Chelonia 2002

The World's Extant Turtle Species 2013










Numerous turtle checklists have been published, but all recent ones like FRITZ & HAVAS (2006, 2007), RHODIN et al. (2008), the TURTLE TAXONOMY WORKING GROUP (2007, 2009, 2010, 2011, 2012) were compiled by splitters meaning that they do not reflect the real species numbers. Oversplitting is a very common phenomenon in modern taxonomy. For most of these authors, the biological species concept is of no use as it strongly favours the subspecies concept and doesn't facilitate to make a full species out of every local population of turtles.


As it is extremely difficult to find new species nowadays, many taxonomists tend to split up existing species into two or even more new species, to convert well-defined subspecies into poorly defined full species or even to describe new forms differing in minor aspects as full species instead of subspecies.


The phylogenetic and the evolutionary species concepts stand tall today and together with the rapidly evolving genetic tools and newly developed statistical methods, they are constantly misused for more and more splitting.


There are some major setbacks in doing this. As the modern species concepts practically do not recognize subspecies, the elevation of nearly all newly recognized taxa to species level produces some really bizarre and grotesque results:


(1) STARKEY et al. (2003) recognize Chrysemys picta dorsalis as a full species. In most herpetologists' opinions Chrysemys picta is the classic example of a well-defined species with numerous subspecies. Wherever these subspecies meet there is a zone of intergradation (see ERNST et al. [2006] and ERNST & LOVICH [2009]). So why does the TTWG accept a work, that separates one of these subspecies and simply makes a full species out of it? Well, a closer look towards the other authors of that paper might help. Iverson, Rhodin and Shaffer and thus the majority of the TTWG are all among them!


(2) STUART & PARHAM (2004) elevated the subspecies Cuora galbinifrons bourreti OBST & REIMANN, 1994 and Cuora galbinifrons picturata LEHR, FRITZ & OBST, 1998 to species level. Following their recognition would mean that all three subspecies of Cuora galbinifrons (the third being the nominate form, C. g. galbinifrons BOURRET, 1939) had the same rank as the morphologically very distinct Cuora flavomarginata GRAY, 1863. So although even a beginner could easily tell the two species C. flavomarginata and C. galbinifrons apart and could also easily see the similarities between the three subspecies of C. galbinifrons, these authors want us to accept that all four taxa shared the same taxonomic level!


(3) SERB et al. (2001) elevated Kinosternon flavescens arizonense GILMORE, 1922 and K. f. durangoense IVERSON, 1979 to full species level. Even for an experienced turtle specialist it is very difficult to differentiate between these two forms and the nominate form K. f. flavescens (AGASSIZ, 1857), especially when they are still young. But how different are all three subspecies compared to all other species of Kinosternon! This, however, is not at all reflected by such kind of taxonomy!


(4) FRITZ et al. (2008), although explicitly stating that it is virtually impossible to tell the two taxa Cyclemys atripons IVERSON & MCCORD, 1997 and Cyclemys pulchristriata FRITZ, GAULKE & LEHR, 1997 apart morphologically, keep insisting that they are full species!


(5) The probably most striking examples of taxonomic inflation are the works of PERAELAE (2002a, 2002b, 2002c). Not only did this author elevate countless previously recognized subspecies within the genus Testudo LINNAEUS, 1758 to species rank, he even described numerous new "species". More recent studies (VAN DER KUYL et al. 2005, PARHAM et al. 2006, FRITZ et al. 2007) have shown that this was all nonsense but for years turtle books and journals uncritically spread this type of information.


Due to these decade-long procedures some turtle genera are highly oversplit. Examples include the North American genus Graptemys AGASSIZ, 1857, the Mesoamerican members of the genus Trachemys MERREM, 1820, the Mediterranean tortoises of the genus Testudo LINNAEUS, 1758, and the Australasian genera Chelodina FITZINGER, 1826, Elseya GRAY, 1867, and Macrochelodina WELLS & WELLINGTON, 1985.


 I truly regret this as all of the authors of the TTWG have provided excellent masterpieces of taxonomy as well. But this kind of splitting is just too much. Turtles are one of the most endangered species on this planet, but according to the TTWG the number of turtle species increases every year!


With this updated checklist (also accessible on my homepage, I try to provide a counterweight to the most recent checklist published by the TURTLE TAXONOMY WORKING GROUP (2012) that recognizes no less than 331 species of turtles! My checklist recognizes 255 well-defined species and 241 subspecies of turtles and seems to be much more balanced. I strongly hope that it will contribute to counterweigh the impact of the checklists produced by professional and semi-professional splitters.


So after having ousted myself as having a very conservative approach towards taxonomy, I will try to present a checklist respecting guidelines that are essential for creating an equally balanced list of all extant turtle taxa:


(1) The biological species concept is favored in this checklist, being the only one truly encouraging subspecies classification.


(2) As subspecies intergrade wherever they occur in sympatry, finding natural hybrids producing viable offspring usually is a clear sign of subspecific status. There are some exceptions, though, especially in northern Australia (GEORGES, pers. comm. to the author).


(3) Similarly to KUCHLING et al. (2007) I believe that even in cases of allopatry and parapatry the subspecies category should be used when morphological and/or genetic data suggest close relationship, and reproductive incompatibility is unlikely. Good examples are the subspecies of the Galápagos Tortoise Chelonoidis nigra (QUOY & GAIMARD, 1824).


(4) Within the range of a given turtle species clinal variation of morphology can sometimes be observed. Populations at the far ends of the distribution area can, however, can be separated well by morphological means. Only these populations should be given subspecies status to avoid oversplitting of very similar populations. Examples include Emydura macquarii (GRAY, 1831), Emys orbicularis (LINNAEUS, 1758), Malaclemys terrapin (SCHOEPFF, 1793) and Testudo graeca LINNAEUS, 1758.


(5) Genetics are a great tool in modern taxonomy but only as long as used with extreme caution and never as the dominant or even exclusive method. Papers suggesting taxonomic changes relying exclusively on genetics are thus usually not accepted. This also means that "cryptic" or "enigmatic" taxa are not recognized. I have difficulties believing that a species is valid if morphological separation using all our sophisticated statistical methods is impossible. Examples include Cyclemys enigmatica FRITZ, GUICKING, AUER, SOMMER, WINK & HUNDSDOERFER, 2008 and Emys trinacris FRITZ, FATTIZZO, GUICKING, TRIPEPI, PENNISI, LENK, JOGER & WINK 2005. If, however, genetic results are supported by good morphological data and – even better – a reliable key to species/subspecies determination within the genus or species is provided, I have no objections against recognizing new taxa. Such a paper is the one by PRASCHAG et al. (2009). It's a very good example how good taxonomy can be done nowadays. Unfortunately, it's hard to find such papers today.


(6) Turtles thought to be extinct are not included in this checklist. Examples include Pelusios seychellensis (SIEBENROCK, 1906), Cuora yunnanensis (BOULENGER, 1906), and Kinosternon hirtipes megacephalum IVERSON, 1981.


I would love to see more people favoring the biological species concept getting involved in this project. Both, professional herpetologists and turtle enthusiasts are welcome. Let's stop this taxonomic trend towards more and more species and less and less subspecies, mostly driven by the excessive use of genetics and the phylogenetic and evolutionary species concepts. If we can't stop this trend, we will end up with a 1,000 turtle genera all containing just a single species without any subspecies!



































Suborder Pleurodira



Family Chelidae—Snake-necked Turtles



Acanthochelys GRAY, 1873—South American Swamp Turtles


Acanthochelys macrocephala (RHODIN, MITTERMEIER & MCMORRIS, 1984)—Pantanal Swamp Turtle

Acanthochelys pallidipectoris (FREIBERG, 1945)—Chaco Swamp Turtle

Acanthochelys radiolata (MIKAN, 1820)—Radiolated Swamp Turtle

Acanthochelys spixii (DUMERIL & BIBRON, 1835)—Spiny-necked Swamp Turtle



Chelodina FITZINGER, 1826—Thin-necked Snake-necked Turtles


Chelodina longicollis (SHAW, 1794)—Australian Snake-necked Turtle

Chelodina mccordi RHODIN, 1994—McCord's Snake-necked Turtle

            Chelodina mccordi mccordi RHODIN, 1994—McCord's Snake-necked Turtle

            Chelodina mccordi roteensis MCCORD, JOSEPH-OUNI & HAGEN, 2007—Roti Snake-necked Turtle

            Chelodina mccordi timorensis MCCORD, JOSEPH-OUNI & HAGEN, 2007—Timor Snake-necked Turtle

Chelodina novaeguineae BOULENGER, 18881—New Guinea Snake-necked Turtle

            Chelodina novaeguineae novaeguineae BOULENGER, 1888—New Guinea Snake-necked Turtle

            Chelodina novaeguineae canni MCCORD & THOMSON, 2002—Cann's Snake-necked Turtle

            Chelodina novaeguineae gunaleni MCCORD & JOSEPH-OUNI, 2007—Gunalen's Snake-necked Turtle

            Chelodina novaeguineae reimanni PHILIPPEN & GROSSMANN, 1990—Reimann's Snake-necked Turtle

Chelodina pritchardi RHODIN, 1994—Pritchard's Snake-necked Turtle

Chelodina steindachneri SIEBENROCK, 1914—Steindachner's Snake-necked Turtle



Chelus DUMÉRIL, 1806—Matamata Turtles


Chelus fimbriata (SCHNEIDER, 1783)—Matamata



Elseya GRAY, 18672—Australian Snapping Turtles


Elseya branderhorsti (OUWENS, 1914)—Branderhorst's Snapping Turtle

Elseya dentata (GRAY, 1863)3—Northern Australian Snapping Turtle

            Elseya dentata dentata (GRAY, 1863)—Victoria River Snapping Turtle

            Elseya dentata jukesi WELLS, 20074—Alligator River Snapping Turtle

Elseya lavarackorum (WHITE & ARCHER, 1994)5—Gulf Snapping Turtle

            Elseya lavarackorum lavarackorum (WHITE & ARCHER, 1994)—Gulf Snapping Turtle 

            Elseya lavarackorum albagula THOMSON, GEORGES & LIMPUS, 2006—Southern Snapping Turtle

