that of the widely accepted emydid subfamilies (dated at 37 mya by Dornburg et al. 2011; 57 mya by Lourenço et al.
2012), we support the subfamilial status of the genus Rhinoclemmys. However,
we amend Le and McCord’s name to Rhinoclemmydinae to correspond to the
proper case for its ending -emys (genitive emydis), and we note that the group
may also include North American fossil species of the genera Echmatemys
(Hervet 2004) and Bridgeremys (Hutchison 2006). Correspondingly, we apply
the subfamily name Geoemydinae to all Eurasian geoemydid taxa.
12:22. Cuora: Phylogeny and species boundaries were studied for the
genus Cuora by Spinks et al. (2012a) using a combination of mitochondrial
and nuclear genes. While noting discrepancies between the results of the
mitochondrial and nuclear data sets, concordance with traditional, morphologybased phylogenies for Cuora was greatest with the nuclear dataset, which they
considered to represent the most reliable estimates of phylogeny and species
boundaries. The species amboinensis, flavomarginata, mouhotii, and yunnanensis emerged clearly resolved, whereas the trifasciata and galbinifrons taxon
clusters were less well resolved. Cuora aurocapitata and C. pani showed signs
of introgression of trifasciata mtDNA, but were reciprocally monophyletic
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
based on nuclear markers. The position of animals attributed to 'cyclornata'
was highly variable depending on what markers were analyzed, a result that
was interpreted as being most consistent with introgression, and leading to the
recommendation to consider animals with 'cyclornata' phenotype as part of
C. trifasciata rather than a distinct separate taxon. Cuora zhoui emerged as a
strongly divergent species based on nuclear DNA, but its mtDNA association
with trifasciata may indicate mitochondrial introgression. The monophyly of
C. mccordi was well supported, though its phylogenetic position was different
when nuclear and mtDNA datasets were analyzed. While some doubts were
expressed about the species status of mccordi, the authors recommended
continued treatment as a valid species for the time being. The members of the
galbinifrons group (bourreti, galbinifrons, and picturata) were strongly supported as a clade of three genetically and morphologically diagnosable taxa, and
while some potential gene flow was detected between these taxa, the authors
recommended that they continue to be recognized as valid separate species.
12:23. Cuora amboinensis: Ernst et al. (2011) evaluated the potential to
use shell morphology and postorbital stripe to separate the subspecies amboinensis and kamaroma in the Philippines, as alternative approaches to the usual
plastron pattern character used to differentiate these subspecies. They found
that shell shape and postorbital stripe data do not necessarily correspond to
plastron pattern, suggesting that animals previously identified as kamaroma,
and as such representing a potential co-occurrence of two different subspecies in some areas of the Philippines (i.e., Palawan and Sulu), may in effect
be amboinensis animals with a plastral pattern resembling that of kamaroma
through introgression or phenotypic plasticity.
12:24. Cuora galbinifrons: Bourret described this species in a work dated
1939 on the title page, but it was not actually published until 1940 (R. Bour, pers.
comm.). We have therefore changed the date of publication from 1939 to 1940.
12:25. Heosemys spinosa: Spinks et al. (2012b) examined variation in
mitochondrial (cyt b) and nuclear (11 loci) DNA in a large sample of Heosemys
spinosa that lacked locality data. Two clades were clearly identified by the
mtDNA analysis, and were supported by preliminary morphological analysis,
but not recovered in the nuDNA analysis. Future identification of the geographic
provenance of these two clades is critical for conservation management as well
as establishing the taxonomic implications of this work.
12:26. Mauremys caspica: Vamberger et al. (2013) investigated population structure and history of Mauremys caspica based on an analysis of 14
microsatellite loci and cytochrome b mtDNAsequences from nearly range-wide
samples (but lacking Iraq). Their results found two clusters of mitochondrial
haplotypes, and four microsatellite clusters, with each mtDNA haplotype
cluster comprising two of the microsatellite clusters. Specimens from Bahrain
and Saudi Arabia were found to constitute a distinct microsatellite cluster, and
were thus viewed as representing native (and endangered) populations. The
authors proposed that each of the four identified phylogenetic clusters (i.e.,
Central Anatolia; eastern Turkey and Syria; Dagestan, Azerbaijan, and Iran; and
Bahrain and Saudi Arabia) be treated as distinct management units. Their results
did not support the validity of any of the three previously morphologicallydefined subspecies, and they therefore proposed that the subspecies no longer
be recognized, a recommendation that we follow here.
12:27. Orlitia borneensis: Palupcikova et al. (2012) analyzed mitochondrial (cyt b) and nuclear (R35) sequences, and shell and scute morphometrics
among Orlitia borneensis specimens in European collections. Most specimens
lacked precise locality data because they originated from a single confiscation in 2001, but three known-locality specimens from Borneo and Sumatra
were added to the sample series (no reference specimens from West Malaysia
were available). Haplotype diversity in cyt b was found to be relatively high,
with three main haplotype groups identified; nucleotide diversity was low,
and phylogenetic structure was poorly supported. The three known-origin
animals clustered within one of the main haplotype groups, suggesting that
the confiscated animals covered much of the species' genetic diversity. Only
minimal variation was found in R35 sequences. Geometric morphometrics
demonstrated morphological similarity of all examined specimens. These
results led the authors to conclude that all examined animals represented a
single conservation unit.
12:28. Aldabrachelys gigantea or Dipsochelys dussumieri: The latest
contribution to the ongoing debate in the ICZN (case #3463) on the scientific
name for the Aldabra Tortoise came from Ceríaco and Bour (2012), who traced
the history of the specimen claimed to be the type of Testudo gigantea Schweigger
from the MNHN Paris via the defunct Royal Cabinet of Natural History ofAjuda
in Lisbon to the collections made in Brazil and other parts of South America
by Alexandre Rodrigues Ferreira during his travels there during 1783–1792.
The authors thus provided further evidence that Schweigger’s type specimen of
Testudo gigantea originated in Brazil and represents a Chelonoidis denticulata,
providing support for possibly precluding the use of the name gigantea for the
Aldabra Tortoise. At the time of writing of this annotation, the ICZN is voting
regarding the use of either gigantea or dussumieri, and a result is expected to
be published in the next issue of the Bulletin of Zoological Nomenclature.
12:29. Geochelone or Centrochelys sulcata: Bour and Henkel (2012)
described a captive colony of tortoises of different species maintained in extensive
enclosures in southern France and documented a female Centrochelys sulcata
producing viable, morphologically aberrant, hatchlings that were interpreted
as hybrids between this sulcata female and a male Dipsochelys dussumieri [or
Aldabrachelys gigantea], with which copulation had been observed. From each
of three clutches of 20 eggs each, normal sulcata hatched from 15 eggs after
75–90 days, one hatchling emerged after 140–170 days and was a suspected
sulcata-dussumieri hybrid, and 4 eggs failed to develop. The authors also
reported the repeated production of hybrid hatchlings between C. sulcata and
Astrochelys radiata at the A Cupulatta facility in Corsica; these either died just
before emergence from the egg, or within days after hatching. These records
add to the growing catalog of documented intergeneric turtle hybrids (see also
annotation number 8 above).
12:30. Chelonoidis chilensis: The number of species in the Chelonoidis chilensis complex recognized by various authors has ranged from one to
three (see TTWG 2010, annotation 21). However, based on an examination
of a mitochondrial gene (cyt b) and 10 microsatellite loci, Fritz et al. (2012a)
found negligible genetic variation among populations and concluded that this
complex represents only a single species, C. chilensis. Previously described
morphological variation among populations is apparently explained by Bergmann’s Rule, with body size increasing with latitude. Hence, C. petersi and C.
donosobarrosi were placed in the synonymy of C. chilensis.
12:31. Chelonoidis nigra species complex: Only two papers published
in the past year bear on the taxonomy of the Galapagos tortoises, and both
continue to recognize the various taxa at the species level (see TTWG 2009,
annotation 32). Garrick et al. (2012) reported the discovery of hybrid tortoises
on Volcan Wolf that were F1 hybrids between a purebred Floreana tortoise (C.
nigra; erroneously referred to in their paper as C. elephantopus) and the local
Volcan Wolf tortoise (C. becki). This led the authors to speculate that translocated
Floreana tortoises might still exist on northern Isabela. Actual rediscovery of
genetically pure individuals of Floreana tortoises would reverse their current
IUCN status as Extinct, and be cause for great celebration.
Based on DNAsequences from three mitochondrial genes from extant and
museum specimens of Galapagos tortoises, Poulakakis et al. (2012) attempted to
resolve the phylogenetic relationships and reconstruct the biogeographic history
of the living and extinct taxa. Prior to their study, they recognized eleven extant
and four extinct species of Galapagos tortoises, though one of the latter has not
yet been described. It should be noted that the taxa they recognized a priori
did not completely agree with those recognized by the TTWG. In any case,
the results of Poulakakis et al. (2012) suggested that the following populations
represent “independent evolutionary units” and they applied the term “species”
to them: abingdonii (extinct in 2012, Pinta), becki (Volcan Wolf, northern Isabela), chathamensis (San Cristobal), darwini (Santiago), elephantopus (extinct,
Floreana; recognized as nigra by the TTWG), ephippium (Pinzón; recognized as
duncanensis by the TTWG), hoodensis (Española), porteri (La Reserva, Santa
Cruz), and vicina (central and southern Isabela, and including the names microphyes, guentheri, and vandenburghi; matching previous TTWG checklists),
an unnamed extant species (Cerro Fatal, Santa Cruz), and an unnamed extinct
species (Santa Fe). They explicitly noted that two taxa were not likely to be
independent evolutionary units: wallacei (Rabida) being subsumed under vicina
(as already reflected in previous TTWG checklists), and phantastica (extinct,
Fernandina; possibly introduced by humans) under porteri. However, in the
face of pronounced morphological differences between the extreme saddleback
phantastica and the greatly domed porteri, the TTWG believes that additional
support must be presented before synonymization of phantastica is warranted.
Finally, while Poulakakis et al. (2012) laudably extracted DNA from bones of
museum specimens from extinct lineages, it is critical that the genotyping of
actual type specimens of all possible named Galapagos tortoise taxa (including
those of synonyms) be completed before final allocations of names to existing
populations can be done with full confidence.
12:32. Gopherus flavomarginatus: Ureña-Aranda and Espinosa de los
Monteros (2012) examined variation in a mitochondrial gene in the Bolson
249
250
Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Tortoise and not only found no geographic structure in that variation, but also
noted that the existing variation was the lowest ever reported for a tortoise.
They attributed this low variability to the post-Pliocene collapse of this once
more widely distributed (New Mexico to central Mexico) species.
12:33. Gopherus polyphemus: Based on a single mitochondrial gene
fragment (ND4), Ennen et al. (2012) examined geographic variation in Gopher
Tortoises and found two major haplotype assemblages that overlapped in distribution in the Appalachicola-Chattahoochee River basin. They also found some
(albeit weak) support for the distinction of the Federally Threatened portion of
the western assemblage to the west of the Mobile River Basin (USFWS 1987).
In a more comprehensive study of both mitochondrial (cyt b) and nuclear microsatellite markers, Clostio et al. (2012) confirmed the Appalachicola basin
as the transition region between the distinctive western and eastern lineages.
In addition, based on both mtDNA and nuDNA, they noted the distinction of
the populations west of the Mobile River and those in western Georgia. They
concluded that the tortoises in each of these four regions should be managed
independently, but they made no specific taxonomic recommendations.
12:34. Gopherus species indeterminate Testudo australis Girard 1858
was described based on a specimen reputedly originating from New Zealand.
Clearly this specimen must have been transported by humans, as no testudinids
or other non-marine turtles are known from New Zealand, living or fossil. The
name has generally been overlooked or ignored, except by Boulenger (1889)
who attributed it with doubt to Gopherus polyphemus. Based on communication with Robert Murphy and Steve Gotte, the specimen cannot be found in the
USNM collection, if indeed it ever was there. Girard’s description of a uniform
near-black tortoise agrees nearly perfectly with Gopherus, except for the small
nuchal (cervical) scute, which is usually wide in Gopherus. The scutellation
on the top of the head and on the forearms, the absence of thigh spurs, as well
as the shape of the caudal (supracaudal) scute are all characters that eliminate
Manouria. Chelonoidis species are eliminated from consideration by the
presence of a nuchal in australis. Therefore, we tentatively attribute Testudo
australis Girard 1858 to the genus Gopherus. Attribution to any particular
species is challenging: based on size and coloration, G. berlandieri and G.
flavomarginatus are easily excluded, and while the Pacific location of New
Zealand suggests possible seafaring or trade links with California and therefore
G. agassizii and/or G. morafkai, morphological details such as the large head
with rounded snout, and upward curving of the gulars, are more reminiscent
of G. polyphemus. We therefore include Testudo australis Girard 1858 as a
nomen dubium and nomen oblitum under Gopherus species indeterminate.
12:35. Kinixys: Kindler et al. (2012) examined the phylogeography,
phylogeny, and taxonomy of all currently recognized Kinixys taxa, based on
examination of sequence data for three mtDNA fragments (12S, ND4, cyt b)
and three nuclear loci (C-mos, ODC, R35). Their findings indicated that the
savannah taxa, traditionally recognized as subspecies or affiliated species of
K. belliana, represent three deeply divergent clades, which are paraphyletic
with respect to the rainforest species K. erosa and K. homeana.