            Elseya lavarackorum irwini CANN, 1997—Burdekin River Snapping Turtle



Elusor CANN & LEGLER, 1994—Mary River Turtles


Elusor macrurus CANN & LEGLER, 1994—Mary River Turtle



Emydura BONAPARTE, 1836—Short-necked Turtles


Emydura macquarii (GRAY, 1831)—Murray River Short-necked Turtle

            Emydura macquarii macquarii6 (GRAY, 1831)—Murray River Short-necked Turtle

            Emydura macquarii kreffti (GRAY, 1871)—Krefft's Short-necked Turtle

Emydura subglobosa (KREFFT, 1876)7—Red-bellied Short-necked Turtle

            Emydura subglobosa subglobosa (KREFFT, 1876)—Red-bellied Short-necked Turtle

            Emydura subglobosa tanybaraga (CANN, 1997)—Yellow-faced Short-necked Turtle

            Emydura subglobosa worrelli (WELLS & WELLINGTON, 1985)—Worrell's Short-necked Turtle

Emydura victoriae (GRAY, 1842)8—Red-faced Short-necked Turtle



Hydromedusa WAGLER, 1830—South American Snake-necked Turtles


Hydromedusa maximiliani (MIKAN, 1820)—Brazilian Snake-necked Turtle

Hydromedusa tectifera COPE, 1869—South American Snake-necked Turtle



Macrochelodina WELLS & WELLINGTON, 19859—Thick-necked Snake-necked Turtles


Macrochelodina expansa (GRAY, 1857)—Giant Snake-necked Turtle

Macrochelodina parkeri (RHODIN & MITTERMEIER, 1976)—Parker's Snake-necked Turtle

Macrochelodina rugosa (OGILBY, 1890)10—Northern Australian Snake-necked Turtle

            Macrochelodina rugosa rugosa (OGILBY, 1890)—Northern Australian Snake-necked Turtle

            Macrochelodina rugosa burrungandjii (THOMSON, KENNETT & GEORGES, 2000)—Arnhem Land Snake-necked Turtle

            Macrochelodina rugosa walloyarrina—MCCORD & JOSEPH-OUNI, 2007—Kimberley Longneck Turtle



Macrodiremys MCCORD & JOSEPH-OUNI, 200711—Narrow-breasted Snake-necked Turtles


Macrodiremys colliei (GRAY, 1856)12—Narrow-breasted Snake-necked Turtle



Mesoclemmys GRAY, 1873—Toad-headed Turtles


Mesoclemmys dahli (ZANGERL & MEDEM, 1958)13—Dahl's Toad-Headed Turtle

            Mesoclemmys dahli dahli (ZANGERL & MEDEM, 1958)—Dahl's Toad-Headed Turtle

            Mesoclemmys dahli zuliae (PRITCHARD & TREBBAU, 1984)—Zulia Toad-headed Turtle

Mesoclemmys gibba (SCHWEIGGER, 1812)14—Gibba Turtle

            Mesoclemmys gibba gibba (SCHWEIGGER, 1812)—Gibba Turtle

            Mesoclemmys gibba perplexa BOUR & ZAHER, 2005—Piaui Toad-headed Turtle

Mesoclemmys hogei (MERTENS, 1967)—Hoge's Side-necked Turtle

Mesoclemmys nasuta (SCHWEIGGER, 1812)—Common Toad-headed Turtle

Mesoclemmys raniceps (GRAY, 1855)15—Amazon Toad-headed Turtle

            Mesoclemmys raniceps raniceps (GRAY, 1855)—Amazon Toad-headed Turtle

            Mesoclemmys raniceps heliostemma MCCORD, JOSEPH-OUNI & LAMAR, 2001—Peruvian Toad-headed Turtle

Mesoclemmys tuberculata (LUEDERWALDT, 1926)—Tuberculate Toad-headed Turtle

Mesoclemmys vanderhaegei (BOUR, 1973)—Vanderhaege's Toad-headed Turtle



Phrynops WAGLER, 1830—South American Side-necked Turtles


Phrynops geoffroanus (SCHWEIGGER, 1812)16—Geoffroy's Side-necked Turtle

            Phrynops geoffroanus geoffroanus (SCHWEIGGER, 1812)—Geoffroy's Side-necked Turtle

            Phrynops geoffroanus tuberosus (PETERS, 1870)—Peters' Side-necked Turtle

Phrynops hilarii (DUMERIL & BIBRON, 1835)—Saint Hilaire's Side-necked Turtle

Phrynops williamsi RHODIN & MITTERMEIER, 1983—Williams' Side-necked Turtle



Platemys WAGLER, 1830—Twist-necked Turtles


Platemys platycephala (SCHNEIDER, 1792)—Twist-necked Turtle

            Platemys platycephala platycephala (SCHNEIDER, 1792)—Twist-necked Turtle

            Platemys platycephala melanonota ERNST, 1984—Dark Twist-necked Turtle



Pseudemydura SIEBENROCK, 1901—Western Swamp Turtles


Pseudemydura umbrina SIEBENROCK, 1901—Western Swamp Turtle



Rheodytes LEGLER & CANN, 1980—Fitzroy River Turtles


Rheodytes leukops LEGLER & CANN, 1980—Fitzroy River Turtle



Rhinemys MCCORD, JOSEPH-OUNI & LAMAR, 2001—Red Side-necked Turtles


Rhinemys rufipes (SPIX, 1824)—Red Side-necked Turtle



Wollumbinia WELLS 2007 or Myuchelys GEORGES & THOMSON 201017—Saw-shelled Snapping Turtles


Wollumbinia or Myuchelys latisternum GRAY, 186718—Saw-shelled Snapping Turtle

            Wollumbinia or Myuchelys latisternum latisternum GRAY, 1867—Saw-shelled Snapping Turtle

            Wollumbinia or Myuchelys latisternum bellii (GRAY, 1844)—Bell's Snapping Turtle

Wollumbinia or Myochelys purvisi WELLS & WELLINGTON, 198519—Purvis' Snapping Turtle    

            Wollumbinia or Myuchelys purvisi purvisi WELLS & WELLINGTON, 1985—Purvis' Snapping Turtle

            Wollumbinia or Myuchelys purvisi georgesi CANN, 1997—Georges' Snapping Turtle  

Wollumbinia or Myuchelys novaeguineae (MEYER, 1874)20—New Guinea Snapping Turtle




Family Pelomedusidae—African Side-necked Turtles



Pelomedusa WAGLER, 1830—Helmeted Turtles


Pelomedusa subrufa (LACEPEDE, 1788)—Helmeted Turtle



Pelusios WAGLER, 1830—African Mud Turtles


Pelusios adansonii (SCHWEIGGER, 1812)—Adanson's Mud Turtle

Pelusios bechuanicus FITZSIMONS, 1932—Okavango Mud Turtle

Pelusios broadleyi BOUR, 1986—Turkana Mud Turtle

Pelusios carinatus LAURENT, 1956—African Keeled Mud Turtle

Pelusios castaneus (SCHWEIGGER, 1812)21—West African Mud Turtle

            Pelusios castaneus castaneus (SCHWEIGGER, 1812)—West African Mud Turtle

            Pelusios castaneus chapini LAURENT, 1965—Central African Mud Turtle

Pelusios castanoides HEWITT, 193122—Yellow-bellied Mud Turtle

            Pelusios castanoides castanoides HEWITT, 1931—Yellow-bellied Mud Turtle

            Pelusios castanoides intergularis BOUR, 1983—Yellow-bellied Mud Turtle

Pelusios cupulatta BOUR & MARAN, 2003—Côte d'Ivoire Mud Turtle

Pelusios gabonensis (DUMERIL, 1856)—African Forest Turtle

Pelusios marani BOUR, 2000—Maran's Mud Turtle

Pelusios nanus LAURENT, 1956—African Dwarf Mud Turtle

Pelusios niger (DUMERIL & BIBRON, 1835)—West African Black Mud Turtle

Pelusios rhodesianus HEWITT, 1927—Variable Mud Turtle

Pelusios sinuatus (SMITH, 1838)—East African Serrated Mud Turtle

Pelusios subniger (LACEPEDE, 1788) 22—East African Black Mud Turtle

            Pelusios subniger subniger (LACEPEDE, 1788)—East African Black Mud Turtle

            Pelusios subniger parietalis BOUR, 1983—Seychelles Black Mud Turtle

Pelusios upembae BROADLEY, 1981—Upemba Mud Turtle

Pelusios williamsi LAURENT, 1965—Williams' Mud Turtle

            Pelusios williamsi williamsi LAURENT, 1965—Lake Victoria Mud Turtle

            Pelusios williamsi laurenti BOUR, 1984—Ukerewe Island Mud Turtle

            Pelusios williamsi lutescens LAURENT, 1965—Albert Nile Mud Turtle




Family Podocnemidae—American Side-necked Turtles



Erymnochelys BAUR, 1888—Madagascan Big-headed Turtles


Erymnochelys madagascariensis (GRANDIDIER, 1867)—Madagascan Big-headed Turtle



Peltocephalus DUMERIL & BIBRON, 1835—Big-headed Amazon River Turtles


Peltocephalus dumerilianus DUMÉRIL & BIBRON, 1835—Big-headed Amazon River Turtle



Podocnemis WAGLER, 1830—South American River Turtles


Podocnemis cayennensis (SCHWEIGGER, 1812)23—Yellow-spotted River Turtle

Podocnemis erythrocephala (SPIX, 1824)—Red-headed River Turtle

Podocnemis expansa (SCHWEIGGER, 1812)—South American Giant River Turtle

Podocnemis lewyana DUMERIL, 1852—Magdalena River Turtle

Podocnemis sextuberculata CORNALIA, 1849—Six-tubercled River Turtle

Podocnemis vogli MUELLER, 1935—Savanna Side-necked Turtle





Suborder Cryptodira



Family Carettochelyidae—Pig-nosed Turtles



Carettochelys RAMSAY, 1886—Pig-nosed Turtles


Carettochelys insculpta RAMSAY, 1886—Pig-nosed Turtle




Family Cheloniidae—Marine Turtles



Caretta RAFINESQUE, 1814—Loggerhead Turtles


Caretta caretta (LINNAEUS, 1758)—Loggerhead Turtle



Chelonia BROGNIART, 1800—Green Turtles


Chelonia mydas (LINNAEUS, 1758)—Green Turtle



Eretmochelys FITZINGER, 1843—Hawksbill Seaturtles


Eretmochelys imbricata (LINNAEUS, 1766)24—Hawksbill Turtle



Lepidochelys FITZINGER, 1843—Ridley Seaturtles


Lepidochelys kempii (GARMAN, 1880)—Kemp's Ridley Turtle

Lepidochelys olivacea (ESCHSCHOLTZ, 1829)—Olive Ridley Turtle



Natator MCCULLOCH, 1908—Flatback Seaturtles


Natator depressus (GARMAN, 1880)—Flatback Seaturtle




Family Chelydridae—Snapping Turtles



Chelydra SCHWEIGGER, 1812—Snapping Turtles


Chelydra acutirostris PETERS, 1862—South American Snapping Turtle

Chelydra serpentina (LINNAEUS, 1758)—Snapping Turtle

            Chelydra serpentina serpentina (LINNAEUS, 1758)—Common Snapping Turtle

            Chelydra serpentina osceola STEJNEGER, 191825—Florida Snapping Turtle

Chelydra rossignonii (BOCOURT, 1868)—Central American Snapping Turtle



Macrochelys GRAY, 1856—Alligator Snapping Turtles


Macrochelys temminckii (TROOST, 1835)—Alligator Snapping Turtle




Family Dermatemydidae—River Turtles



Dermatemys GRAY, 1847—Central American River Turtles


Dermatemys mawii GRAY, 1847—Central American River Turtle




Family Dermochelyidae—Leatherback Turtles



Dermochelys BLAINVILLE, 1816—Leatherback Turtles


Dermochelys coriacea (VANDELLI, 1761)—Leatherback Turtle




Family Emydidae—New World Pond Turtles



Chrysemys GRAY, 1844—Painted Turtles


Chrysemys picta (SCHNEIDER, 1783)—Painted Turtle

            Chrysemys picta picta (SCHNEIDER, 1783)—Eastern Painted Turtle

            Chrysemys picta bellii (GRAY, 1831)—Western Painted Turtle

            Chrysemys picta dorsalis AGASSIZ, 185726—Southern Painted Turtle

            Chrysemys picta marginata AGASSIZ, 1857—Midland Painted Turtle



Clemmys RITGEN, 1828—Pond Turtles


Clemmys guttata (SCHNEIDER, 1792)—Spotted Turtle



Deirochelys AGASSIZ, 1857—Chicken Turtles


Deirochelys reticularia (LATREILLE, 1801)—Chicken Turtle

            Deirochelys reticularia reticularia (LATREILLE, 1801)—Eastern Chicken Turtle

            Deirochelys reticularia chrysea SCHWARTZ, 1956—Florida Chicken Turtle

            Deirochelys reticularia miaria SCHWARTZ, 1956—Western Chicken Turtle



Emys DUMERIL, 180627—European Pond Turtles


Emys blandingii (HOLBROOK, 1838)—Blanding's Turtle

Emys marmorata (BAIRD & GIRARD, 1852)—Pacific Pond Turtle

Emys orbicularis (LINNAEUS, 1758)28—European Pond Turtle

            Emys orbicularis orbicularis (LINNAEUS, 1758)—Common European Pond Turtle

            Emys orbicularis galloitalica FRITZ, 199529—Franco-Italian Pond Turtle

            Emys orbicularis hellenica (VALENCIENNES, 1832)—Hellenic Pond Turtle

            Emys orbicularis luteofusca FRITZ, 198930—Central Turkey Pond Turtle

            Emys orbicularis occidentalis FRITZ, 1993—Magrab Pond Turtle

            Emys orbiculairs persica EICHWALD, 1831—Persian Pond Turtle



Glyptemys AGASSIZ, 1857—Wood and Bog Turtles


Glyptemys insculpta (LECONTE, 1830)—Wood Turtle

Glyptemys muhlenbergii (SCHOEPFF, 1801)—Bog Turtle



Graptemys AGASSIZ, 185731—Map Turtles


Graptemys caglei HAYNES & MCKOWN, 1974—Cagle's Map Turtle

Graptemys geographica (LESUEUR, 1817)—Common Map Turtle

Graptemys nigrinoda CAGLE, 1954—Black-knobbed Map Turtle

            Graptemys nigrinoda nigrinoda CAGLE, 1954—Northern Black-knobbed Map Turtle

            Graptemys nigrinoda delticola FOLKERTS & MOUNT, 1969—Southern Black-knobbed Map Turtle

Graptemys oculifera (BAUR, 1890)32—Ringed Map Turtle

            Graptemys oculifera oculifera (BAUR, 1890)—Ringed Map Turtle

            Graptemys oculifera flavimaculata CAGLE, 1954—Yellow-blotched Map Turtle

Graptemys pseudogeographica (GRAY, 1831)33—False Map Turtle

            Graptemys pseudogeographica pseudogeographica (GRAY, 1831)—False Map Turtle

            Graptemys pseudogeographica kohnii (BAUR, 1890)—Mississippi Map Turtle

            Graptemys pseudogeographica ouachitensis CAGLE, 1953—Ouachita Map Turtle

            Graptemys pseudogeographica sabinensis CAGLE, 1953—Sabine Map Turtle

            Graptemys pseudogeographica versa STEJNEGER, 1925—Texas Map Turtle

Graptemys pulchra BAUR, 189334—Alabama Map Turtle

            Graptemys pulchra pulchra BAUR, 1893—Alabama Map Turtle

            Graptemys pulchra barbouri CARR & MARCHAND, 1942—Barbour's Map Turtle

            Graptemys pulchra ernsti LOVICH & MCCOY, 1992—Escambia Map Turtle

            Graptemys pulchra gibbonsi LOVICH & MCCOY, 1992—Pascagoula Map Turtle

            Graptemys pulchra pearlensis ENNEN, LOVICH, KREISER, SELMAN & QUALLS (2010)—Pearl Map Turtle



Malaclemys GRAY, 1844—Diamondback Terrapins


Malaclemys terrapin (SCHOEPFF, 1793)35—Diamondback Terrapin

            Malaclemys terrapin terrapin (SCHOEPFF, 1793)36—Northern Diamondback Terrapin

            Malaclemys terrapin pileata (WIED, 1865)37—Mississippi Diamondback Terrapin



Pseudemys GRAY, 185538—Cooters


Pseudemys concinna (LECONTE, 1830)39—River Cooter

            Pseudemys concinna concinna (LECONTE, 1830)—Eastern River Cooter

            Pseudemys concinna floridana (LECONTE, 1830)—Florida River Cooter

            Pseudemys concinna peninsularis CARR, 1938—Peninsula Cooter

            Pseudemys concinna suwanniensis CARR, 1937—Suwannee Cooter

Pseudemys gorzugi WARD, 1984—Rio Grande Cooter

Pseudemys rubriventris (LECONTE, 1830)40—Red-bellied Cooter

            Pseudemys rubriventris rubriventris (LECONTE, 1830)—Northern Red-bellied Cooter

            Pseudemys rubriventris alabamenis BAUR, 1893—Alabama Red-bellied Turtle

            Pseudemys rubriventris nelsoni CARR, 1938—Florida Red-bellied Cooter

Pseudemys texana BAUR, 1893—Texas Cooter



Terrapene MERREM, 1820—North American Box Turtles


Terrapene carolina (LINNAEUS, 1758)41—Eastern Box Turtle

            Terrapene carolina carolina (LINNAEUS, 1758)—Eastern Box Turtle

            Terrapene carolina bauri TAYLOR, 1895—Florida Box Turtle

            Terrapene carolina mexicana (GRAY, 1849)—Mexican Box Turtle

            Terrapene carolina triunguis (AGASSIZ 1857)—Three-toed Box Turtle

Terrapene coahuila SCHMIDT & OWENS 1944—Coahuila box turtle

Terrapene nelsoni STEJNEGER, 1925—Spotted Box Turtle

            Terrapene nelsoni nelsoni STEJNEGER, 1925—Southern Spotted Box Turtle

            Terrapene nelsoni klauberi BOGERT, 1943—Northern Spotted Box Turtle

Terrapene ornata (AGASSIZ, 1857)—Ornate Box Turtle

            Terrapene ornata ornata (AGASSIZ, 1857)—Ornate Box Turtle

            Terrapene ornata luteola SMITH & RAMSAY, 1944—Desert Box Turtle

Terrapene yucatana (BOULENGER, 1895)—Yucatan Box Turtle



Trachemys AGASSIZ, 185742—Slider Turtles


Trachemys decorata (BARBOUR & CARR, 1940)—Hispaniolan Slider

Trachemys decussata (GRAY, 1831)—Cuban Slider

            Trachemys decussata decussata (GRAY, 1831)—Eastern Cuban Slider

            Trachemys decussata angusta (BARBOUR & CARR, 1940)—Western Cuban Slider

Trachemys dorbigni (DUMERIL & BIBRON, 1835)—d'Orbigny's Slider

            Trachemys dorbigni dorbigni (DUMERIL & BIBRON, 1835)—d'Orbigny's Slider

            Trachemys dorbigni adiutrix VANZOLINI, 1995—Brazilian Slider

Trachemys gaigeae (HARTWEG, 1939)—Big Bend Slider

            Trachemys gaigeae gaigeae (HARTWEG, 1939)—Big Bend Slider

            Trachemys gaigeae hartwegi (LEGLER, 1990)—Nazas Slider

Trachemys grayi (BOCOURT, 1868)—Gray's Slider

            Trachemys grayi grayi (BOCOURT, 1868)—Gray's Slider

            Trachemys grayi emolli (LEGLER, 1990)—Nicaraguan Slider

            Trachemys grayi panamensis McCORD, JOSEPH-OUNI, HAGEN & BLANCK 2010—Panamanian Slider