To reflect this phylogeny, the authors recognized their East African cluster
as a valid species, for which they followed the first reviser (Bour 1979) to apply
the name zombensis in preference over zuluensis. They also elevated nogueyi
to full species status, and reconfirmed lobatsiana, natalensis, and spekii as
evolutionarily distinct and ranked as full species.
They did not, however, present a clear arrangement of recognized taxa;
their new delineation of species is presented in their Fig. 2, implying that no
subspecies are recognized. They noted that their samples of domerguei from
Madagascar clustered with zombensis, and it appears that they intended to
transfer domerguei from the synonymy of K. b. belliana (following Broadley
1992, 1993; Fritz and Havas 2007; Branch 2008) to that of K. zombensis. In
their text (Kindler et al. 2012:198), the five-clawed tortoises of the Central
African Republic are specifically combined with the four-clawed West African
animals to form the distinct species K. nogueyi; however, these Central African
Republic records are mapped (Fig. S1) and listed (Table S1) as ‘belliana belliana’ in the online supporting material.
While the TTWG generally believes that the results presented by Kindler
et al. (2012) represent major advances in our understanding of Kinixys phylogeny, we prefer to retain a slightly more traditional arrangement for domerguei,
pending further data. While domerguei may not be genetically recognizable
based on the examined genes, it is morphologically well established (e.g., Bour
2006), and we continue to recognize it as a valid taxon at subspecies rank under
zombensis (to which it appears most closely related). Hence, we recognize the
new combinations Kinixys zombensis zombensis and K. z. domerguei.
12:36. Testudo graeca: Parham et al. (2012) extracted mtDNA sequence
data from Iranian tortoises of the T. graeca complex, specifically the holotype
of T. g. zarudnyi and topotypes of T. g. buxtoni and T. g. perses. Their results
confirmed the previous work of Fritz et al. (2007, 2009) regarding the existence
of two distinctive mitochondrial haplotypes in Iran, one in the northwest, and
one in eastern and central regions.
Using AFLPs (amplified fragment length polymorphisms) from samples
across the range of the T. graeca species complex, Mikulíček et al. (2013)
identified four geographically defined genetic groups: 1) western Mediterranean
(Morocco and Spain to Libya); 2) Balkans-Middle East (Bulgaria and Romania
to southwestern Russia, Azerbaijan, eastern Turkey, and southern Israel); 3)
Caucasian (extreme eastern Turkey, Russia, and southeastern Azerbaijan to
northwestern Iran); and 4) central and eastern Iran. These groups generally
correspond to the mtDNA haplotype lineages identified by Fritz et al. (2007,
2009), but with one to four mitochondrial lineages per AFLP group.
Considered together, these two data sets suggest that the western Mediterranean and central-eastern Iran groups are both divergent and allopatric,
and thus could be considered as distinct species. The oldest available names
for these groups are T. graeca (including cyrenaica, lamberti, marokkensis,
nabuelensis, soussensis, and whitei) and T. zarudnyi, respectively. The remaining two groups come into contact in the eastern Caucasus, with some mtDNA
evidence of gene flow between them. Hence, there remains uncertainty as to
whether they represent distinct evolutionary units worthy of species status. The
oldest available names for those two groups are T. terrestris (Balkans-Middle
East) and T. buxtoni (Caucasus; including armeniaca, perses, and pallasi).
Unfortunately, variation in neither set of markers corresponds well with
described morphological variation in this complex, on which the traditional
taxonomy has been based.
Pending future studies of variation in nuclear markers and a re-examination of morphological variation to determine minimally plastic characters that
correspond to the currently defined genetic groups (i.e., dismissing color and
general body size and shape), we prefer to retain T. graeca as a single species,
with no changes in the subspecies recognized from our previous checklist.
12:37. Apalone spinifera: McGaugh (2012) examined variation at ten
microsatellite loci for populations of Apalone spinifera within and outside of the
Cuatro Ciénegas basin in Mexico. She found considerable divergence among
all sampled populations (particularly eastern versus western basin localities),
but found no genetic variation associated with carapacial color variation within
the basin (the basis for the original description of A. s. atra). She reported negligible differentiation between softshells sampled inside the basin compared to
those outside, but provided no indication whether additional A. s. emoryi were
sampled since the range-wide analysis of A. spinifera reported by McGaugh et
al. (2008) and did not provide a detailed comparison between Apalone within
the Cuatro Cienegas basin versus outside the basin. As McGaugh did not make
an explicit taxonomic recommendation to synonymize atra with emoryi, we
conservatively continue to recognize atra as before, at subspecies level.
12:38. Nilssonia gangetica: An analysis of mitochondrial and nuclear
DNA sequence data by Liebing et al. (2012) confirmed the monophyly of the
genus, resolved the placement of N. formosa as sister to other Nilssonia, and
identified significant intraspecific genetic variation within N. gangetica corresponding to river basin of origin. Populations from the Brahmaputra, Mahanadi,
and combined Indus and Ganges basins were each genetically distinct, and
worthy of separate management. The authors did not support species recognition
of the three identified units, but noted that if these units were given subspecies
status, the name mahanaddica (Annandale 1912b) is available for the Mahanadi
basin population, and the nominate trinomial would apply to the Indus-Ganges
population. No name has been applied to the Brahmaputra population. Until
the three intra-specific populations are adequately characterized and named,
we continue to recognize N. gangetica as monotypic.
12:39. Chelidae: It was assumed by Gaffney (1977) that the three longnecked taxa in the Chelidae (Chelodina, Hydromedusa and Chelus) formed a
monophyletic lineage. However, Pritchard (1984) proposed that these three taxa
were not necessarily closely related, based on the major structural differences in
how they arrived at their long-necked condition, a position generally consistent
with the phylogenetic arrangement previously proposed by Baur (1893a). Sequencing of 12S mtDNA was used to demonstrate that the Australian radiation
of the Chelidae formed a monophyletic group (Seddon et al. 1997; Georges et
al. 1998) and that the shorter-necked South American taxa (including Chelus)
were also a monophyletic group, with Hydromedusa a third lineage (Georges
et al. 1998). At that time Georges et al. (1998) proposed names for the three
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
lineages; Chelinae [as Chelidinae] for Chelus and its South American relatives;
Chelodininae for the Australasian species; and Hydromedusinae for the genus
Hydromedusa, a clade that also includes the fossil genus Yaminuechelys (de la
Fuente et al. 2001) and hence is not monotypic. Further morphological work
agreed with Pritchard (1984) and supported the molecular results with differences in skull, shell, and cervical structures among the three monophyletic
lineages (Thomson 2000), indicating that what gross similarity there is between
the long-necked forms was a result of convergence and not of shared ancestry.
In the ten or so years since these publications, numerous studies have added
further support to this phylogenetic arrangement, which was summarized and
further supported by Guillon et al. (2012). Therefore we now include these
three subfamily assignments in this edition of the checklist.
We are aware of the family-group name Hydraspidina Bonaparte 1836,
based on the genus Hydraspis Bell 1828, whose type species is Testudo [now
Chelodina] longicollis, as a potential source for the subfamily name for chelids
of the Australia-New Guinea region. However, as Bonaparte’s concept of
Hydraspis differed fundamentally from that of Bell, to the point of excluding
longicollis from Hydraspis by properly placing it in Chelodina Fitzinger
1826, revival of the family-group name Hydraspidina (as Hydraspidinae for
this subfamily) would lead to unnecessary confusion. In the absence of strict
ICZN priority requirements for family-group names, we therefore use the name
Chelodininae Baur 1893b for the chelids of the Australia-New Guinea region.
12:40. Mesoclemmys dahli and M. zuliae: Based on both mitochondrial
and nuclear gene sequence variation, Vargas-Ramírez et al. (2012a) confirmed
the species distinction of Mesoclemmys dahli and M. zuliae, and identified two
geographically isolated, weakly divergent populations within M. dahli (Córdoba
and Cesar Departments, Colombia). Further sampling will be necessary to
determine if this divergence has taxonomic implications.
12:41. Mesoclemmys heliostemma and M. raniceps: Because some
authors (e.g., Rueda-Almonacid et al. 2007) have questioned whether M.
heliostemma is distinct from M. raniceps, Molina et al. (2012) undertook a
multivariate morphometric study of those two taxa (and M. gibba). Their results
supported the validity of M. heliostemma, and identified eight new localities that
significantly expand the known range of the species across the Amazon basin.
12:42. Pelusios castanoides: Analysis by Fritz et al. (2012b) of mitochondrial genes of P. castanoides revealed that samples from Madagascar and
the Seychelles were weakly differentiated from each other, and significantly
different from the sampled mainland populations. However, sparse sampling
from the mainland precluded any final conclusions about the origins of the
Malagasy and Seychelles populations, and therefore the validity of the currently
named, endemic subspecies of the Seychelles. As the population of Madagascar
is related more closely to that of the Seychelles than to the continental African
populations, a subspecies designation for these combined populations might
be warranted. The oldest available name is kapika Bour 1979; however, we
defer any changes to our checklist at this time.
12:43. Pelusios subniger: Parallel analysis by Fritz et al. (2012b) of
mtDNA samples of Pelusios subniger from mainland Africa, Madagascar, and
the Seychelles demonstrated the existence of a cryptic, unnamed taxon from
the Democratic Republic of the Congo, but no significant variation among all
other sampled populations. Based on these results, the authors suggested that
subniger was introduced by humans to both Madagascar and the Seychelles, and
recommended that the supposed endemic Seychelles subspecies P. s. parietalis
be placed into the synonymy of a monotypic P. subniger. However, reanalysis
of the morphological differentiation that originally distinguished these taxa
has not been carried out, and we are reluctant to make a definitive change at
this time, particularly for a taxon with such significant regional conservation
concerns. We await further analysis before amending this taxonomy.
12:44. Podocnemis lewyana: Based on 10 microsatellite loci and a mitochondrial DNA fragment, Vargas-Ramírez et al. (2012b) found low genetic
variability across the range of Podocnemis lewyana, consistent with a significant
historic population bottleneck. They did identify three weakly differentiated
genetic management units: 1) the Upper Magdalena River Basin; 2) the Lower
Magdalena, Lower Cauca, and San Jorge Basins; and 3) the Sinú River Basin,
but made no taxonomic recommendations.
2014 Checklist Annotations
TTWG 2014 (000.329-479.checklist.v.7) (14:1-50)
14:1. Macrochelys temminckii: Two new names for Alligator Snapping
Turtles were coined by Hoser (2013). There are significant nomenclatural,
technical, and biological problems inherent in these descriptions, and at this time
it seems appropriate to treat his names, Macrochelys temminckii muscati and
Macrochelys maxhoseri, as unavailable synonyms of Macrochelys temminckii.
14:2. Caretta caretta: Considerable confusion has surrounded the names
and authors and dates of publication of the turtle descriptions published in
various outputs of the Expédition Scientifique de Morée, currently cited in our
checklist as Valenciennes (1833), Bibron and Bory de Saint-Vincent (1833), and
Bory de Saint-Vincent (1835). See the detailed clarification below in annotation
25 for Emys orbicularis hellenica. The name Chelonia pelasgorum was first
published on plate 6 by Valenciennes (1833), but rendered as C. pelasgica in
the subsequent text by Bibron and Bory de Saint-Vincent (1833), where they
synonymized it with Chelonia caouanna (= Caretta caretta).
14:3. Eretmochelys: In last year’s checklist, in annotation 7, we outlined
our reasoning for recognizing the subfamilies Cheloniinae and Carettinae.
Unfortunately, we made an editorial error in listing Eretmochelys under the
Cheloniinae when, in fact, studies have shown that it is more closely related
to Caretta and Lepidochelys, and belongs in the Carettinae, as we pointed out
in our annotation. We correct the error in this year’s checklist.
14:4. Eretmochelys imbricata: The junior synonym Chelonia grisea
Eschscholtz 1829b has for many years been incorrectly cited as Chelonia griseam
in our previous checklists and in Fritz and Havaš (2007) and extensively on
the web. Having finally successfully accessed the obscure original publication,
we now note this long-standing error and correct it.
14:5. Chelonia mydas: We note also that in the same obscure publication cited above, Eschscholtz (1829b) also described Chelonia castanea from
Surinam as a new species. The name has been overlooked since its description
and is a nomen oblitum and junior synonym of C. mydas.
14:6. Dermochelys coriacea: The original citation for the junior synonym
Dermatochelys porcata is actually Wagler (1830b), not Wagler (1833) as listed
in our previous checklists, but which contained no new turtle descriptions.
14:7. Dermochelys coriacea: In our previous checklists we had included
the name Testudo marina Wilhelm 1794 in the synonymy of D. coriacea, based
on its inclusion in older checklists (e.g., Fritz and Havaš 2007). However, examination of Wilhelm’s (1794) work indicates that his use of the name Testudo
marina was as an incorrect collective group name for “marine” species (sea
turtles and softshells), as he also grouped most “terrestrial” turtles (testudinids
and kinosternids) under the incorrect group name Testudo terrestris, and all
other freshwater turtles under the group name Testudo fluviatilis. In discussing
separate species under these group names, he used names previously described
by other authors (including a description of the Leatherback, using the name
Testudo coriacea), but not in a consistently binomial manner. His work therefore has no standing nomenclaturally, and we have removed the name Testudo
marina Wilhelm 1794 from the synonymy of D. coriacea.