Trachemys nebulosa (VAN DENBURGH, 1895)—Baja California Slider

            Trachemys nebulosa nebulosa (VAN DENBURGH, 1895)—Baja California Slider

            Trachemys nebulosa hiltoni (CARR, 1942)—Fuerte River Slider

Trachemys ornata (GRAY, 1831)—Ornate Slider

            Trachemys ornata ornata (GRAY, 1831)—Ornate Slider

            Trachemys ornata callirostris (GRAY, 1856)—Colombian Slider

            Trachemys ornata cataspila (GUENTHER, 1885)—Huestecan Slider

            Trachemys ornata chichiriviche (PRITCHARD & TREBBAU, 1984)—Venezuelan Slider

            Trachemys ornata taylori (LEGLER, 1960)—Cuatrocienegas Slider

            Trachemys ornata venusta (GRAY, 1856)—Mesoamerican Slider

            Trachemys ornata yaquia (LEGLER & WEBB, 1970)—Yaqui Slider

Trachemys scripta (SCHOEPFF, 1792)—Slider Turtle

            Trachemys scripta scripta (SCHOEPFF, 1792)—Yellow-bellied Slider

            Trachemys scripta elegans (WIED, 1839)—Red-eared Slider

            Trachemys scripta troosti (HOLBROOK, 1836)—Cumberland Slider

Trachemys stejnegeri (SCHMIDT, 1928)—Central Antillean Slider

            Trachemys stejnegeri stejnegeri (SCHMIDT, 1928)—Puerto Rican Slider

            Trachemys stejnegeri malonei (BARBOUR & CARR, 1938)—Inagua Slider

            Trachemys stejnegeri vicina (BARBOUR & CARR, 1938)—Dominican Slider

Trachemys terrapen (LACEPEDE, 1788)—Jamaican Slider




Family Geoemydidae—Old World Pond Turtles



Batagur GRAY, 185643—River Terrapins


Batagur affinis (CANTOR, 1847)—Malay River Terrapin

         Batagur affinis affinis (CANTOR, 1847)—Western Malay River Terrapin

         Batagur affinis edwardmolli PRASCHAG, HOLLOWAY, GEORGES, PAECKERT, HUNDSDOERFER & FRITZ 2009—Eastern Malay River Terrapin

Batagur baska (GRAY, 1830)—River Terrapin

Batagur borneoensis (SCHLEGEL & MUELLER, 1845)—Painted River Terrapin

Batagur dhongoka (GRAY, 1835)—Three-striped Roof Turtle

Batagur kachuga (GRAY, 1831)—Red-crowned Roof Turtle

Batagur trivittata (DUMERIL & BIBRON, 1835)—Burmese Roof Turtle



Cuora GRAY, 185644—Asian Box Turtles


Cuora amboinensis (DAUDIN, 1801)45—Ambon Box Turtle

            Cuora amboinensis amboinensis (DAUDIN, 1801)—Eastern Ambon Box Turtle

            Cuora amboinensis kamaroma RUMMLER & FRITZ 1991—Western Ambon Box Turtle

Cuora flavomarginata (GRAY, 1863)—Yellow-margined Box Turtle

            Cuora flavomarginata flavomarginata (GRAY, 1863)—Yellow-margined Box Turtle

            Cuora flavomarginata evelynae ERNST & LOVICH, 1990—Ryukyu Box Turtle

Cuora galbinifrons BOURRET, 193946—Indochinese Box Turtle

            Cuora galbinifrons galbinifrons BOURRET, 1939—Chinese Box Turtle

            Cuora galbinifrons bourreti OBST & REIMANN, 1994—Bourret's Box Turtle

            Cuora galbinifrons picturata LEHR, FRITZ & OBST, 1998—Southern Vietnam Box Turtle

Cuora mccordi ERNST, 1988—McCord's Box Turtle

Cuora mouhotii (GRAY, 1862)—Keeled Box Turtle

            Cuora mouhotii mouhotii (GRAY, 1862)—Northern Keeled Box Turtle

            Cuora mouhotii obsti (FRITZ, ANDREAS & LEHR, 1998)—Southern Keeled Box Turtle

Cuora pani SONG, 198447—Pan's Box Turtle

            Cuora pani pani SONG, 1984—Pan's Box Turtle

            Cuora pani aurocapitata LUO & ZONG, 1988—Yellow-headed Box Turtle

Cuora trifasciata (BELL, 1825)—Three-striped Box Turtle

Cuora zhoui ZHAO, 1990—Zhou's Box Turtle



Cyclemys BELL, 183448—Asian Leaf Turtles


Cyclemys atripons IVERSON & MCCORD, 199749—Black-bridged Leaf Turtle

Cyclemys dentata (GRAY, 1831)—Asian Leaf Turtle

Cyclemys oldhamii GRAY, 186350—Oldham's Leaf Turtle



Geoclemys GRAY, 1856—Spotted Pond Turtles


Geoclemys hamiltonii (GRAY, 1831)—Spotted Pond Turtle



Geoemyda GRAY, 1834—Leaf Turtles


Geoemyda japonica FAN, 1931—Ryukyu Leaf Turtle

Geoemyda spengleri (GMELIN, 1789)—Black-breasted Leaf Turtle



Hardella GRAY, 1870—Crowned River Turtles


Hardella thurjii (GRAY, 1831)—Crowned River Turtle



Heosemys STEJNEGER, 1902—Forest Turtles


Heosemys annandalii (BOULENGER, 1903)—Yellow-headed Temple Turtle

Heosemys depressa (ANDERSON, 1875)—Myanmar Forest Turtle

Heosemys grandis (GRAY, 1860)—Giant Asian Pond Terrapin

Heosemys spinosa (GRAY, 1831)—Spiny Turtle



Malayemys LINDHOLM, 1931—Snail-eating Turtles


Malayemys subtrijuga (SCHLEGEL & MUELLER, 1845)51—Malayan Snail-eating Turtle

            Malayemys subtrijuga subtrijuga (SCHLEGEL & MUELLER, 1845)—Eastern Malayan Snail-eating Turtle

            Malayemys subtrijuga macrocephala (GRAY, 1859)—Western Malayan Snail-eating Turtle



Mauremys GRAY, 1869—Stripe-necked Turtles


Mauremys annamensis (SIEBENROCK, 1903)—Annam Leaf Turtle

Mauremys caspica (GMELIN, 1774)52—Caspian Turtle

            Mauremys caspica caspica (GMELIN, 1774)—Eastern Caspian Turtle

            Mauremys caspica rivulata (VALENCIENNES, 1833)—Western Caspian Turtle

Mauremys japonica (TEMMINCK & SCHLEGEL, 1835)—Japanese Pond Turtle

Mauremys leprosa (SCHWEIGGER, 1812)—Mediterranean Pond Turtle

            Mauremys leprosa leprosa (SCHWEIGGER 1812)—Spanish Turtle

            Mauremys leprosa saharica SCHLEICH, 1996—Sahara Pond Turtle

            Mauremys leprosa vanmeerhaeghei BOUR & MARAN, 199853—Oued Draa Pond Turtle

Mauremys mutica (CANTOR, 1842)—Yellow Pond Turtle

            Mauremys mutica mutica (CANTOR, 1842)—Asian Yellow Pond Turtle

            Mauremys mutica kami YASUKAWA, OTA & IVERSON, 1996—Ryukyu Yellow Pond Turtle

Mauremys nigricans (GRAY, 1834)—Red-necked Pond Turtle

Mauremys reevesii (GRAY, 1831)—Reeves' Turtle

Mauremys sinensis (GRAY, 1834)—Chinese Striped-necked Turtle



Melanochelys GRAY, 1869—Indian Black Turtles


Melanochelys tricarinata (BLYTH, 1856)—Tricarinate Hill Turtle

Melanochelys trijuga (SCHWEIGGER, 1812)54—Indo-Burmese Black Turtle

            Melanochelys trijuga trijuga (SCHWEIGGER, 1812)—Peninsula Black Turtle

            Melanochelys trijuga coronata (ANDERSON, 1878)—Malabar Black Turtle

            Melanochelys trijuga edeniana THEOBALD, 1876—Burmese Black Turtle

            Melanochelys trijuga thermalis (LESSON, 1830)55—Red-spotted Black Turtle




Morenia GRAY, 187056—Indo-Burmese Eyed Turtles


Morenia ocellata (DUMERIL & BIBRON, 1835)—Burmese Eyed Turtle

Morenia petersi (ANDERSON, 1879)—Indian Eyed Turtle



Notochelys GRAY, 186357—Flat-shelled Turtles


Notochelys platynota (GRAY, 1834)—Flat-shelled Turtle

Notochelys yuwonoi (MCCORD, IVERSON, SPINKS & SHAFFER, 2000)—Sulawesi Forest Turtle



Orlitia GRAY, 1873—Malaysian Giant Turtles


Orlitia borneensis GRAY, 1873—Malaysian Giant Turtle



Pangshura GRAY, 1856—Asian Roof Turtles


Pangshura smithii (GRAY, 1863)—Brown Roof Turtle

            Pangshura smithii smithii (GRAY, 1863)—Brown Roof Turtle

            Pangshura smithii pallidipes MOLL, 1987—Pale-footed Roof Turtle

Pangshura sylhetensis JERDON, 1870—Assam Roof Turtle

Pangshura tecta (GRAY, 1831)—Indian Roof Turtle

Pangshura tentoria (GRAY, 1834)—Indian Tent Turtle

            Pangshura tentoria tentoria (GRAY, 1834)—Indian Tent Turtle

            Pangshura tentoria circumdata (MERTENS, 1969)—Pink-ringed Tent Turtle

            Pangshura tentoria flaviventer (GUENTHER, 1864)—Yellow-bellied Tent Turtle



Rhinoclemmys FITZINGER, 1835—Neotropical Wood Turtles


Rhinoclemmys annulata (GRAY, 1860)—Brown Wood Turtle

Rhinoclemmys areolata (DUMERIL & BIBRON in DUMERIL & DUMERIL, 1851)—Furrowed Wood Turtle

Rhinoclemmys funerea (COPE, 1875)—Black Wood Turtle

Rhinoclemmys nasuta (BOULENGER, 1902)—Large-nosed Wood Turtle

Rhinoclemmys pulcherrima (GRAY, 1856)—Painted Wood Turtle

            Rhinoclemmys pulcherrima pulcherrima (GRAY, 1855)—Mexican Red Wood Turtle

            Rhinoclemmys pulcherrima incisa (BOCOURT, 1868)—Incised Wood Turtle

            Rhinoclemmys pulcherrima manni (DUNN, 1930)—Central American Wood Turtle

            Rhinoclemmys pulcherrima rogerbarbouri (ERNST, 1978)—Western Mexican Wood Turtle

Rhinoclemmys punctularia (DAUDIN, 1801)58—Spot-legged Wood Turtle

            Rhinoclemmys punctularia punctularia (DAUDIN, 1801)—Eastern Spot-legged Wood Turtle

            Rhinoclemmys punctularia diademata (MERTENS, 1954)—Maracaibo Wood Turtle

            Rhinoclemmys punctularia flammigera PAOLILLO, 1985—Orinoco Spot-legged Wood Turtle

            Rhinoclemmys punctularia melanosterna (GRAY, 1861)—Colombian Wood Turtle

Rhinoclemmys rubida (COPE, 1870)—Mexican Spotted Wood Turtle

            Rhinoclemmys rubida rubida (COPE, 1870)—Oaxaca Wood Turtle

            Rhinoclemmys rubida perixantha (MOSIMANN & RABB, 1953)—Colima Wood Turtle



Sacalia GRAY, 1870—Asian Eyed Turtles


Sacalia bealei (GRAY, 1831)—Beale's Eyed Turtle

Sacalia quadriocellata (SIEBENROCK, 1903)59—Four-eyed Turtle

            Sacalia quadriocellata quadriocellata (SIEBENROCK, 1903)—Four-eyed Turtle

            Sacalia quadriocellata insulensis (ADLER, 1962)—Hainan Four-eyed Turtle



Siebenrockiella LINDHOLM, 1929—Black Marsh Turtles


Siebenrockiella crassicollis (GRAY, 1831)—Black Marsh Turtle

Siebenrockiella leytensis (TAYLOR, 1920)—Philippine Pond Turtle




—Cochin Forest Cane Turtles


Vijayachelys silvatica (HENDERSON, 1912)—Cochin Forest Cane Turtle




Family Kinosternidae—American Mud and Musk Turtles



Claudius COPE, 1865—Narrow-bridged Musk Turtles


Claudius angustatus COPE, 1865—Narrow-bridged Musk Turtle



Kinosternon SPIX, 1824—American Mud Turtles


Kinosternon acutum GRAY, 1831—Tabasco Mud Turtle

Kinosternon alamosae BERRY & LEGLER, 1980—Alamos Mud Turtle

Kinosternon angustipons LEGLER, 1965—Narrow-bridged Mud Turtle

Kinosternon baurii GARMAN, 1891—Striped Mud Turtle

Kinosternon creaseri HARTWEG, 1934—Creaser's Mud Turtle

Kinosternon dunni SCHMIDT, 1947—Dunn's Mud Turtle

Kinosternon flavescens (AGASSIZ, 1857)60—Yellow Mud Turtle

            Kinosternon flavescens flavescens (AGASSIZ, 1857)—Yellow Mud Turtle

            Kinosternon flavescens arizonense GILMORE, 1922—Arizona Mud Turtle

            Kinosternon flavescens durangoense IVERSON, 1979—Durango Mud Turtle

Kinosternon herrerai STEJNEGER, 1925—Herrera's Mud Turtle

Kinosternon hirtipes WAGLER, 183061—Rough-footed Mud Turtle

            Kinosternon hirtipes hirtipes WAGLER, 1830—Valley of Mexico Mud Turtle

            Kinosternon hirtipes chapalense IVERSON, 1981—Lake Chapala Mud Turtle

            Kinosternon hirtipes magdalense IVERSON, 1981—San Juanico Mud Turtle

            Kinosternon hirtipes murrayi GLASS & HARTWEG, 1951—Mexican Plateau Mud Turtle

            Kinosternon hirtipes tarascense IVERSON, 1981—Pátzcuaro Mud Turtle

Kinosternon integrum LECONTE, 185462—Mexican Mud Turtle

            Kinosternon integrum integrum LECONTE, 1854—Mexican Mud Turtle

            Kinosternon integrum chimalhuaca BERRY, SEIDEL & IVERSON, 1997—Jalisco Mud Turtle

            Kinosternon integrum oaxacae BERRY & IVERSON, 1980—Oaxaca Mud Turtle

Kinosternon leucostomum DUMERIL & BIBRON in DUMERIL & DUMERIL, 1851—White-lipped Mud Turtle

            Kinosternon leucostomum leucostomum DUMERIL & BIBRON in DUMERIL & DUMERIL,1851—Northern White-lipped Mud Turtle

            Kinosternon leucostomum postinguinale COPE, 1887—Southern White-lipped Mud Turtle

Kinosternon scorpioides (LINNAEUS, 1766)—Scorpion Mud Turtle

            Kinosternon scorpioides scorpioides (LINNAEUS, 1766)—Scorpion Mud Turtle

            Kinosternon scorpioides abaxillare BAUR, 1925—Central Chiapas Mud Turtle

            Kinosternon scorpioides albogulare (DUMERIL & BOCOURT, 1870)—White-throated Mud Turtle

            Kinosternon scorpioides cruentatum (DUMERIL & BIBRON in DUMERIL & DUMERIL, 1851)—Red-cheeked Mud Turtle

Kinosternon sonoriense LECONTE, 1854—Sonora Mud Turtle

            Kinosternon sonoriense sonoriense LECONTE, 1854—Sonora Mud Turtle

            Kinosternon sonoriense longifemorale IVERSON, 1981—Sonoyta Mud Turtle

Kinosternon subrubrum (LACEPEDE, 1788)—Eastern Mud Turtle

            Kinosternon subrubrum subrubrum (LACEPEDE, 1788)—Eastern Mud Turtle

            Kinosternon subrubrum hippocrepis GRAY, 1856—Mississippi Mud Turtle

            Kinosternon subrubrum steindachneri SIEBENROCK, 1906—Florida Mud Turtle



Staurotypus WAGLER, 1830—Giant Musk Turtles


Staurotypus salvinii GRAY, 1864—Pacific Coast Giant Musk Turtle

Staurotypus triporcatus (WIEGMANN, 1828)—Mexican Giant Musk Turtle



Sternotherus GRAY, 1825—Musk Turtles


Sternotherus carinatus (GRAY, 1856)—Razor-backed Musk Turtle

Sternotherus minor (AGASSIZ, 1857)63—Loggerhead Musk Turtle

            Sternotherus minor minor (AGASSIZ, 1857)—Loggerhead Musk Turtle

            Sternotherus minor depressus TINKLE & WEBB, 1955—Flattened Musk Turtle

            Sternotherus minor peltifer SMITH & GLASS, 1947—Stripe-necked Musk Turtle

Sternotherus odoratus (LATREILLE, 1801)—Common Musk Turtle




Family Platysternidae—Big-headed Turtles



Platysternon GRAY, 1831—Big-headed Turtles


Platysternon megacephalum GRAY, 183164—Big-headed Turtle




Family Testudinidae—Tortoises



Astrochelys GRAY, 1873—Madagascar Tortoises


Astrochelys radiata (SHAW, 1802)—Radiated Tortoise

Astrochelys yniphora (VAILLANT, 1885)—Angonoka Tortoise



Chelonoidis FITZINGER, 1835—South American Tortoises


Chelonoidis carbonaria (SPIX, 1824)—Red-footed Tortoise

Chelonoidis chilensis (GRAY, 1870)—Chaco Tortoise

Chelonoidis denticulata (LINNAEUS, 1766)—Yellow-footed Tortoise

Chelonoidis nigra (QUOY & GAIMARD, 1824)65—Galápagos Tortoise

            Chelonoidis nigra becki (ROTHSCHILD, 1901)—Volcán Wolf Tortoise

            Chelonoidis nigra chathamensis (VAN DENBURGH, 1907)—Chatham Island Tortoise

            Chelonoidis nigra darwinii (VAN DENBURGH, 1907)—James Island Tortoise

            Chelonoidis nigra duncanensis (GARMAN, 1917)—Duncan Island Tortoise

            Chelonoidis nigra hoodensis (VAN DENBURGH, 1907)—Hood Island Tortoise

            Chelonoidis nigra porteri (ROTHSCHILD, 1903)—Indefatigable Island Tortoise

            Chelonoidis nigra vicina (GUENTHER, 1875)—Iguana Cove Tortoise



Chersina GRAY, 1831—Bowsprit Tortoises


Chersina angulata (SCHWEIGGER, 1812)—Bowsprit Tortoise



Dipsochelys BOUR, 198266—Indian Ocean Tortoises


Dipsochelys dussumieri (GRAY, 1831)—Aldabra Tortoise



Geochelone FITZINGER, 183567—Star Tortoises


Geochelone elegans (SCHOEPFF, 1794)68—Star Tortoise

            Geochelone elegans elegans (SCHOEPFF, 1794)—Indian Star Tortoise

            Geochelone elegans platynota (BLYTH, 1863)—Burmese Star Tortoise

Geochelone sulcata (MILLER, 1779)—African Spurred Tortoise



Gopherus RAFINESQUE, 183269—Gopher Tortoises


Gopherus agassizii (COOPER, 1863)—Desert Tortoise

            Gopherus agassizii agassizii (COOPER, 1863)—Agassiz's Desert Tortoise

            Gopherus agassizii morafkai (MURPHY, BERRY, EDWARDS, LEVITON, LATHROP & RIEDLE, 2011)—Morafka's Desert Tortoise