On the other hand, Ranzani (1832) published a description in Latin of the
Leatherback Turtle in which he described it as Testudine coriacea marina. This
description is valid (as the trinomen Testudo coriacea marina), as per ICZN
Code Article 11(h)(ii) allowing for the use of adjectival Latin descriptions, as
previously noted by Smith and Rhodin (1986) in regard to the validity of the
original authorship of Testudo coriacea Vandelli 1761.
14:8. Dermatemys mawii: González-Porter et al. (2013) presented
microsatellite data that supported their previous mitochondrial DNA studies
(González-Porter et al. 2011) in recognizing populations of Dermatemys mawii
in the Papaloapan River drainage as genetically distinctive. However, they made
no taxonomic recommendations based on their results. In addition, they also
identified a small sample of genetically divergent individuals in the Sarstun
and Salinas River basins along the southeastern distribution of the species that
they speculated might represent a cryptic taxon.
14:9. Kinosternidae: Iverson et al. (2013) sequenced three mtDNA
and three nuclear markers for every recognized species and most subspecies
of kinosternids. Their analyses revealed three well-resolved clades within the
Kinosterninae, corresponding to Sternotherus, a previously unnamed clade that
they described as the new genus Cryptochelys, and Kinosternon sensu stricto.
Their molecular data support for Cryptochelys was strong, but data support for
non-monophyly of Kinosternon with respect to Sternotherus was weak. The
identified groups are broadly consistent with morphological and biogeographical
features. Their new genus Cryptochelys was diagnosed based on an extensive
set of morphological and molecular characters, and contains the designated
type species leucostoma, as well as acuta, angustipons, creaseri, dunni, and
herrerai.
As we are aware of a parallel study of kinosternid phylogenetics, currently
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in review, that reaches different taxonomic conclusions, we present the recommended taxonomy of Iverson et al. (2013) as an additional alternative to
the traditional arrangement, in the knowledge that we will revisit kinosternid
taxonomy again in our next edition, and hopefully come to a consensus position
then.
14:10. Kinosternon abaxillare: A multivariate analysis of morphometric
data by Berry (1978) demonstrated the distinctiveness of the endemic, allopatric taxon K. scorpioides abaxillare from the parapatric K. s. cruentatum.
In addition, preliminary molecular sampling of the K. scorpioides complex by
Iverson et al. (2013) suggested that K. s. abaxillare was more closely related
to K. oaxacae than to K. s. cruentatum (or any other K. scorpioides). Given
both the morphometric and molecular evidence, the latter authors followed
Alvarez del Toro (1972, among many others) and suggested that K. abaxillare
be recognized as a full species. Until more thorough geographic and molecular
sampling is completed, we acknowledge both options in this checklist, but treat
the taxon as more likely a species.
14:11. Kinosternon chimalhuaca: In 1996, while the original, full description by Berry et al. (1997) of K. chimalhuaca was in press, it was shared
with Manfred Rogner for inclusion in his forthcoming book. However, though
unintended, Rogner’s abbreviated version, clearly attributed to Berry et al.,
was published first (in 1996). Hence, although many authors have cited Berry
et al. (1997) as the original description for this taxon, the proper attribution
should be Berry, Seidel, and Iverson in Rogner (1996). The ICZN has now
been petitioned (Rogner et al. 2013) to officially confirm this proper authorship
and date, which has already been used in all previous TTWG checklists.
14:12. Kinosternon subrubrum steindachneri: Based on osteology,
Bourque (2012) recommended that K. s. steindachneri be elevated to full
species status, as it was originally described. Preliminary molecular data
provided by Iverson et al. (2013) supported this conclusion. However, until a
more complete, range-wide study of molecular and morphological variation
of the K. subrubrum-baurii complex is available, we here retain steindachneri
as a subspecies of K. subrubrum.
14:13. Staurotypinae or Staurotypidae: Highlighting the extensive
divergence of the staurotypines from the kinosternines based on morphology
(Hutchison 1991), genetics (Iverson et al. 2103), karyotype (Bickham and Carr
1983), and sex determination mechanisms (Ewert et al. 2004), Iverson et al.
(2013) followed Bickham and Carr (1983) in recognizing the Staurotypidae
as a separate family. Within the TTWG we have differing opinions on the
appropriate ranking of this taxonomic node, and recognize that the views and
actions of the wider turtle taxonomic community will determine its eventual
accepted ranking; until consensus emerges, we provide alternative rankings
in the checklist.
14:14. Graptemys: In a historical review of the taxonomic history of the
genus Graptemys, Lindeman (2013:20) mentioned two genus names from an
unpublished manuscript by Georg Baur: Neoclemmys (intended to include
pseudogeographica and oculifera) and Megaloclemmys (for pulchra), while
Graptemys would have been retained for geographica and kohnii. However,
Lindeman only used these names in a conditional manner, without formal
status as valid taxa, and hence the names Neoclemmys and Megaloclemmys
were not made available according to Article 15.1 of the International Code of
Zoological Nomenclature (see also annotation 19 for Graptemys intermedia).
14:15. Graptemys flavimaculata: Using microsatellite loci, Selman et
al. (2013) demonstrated a significant degree of genetic structure across the
range of the species in the Pascagoula River basin, with the greatest divergence
between the main Pascagoula basin and the lowland Escatawpa River tributary,
historically separate drainages. Although they urged that at least these two units
be managed separately, they made no taxonomic recommendations.
14:16. Graptemys ouachitensis or G. o. ouachitensis: See annotation
number 19 below regarding the taxon sabinensis, previously listed as a subspecies of ouachitensis, but now conditionally elevated to full species status,
therefore also necessitating listing the Ouachita Map Turtle as a full species,
rather than the nominate subspecies.
14:17. Graptemys pulchra: In his historical review of the taxonomic history
of Graptemys, Lindeman (2013:20) also made reference to Baur’s manuscript
names for the species he eventually described as Graptemys pulchra; we hereby
designate these names, G. alabamensis and G. grandis, as nomina nuda, and
associate them with the synonymy of G. pulchra, as they were considered for
application to that taxon.
14:18. Graptemys sabinensis or G. o. sabinensis: Originally described
as a subspecies of Graptemys pseudogeographica by Cagle (1953a), the Sabine
Map Turtle (G. sabinensis) was later classified by Vogt (1980) as a subspecies
of G. ouachitensis, and most subsequent authors have followed that arrangement. However, based on a small sample of skulls, Ward (1980) believed that
sabinensis was so distinctive that it warranted species status. Recent analyses
of morphology, mitochondrial DNA, and nuclear DNA have generally failed
to resolve the relationships of sabinensis with confidence, and the interrelationships of the “narrow-headed Graptemys” remain largely unresolved (Stephens
1998; Stephens and Wiens 2003; Myers 2008; Wiens et al. 2010; Brown et al.
2012). Based on these previous studies and his own extensive examinations of
Gulf Coast Graptemys specimens, Lindeman (2013) noted that sabinensis is
allopatric, non-intergrading, and diagnosable morphologically, and concluded
that it should be recognized as a full species. While subspecific as well as specific recognition can each be supported, we conclude that enough uncertainty
remains regarding this lineage to list it as either a species or subspecies. Further
sampling of the nuclear genome and more strongly supported phylogenetic
trees will be necessary to settle this issue as well as the relationships across
the entire genus Graptemys.
14:19. Graptemys sabinensis or G. o. sabinensis: In a checklist of turtles
of Louisiana, Beyer (1900) listed “Malacoclemmys intermedia Baur” from the
“southern and southwestern parts”, referring to a manuscript name by Georg
Baur for the taxon subsequently described as Graptemys pseudogeographica
sabinensis by Cagle (1953a). The name was based on specimens from the
Mermentau River basin, now in the Tulane University Museum collection, sent
to Baur by Joseph Gustave Kohn (Lindeman 2013). The original publication
of the name Malacoclemmys intermedia is therefore attributable to Beyer and
pre-dates the name sabinensis Cagle by 53 years, but is clearly a nomen nudum.
Furthermore, it has not been used in over 113 years and would have the status
of nomen oblitum. In his discussion of the history of the name Graptemys
intermedia used by Baur in his unpublished manuscript, Lindeman (2013)
published Baur’s original manuscript drawings (Fig. 2.4) and a photograph of
the Kohn specimens on which Baur had intended his diagnosis to rest (Fig.
8.18). Lindeman also clearly identified intermedia as a synonym of the taxon
he recognized by the name of G. sabinensis. However, Lindeman only, and
consistently, used the name G. intermedia in a conditional manner, without
formal status of valid taxon, and the name intermedia as used by Lindeman,
published after 1960, has not been made available according to Article 15.1
of the International Code of Zoological Nomenclature.
14:20. Pseudemys: Based on three mitochondrial and ten nuclear gene
loci, Spinks et al. (2013) examined variation across all recognized taxa of
the genus Pseudemys. Their analyses revealed essentially no support for currently recognized species groups, species, or subspecies. Only P. gorzugi was
consistently recovered as monophyletic across all their analyses, while their
molecular evidence identified three geographically cohesive groups that do
not correspond to current species boundaries. They concluded that the genus
Pseudemys has probably been oversplit taxonomically. However, they made no
explicit recommendations for change until a much larger and more definitive,
multi-character data set is brought to bear on this complex. Thus we retain
Pseudemys essentially unchanged from the previous checklist.
14:21. Trachemys: In their book on Mexican turtles, Legler and Vogt
(2013) continued to follow the taxonomy for Trachemys as used by Legler (1990)
(i.e., all Mexican taxa as subspecies of T. scripta), and did not provide data
or rationale to refute the phylogenetic data and taxonomic opinions published
since 1990. Because implementing that taxonomy here would reverse 23 years
of increased understanding and progress toward a stable classification of this
complex genus, we have not incorporated the Trachemys taxonomy presented
by Legler and Vogt (2013) in our current checklist.
14:22. Trachemys (Caribbean): Parham et al. (2013) examined variation
in mitochondrial and nuclear DNA markers for Trachemys populations across
the Greater Antilles. They identified the morphologically distinct population
reported by Tuberville et al. (2005) in northwestern Jamaica as T. d. decussata,
representing a significant range extension from eastern Cuba. Parham et al.
(2013) demonstrated the monophyly of West Indian taxa, as well as evidence of
hybridization between T. decorata and T. stejnegeri in the southern Dominican
Republic, and between T. terrapen and T. d. decussata in northwestern Jamaica.
The authors were unable to determine whether the presence of decussata on
Jamaica and localized hybridization with terrapen was the result of natural or
human-mediated dispersal.
Their data also supported the continued recognition of the subspecies T.
s. stejnegeri (Puerto Rico) and T. s. vicina (Hispaniola), with occasional gene
flow (natural or human-mediated) between them. They also acknowledged
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
that morphological and genetic data suggest the recognition of T. d. decussata
(eastern Cuba) and T. d. angusta (western Cuba) as full species, but declined
to make that recommendation pending further sampling in Cuba.
Finally, they speculated that the occurrence of T. d. angusta on the Cayman Islands was “non-native”. Given that Grand Cayman was periodically
inundated even in the latest Pleistocene (20–25 thousand years ago; IturraldeVinent 2006), and that the prevailing winds and currents would make a natural
colonization from the northwest difficult, we concur that the Grand Cayman
populations of T. d. angusta are likely the result of human introduction. This
is further supported by Echternacht et al. (2011) who, in their review of the
herpetofauna of the Cayman Islands, explicitly stated that since no Trachemys
fossils have been found in peat deposits on the island (which contained many
other vertebrates), they presumed T. decussata to be introduced.
In addition, Parham et al. (2013) analyzed a small sample of Trachemys
from Central America that yielded results indicating genetic similarity of T.
venusta and T. emolli, demonstrating the need for further sampling and analysis
to evaluate the sweeping taxonomic changes proposed by Fritz et al. (2012)
and the subspecies described by McCord et al. (2010). Until such additional
information becomes available, Parham et al. (2013) recommended taxonomic
conservatism and cautious interpretation of preliminary results, and proposed
no taxonomic changes.
14:23. Trachemys (Central America): Using mitochondrial DNA
sequence data for Trachemys downloaded from the European Nucleotide
Archive and new data from two Honduran specimens, McCranie et al. (2013)
confirmed that the range of the taxon emolli extends from northwestern Costa
Rica to southeastern El Salvador (see Ibarra Portillo et al. 2009). Because of
the extensive overlap in their analysis with the data used by Fritz et al. (2012),
McCranie et al. (2013) supported their earlier taxonomic recommendations.
McCranie et al. (2013) also commented on the status of the taxon T. v. uhrigi,
originally described from Honduras, but subsequently reported from Colombia,
Costa Rica, Nicaragua, and Panama, and argued that the diagnostic coloration
of uhrigi is not exhibited consistently by Caribbean Honduran specimens (and
presumably occurs in individuals of T. venusta as far away as Colombia), and
suggested that T. v. uhrigi has no taxonomic validity. In contrast, Páez et al.
(2012) listed T. v. uhrigi as being the subspecies occurring in the Colombian
Departments of Antioquia and Chocó. However, until additional molecular data
are forthcoming from the Caribbean versant of Central America, particularly
from the Yucatan peninsula, Honduras, Nicaragua, Costa Rica, Panama, and
Colombia, we adhere mainly to the alternative taxonomies of Seidel (2002)
and Fritz et al. (2012).