Gopherus berlandieri (AGASSIZ, 1857)—Berlandier's Tortoise

Gopherus flavomarginatus LEGLER, 1959—Bolson Tortoise

Gopherus polyphemus (DAUDIN, 1801)—Gopher Tortoise



Homopus DUMERIL & BIBRON, 1834—Cape Tortoises


Homopus areolatus (THUNBERG, 1787)—Beaked Cape Tortoise

Homopus boulengeri DUERDEN, 1906—Boulenger's Cape Tortoise

Homopus femoralis BOULENGER, 1888—Karoo Cape Tortoise

Homopus signatus (GMELIN, 1789)70—Speckled Cape Tortoise

Homopus solus BRANCH, 2007—Namibian Cape Tortoise



Indotestudo LINDHOLM, 1929—Asian Tortoises


Indotestudo elongata (BLYTH, 1853)71—Elongated Tortoise

            Indotestudo elongata elongata (BLYTH, 1853)—Elongated Tortoise

            Indotestudo elongata travancorica (BOULENGER, 1907)—Travancore Tortoise

Indotestudo forstenii (SCHLEGEL & MUELLER, 1845)—East Indian Tortoise



Kinixys BELL, 182772—Hingeback Tortoises


Kinixys belliana GRAY, 1831—Bell's Hingeback Tortoise

            Kinixys belliana belliana GRAY, 1831—Eastern Bell's Hingeback Tortoise

            Kinixys belliana domerguei (VUILLEMIN, 1972)—Madagascar Hingeback Tortoise

            Kinixys belliana nogueyi (LATASTE, 1886)—Western Hingeback Tortoise

            Kinixys belliana zombensis HEWITT, 1931—Star Hingeback Tortoise

Kinixys erosa (SCHWEIGGER, 1812)—Forest Hingeback Tortoise

Kinixys homeana BELL, 1827—Home's Hingeback Tortoise

Kinixys spekii GRAY, 186373—Speke's Hingeback Tortoise

            Kinixys spekii spekii GRAY, 1863—Speke's Hingeback Tortoise

            Kinixys spekii lobatsiana POWER, 1927—Lobatse Hingeback Tortoise

            Kinixys spekii natalensis HEWITT, 1935—Natal Hingeback Tortoise



Malacochersus LINDHOLM, 1929—Pancake Tortoises


Malacochersus tornieri (SIEBENROCK, 1903)—Pancake Tortoise



Manouria GRAY, 1852—Indochinese Tortoises


Manouria emys (SCHLEGEL & MUELLER, 1841)—Asian Giant Tortoise

            Manouria emys emys (SCHLEGEL & MUELLER, 1841)—Brown Tortoise

            Manouria emys phayrei (BLYTH, 1853)—Burmese Black Tortoise

Manouria impressa (GUENTHER, 1882)—Impressed Tortoise



Psammobates FITZINGER, 1835—South African Star Tortoises


Psammobates geometricus (LINNAEUS, 1758)—Geometric Tortoise

Psammobates oculifer (KUHL, 1820)—Serrated Tortoise

Psammobates tentorius (BELL, 1828)—Tent Tortoise

            Psammobates tentorius tentorius (BELL, 1828)—Common Tent Tortoise

            Psammobates tentorius trimeni (BOULENGER, 1886)—Western Tent Tortoise

            Psammobates tentorius verroxii (SMITH, 1839)—Northern Tent Tortoise



Pyxis BELL, 1827—Spider Tortoises


Pyxis arachnoides BELL, 1827—Spider Tortoise

            Pyxis arachnoides arachnoides BELL, 1827—Common Spider Tortoise

            Pyxis arachnoides brygooi (VUILLEMIN & DOMERGUE, 1972)—Northern Spider Tortoise

            Pyxis arachnoides oblonga GRAY, 1869—Southern Spider Tortoise

Pyxis planicauda (GRANDIDIER, 1867)—Flat-shelled Spider Tortoise



Stigmochelys GRAY, 1873—Leopard Tortoises


Stigmochelys pardalis (BELL, 1828)74—Leopard Tortoise



Testudo LINNAEUS, 175875—Palearctic Tortoises


Testudo graeca LINNAEUS, 175876—Spur-thighed Tortoise

            Testudo graeca graeca LINNAEUS, 175877—Algerian Tortoise

            Testudo graeca armeniaca CHKHIKVADZE & BAKRADZE, 1991—Araxes Spur-thighed Tortoise

            Testudo graeca buxtoni BOULENGER, 1920—Caspian Tortoise

            Testudo graeca ibera PALLAS, 1814—Spur-thighed Tortoise

            Testudo graeca terrestris FORSSKAL, 1775—Mesopotamian Tortoise

            Testudo graeca zarudnyi NIKOLSKIY, 1896—Iranian Spur-thighed Tortoise

Testudo hermanni GMELIN, 1789—Hermann's Tortoise

            Testudo hermanni hermanni GMELIN, 1789—Greek Tortoise

            Testudo hermanni boettgeri MOJSISOVICS, 1889—Greek Tortoise

Testudo horsfieldii (GRAY, 1844)78—Central Asian Tortoise

            Testudo horsfieldii horsfieldii (GRAY, 1844)—Central Asian Tortoise

            Testudo horsfieldii kazachstanica CHKHIKVADZE, 1988—Kazachstan Tortoise

Testudo kleinmanni LORTET, 1883—Egyptian Tortoise

Testudo marginata SCHOEPFF, 1793—Marginated Tortoise

            Testudo marginata marginata SCHOEPFF, 1793—Marginated Tortoise

            Testudo marginata weissingeri BOUR, 199579—Southern Peloponnese Tortoise




Family Trionychidae—Softshell Turtles



Amyda GEOFFROY SAINT-HILAIRE, 1809—Asiatic Softshell Turtles


Amyda cartilaginea (BODDAERT, 1770)—Asiatic Softshell Turtle



Apalone RAFINESQUE, 1832—North American Softshell Turtles


Apalone ferox (SCHNEIDER, 1783)—Florida Softshell Turtle

Apalone mutica (LESUEUR, 1827)—Smooth Softshell Turtle

            Apalone mutica mutica (LESUEUR, 1827)—Smooth Softshell Turtle

            Apalone mutica calvata (WEBB, 1959)—Gulf Coast Smooth Softshell Turtle

Apalone spinifera (LESUEUR, 1827)—Spiny Softshell Turtle

            Apalone spinifera spinifera (LESUEUR, 1827)—Eastern Spiny Softshell Turtle

            Apalone spinifera aspera (AGASSIZ, 1857)—Gulf Coast Spiny Softshell Turtle

            Apalone spinifera atra (WEBB & LEGLER, 1960)—Black Spiny Softshell Turtle

            Apalone spinifera emoryi (AGASSIZ, 1857)—Texas Spiny Softshell Turtle

            Apalone spinifera guadalupensis (WEBB, 1962)—Guadalupe Spiny Softshell Turtle

            Apalone spinifera pallida (WEBB, 1962)—Pallid Spiny Softshell Turtle



Chitra GRAY, 1844—Narrow-headed Softshell Turtles


Chitra chitra NUTAPHAND, 1986—Asian Narrow-headed Softshell Turtle

            Chitra chitra chitra NUTAPHAND, 1986—Thai Narrow-headed Softshell Turtle

            Chitra chitra javanensis MCCORD & PRITCHARD, 2003—Java Narrow-headed Softshell Turtle

            Chitra chitra vandijki MCCORD & PRITCHARD, 200380—Myanmar Narrow-headed Softshell Turtle

Chitra indica (GRAY, 1831)—Indian Narrow-headed Softshell Turtle



Cyclanorbis GRAY, 1852—Sub-Saharan Flapshell Turtles


Cyclanorbis elegans (GRAY, 1869)—Nubian Flapshell Turtle

Cyclanorbis senegalensis (DUMERIL & BIBRON, 1835)—Senegal Flapshell Turtle



Cycloderma PETERS, 1854—Central African Flapshell Turtles


Cycloderma aubryi (DUMERIL, 1856)—Aubry's Flapshell Turtle

Cycloderma frenatum PETERS, 1854—Zambezi Flapshell Turtle



Dogania GRAY, 1844—Malayan Softshell Turtles


Dogania subplana (GEOFFROY SAINT-HILAIRE,1809)—Malayan Softshell Turtle



Lissemys SMITH, 193181—Indian Flapshell Turtles


Lissemys ceylonensis (GRAY 1856)—Sri Lankan Flapshell Turtle

Lissemys punctata (LACEPEDE, 1788)—Indian Flapshell Turtle

            Lissemys punctata punctata (LACEPEDE, 1788)—South Indian Flapshell Turtle

            Lissemys punctata andersoni WEBB, 1980—North Indian Flapshell Turtle

            Lissemys punctata vittata (PETERS 1854)—West Indian Flapshell Turtle

Lissemys scutata (PETERS, 1868)—Burmese Flapshell Turtle



Nilssonia GRAY, 1872—Asian Softshell Turtles


Nilssonia formosa (GRAY, 1869)—Burmese Peacock Softshell Turtle

Nilssonia gangetica (CUVIER, 1824)—Indian Softshell Turtle

Nilssonia hurum (GRAY, 1831)82—Peacock Softshell Turtle

            Nilssonia hurum hurum (GRAY, 1831)—Indian Peacock Softshell Turtle

            Nilssonia hurum nigricans (ANDERSON, 1875)—Black Peacock Softshell Turtle

Nilssonia leithii (GRAY, 1872)—Leith's Softshell Turtle



Pelochelys GRAY, 1864—Asian Giant Softshell Turtles


Pelochelys bibroni (OWEN, 1853)—South New Guinea Giant Softshell Turtle

Pelochelys cantorii GRAY, 1864—Asian Giant Softshell Turtle

Pelochelys signifera WEBB, 2003—North New Guinea Giant Softshell Turtle



Pelodiscus FITZINGER, 183583—East Asian Softshell Turtles


Pelodiscus sinensis (WIEGMANN, 1835)84—Chinese Softshell Turtle

            Pelodiscus sinensis sinensis (WIEGMANN, 1835)—Chinese Softshell Turtle

            Pelodiscus sinensis axenaria (ZHOU, ZHANG & FANG, 1991)—Hunan Softshell Turtle

            Pelodiscus sinensis maackii (BRANDT, 1858)—Amur Softshell Turtle

            Pelodiscus sinensis parviformis TANG, 1997—Guangxi Softshell Turtle

Pelodiscus steindachneri (SIEBENROCK, 1906)—Wattle-necked Softshell Turtle



Rafetus GRAY, 1864—Bicallosite Softshell Turtles


Rafetus euphraticus (DAUDIN, 1801)—Euphrates Softshell Turtle

Rafetus swinhoei (GRAY, 1873)—Swinhoe's Softshell Turtle



Trionyx GEOFFROY SAINT-HILAIRE, 1809—African Softshell Turtles


Trionyx triunguis (FORSSKAL, 1775)—Nile Softshell Turtle










The following comments show all the differences compared to the checklist of ARTNER (2008) and the most recent checklist of the TURTLE TAXONOMY WORKING GROUP (2012).


(1) Chelodina novaeguineae: morphological similarities between C. novaeguineae, C. canni, C. gunaleni, and C. reimanni are striking, so they are all considered subspecies of C. novaeguineae here. I have observed intergrades between C. novaeguineae and C. reimanni near the border between Papua New Guinea and Papua Province, Indonesia (ARTNER 1995).

(2) Elseya: compared to the closely related and quite homogeneous genus Emydura, the genus Elseya is somewhat more heterogeneous. This fact encouraged some authors to split it into Elseya (the "Elseya dentata complex" sensu GEORGES & THOMSON 2006) and Wollumbinia (WELLS 2007) or Myuchelys (THOMSON & GEORGES 2009), the "Elseya latisternum complex" sensu GEORGES & THOMSON (2006). GEORGES & THOMSON (2010) present their opinion that the work of WELLS (2007) didn't meet the IUZN criteria and thus the name Wollumbinia should not be recognized. Their arguments are being followed by the TURTLE TAXONOMY WORKING GROUP (2011, 2012). I think this to be premature as no independent authorities have ruled on that subject yet and it would seem fair to me that other scientists rather than the ones proposing a different name for the very same genus should check this out. GEORGES & THOMSON (2010), however, do not recognize Macrochelodina and Macrodiremys as valid genera but consider them to be subgenera at best. I think that the morphological differences between these two genera and the genus Chelodina are far more obvious than the ones between Elseya and Wollumbinia / Myuchelys. So I tentatively recognize Wollumbinia / Myuchelys until further data especially about the New Guinean forms (at present all included in the polymorph Wollumbinia / Myuchelys novaeguineae) will be available. Contrary to the TURTLE TAXONOMY WORKING GROUP (2011) I agree with GEORGES & THOMSON (2010) in placing Elseya novaeguineae into the new genus Wollumbinia / Myuchelys as this species has much more in common with the "Elseya latisternum complex" sensu GEORGES & THOMSON (2006).