14:24. Emys (sensu lato): In a molecular analysis of emydine turtles,
Angielczyk and Feldman (2013) found strong support for a monophyletic Emys
(including orbicularis, blandingii, and marmorata) using mtDNA sequence
data, but strong support for a paraphyletic Emys using 14 nuclear genes. The
combined data set resolved a monophyletic Emys, but the results were apparently driven by the much more variable mitochondrial genome. Despite some
uncertainty about the monophyly of Emys sensu lato (see also Wiens et al.
2010), we retain both options of a narrow and a broad definition of the genus
Emys, pending even more genetic data.
14:25. Emys orbicularis hellenica and Mauremys rivulata: Considerable confusion has surrounded the names and authors and dates of publication of the turtle descriptions published in various outputs of the Expédition
Scientifique de Morée, currently cited in our checklist as Valenciennes (1833),
Bibron and Bory de Saint-Vincent (1833), and Bory de Saint-Vincent (1835).
Sherborn and Woodward (1901) documented that the zoology sections dealing
with vertebrates, in which turtle descriptions (Chelonia pelasgorum, Emys
hellenica, Emys iberica, and Emys rivulata) appear, were all first published
sequentially in 1833 in looseleaf “livraisons” with sets of plates (“planches”)
and text, and later all the plates were re-published as a bound volume in 1835.
Confusion about the sequence of publication of the turtle plates and the text
has arisen due to the imprint of “1832” on the frontispiece of the text, but it
actually appeared in 1833, after the plates. That the unbound turtle plates by
Valenciennes (1833:pls.6–9) were published first was clearly documented in
the subsequent text by Bibron and Bory de Saint-Vincent (1833:61, lines 5–9,
21–23, footnote 2), who referred to the specifically numbered plates as coming
from the “troisième séries”.
The name Emys hellenica was first published in the third series of planches
(plate 8, figures 2–2a), where it was attributed to Valenciennes in the legend,
and then subsequently described (as Cistuda hellenica) on pages 61–62 of
the text, where it was attributed to Bibron and Bory de Saint-Vincent. Proper
original attribution of the name is therefore Emys hellenica Valenciennes in
Bory de Saint-Vincent 1833:pl.8. That name was subsequently synonymized,
as “Cistude hellénique”, with Cistudo europaea, another synonym of Emys
orbicularis, by Duméril and Bibron (1835:227), but is today recognized as a
valid subspecies of that taxon (Fritz et al. 2005; Fritz and Havaš 2007).
According to Bibron and Bory de Saint-Vincent (1833), in their text
on page 61, lines 5–9, under the synonymy of Cistuda europaea (= Emys
orbicularis), they noted that Valenciennes (1833) had “for unknown reasons”
(“on ne sait pourquoi”) named the juvenile specimen of Emys on plate 9 as
iberica [not Emys iberica Eichwald 1831]. The original typeface in the legend
of plate 9 reads “Emyde ibérienne. Emys iberica. Val.”; however, in at least
some contemporary copies of the subsequently bound atlas (Bory de SaintVincent 1835), a small printed label in similar text reading “des anciens” has
been pasted over “ibérienne”, and a second label reading “antiquorum” pasted
over “iberica” and the first part of “Val.” We do not know if these labels were
originally added as an “erratum” to all looseleaf copies at the time of their
original publication in 1833, or more likely only to some of them when bound
into the atlas in 1835, because some copies today lack the labels (e.g., that in the
Paris Museum, but apparently not those in the British Museum [Gray 1844:31]
or the Museum of Comparative Zoology at Harvard [Loveridge and Williams
1957:213]). It must also be noted that Bibron and Bory de Saint-Vincent (1833)
commented on the name iberica printed on the original plate, but made no
mention of the name antiquorum. This uncertainty greatly complicates these
names and their authorship. If the “antiquorum” labels were added to all copies
of this work by the publisher, then Emys iberica Valenciennes was technically
never described, and Emys antiquorum would presumably be attributable to
Bory de Saint-Vincent, the editor of the 1835 atlas, although the remaining
partial exposure of the name “Val.” led Gray (1844:31) to attribute the name
antiquorum to Valenciennes when he (Gray) synonymized that name with
Cistudo europaea [= Emys orbicularis]. If the labels were inconsistently added
to only some copies of the original work, then the name antiquorum would
have no nomenclatural status. Pending the availability of additional historic
information about the consistency of this labeling and the reasons behind it, we
here attribute the name Emys iberica to Valenciennes in Bory de Saint-Vincent
(1833), and declare Emys antiquorum Bory de Saint-Vincent (1835) a nomen
novum and nomen nudum; and interpret both names as junior synonyms of
Emys orbicularis hellenica (Valenciennes in Bory de Saint-Vincent 1833).
14:26. Emys orbicularis, ssp. indet.: Rook et al. (2013) synonymized
the two Upper Pliocene fossil taxa, E. major and E. latens, described by Portis
(1890), with Emys orbicularis, citing an unpublished thesis by Chesi (2009).
The fossils were from Valdarno (d’Arno valley) along the northwest Italian
Ligurian coast, within the range of the present-day subspecies E. o. galloitalica.
However, given the subsequent Pleistocene and Holocene climate-associated
range shifts of E. orbicularis, we do not associate these fossil names with any
current subspecies at this time.
14:27. Terrapene: Martin et al. (2103) examined variation in two mitochondrial genes and one nuclear gene across all previously recognized taxa of
Terrapene (except T. nelsoni klauberi). Both mtDNA and a single nuclear gene
supported the monophyly of T. ornata, T. carolina (including T. coahuila), and
T. nelsoni. All analyses confirmed the distinctiveness of T. nelsoni and T. ornata,
but found no support for distinction between T. o. ornata and T. o. luteola, and
hence, they recommended the synonymy of the latter. For mtDNA only, they
found significant divergence within the previously recognized T. carolina
group taxa (including T. coahuila), and identified a western clade (including
triunguis, mexicana, and yucatana) and an eastern clade (all others, including
coahuila); the western clade was strongly supported based on mtDNA, but
the eastern clade had only very weak support. The authors recommended
the recognition of the western clade as a full species, T. mexicana, with three
subspecies (mexicana, triunguis, and yucatana). They were not able to resolve
the relationships among bauri, major, and carolina, and retained them as subspecies of T. carolina. Similarly, the relationship of T. coahuila to Gulf Coast
T. carolina was suggested but not resolved, and they recommended continued
recognition of T. coahuila at the species level. Meanwhile, Legler and Vogt
(2013) treated T. mexicana and T. yucatana as full, monotypic species, and
continued to recognize T. o. luteola as the subspecies of T. ornata inhabiting
Mexico.
As the suggested taxonomic rearrangements of Martin et al. (2013)
and Legler and Vogt (2013) are not based on a comprehensive analysis of
morphology, mitochondrial DNA, and nuclear genes, and show disagreement
with both the traditional taxonomy of Terrapene and the molecular phylogeny
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presented by Butler et al. (2011), we consider that the phylogenetic relationships
within this genus remain insufficiently resolved for us to adopt such significant
taxonomic changes, especially in light of the desire for minimal fluctuations
in taxonomy of this genus, given its extensive inclusion in State, Federal, and
CITES legislation.
14:28. Terrapene putnami: This species was described by Hay (1906)
based on a single fossil hypoplastron from the Alafia River basin in Florida
with imprecise stratigraphic data, but presumably Late Pleistocene. The taxon
has been widely assumed to represent virtually all eastern North American
fossil Terrapene material from the Miocene to the late Pleistocene, which is
clearly an exaggerated concept of the taxon (Ehret et al. 2011). This has become
increasingly problematic because of recent molecular analyses of extant taxa
that suggested that T. putnami should be synonymized with T. carolina major
(Butler et al. 2011; followed in TTWG 2012) or that argued that putnami be
retained as an extinct subspecies of T. carolina (Martin et al. 2013). To facilitate
future attempts to resolve the relationships among living and fossil turtles of the
T. carolina complex, Ehret et al. (2013) proposed to the ICZN the designation
of a neotype for T. putnami with precise locality and stratigraphic data, and
consisting of a nearly complete carapace and plastron with numerous associated
postcranial elements.
We here maintain T. putnami as a synonym of T. carolina major as
recommended by Butler et al. (2011), until the ICZN makes a ruling and additional research clarifies the relationships of extant and fossil members of the
T. carolina group.
14:29. Mauremys annamensis: Clemmys guangxiensis was described
by Qin (1992) based on two market specimens supposedly originating from
Guangxi, China. Iverson and McCord (1994) speculated that the type series
might be a composite of Mauremys mutica and M. iversoni (the latter now
known to be of hybrid origin between Cuora trifasciata and M. mutica; Parham
et al. 2001). As a result, we have previously included C. guangxiensis in the
synonymies of both C. trifasciata and M. mutica. However, Hu et al. (2013)
provided sequence data from a single mitochondrial gene for four specimens
of Mauremys from Guangxi purported to be M. guangxiensis, along with two
M. mutica from the same province. When included in a phylogenetic analysis
with sequences of Mauremys downloaded from GenBank, they determined
that their four specimens of M. guangxiensis were nearly identical to M. annamensis (which is endemic to Vietnam) and not M. mutica. Assuming their
four specimens represented the same taxon as described by Qin (1992), Hu et
al. (2013) interpreted their results as indicating that M. guangxiensis was either
synonymous with M. annamensis or a subspecies of the latter. However, their
analysis did not address the possibility of a hybrid origin for guangxiensis.
Pending further study of specimens being referred to M. guangxiensis, we
add guangxiensis to the synonymy of annamensis (as partim, hybrid), while
retaining its inclusion in the synonymy of M. mutica and C. trifasciata.
14:30. Mauremys japonica and Pelodiscus sinensis: Temminck and
Schlegel’s publications in Fauna Japonica are usually recorded as having been
published in 1835. However, their chapter on “Les Cheloniens” (pp. 1–80,
plates 1–9) was actually published in 1834 (see Hoogmoed et al. 2010) and
only contained invalid vernacular names. Their valid names Trionyx japonica
= Pelodiscus sinensis and Emys vulgaris japonica = Mauremys japonica were
not published until 1838 when Schlegel wrote and published (on p. 139) his
dated explanation of the previously published plates and for the first time provided Latin names for the two new species described earlier in French (Trionyx
stellatus Var. Japon [pls. 5 and 7] = Trionyx japonica and Emys palustris Var.
Japon [pls. 8 and 9] = Emys vulgaris japonica). Although the species name
“palustris” was used on the plate, in the text it was corrected to “vulgaris”, but
never with a specific “var. Japon” modifier attached to it.
14:31. Rhinoclemmys: Based on both mitochondrial and nuclear DNA
data, Vargas-Ramírez et al. (2103) identified significant phylogeographic structuring within R. melanosterna, but found conflicting phylogenetic relationships
among the allopatric/parapatric members of the R. punctularia group (including R. funerea, R. diademata, and R. melanosterna). They recommended no
taxonomic changes without further geographic and genome sampling.
14:32. Aldabrachelys gigantea: After several years of vigorous debate,
the ICZN (2013b) published their decision (Opinion 2316) regarding the appropriate scientific name for the Aldabra Tortoise (Case 3463; Frazier 2008,
2009). The Commission ruled to conserve the long-term use of the specific
name Testudo gigantea Schweigger (1812) for this tortoise, to affirm the
neotype designation of Frazier (2006), and to suppress the more recently used
name Testudo dussumieri Gray (1831d). One effect of this action was also to
validate the genus name Aldabrachelys Loveridge and Williams (1957) over
Dipsochelys Bour (1982a). Comments were published in BZN 66:80–87,
169–186, 274–290, 352–357; 67: 71–90, 170–178, 246–254, 319–331; 68:
72–77, 140–143, 294–300. With 83 published comments, this represented the
most extensive correspondence received by the Commission on a Case to date.
14:33. Chelonoidis carbonaria: The original citation for the junior synonym Testudo boiei is actually Wagler (1830a), not Wagler (1833) as listed in
our previous checklists; the latter contained no new turtle descriptions.
14:34. Gopherus berlandieri: In 1850, Berlandier described two terrestrial turtles from the “llanos” of Tamaulipas, Mexico: Testudo bicolor (not to
be confused with Testudo bicolor Schweigger 1812 or Terrapene bicolor Bell
1826, both synonyms for Testudo [= Cuora] amboinensis Daudin 1801) and
Testudo tuberculatu [sic] (not to be confused with Testudo tuberculata Schoepff
1801 [= Dermochelys coriacea]). His ample description leaves little doubt that
the names referred to a juvenile and adult male, respectively, of Xerobates [=
Gopherus] berlandieri Agassiz 1857a, and hence should be considered senior
subjective synonyms of the latter. However, since 1850, T. tuberculatu has
only been mentioned by True (1882), as T. tuberculata. In 1980 Berlandier’s
manuscript was translated and republished, with both species recorded again,
as T. bicolor and T. tuberculata, with distinct diagnostic characters, “and they
are common on both banks of the Rio Bravo.” No other publications seem to
include these names, both considered here as being nomina oblita.
14:35. Testudo graeca: Based strictly on morphology Chkhikvadze et
al. (2013) continued to argue for the recognition of six taxa of tortoises in the
Caucasus (Testudo graeca ibera, T. g. nikolskii, T. g. armeniaca, T. marginata
pallasi, T. m. buxtoni, and T. dagestanica). However, genetic sampling by Fritz
et al. (2007), Mashkaryan et al. (2013), and Mikulíček et al. (2013), including
specimens from within the ranges of each of those six purported taxa, supports
only the recognition of three taxa in the area (T. g. ibera, T. g. buxtoni, and T.
g. armeniaca). Because the unreliability of morphology in establishing species
boundaries within the genus Testudo has been well documented (Parham et
al. 2006; Fritz et al. 2007, 2009; Mikulíček et al. 2013; Danilov et al. 2013;
among others), we have not followed Chkhikvadze et al. (2013), pending
further genetic sampling.
14:36. Testudo or Chersine hermanni: Perez et al. (2014) examined
variation in mitochondrial DNA and nuclear microsatellites across the range
of T. hermanni, and found substantial geographic differentiation based on
distance between sites. They documented the greatest divergence between
the recognized subspecies, with the eastern subspecies (T. h. boettgeri) ranging westward to and including the Po River valley in northeastern Italy, Their
data also demonstrated the effects of thousands of years of human-mediated
dispersal of these tortoises. Although they noted that the isolated population in
France could have been established via natural or human dispersal, their data
supported the hypothesis that the Spanish, Corsican, and Sardinian populations
were likely the result of prehistoric human introductions of animals with Sicilian
genotypes.
14:37. Testudo or Chersine h. hermanni: Lapparent de Broin et al.
(2006b) analyzed all known fossil specimens of T. globosa, T. oriens, and T.
seminota, and concluded that oriens and seminota were synonymous with
globosa and that globosa was apparently synonymous with western T. h.
hermanni.
14:38. Testudo or Agrionemys h. kazachstanica: Agrionemys kazachstanica terbishi was described by Chkhikvadze (2009) based on a mummified
specimen, supposedly from Mongolia, in the Kohovd University collection
(Kohovd City, Mongolia). Ansorge et al. (2012) reported that the type specimen has been lost, that the herpetologist who collected the specimen believed
that it was a pet brought from Kazachstan, and that there is no confirmed
record of an extant tortoise indigenous to Mongolia. They recommended that
Testudo horsfieldii terbishi (Chkhikvadze 2009) should be regarded as a nomen dubium and allocated to the synonymy of Testudo horsfieldii; they also
recommended that Mongolia should be excluded from its distribution range.
We adopt these recommendations and associate the taxon with T. or A. h.
kazachstanica by virtue of it having originally been described as a subspecies
of that taxon.
14:39. Acanthochelys and Platemys: Sequence data from two mitochondrial genes analyzed by Huebinger et al. (2013) supported the sister
group relationship between Platemys and Acanthochelys, the monophyly of
the latter, and the possibility that A. radiolata as currently defined morphologically may be polyphyletic. We continue to affirm the recognition of both
genera, even though Platemys is monotypic.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
14:40. Phrynops geoffroanus: The name Emys tritentaculata was
listed by Cuvier (1829) as attributed to Auguste de Saint-Hilaire, a botanist
who traveled in Brazil and subsequently deposited several chelid turtles in
the Paris Museum (Bour, unpubl. data). Based on the name, suggestive of
several barbels, we assign it tentatively to the synonymy of P. geoffroanus
pending further study of Saint-Hilaire’s original specimens. The name E. tritentaculata does not refer to the American Box Turtle, Terrapene carolina,
as originally synonymized by Wermuth and Mertens (1961) and followed by
several others since then.
14:41. Platemys platycephala: The name Emys carunculata Cuvier
1829 was listed by Wermuth and Mertens (1961, 1977) and Fritz and Havaš
(2007) as an ex errore name for E. canaliculata Spix 1824, itself a synonym
of P. platycephala. However, the name E. carunculata was attributed by Cuvier (1829) to Auguste de Saint-Hilaire, while in the same paragraph also
listing E. canaliculata as attributed to Spix. The two names clearly represent
different nomina nuda. Pending further studies of the chelid turtles that SaintHilaire collected in Brazil (Bour, unpubl. data), we leave E. carunculata in
the synonymy of P. platycephala for the present.
14:42. Chelodina (Macrochelodina) kuchlingi: This species, originally described by Cann (1997d), was synonymized under “Chelodina rugosa”
(now Chelodina oblonga) by Georges and Thomson (2010). Their original
basis for the synonymization was “that names that are available under the
Code, but that apply to supposed taxa, unsupported by scientific evidence
either in the original account or subsequently, are placed in synonymy.” They
also indicated that C. kuchlingi was described from a single specimen of uncertain origin with a long history of captivity and so was treated as a junior
synonym of “C. rugosa” (now C. oblonga), citing Georges and Thomson
(2006) who had questioned the distinction between C. kuchlingi and C. rugosa, but did not synonymize them. The synonymization by Georges and
Thomson (2010) was subsequently followed by us (TTWG 2010) and Kennett et al. (2014) in their recent CBFTT species account for C. oblonga (see
link under that species). However, the synonymization has recently been challenged by Kuchling (CCB, in review, and in litt.), who has provided data that
C. kuchlingi is an apparently demonstrably valid and distinct range-restricted
species, with a more extensive distribution in northeastern Western Australia
(including the Ord River basin) than noted in the original description. Kuchling also raised serious concerns about the conservation status and potential
regional development threats to C. kuchlingi. Georges (in litt.) has acknowledged the difference in opinion, but stands by his opinion that the original
description was deficient, and that there has been insufficient evidence presented to date that C. kuchlingi is a valid taxon.
We take note of this on-going controversy here and, based on our own
principles of making only data-driven taxonomic changes in the checklist,
acknowledge that our original decision to follow the hypotheses of Georges
and Thomson (2006, 2010) to synonymize C. kuchlingi was likely premature,
and also inconsistent with our continued recognition at that time of other species also synonymized by Georges and Thomson at the same time (e.g., C.
gunaleni and C. walloyarrina). Given the potential conservation threats to
this range-restricted species and the lack of data supporting the prior synonymization, we therefore now reverse our earlier decision and resurrect C.
kuchlingi from its synonymy with “C. rugosa” (now C. oblonga) and await
further data-driven analyses from Kuchling, Georges, and others.
14:43. Chelodina oblonga (formerly C. rugosa): Thomson (2000)
demonstrated that the holotype of Chelodina oblonga Gray 1841 is actually
a specimen of what had over the last ca. 40 years been referred to as Chelodina rugosa Ogilby 1890 from northern Australia. The ICZN was petitioned
(Thomson 2006, 2007) to conserve current usage of the name C. rugosa for
the Northern Snake-necked Turtle and to apply the next available name, Chelodina colliei Gray 1856a, to the Southwestern Snake-necked Turtle, instead
of the commonly and erroneously used name C. oblonga. We previously discussed this ICZN case in our second checklist (Rhodin et al. 2008). Recently,
in their Opinion 2315, the ICZN (2013a) declined to support the petition to
give precedence to the younger, recently used name C. rugosa over the older
name C. oblonga for the Northern Snake-necked Turtle. Although the latter
species has been known as C. rugosa since 1974 and was listed as such in
previous editions of this checklist, we now follow the ruling of the ICZN
and use the name Chelodina (Macrochelodina) oblonga Gray 1841 for the
Northern Snake-necked Turtle, although the name Chelodina (Macrochelodina) rugosa Ogilby 1890 remains an available name in the synonymy of
C. oblonga. The decision by the ICZN has also been followed by Kennett et
al. (2014) in their recently published account on the Northern Snake-necked
Turtle in this CBFTT monograph series.
14:44. Macrodiremys: In an attempt to conserve usage of the name
Chelodina oblonga for the Southwestern Snake-necked Turtle, McCord and
Joseph-Ouni (2007b) designated the lectotype of Chelodina colliei (set by
Thomson 2000) as the neotype of Chelodina oblonga. At the time this was
done, there was already an open case before the ICZN (Thomson 2006) concerning whether to use the name C. oblonga or C. rugosa for the Northern
Snake-necked Turtle (see annotation 43). It should also be noted that the setting of a neotype where an extant holotype (or lectotype) already exists can
only be done by the ICZN. In their subsequent Opinion (ICZN 2013a), it was
ruled that, considering the confusion over these names and the potential for
further confusion, that the Principle of Priority should be followed, and that
C. oblonga should maintain priority over C. rugosa for the northern taxon. By
associating the new name Macrodiremys oblonga to the lectotype of C. colliei, McCord and Joseph-Ouni (2007b) had effectively erected a new nominal
species as a junior objective synonym of C. colliei. Thus, since M. oblonga
was the type species for the new genus Macrodiremys, then in effect so was
also its senior objective synonym, C. colliei. Fortunately, the latter was not already a type species for another genus. Georges and Thomson (2010) reduced
the various genera of snake-necked turtles to subgeneric status, all under the
oldest genus name Chelodina; this has been recognized in previous editions
of the checklist (TTWG 2012), however, the subgenus name for C. colliei
was left undetermined because of the uncertainty surrounding the case. In this
checklist edition, now that the ICZN Opinion has been published, this matter
can be rectified by restoring the subgeneric name Macrodiremys, which follows the intent of McCord and Joseph-Ouni (2007b).
14:45. Elseya and E. schultzei: Based primarily on mtDNAdata, Georges
et al. (2014) identified three reciprocally monophyletic, deeply divergent
clades within the taxon formerly recognized as Elseya novaeguineae: 1) the
Birds Head (Kepala Burung, Vogelkop, or Doberai Peninsula) population
of western Indonesian New Guinea, 2) the population on the New Guinea
mainland north of the Central Range, and 3) the mainland population south of
the Central Range. They also demonstrated some phylogeographic structure
within each of those three clades, and confirmed the genetic distinction of
E. branderhorsti of the southern lowlands / Fly River floodplain as separate
from the E. novaeguineae clades. They suggested that these three clades each
deserved species rank, and they followed Rhodin and Genorupa (2000) in
noting that the southern form is distinct and undescribed and that the name
E. schultzei (Vogt 1911) is available for the northern population. They also
implied that the name E. novaeguineae should be applied to the Birds Head
population (the source of the type). We follow these recommendations and
now recognize E. schultzei as a full species (again), and await additional work
in progress to determine the appropriate name for the southern form, whose
populations we retain under E. novaeguineae pending further work.
14:46. Flaviemys and F. purvisi: Using molecular data only, Le et
al. (2013) concluded that the species known as Myuchelys purvisi is the
sister taxon to all other taxa that were included in Emydura, Elseya and/or
Myuchelys. To correct this paraphyly, they erected a new monotypic genus,
Flaviemys, with type species Elseya purvisi Wells and Wellington 1985, by
original designation and monotypy. There is also support for this in previous
studies, where Flaviemys purvisi and Myuchelys georgesi were perceived as
a cryptic species pair, very similar by appearance, but on analysis were found
not to be sister taxa (Georges and Adams 1992; Georges et al. 1998; Thomson
and Georges 2009; Georges and Thomson 2010; Fielder et al. 2012; Fielder
2013). We follow this new taxonomy here.
14:47. Pelusios castaneus seychellensis: Based on mitochondrial gene
sequence data from all known lineages of Pelusios, Stuckas et al. (2013)
found that the lectotype of P. seychellensis was nested among specimens
of the West African P. castaneus. They concluded that P. seychellensis was
most likely based on specimens of P. castaneus not native to the Seychelles
Islands, and recommended the synonymy of P. seychellensis with P. castaneus. However, Bour (1983) identified significant morphological differences
between these two taxa, and recently (Bour 2013) argued that seychellensis
might represent an ancient prehistoric introduction of castaneus to the islands
by humans that has subsequently diverged morphologically from the ancestral population. He recommended the use of the subspecific designation P.
castaneus seychellensis until additional comparisons (especially morphological) can be made between castaneus and seychellensis, a recommendation we
have adopted. See also the pertinent discussion of the geographic occurrence
255
256
Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
of this species [P. castaneus] in the distributional data appendix below.
14:48. Podocnemis erythrocephala: The forgotten names Emys bitentaculata and Hydraspis bitentaculata were not listed or synonymized by Wermuth and Mertens (1961, 1977) or Fritz and Havaš (2007). Gray (1830e) first
placed the Cuvier manuscript name Emys bitentaculata under his concept
of Chelys (Hydraspis) and subsequently (Gray 1831d) described it himself
as Hydraspis bitentaculata: “Testa rufa, subtus pallide lutea nigro maculate,
scutello nuchal nullo.” Fitzinger (1835) synonymized both names under his
concept of Hydraspis (Podocnemis) tracaxa (which also included Podocnemis expansa in part and some chelid turtles). Gray’s description did not identify the species very well, but the combination of a red shell (testa rufa), two
barbels (bitentaculata), and lack of a nuchal scute (scutello nuchal nullo) suggests that it is indeed a Podocnemis, and we tentatively place it as most likely
synonymous with P. erythrocephala, which shares those features, including a
red shell margin in juveniles.
14:49. Humboldt Podocnemis names: Alexandre de Humboldt first
published the names Testudo arrau (= P. expansa) and Testudo terekay (= P.
unifilis) in the French version of his original work (Humboldt 1819a:243).
This work was subsequently translated into English (Humboldt 1819b:482),
and later (Humboldt 1820:415) into German. These various translations have
caused some confusion in the literature, and some authors (including our
previous TTWG checklists) have even attributed the names to Humboldt in
Gray (1831d:77). However, the French version remains the original source
for these names. Both names are considered nomina oblita (see also annotation for P. unifilis).
14:50. Podocnemis unifilis: Emys cayennensis was described from
French Guiana by Schweigger (1812), but was incorrectly applied to
Podocnemis erythrocephala for most of its history (reviewed in Pritchard and
Trebbau 1984; but see David 1994 and Bour 2006). In 1819 Humboldt (see
annotation 14:49) described Testudo terekay from Venezuela; however, this
obscure work was ignored by most subsequent authors (but see Schinz 1833).