(3) Elseya dentata: GEORGES & THOMSON (2010) have clearly shown that E. dentata is different from E. albagula, E. irwini, and E. lavarackorum, so that it should be considered a species of its own.

(4) Elseya dentata jukesi: GEORGES & THOMSON (2010) present their opinion that the work of WELLS (2007) did not meet the IUZN criteria and thus his proposed names shouldn't be recognized. Their arguments are being followed by the TURTLE TAXONOMY WORKING GROUP (2011, 2012). I think this to be premature, as no independent authorities have ruled on that subject yet. Also see comments in (2).

(5) Elseya lavarackorum: in their description of Elseya albagula THOMSON et al. (2006) clearly state that "the affinities of E. albagula lie with a well-defined clade within the E. dentata subgeneric group comprising E. irwini, E. lavarackorum,... " and that "we consider the closest living relative to be an undescribed taxon from the Johnstone Rivers region near Cairns, but among described taxa, it is E. lavarackorum (White and Archer 1994) from the Nicholson Drainage, Queensland". So if E. dentata forms a clade of its own and all the others form another "well-defined" clade, why not consider them two separate species? Consequently I am recognizing E. albagula and E. irwini as subspecies of E. lavarackorum.

(6) Emydura macquarii macquarii: includes E. m. binjing CANN, 1998, E. m. dharra CANN, 1998, E. m. dharuk CANN, 1998, E. m. gunabarra CANN, 1998, and E. m. signata AHL, 1932 according to GEORGES & THOMSON (2010). ARTNER (2008) had already argued for inclusion of all of the above except E. m. signata into one subspecies based on morphological grounds. GEORGES & THOMSON (2010) further cite GEORGES & ADAMS (1996) stating that the two forms from Cooper Creek (later described as E. m. emmotti CANN, McCord & Joseph-Ouni in McCord, Cann & Joseph-Ouni 2003) and from Fraser Island (later described as E. m. nigra McCord, CANN & Joseph-Ouni 2003) are  "not genetically distinct from other populations of Emydura macquarii based on allozyme electrophoresis". So I do not recognize these two forms either, leaving E. m. macquarii and E. m. kreffti to be the only remaining subspecies of Emyura macquarii.

(7) Emydura subglobosa: includes E. tanybaraga CANN, 1997 due to evident morphological similarities as confirmed by GEORGES & THOMSON (2010), who state that "morphological characters to diagnose E. tanybaraga from E. victoriae are substantial (refer to the key above), but less so between E. tanybaraga and E. subglobosa worrelli. The latter two species are very difficult to distinguish consistently in the field, and further examination of their status using a combination of DNA sequence data and morphological data is warranted." In their paper, however, E. tanybaraga is kept as a species. The TURTLE TAXONOMY WORKING GROUP (2011, 2012) doesn't challange this placement either. I do so, however, and keep the form as a subspecies of E. subglobosa like I did in the previous edition (ARTNER 2008).

(8) Emydura victoriae: includes all forms previously attributed to E. australis, which is a junior synonym of E. macquarii according to COGGER et al. (1983) fide GEORGES & THOMSON (2010). I am following their arguments here.

(9) Macrochelodina: morphological differences between Chelodina FITZINGER, 1826 and Macrochelodina WELLS & WELLINGTON, 1985 are enormous, and thus they are best considered as different genera (ARTNER 2008). The arguments of GEORGES & THOMSON (2010) for Macrochelodina being merely a subgenus of Chelodina are partly convincing (i.e. that "Chelodina" (i.e. Macrochelodina) rugosa and Chelodina "canni" (i.e Chelodina novaeguineae canni) "...undergo widespread and common natural hybridization in Australia to yield viable and fertile offspring in the wild") but the morphological differences are simply too obvious to be ignored.

(10) Macrochelodina rugosa: I agree with GEORGES & THOMSON (2010) to include M. kuchlingi (CANN, 1997) and M. siebenrocki (WERNER, 1901) due to the fact that the former was described from a single specimen of uncertain origin with a long history of captivity and the latter is morphologically and genetically indistinguishable from Cape York M. rugosa. I also include M. burrungandjii (THOMSON, KENNETT & GEORGES, 2000) and M. walloyarrina MCCORD & JOSEPH-OUNI, 2007 as subspecies. Besides strong morpholgical similarities I quote GEORGES et al. (2002): "C. burrungandjii hybridizes with C. rugosa yielding fertile offspring where they come in contact... ".

(11) Macrodiremys: MCCORD & JOSEPH-OUNI (2007) placed this species in a new genus, which is justified due to its different morphology compared to Chelodina FITZINGER, 1826 and Macrochelodina WELLS & WELLINGTON, 1985.

(12) Macrodiremys colliei: THOMSON (2000) showed that the holotype of Chelodina oblonga GRAY, 1841 is in fact a specimen belonging to the species Macrochelodina rugosa. So this species name is not available for the turtle form from Perth, Western Australia anymore. The next available name is Chelodina colliei GRAY, 1856 (THOMSON 2000) thus being used here.

(13) Mesoclemmys dahli: includes Phrynops (Batrachemys) zuliae PRITCHARD & TREBBAU, 1984 as second subspecies due to similar morphology.

(14) Mesoclemmys gibba: includes Mesoclemmys perplexa BOUR & ZAHER, 2005 due to close morphology.

(15) Mesoclemmys raniceps: includes Batrachemys heliostemma MCCORD, JOSEPH-OUNI & LAMAR, 2001 as second subspecies due to striking similarities in morphology when adult.

(16) Phrynops geoffroanus: the formerly recognized species Phrynops tuberosus (PETERS, 1870) is very similar to P. geoffroanus (SCHWEIGGER, 1812). Therefore both forms are considered subspecies of the same species here.

(17) Wollumbinia or Myuchelys: see comments under (2).

(18) Wollumbinia / Myuchelys latisternum: morphological similarities between W. / M. latisternum and W. / M. bellii, are striking. This is confirmed by GEORGES & THOMSON (2010). Thus they are both considered subspecies of W. / M. latisternum here.

(19) Wollumbinia / Myuchelys purvisi: morphological similarities between W. / M. purvisi and W. / M. georgesi are striking. This is confirmed by GEORGES & THOMSON (2010). Thus they are both considered subspecies of W. / M. purvisi here.

(20) Wollumbinia / Myuchelys novaeguineae: there are a number of still undescribed taxa of the genus Wollumbinia / Myuchelys on New Guinea (W.P. MCCORD, A.G.J. RHODIN and D. WELLS pers. comm. to the author and own observations in 1995). As for now they are all included within W. / M. novaeguineae awaiting further study.

(21) Pelusios castaneus: Pelusios chapini LAURENT, 1965 is morphologically very similar to P. castaneus and thus considered a subspecies of the latter. Although only relying on genetics and therefore taxonomically doubtful, it is noted that the work of FRITZ et al. (2011a) also found only little differences between the two.

(22) Pelusios castanoides, Pelusios subniger: as clearly shown by the works of SILVA et al. (2010) and FRITZ et al. (2011a) Pelusios castanoides intergularis BOUR, 1983 and Pelusios subniger parietalis BOUR, 1983 from the Seychelles are not reliably distinguishable genetically from the nominate subspecies. Both papers, however, completely ignore the morphological differences described by BOUR. Until further evidence I still consider both subspecies valid.

(23) Podocnemis cayennensis: as suggested by MITTERMEIER & WILSON (1974) and WERMUTH & MERTENS (1977), and finally clearly shown by BOUR (in PRITCHARD & TREBBAU 1984), the correct name for this species is P. cayennensis (SCHWEIGGER, 1812) and not P. unifilis TROSCHEL, 1848. So unless a ruling is available by the ICZN, P. cayennensis is the only valid name for the species.

(24) Eretmochelys imbricata: the TURTLE TAXONOMY WORKING GROUP (2011) agreed on not further recognizing E. i. bissa. They argued that "no genetic studies or reviews (OKAYAMA et al. 1999; BOWEN & KARL 2007) have argued for continued recognition of the subspecies bissa. " I follow their arguments here.

(25) Chelydra serpentina osceola: SHAFFER et al. (2008) implicate that genetically speaking C. s. osceola does not show enough differences to deserve subspecies status and thus they recommend to regard C. serpentina as monotypic. ARESCO & GUNZBURGER (2007), however, show that the two subspecies intergrade in northwestern Florida thus proving that the subspecies status is clearly warranted for C. s. osceola. Their results are just another proof that genetics alone are at present useless for taxonomic changes.

(26) Chrysemys picta dorsalis: this is probably the very best example for how the phylogenetic species concept interpreted nowadays by "modern" taxonomists based almost exclusively on genetics, is wrong. Despite intergradiation between the four subspecies of Chrysemys picta having been repeatedly documented in the wild, the last of many works being that of ERNST et al. (2006), the TURTLE TAXONOMY WORKING GROUP's latest editions (2011, 2012), after refusing to do so earlier, now recognize Chrysemys dorsalis as a separate species merely because of genetic results in the work of STARKEY et al. (2003). I strongly agree with ERNST & LOVICH (2009) in not recognizing this and keeping C. p. dorsalis as a clearly defined subspecies of C. picta.

(27) Emys: this is a tough decision, but due to morphological similarities I think that Emys DUMERIL, 1806 also includes Emydoidea GRAY, 1870 and Actinemys AGASSIZ, 1857. This view is supported by the mainly genetic works of BICKHAM et al. (1996), FELDMAN & PARHAM (2002), and SPINKS & SHAFFER (2009). WIENS et al. (2010) and FRITZ et al. (2011b) argue in favor of regarding all three genera as valid. The TURTLE TAXONOMY WORKING GROUP (2012) is undecided. I do not recognize Emys trinacris FRITZ, FATTIZZO, GUICKING, TRIPEPI, PENNISI, LENK, JOGER & WINK, 2005 as it is a so-called "cryptic species" with its description based exclusively on genetic data. The authors themselves state that it is virtually impossible to distinguish this form from its neighboring ones without genetic testing.