In 1830 Bell (in Gray 1830e) described Chelys (Hydraspis) lata from Guyana
and this name was also ignored by most subsequent authors until Rhodin et
al. (2008) declared it a nomen oblitum (see also Schneider et al. 2012). All
three of these names apply to the taxon Podocnemis unifilis that was finally
described from Guyana by Troschel (1848), and the latter name has been applied to the Yellow-spotted Amazon Turtle by most (but not all) authors over
the last 165 years. In light of this complicated nomenclatural history, and in an
effort to ensure the stability of Troschel’s name, Vogt et al. (2013) petitioned
the ICZN to conserve the name Podocnemis unifilis Troschel 1848 for the
Yellow-spotted Amazon Turtle, giving it precedence over Emys cayennensis
whenever the two are considered synonymous. Furthermore, they declared
the names Testudo terekay Humboldt 1819a and Chelys (Hydraspis) lata
Bell in Gray 1830e as nomina oblita. Our checklist reflects this arrangement,
pending an ICZN ruling.
appendix – disTRibuTional daTa
Specific distributional updates were only recorded in 2011, 2012, and
2014, and have not been continued in the current checklist except as reflected
in the distributional information under each taxonomic entry.
Distribution Updates 2011
Dermatemys mawii: Honduras (?) deleted from the range as per CBFTT
account (Vogt et al. 2011).
Kinosternon scorpioides albogulare: Population in San Andrés, Colombia, indicated as possible prehistoric or modern introduction, as per CBFTT
accounts (Berry and Iverson 2011, Forero-Medina and Castaño-Mora 2011).
Chrysemys dorsalis:Apparently established in Florida (Krysko et al. 2011).
Graptemys pseudogeographica: Apparently established in Florida
(Krysko et al. 2011).
Malaclemys terrapin and M. t. centrata: Bermuda added to native range
(Parham et al. 2008).
Pseudemys nelsoni: Introduced to Tortola, British Virgin Islands (Owen
et al. 2005).
Trachemys scripta scripta: Apparently established in Florida (Krysko
et al. 2011).
Emys orbicularis orbicularis: Spain deleted from range; Spanish populations are attributable to subspecies E. o. fritzjurgenobsti and E. o. galloitalica,
but not to the nominate subspecies.
Terrapene nelsoni: Chihuahua, Mexico (T. n. klauberi) and Jalisco,
Mexico (T. n. nelsoni) added to the range, as per CBFTT account (Buskirk
and Ponce-Campos 2011).
Batagur kachuga: Occurrence in Nepal confirmed.
Cuora bourreti: Occurrence in Laos indicated as uncertain.
Cuora picturata: Occurrence in central Vietnam confirmed (Ly et al. 2011).
Cuora zhoui: Vietnam added as uncertain occurrence.
Geoemyda spengleri: Laos added to the range (Stuart et al. 2011).
Mauremys caspica caspica: Israel, Jordan and Lebanon deleted from
the range; populations there attributable to M. rivulata. Turkmenistan added
to the range.
Mauremys caspica siebenrocki: Turkmenistan deleted from the range;
population there attributable to M. c. caspica.
Sacalia bealei: Occurrence in Guangxi, China, indicated as uncertain.
Aldabrachelys/Dipsochelys gigantea/dussumieri hololissa: Cousine Island added to extirpated range, Round and Cousine islands added to introduced
range, as per CBFTT account (Gerlach 2011b).
Stigmochelys pardalis: Sudan deleted from the range, and the new nation
of South Sudan added to the range.
Pelodiscus parviformis: Vietnam added to the range.
Acanthochelys pallidipectoris: Mendoza,Argentina, corrected from native
range to introduced, as per CBFTT account (Vinke et al. 2011).
South Sudan: This newly independent nation has seven taxa of freshwater turtles and tortoises: Kinixys belliana belliana, Stigmochelys pardalis,
Cyclanorbis elegans, Cyclanorbis senegalensis, Trionyx triunguis, Pelomedusa
subrufa, and Pelusios adansonii.
Distribution Updates 2012
Cuora bourreti: Occurrence in Laos was confirmed (Stuart et al. 2011),
and possible occurrence in Cambodia was deleted from the checklist following
consultation with range-state biologists.
Mauremys reevesii: Based on analysis of mitochondrial DNA, Suzuki
et al. (2011) indicated that Japanese populations of M. reevesii were possibly
derived from multiple historical introductions from nearby countries, and thus
questioned its traditional status as a presumed native species. They noted the
ability of M. reevesii to hybridize with native M. japonica as a threat, but also
recognized these populations to be valuable in the context of depleted populations elsewhere in its range.
Kinixys: Ranges adjusted according to species ranges outlined by Kindler
et al. (2012).
Nilssonia formosa: Liebing et al. (2012) referred to a record of N. formosa
from Shuangbai (Yunnan, China), and photographs of a specimen from the
Lancang River [= Mekong] in the Xishuanbanna region of Yunnan, as suggesting
that the species has crossed the watershed divide into the Mekong River basin
of Yunnan, China. However, given the substantial documented trade volumes
of live turtles from Myanmar into Yunnan and onwards, and the propensity of
turtles to escape or be intentionally released by humans, combined with the
great biogeographical barriers (despite their very close proximity) between
the Salween, Mekong, and Yangtze, we consider it doubtful that these records
represent natural occurrences.
Mesoclemmys heliostemma: Additional occurrences across the Amazon
basin of Brazil were reported by Molina et al. (2012).
Italy (Sardinia): Vamberger et al. (2011) compared mitochondrial and
nuclear DNA of Testudo graeca from Sardinia with that of T. graeca from
North Africa, and concluded that the near-absence of differentiation from other
graeca populations, and reduced variation within the Sardinian population,
indicated prehistoric introduction into Sardinia by humans. They also reviewed
recent studies of the other non-marine turtle species occurring in Sardinia and
concluded that Sardinia's populations of Testudo hermanni hermanni, T. marginata, and Emys orbicularis galloitalica likely each represented prehistoric
or early historic human introductions.
Latvia: Pupins and Pupina (2011) recorded introduced populations or
individuals of Trachemys scripta elegans, T. s. troostii, Mauremys caspica, M.
rivulata, Testudo horsfieldii, and Pelodiscus sinensis in Latvia. However, they
did not document successful reproduction in the wild, and it remains uncertain
whether these records represent established populations or isolated individuals.
South Korea: Chang et al. (2012) noted that the native softshells in South
Korea are attributable to Pelodiscus maackii, and reported P. sinensis, Trachemys
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
scripta elegans, T. s. scripta, Pseudemys rubriventris, and Mauremys [Ocadia]
sinensis as introduced into the wild.
Distribution Updates 2014
Chelydra acutirostris: Páez et al. (2012) listed the occurrence of this
species in the Colombian Departments of Caldas and Quindio, but did not list
records from Atlántico, Bolivar, Magdalena, or Sucre.
Trachemys c. or o. callirostris: Páez et al. (2012) reported the occurrence
of this taxon in the Colombian Department of Cundinamarca.
Trachemys d. decussata: Parham et al. (2013) documented the occurrence of this taxon in northwestern Jamaica, including hybridization with T.
terrapen; whether the occurrence is native or introduced remains unknown.
Trachemys emolli or grayi emolli: McCranie et al. (2013) documented
the occurrence of this taxon in Honduras, while Ibarra Portillo et al. (2009)
documented its occurrence in eastern El Salvador.
Cuora amboinensis: Wangyal et al. (2012) reported this and four other
species from southern Bhutan, the first turtles reported from that country.
Cuora mouhotii: Rahman (2012) reported the occurrence of this species
in the southern Chittagong Hill Tracts of Bangladesh, Wangyal et al. (2012)
reported it from southern Bhutan, and Ly et al. (2013) extended its range in
southern Vietnam.
Cyclemys gemeli: Wangyal et al. (2012) reported this species from
southern Bhutan.
Melanochelys tricarinata: Wangyal et al. (2012) reported this species
from southern Bhutan.
Aldabrachelys gigantea: The historic and present distribution of native
and introduced populations of the various morphotypes or subspecies (gigantea, arnoldi, and hololissa) of giant tortoises in the Seychelles (including
all granitic and coralline islands) has been analyzed in detail and updated by
Gerlach et al. (2013).
Centrochelys sulcata: Participants at the IUCN/TFTSG Sub-Saharan
African Red List workshop in 2013 noted that C. sulcata occurs in Benin,
Cameroon, and Togo, and may possibly occur in Djibouti, Somalia, Saudi
Arabia, and Yemen. Its presence in Yemen and Saudi Arabia was also previously noted by Gasperetti et al. (1993).
Indotestudo elongata: Wangyal et al. (2012) reported this species from
southern Bhutan.
Kinixys erosa: Participants at the Sub-SaharanAfrican Red List workshop
considered that Kinixys erosa certainly occurs in Benin and Togo, but is absent
from Burkina Faso.
Kinixys homeana: In their CBFTT species account, Luiselli and Diagne
(2013) noted that K. homeana occurs in the Central African Republic. They
questioned its occurrence in Gabon and noted that it does not occur in Congo
(ROC) and that old historical records from the eastern Congo (DRC) were likely
based on misidentified K. erosa. These historical records need further evaluation.
Kinixys nogueyi: Participants at the Sub-Saharan African Red List
workshop considered the distribution of Kinixys nogueyi to include the Central
African Republic, but that the species does not range as far south as Equatorial
Guinea or Gabon, and that records from Mauritania are likely historic, but that
the species no longer occurs there.
Cyclanorbis senegalensis: Participants at the Sub-Saharan African Red
List workshop considered that the occurrence of C. senegalensis is uncertain
in Cameroon, Central African Republic, and Liberia, and that the species is
likely extirpated from Mauritania.
Rafetus swinhoei: Wang et al. (2013) extended and defined the known
recent historic range of this Critically Endangered species in the upper Red
River of southern Yunnan, China.
Mesoclemmys dahli: Páez et al. (2012) and Forero-Medina et al. (2013,
CBFTT account) documented the occurrence of M. dahli in the Colombian
Department of Magdalena.
Mesoclemmys gibba: Páez et al. (2012) recorded the occurrence of M.
gibba in the Colombian Departments of Arauca and Guaviare.
Mesoclemmys perplexa: Campos et al. (2011) recorded the occurrence
of M. perplexa in the Brazilian State of Goiás.
Mesoclemmys vanderhaegei: Vinke et al. (2013) reviewed the distribution
of M. vanderhaegei and concluded that there are no confirmed records for Bolivia.
Platemys platycephala: Páez et al. (2012) recorded the occurrence of
P. platycephala in the Colombian Departments of Guainía, Guaviare, Meta,
and Vichada.
Peltocephalus dumerilianus: Páez et al. (2012) reported P. dumerilianus
to inhabit the Colombian Department of Guaviare.
Pelusios bechuanicus: In earlier versions of this checklist, we included
Congo (DRC) as part of the range of P. bechuanicus. However, we have been
unable to verify this occurrence, and consider that this was based on old literature records of P. upembae, which was originally described as a subspecies
of P. bechuanicus.
Pelusios castaneus: Stuckas et al. (2013) questioned earlier records of
occurrence of P. castaneus on Cape Verde and suggested that our recording of
that presence on our previous checklists was outdated. We have investigated
this further and agree with them. Although Boulenger (1906b) documented the
collection of a specimen of “Sternothaerus derbianus” (= P. castaneus) from
a “small island in Praja Bay, S. Jago” (= Santiago), Chevalier (1935) noted
that the specimen was most likely introduced from West Africa and that no
Caboverdians were aware of any freshwater turtles in the islands. However,
Boulenger’s record (mapped among others by Iverson 1992) led to the assumption
for a long time that the species occurred in the islands; but surveys of the local
herpetofauna have failed to record its presence (Schleich 1982, 1987, 1996b;
Vasconcelos et al. 2013), even as an introduced population. The small island
where it was originally collected housed a prison where turtles had evidently
been released into a small pond at some point in the past. We therefore remove
Cape Verde from the distribution of P. castaneus.
For this same species, we also question whether it occurs natively on
São Tomé. Although it has been recorded from there, and specimens have been
collected and genetically analyzed (Stuckas et al. 2013), the species does not
occur on either nearby Príncipe or the other volcanic oceanic islands in the
same archipelago (Manaças 1956; Jones 1994), nor is there any record of the
species occurring on the nearby continental island of Bioko (Equatorial Guinea)
off the coast of Cameroon. São Tomé was first settled by the Portuguese, who
brought African slaves to the island, so it appears most likely that West African
P. castaneus were introduced to São Tomé in conjunction with the slave trade.
In fact, the genetic analysis by Stuckas et al. (2013) demonstrated that their
São Tomé specimen was essentially indistinguishable from an Ivory Coast
specimen, lending further strength to this theory.
Podocnemis erythrocephala: Páez et al. (2012) reported that P. erythrocephala occurs in the Colombian Department of Guaviare, and perhaps in
Vichada.
Podocnemis lewyana: Páez et al. (2012) reported that the range of P.
lewyana extends into the Colombian Department of Tolima.
Podocnemis sextuberculata: Páez et al. (2012) reported that P. sextuberculata occurs in the Colombian Departments of Caquetá and Putomayo.
Podocnemis vogli: Páez et al. (2012) reported that P. vogli occurs in the
Colombian Department of Guaviare, but did not indicate occurrence in Boyacá.
LITERATURE CITED
This bibliography is divided into two sections: 1) all primary taxonomic citations
noted in this checklist, plus all secondary literature cited in the introduction and
annotations of any of the checklists, and 2) all CBFTT species accounts published
in this monograph project and referred to in the checklist. Many of the citations
listed here are available online as downloadable pdf’s at www.iucn-tftsg.org/
taxonomic-literature-database/.
adler, k.K. 1962. A new name for a Chinese turtle, genus Clemmys. Natural
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adler, K. 2007. The development of systematic reviews of the turtles of the
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adler, K. 2016. Theodore Cantor (1809–1860): pioneer scientific herpetologist
in China. In: Cantor, T. Zoology of Chusan. Society for the Study of Amphibians and Reptiles, Facsimile Reprint, pp. v–xi.
aGaSSiz, l. 1846. Nomenclatoris Zoologici Index Universalis. Solothurn: Jent
and Gassmann, 393 pp.
aGaSSiz, l. 1857a. Contributions to the Natural History of the United States
of America. First Monograph. Vol. I. Part I. Essay on Classification. Part II.
North American Testudinata. Boston: Little, Brown and Co., Vol. I, pp. 1–452.
aGaSSiz, l. 1857b. Contributions to the Natural History of the United States of
America. First Monograph. Vol. II. Part III. Embryology of the Turtle. Boston:
Little, Brown and Co., Vol. II, pp. 453–643.
aHl, e. 1932. Beschreibung einer neuen Schildkröte ausAustralien. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1932(1/3):127–129.
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alvarez del toro, M. 1972. Los Reptiles de Chiapas, 2nd. Ed. Tuxtla Gutierrez,
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aMeGHino, f. 1882. Catálogo explicativo de las colecciones de Antroplogía, prehistoria y de paleontología de Florentino Ameghino. Buenos Aires: Catálogo
de la Sección de la Provincia de Buenos Aires en la Exposición Continental
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anderSon, j. 1875. Description of some new Asiatic mammals and Chelonia.
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anderSon, j. 1876. Note on the plastron of the Gangetic mud-turtle (Emyda dura
of Buchanan-Hamilton). Journal of the Proceedings of the Linnean Society
of Zoology 12:514–516.
anderSon, j. 1879 [“1878”]. Anatomical and Zoological Researches, Comprising an Account of the Zoological Results of the Two Expeditions to Western
Yunnan in 1868 and 1875. London: Bernard Quaritch, Vol. I, 985 pp., Vol.
II, 29 pp. + pls.
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annandale, n. 1912a. The Indian mud–turtles (Trionychidae). Records of the
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annandale, n. 1912b. The aquatic chelonia of the Mahanaddi and its tributaries.
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annandale, n. 1913. The tortoises of Chota Nagpur. Records of the Indian
Museum 9(5):63–78.
annandale, n. 1915a. Notes on some Indian Chelonia. Records of the Indian
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annandale, n. 1915b. Herpetological notes and descriptions. Records of the
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annandale, n. 1918. Chelonia and Batrachia of the Inlé Lake. Records of the
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BarBour, t. 1935. A new Pseudemys from Cat Island, Bahamas. Occasional
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BarBour, t. and Carr, a.f., jr. 1938. Another Bahamian fresh-water tortoise.
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Bauer, a.M. and Petit, R.E. 2004. On the herpetology of Perry’s Arcana and two
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Baur, G. 1888c. Notes on the American Trionychidae. American Naturalist
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Baur, G. 1889. The gigantic land tortoises of the Galapagos Islands. American
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Baur, G. 1891a. The very peculiar tortoise, Carettochelys Ramsay, from New
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Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 7
Chelonian ReseaRCh MonogRaphs no. 5 – CbFTT
ConseRvaTion biology oF FReshWaTeR TuRTles and ToRToises
published 2008–2017
Turtles of the World: Annotated Checklist and Atlas of Taxonomy,
Synonymy, Distribution, and Conservation Status (8th Ed.). Chelonian
Research Monographs 7:1–292.
All CBFTT publications (8 checklists and 100 accounts covering 130 taxa) are
available online as hyperlinked open-access downloadable doi-designated pdf's
CbFTT speCies aCCounTs
(alphabeTiCal by auThoR)
rHodin, a.G.j., PritCHard, P.C.H., van dijk, P.P., SauMure, r.a., BuHlMann, k.a., iverSon, j.B., and MitterMeier, R.A. (Eds.). 2008–2016.
Conservation Biology of Freshwater Turtles and Tortoises: A Compilation
Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group.
Chelonian Research Monographs, No. 5 (Installments 1–9a). Lunenburg,
Massachusetts: Chelonian Research Foundation.
rHodin, a.G.j., iverSon, j.B., van dijk, P.P., SauMure, r.a., BuHlMann, k.a.,
PritCHard, P.C.H., and MitterMeier, R.A. (Eds.). 2016. Conservation
Biology of Freshwater Turtles and Tortoises: A Compilation Project of the
IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian
Research Monographs, No. 5 (Installment 9b). Lunenburg, Massachusetts:
Chelonian Research Foundation.
rHodin, a.G.j., iverSon, j.B., van dijk, P.P., SauMure, r.a., BuHlMann, k.a.,
PritCHard, P.C.H., and MitterMeier, R.A. (Eds.). 2017. Conservation
Biology of Freshwater Turtles and Tortoises: A Compilation Project of the
IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian
Research Monographs, No. 7. In: Chelonian Research Monographs, No.
5 (Supplement No. 1). Lunenburg, Massachusetts: Chelonian Research
Foundation and Turtle Conservancy.
anderS, B. and iverSon, J.B. 2012. Mauremys nigricans (Gray 1834) – Rednecked Pond Turtle, Chinese Red-necked Turtle, Kwangtung River Turtle,
Black-necked Pond Turtle. Chelonian Research Monographs 5(5):068.1–9.
auliya, M., van dijk, P.P., Moll, e.o., and Meylan, P.A. 2016. Amyda
cartilaginea (Boddaert 1770) – Asiatic Softshell Turtle, Southeast Asian
Softshell Turtle. Chelonian Research Monographs 5(9a):092.1–17.
Baker, P.j., diaGne, t., and luiSelli, L. 2015. Cyclanorbis elegans (Gray 1869)
– Nubian Flapshell Turtle. Chelonian Research Monographs 5(8):089.1–7.
Berry, j.f. and iverSon, J.B. 2011. Kinosternon scorpioides (Linnaeus 1766)
– Scorpion Mud Turtle. Chelonian Research Monographs 5(4):063.1–15.
Bertolero, a., CHeylan, M., Hailey, a., livoreil, B., and willeMSen, R.E.
2011. Testudo hermanni (Gmelin 1789) – Hermann’s Tortoise. Chelonian
Research Monographs 5(4):059.1–20.
BHuPatHy, S., weBB, r.G, and PraSCHaG, P. 2014. Lissemys punctata (Bonnaterre 1789) – Indian Flapshell Turtle. Chelonian Research Monographs
5(7):076.1–12.
BlankenSHiP, e.l., Butterfield, B.P., and Godwin, J.C. 2008. Graptemys
nigrinoda Cagle 1954 – Black-knobbed Map Turtle, Black-knobbed
Sawback. Chelonian Research Monographs 5(1):005.1–6.
BoCk, B.C., Páez, v.P., and daza, J.M. 2010. Trachemys callirostris (Gray
1856) – Colombian Slider, Jicotea, Hicotea, Galapago, Morrocoy de Agua.
Chelonian Research Monographs 5(3):042.1–9.
Bour, R. 2008. Pelusios adansonii (Schweigger 1812) – Adanson’s Mud
Turtle. Chelonian Research Monographs 5(1):017.1–4.
Bour, r. and GerlaCH, J. 2008. Pelusios seychellensis (Siebenrock 1906) –
Seychelles Mud Turtle. Chelonian Research Monographs 5(1):018.1–3.
Bour, r., luiSelli, l., Petrozzi, f., SeGniaGBeto, G.H., and CHirio, L. 2016.
Pelusios castaneus (Schweigger 1812) – West African Mud Turtle, Swamp
Terrapin. Chelonian Research Monographs 5(9a):095.1–11.
Bower, d.S. and HodGeS, k.M. 2014. Chelodina expansa Gray 1857 –
Broad-Shelled Turtle, Giant Snake-Necked Turtle. Chelonian Research
Monographs 5(7):071.1–8.
BoyCott, r.C. and BourQuin, O. 2008. Pelomedusa subrufa (Lacepède 1788)
– Helmeted Turtle, Helmeted Terrapin. Chelonian Research Monographs
5(1):007.1–6.
Broadley, d.G. and BoyCott, R.C. 2008. Pelusios rhodesianus Hewitt 1927
– Variable Mud Turtle, Variable Hinged Terrapin. Chelonian Research
Monographs 5(1):004.1–3.
Broadley, d.G. and BoyCott, R.C. 2009. Pelusios sinuatus (Smith 1838) –
Serrated Hinged Terrapin. Chelonian Research Monographs 5(2):036.1–5.
Broadley, d.G. and SaCHSSe, W. 2011. Cycloderma frenatum Peters 1854
– Zambezi Flapshell Turtle, Nkhasi. Chelonian Research Monographs
5(4):055.1–5.
BuHlMann, k.a., GiBBonS, j.w., and jaCkSon, D.R. 2008. Deirochelys
reticularia (Latreille 1801) – Chicken Turtle. Chelonian Research Monographs 5(1):014.1–6.
Bury, r.B. and GerMano, D.J. 2008. Actinemys marmorata (Baird and Girard
1852) – Western Pond Turtle, Pacific Pond Turtle. Chelonian Research
Monographs 5(1):001.1–9.
BuSkirk, j.r. and PonCe-CaMPoS, P. 2011. Terrapene nelsoni Stejneger 1925
– Spotted Box Turtle, Tortuga de Chispitas, Tortuga de Monte. Chelonian
Research Monographs 5(4):060.1–9.
Cann, j., SPenCer, r.-j., welSH, M., and GeorGeS, A. 2015. Myuchelys
georgesi (Cann 1997) – Bellinger River Turtle. Chelonian Research
Monographs 5(8):091.1–9.
Carr, j.l. and Giraldo, A. 2009. Rhinoclemmys nasuta (Boulenger 1902)
– Large-nosed Wood Turtle, Chocoan River Turtle. Chelonian Research
Monographs 5(2):034.1–6.
Cerdá-ardura, a., SoBerón-MoBarak, f., MCGauGH, S.e., and voGt,
R.C. 2008. Apalone spinifera atra (Webb and Legler 1960) – Black
Spiny Softshell Turtle, Cuatrociénegas Softshell, Tortuga Concha Blanda, Tortuga Negra de Cuatrociénegas. Chelonian Research Monographs
5(1):021.1–4.
TTWg, CbFTT, and TeWg CheCklisTs
(ChRonologiCal)
turtle taxonoMy workinG GrouP [BiCkHaM, j.w., iverSon, j.B., ParHaM,
j.f., PHiliPPen, H.d., rHodin, a.G.j., SHaffer, H.B., SPinkS, P.Q., and
van dijk, P.P.]. 2007b. An annotated list of modern turtle terminal taxa
with comments on areas of taxonomic instability and recent change.
Chelonian Research Monographs 4:173–199.
rHodin, a.G.j., van dijk, P.P., and ParHaM, J.F. 2008. Turtles of the world:
annotated checklist of taxonomy and synonymy. Chelonian Research
Monographs 5(1):000.1–38.
turtle taxonoMy workinG GrouP [rHodin, a.G.j., ParHaM, j.f., van dijk,
P.P., and iverSon, j.B.]. 2009. Turtles of the world: annotated checklist
of taxonomy and synonymy, 2009 update, with conservation status summary. Chelonian Research Monographs 5(2):000.39–84.
turtle taxonoMy workinG GrouP [rHodin, a.G.j., van dijk, P.P., iverSon, j.B., and SHaffer, H.B.]. 2010. Turtles of the world, 2010 update:
annotated checklist of taxonomy, synonymy, distribution, and conservation status. Chelonian Research Monographs 5(3):000.85–164.
turtle taxonoMy workinG GrouP [van dijk, P.P., iverSon, j.B., SHaffer, H.B., Bour, r., and rHodin, a.G.j.]. 2011. Turtles of the world,
2011 update: annotated checklist of taxonomy, synonymy, distribution, and conservation status. Chelonian Research Monographs
5(4):000.165–242.
turtle taxonoMy workinG GrouP [van dijk, P.P., iverSon, j.B., SHaffer, H.B., Bour, r., and rHodin, a.G.j.]. 2012. Turtles of the world,
2012 update: annotated checklist of taxonomy, synonymy, distribution, and conservation status. Chelonian Research Monographs
5(5):000.243–328.
turtle taxonoMy workinG GrouP [van dijk, P.P., iverSon, j.B., rHodin, a.G.j., SHaffer, H.B., and Bour, r.]. 2014. Turtles of the world,
7th edition: annotated checklist of taxonomy, synonymy, distribution
with maps, and conservation status. Chelonian Research Monographs
5(7):000.329–479.
turtle extinCtionS workinG GrouP [rHodin, a.G.j., tHoMSon, S., GeorGaliS, G.l., karl, H.-v., danilov, i.G., takaHaSHi, a., de la fuente,
M.S., BourQue, j.r., delfino, M., Bour, r., iverSon, j.B., SHaffer,
H.B., and van dijk, P.P.]. 2015. Turtles and tortoises of the world during the rise and global spread of humanity: first checklist and review
of extinct Pleistocene and Holocene chelonians. Chelonian Research
Monographs 5(8):000e.1–66.
turtle taxonoMy workinG GrouP [rHodin, a.G.j., iverSon, j.B., Bour,
r., fritz, u., GeorGeS, a., SHaffer, H.B., and van dijk, P.P.]. 2017.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
ConGdon, j.d., GraHaM, t.e., HerMan, t.B., lanG, j.w., PaPPaS, M.j., and
BreCke, B.J. 2008. Emydoidea blandingii (Holbrook 1838) – Blanding’s
Turtle. Chelonian Research Monographs 5(1):015.1–12.
daS, I. 2008. Pelochelys cantorii Gray 1864 – Asian Giant Softshell Turtle.
Chelonian Research Monographs 5(1):011.1–6.
daS, I. 2009. Melanochelys tricarinata (Blyth 1856) – Tricarinate Hill
Turtle, Three-keeled Land Turtle. Chelonian Research Monographs
5(2):025.1–5.
daS, I. 2010. Morenia ocellata (Duméril and Bibron 1835) – Burmese Eyed
Turtle. Chelonian Research Monographs 5(3):044.1–5.
daS, i. and BHuPatHy, S. 2009a. Hardella thurjii (Gray 1831) – Crowned
River Turtle. Chelonian Research Monographs 5(2):023.1–6.
daS, i. and BHuPatHy, S. 2009b. Melanochelys trijuga (Schweigger 1812)
– Indian Black Turtle. Chelonian Research Monographs 5(2):038.1–9.
daS, i. and BHuPatHy, S. 2010. Geoclemys hamiltonii (Gray 1830) – Spotted Pond Turtle, Black Pond Turtle. Chelonian Research Monographs
5(3):043.1–6.
daS, i. and SenGuPta, S. 2010. Morenia petersi Anderson 1879 – Indian
Eyed Turtle. Chelonian Research Monographs 5(3):045.1–5.
daS, i. and SinGH, S. 2009. Chitra indica (Gray 1830) – Narrow-headed
Softshell Turtle. Chelonian Research Monographs 5(2):027.1–7.
daS, i., BaSu, d., and SinGH, S. 2010. Nilssonia hurum (Gray 1830) –
Indian Peacock Softshell Turtle. Chelonian Research Monographs
5(3):048.1–6.
daS, i., SenGuPta, S., and PraSCHaG, P. 2010. Pangshura sylhetensis Jerdon 1870 – Assam Roofed Turtle. Chelonian Research Monographs
5(3):046.1–6.
daS, i., SirSi, S., vaSudevan, k., and MurtHy, B.H.C.K. 2014. Nilssonia
leithii (Gray 1872) – Leith’s Softshell Turtle. Chelonian Research Monographs 5(7):075.1–5.
daS, i., MCCorMaCk, t.e.M., van dijk, P.P., HoanG, H.v., and Struijk,
r.P.j.H. 2016. Cuora mouhotii (Gray 1862) – Keeled Box Turtle. Chelonian Research Monographs 5(9b):099.1–12.
deePak, v., raMeSH, M., BHuPatHy, S., and vaSudevan, K. 2011. Indotestudo travancorica (Boulenger 1907) – Travancore Tortoise. Chelonian Research Monographs 5(4):054.1–6.
deePak, v., PraSCHaG, P., and vaSudevan, k. 2014. Vijayachelys silvatica
(Henderson 1912) – Cochin Forest Cane Turtle. Chelonian Research
Monographs 5(7):078.1–7.
dieSMoS, a.C., BuSkirk, j.r., SCHoPPe, S., dieSMoS, M.l.l., Sy, e.y., and
Brown, R.M. 2012. Siebenrockiella leytensis (Taylor 1920) – Palawan
Forest Turtle, Philippine Forest Turtle. Chelonian Research Monographs
5(5):066.1–9.
dodd, C.k., jr. 2008. Sternotherus depressus Tinkle and Webb 1955 – Flattened Musk Turtle. Chelonian Research Monographs 5(1):013.1–7.
ennen, j.r., loviCH, j.e., and joneS, R.L. 2016. Graptemys pearlensis Ennen, Lovich, Kreiser, Selman, and Qualls 2010 – Pearl River Map Turtle.
Chelonian Research Monographs 5(9a):094.1–8.
fielder, d., CHeSSMan, B., and GeorGeS, a. 2015. Myuchelys bellii (Gray
1844) – Western Saw-shelled Turtle, Bell’s Turtle. Chelonian Research
Monographs 5(8):088.1–7.
forero-Medina, G. and CaStaño-Mora, O.V. 2011. Kinosternon scorpioides albogulare (Duméril and Bocourt 1870) – White-throated Mud
Turtle, Swanka Turtle. Chelonian Research Monographs 5(4):064.1–5.
forero-Medina, G., CaStaño-Mora, o.v., CárdenaS-arevalo, G., and
Medina-ranGel, G.f. 2013. Mesoclemmys dahli (Zangerl and Medem
1958) – Dahl’s Toad-Headed Turtle, Carranchina, Tortuga Montañera.
Chelonian Research Monographs 5(6):069.1–8.
freeMan, a. and Cann, J. 2014. Myuchelys latisternum (Gray 1867) –
Sawshelled Turtle, Saw-Shell Turtle. Chelonian Research Monographs
5(7):073.1–8.
freeMan, a., tHoMSon, S., and Cann, j. 2014. Elseya lavarackorum (White and
Archer 1994) – Gulf Snapping Turtle, Gulf Snapper, Riversleigh Snapping
Turtle, Lavarack’s Turtle. Chelonian Research Monographs 5(7):082.1–10.
GeorGeS, a., doody, j.S., eiSeMBerG, C., alaCS, e.a., and roSe, M. 2008.
Carettochelys insculpta Ramsay 1886 – Pig-nosed Turtle, Fly River
Turtle. Chelonian Research Monographs 5(1):009.1–17.
GerlaCH, J. 2008a. Pelusios castanoides intergularis Bour 1983 – Seychelles Yellow-bellied Mud Turtle, Seychelles Chestnut-bellied Terrapin. Chelonian Research Monographs 5(1):010.1–4.
GerlaCH, J. 2008b. Pelusios subniger parietalis Bour 1983 – Seychelles
Black Mud Turtle. Chelonian Research Monographs 5(1):016.1–4.
GerlaCH, J. 2009. Aldabrachelys arnoldi (Bour 1982) – Arnold’s Giant Tortoise. Chelonian Research Monographs 5(2):028.1–5.
GerlaCH, J. 2011b. Aldabrachelys hololissa (Günther 1877) – Seychelles
Giant Tortoise. Chelonian Research Monographs 5(4):061.1–5.
HaGen, C., Platt, S.G., and inniS, C.J. 2009. Leucocephalon yuwonoi (McCord, Iverson, and Boeadi 1995) – Sulawesi Forest Turtle, Kura-kura
Sulawesi. Chelonian Research Monographs 5(2):039.1–7.
HofMeyr, M.D. 2009. Chersina angulata (Schweigger 1812) – Angulate
Tortoise, South African Bowsprit Tortoise. Chelonian Research Monographs 5(2):030.1–6.
HowetH, j.G. and Brown, W.S. 2011. Terrapene coahuila Schmidt and
Owens 1944 – Coahuilan Box Turtle. Chelonian Research Monographs
5(4):049.1–13.
iHlow, f., dawSon, j.e., HartMann, t., and SoM, S. 2016. Indotestudo elongata (Blyth 1854) – Elongated Tortoise, Yellow-headed Tortoise, Yellow
Tortoise. Chelonian Research Monographs 5(9a):096.1–14.
iverSon, j.B. and voGt, R.C. 2011. Kinosternon acutum Gray 1831 – Tabasco Mud Turtle, Montera, Chechagua de Monte. Chelonian Research
Monographs 5(4):062.1–6.
iverSon, j.B., Carr, j.l., CaStaño-Mora, o.v., GalviS-rizo, C.a., rentería-Moreno, l.e., and forero-Medina, G. 2012. Kinosternon dunni
Schmidt 1947 – Dunn’s Mud Turtle, Cabeza de Trozo. Chelonian Research Monographs 5(5):067.1–5.
jaCkSon, D.R. 2010. Pseudemys nelsoni Carr 1938 – Florida Red-bellied
Turtle. Chelonian Research Monographs 5(3):041.1–8.
joneS, r.l. and SelMan, W. 2009. Graptemys oculifera (Baur 1890) –
Ringed Map Turtle, Ringed Sawback. Chelonian Research Monographs
5(2):033.1–8.
kennett, r., roe, j., HodGeS, k., and GeorGeS, a. 2009. Chelodina longicollis (Shaw 1784) – Eastern Long-necked Turtle, Common Longnecked Turtle, Common Snake-necked Turtle. Chelonian Research
Monographs 5(2):031.1–8.
kennett, r., fordHaM, d.a., alaCS, e., Corey, B., and GeorGeS, A. 2014.
Chelodina oblonga Gray 1841 – Northern Snake-Necked Turtle. Chelonian Research Monographs 5(7):077.1–13.
kieSter, a.r. and willey, l.l. 2015. Terrapene carolina (Linnaeus 1758) –
Eastern Box Turtle, Common Box Turtle. Chelonian Research Monographs
5(8):085.1–25.
leary, C.j., doBie, j.l., Mann, t.M., floyd, P.S., and nelSon, D.H. 2008.
Pseudemys alabamensis Baur 1893 – Alabama Red-bellied Cooter, Alabama Red-bellied Turtle. Chelonian Research Monographs 5(1):019.1–9.
lindeMan, P.V. 2008. Sternotherus carinatus Gray 1856 – Razorback Musk
Turtle, Razor-backed Musk Turtle. Chelonian Research Monographs
5(1):012.1–6.
lindeMan, P.v., Stuart, j.n., and killeBrew, F.C. 2016. Graptemys versa
Stejneger 1925 – Texas Map Turtle. Chelonian Research Monographs
5(9a):093.1–10.
loviCH, j.e., Godwin, j.C., and MCCoy, C.J. 2011. Graptemys ernsti
Lovich and McCoy 1992 – Escambia Map Turtle. Chelonian Research
Monographs 5(4):051.1–6.
loviCH, j.e., Godwin, j.C., and MCCoy, C.j. 2014. Graptemys pulchra
Baur 1893 – Alabama Map Turtle. Chelonian Research Monographs
5(7):072.1–6.
loviCH, j.e., SelMan, w., and MCCoy, C.J. 2009. Graptemys gibbonsi Lovich and McCoy 1992 – Pascagoula Map Turtle, Pearl River Map Turtle,
Gibbons’ Map Turtle. Chelonian Research Monographs 5(2):029.1–8.
loviCH, j.e., yaSukawa, y., and ota, H. 2011. Mauremys reevesii (Gray
1831) – Reeves’ Turtle, Chinese Three-keeled Pond Turtle. Chelonian
Research Monographs 5(4):050.1–10.
luiSelli, l. and diaGne, t. 2013. Kinixys homeana Bell 1827 – Home’s
Hinge-Back Tortoise. Chelonian Research Monographs 5(6):070.1–10.
luiSelli, l. and diaGne, t. 2014. Kinixys erosa (Schweigger 1812) – Forest
Hinge-back Tortoise, Serrated Hinge-back Tortoise, Serrated Hinged
Tortoise. Chelonian Research Monographs 5(7):084.1–13.
MaGnuSSon, w.e. and voGt, r.C. 2014. Rhinemys rufipes (Spix 1824) –
Red Side-necked Turtle, Red-footed Sideneck Turtle, Perema. Chelonian
Research Monographs 5(7):079.1–7.
Mantziou, G. and rifai, l. 2014. Mauremys rivulata (Valenciennes in Bory de
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Saint-Vincent 1833) – Western Caspian Turtle, Balkan Terrapin. Chelonian
Research Monographs 5(7):080.1–9.
MarQueS, t.S., BöHM, S., Brito, e.S., CaBrera, M.r., and verdade, l.M. 2014.
Mesoclemmys vanderhaegei (Bour 1973) – Vanderhaege’s Toad-headed
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Citation Format for this Publication:
turtle taxonoMy workinG GrouP [rHodin, a.G.j., iverSon, j.B., Bour, r.
fritz, u., GeorGeS, a., SHaffer, H.B., and van dijk, P.P.]. 2017. Turtles
of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy,
Distribution, and Conservation Status (8th Ed.). In: Rhodin, A.G.J.,
Iverson, J.B., van Dijk, P.P., Saumure, R.A., Buhlmann, K.A., Pritchard,
P.C.H., and Mittermeier, R.A. (Eds.). Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC
Tortoise and Freshwater Turtle Specialist Group. Chelonian Research
Monographs 7:1–292. doi: 10.3854/crm.7.checklist.atlas.v8.2017.
Turtles of the World: Annotated Checklist and Atlas (8th Ed.) – 2017
293
294
Conservation Biology of Freshwater Turtles (ISSN
and Tortoises
• Chelonian Research Monographs, No. 7
1088-7105)
Chelonian ReseaRCh MonogRaphs
Number 7 – August 2017
ISBN: 978-1-5323-5026-9 (online)