(28) Emys orbicularis: this species is probably still oversplit. Like in the previously recognized Emydura macquarii subspecies, no reliable key for the determination of subspecies of Emys orbicularis is available, another indication of weak morphological differences. I recognize only the "subspecies groups" given by FRITZ (2001) and E. o. persica EICHWALD 1831.

(29) Emys orbicularis galloitalica: includes E. o. ingauna JESU, PIOMBO, SALVIDIO, LAMAGNI, ORTALE & GENTA, 2004.

(30) Emys orbicularis luteofusca: FRITZ et al. (2009) synonimized E. o. luteofusca under E. o. orbicularis due to lack of sufficient genetic differences (based solely on mitochondrial DNA differentiation). I am not sure if FRITZ'  concept of E. o. luteofusca is correct. In his original description (FRITZ 1989) he describes the subspecies to be bigger and darker than the nominate one. When shown E. orbicularis from the Konya area imported by M. Reimann in 1997, turtles being really small and extremely yellowish (especially the males), FRITZ thought them to be from Iran and never believed us that they originated from the type area of E. o. luteofusca. So there is some doubt, if the locality data of FRITZ' type series are correct. The specimens of E. orbicularis collected at the type locality of E. o. luteofusca (sensu FRITZ) clearly are a very different form, probably the most different of all Emys orbicularis. 

(31) Graptemys: research by STEPHENS & WIENS (2003) proved that Graptemys is heavily oversplit. Without rigorously calling for taxonomic changes due to their work, these authors with their analyses of combined morphological and molecular data nevertheless proved that taxonomic changes are indeed inevitable. FREEDBERG & MYERS (2012) proved hybridization between G. geographica, G. pseudogeographica, and G. ouachitensis in parts of their range.

(32) Graptemys oculifera: STEPHENS & WIENS (2003) clearly showed that G. flavimaculata CAGLE, 1954 differs only slightly from G. oculifera (BAUR, 1890) in a combined morphological and molecular analysis, meaning that there is no justification for species status. They are subspecies that readily intergrade in captivity (R. ENGERTH pers. comm. to the author). Both forms are also very close to G. nigrinoda CAGLE 1954, maybe also resulting in being conspecific.

(33) Graptemys pseudogeographica: I do not agree with the interpretations of VOGT (1993). As intergrades between G. pseudogeographica and G. ouachitensis have been reported from the wild (VOGT 1980, C. LECHOWICZ in litt.) in a number of US states and intergradation also occurs between G. o. ouachitensis and G. o. sabinensis in Louisiana (C. LECHOWICZ in litt.), all these are subspecies of G. pseudogeographica. I also include G. versa here due to its morphological similarities with G. p. pseudogeographica and G. p. sabinensis. Its close relationship with the latter is also confirmed by the combined morphological and molecular data provided by STEPHENS & WIENS (2003).

(34) Graptemys pulchra: due to convincing morphological similarities G. ernsti LOVICH & MCCOY, 1992 and G. gibbonsi LOVICH & MCCOY, 1992 are here considered subspecies of G. pulchra BAUR, 1893. Quite recently, hybridization between G. barbouri CARR & MARCHAND, 1942 and G. ernsti LOVICH & MCCOY, 1992 was observed in southern Alabama (LECHOWICZ 2008), proving G. barbouri to be merely a subspecies of G. pulchra, too. By splitting "G. gibbonsi" even further ENNEN, LOVICH, KREISER, SELMAN & QUALLS (2010) created one more new "species" they named Graptemys pearlensis. It is a weakly defined subspecies at best and as such included provisionally until further research will be done.

(35) Malaclemys terrapin: it is very difficult to tell the seven subspecies apart morphologically (pers. observations). LAMB & AVISE (1992) found only significant differences between the Atlantic Coast complex (comprising three subspecies) and the Gulf Coast complex (with four subspecies) testing mitochondrial DNA variability. The presently recognized subspecies are more clinal variants than subspecies according to these authors. I agree with that.

(36) Malaclemys terrapin terrapin: includes M. t. centrata (LATREILLE, 1801) and M. t. tequesta SCHWARTZ, 1955.

(37) Malaclemys terrapin pileata: includes M. t. littoralis HAY, 1904, M. t. macrospilota HAY, 1904, and M. t. rhizophorarum FOWLER, 1906.

(38) Pseudemys: a lot of reported intergradation occurs in the wild. Multiple reports exist on hybridyzation between P. concinna (LECONTE, 1830) and P. floridana (LECONTE, 1830) (SEIDEL & PALMER 1991, ERNST et al. 1994), between P. floridana and P. peninsularis CARR, 1938 (THOMAS & JANSEN 2006) and even between P. floridana and P. rubriventris (LECONTE 1830) (SEIDEL & PALMER 1991), the latter being an exceptional event, however.

(39) Pseudemys concinna: includes P. floridana (LECONTE, 1830), P. peninsularis CARR, 1938, and P. suwanniensis CARR, 1937 as subspecies due to multiple hybridization in the wild (see comments under Pseudemys). Pseudemys gorzugi WARD, 1984 is also a very similar form, but the problem is that the complete range of the quite different P. texana BAUR, 1893 lies between the easternmost occurrence of P. gorzugi and the westernmost occurrence of P. concinna.

(40) Pseudemys rubriventris: close relationship of this species with the other two red-bellied taxa P. alabamensis BAUR, 1893 and P. nelsoni CARR, 1938 is proven by the results of STEPHENS & WIENS (2003). All three are thus conspecific.

(41) Terrapene carolina: BUTLER et al. (2011) in a combined morphological and genetic analysis proved T. c. major invalid and that box turtles traditionally assigned to T. c. major based on phenotype are the result of introgression between eastern extant (predominantly T. c. carolina) and an extinct subspecies, T. c. putnami. STEPHENS & WIENS (2003) suggest to elevate both Mexican subspecies of T. carolina to specific rank. By taking a closer look at their results it becomes evident that T. c. yucatana (BOULENGER, 1895) is strikingly different in terms of morphology whereas T. c. mexicana (GRAY, 1849) is not. The latter subspecies differs a lot genetically though, demanding further analysis.

(42) Trachemys: this is a genus currently developing new species. So it is logical that there is much controversy in naming some forms full species or merely subspecies. The former concepts with only a few species (i. e. WERMUTH & MERTENS [1977] with Trachemys, however, still included in Pseudemys) were underestimating diversity. On the other hand, SEIDEL (2002) definitely oversplits the genus naming no less than 15(!) species with 11 additional subspecies. Based on the works by SEIDEL (1998, 2002), STEPHENS & WIENS (2003), FRITZ et al. (2011) and my own field work in Mexico, the West Indian Islands (ARTNER 2006), Brazil, Uruguay, and Venezuela, I think that the genus Trachemys consists of 8 to 10 species with a number of additional subspecies. The most recent work of FRITZ et al. (2011) once again relied on genetics only, but at least a combination of mitochondrial and nuclear DNA was used. Their nuclear data, as confirmed by the short summary given by the TURTLE TAXONOMY WORKING GROUP (2011), unfortunately were uninformative and thus they relied heavily on their mitochondrial DNA data, a procedure explicitly criticized by WIENS et al. (2010). Still, their results were impressive and put some light on the highly complicated taxonomy of the genus. According to FRITZ et al. (2011) the Pacific Coast taxa grayi, emolli, and the newly described subspecies Trachemys venusta panamensis McCORD, JOSEPH-OUNI, HAGEN & BLANCK 2010 form a distinct clade that warrants recognition at the species level, as Trachemys grayi, with three subspecies and the taxa cataspila, ornata, venusta, callirostris, and chichiriviche form another one under a single species, Trachemys ornata. According to their results, T. adiutrix should be placed at subspecies level into Trachemys dorbigni, a procedure already proposed on morphological grounds by me (ARTNER 2008). Finally, the newly described subspecies Trachemys venusta uhrigi McCORD, JOSEPH-OUNI, HAGEN & BLANCK 2010 should be synonymized within T. v. venusta. Their analysis did not include samples of the other recently decribed taxon Trachemys venusta iversoni McCORD, JOSEPH-OUNI, HAGEN & BLANCK 2010, which is very weakly defined by McCORD ET AL. (2010) and hence not recognized here. Trachemys gaigeae is a species of its own with two subspecies. The positions of the taxa T. hiltoni, T. nebulosa, T. taylori, and T. yaquia weren't addressed by FRITZ et al. (2011). Until further investigations, I consider them to be separate species (T. nebulosa with T. hiltoni included as a subspecies) or subspecies within T. ornata (taylori and yaquia). More work is urgently needed here.

(43) Batagur: PRASCHAG et al. (2009) provided quite detailed morphological data on how to distinguish B. affinis and B. baska. They also described a new subspecies, naming it B. affinis edwardmolli.

(44) Cuora: after many discussions and considerations Cuora yunnanensis (BOULENGER, 1906) is still not included in this list. HE et al. (2007) compared a small fragment of mitochondrial DNA extracted from three live specimens recently purchased on Chinese markets with the data of a single museum specimen used in the study by PARHAM et al. (2006) and claimed them to be very similar in mitochondrial DNA sequence. The three "wild caught" specimens are morphologically quite different from the type series in the BMNH (R. DE BRUIN pers. comm.) and morphologically could very well be hybrids. So until reliable studies and true wild-caught specimens prove me wrong, I still consider this species extinct or to be a (natural?) hybrid. The same might apply to Cuora mccordi and Cuora zhoui, because both species have never been found in the wild since their descriptions (in both cases based on market specimens and / or pet store specimens).

          Unfortunately, the work by SPINKS et al. (2012) raises more questions than it answers and the authors on one hand suggest to accept Cuora mccordi as a valid species while on the other they doubt its status as a valid species. They found some potential gene flow between the three forms of Cuora galbinifrons and despite this still recommend them to be recognized as valid full species. That's the kind of taxonomic work we don't need anymore.  

(45) Cuora amboinensis: it is very difficult to tell the four subspecies apart morphologically (pers. observations). Only the clearly distinguishable Cuora amboinensis amboinensis (DAUDIN, 1801) from Sulawesi and parts of the Philippines and Cuora amboinensis kamaroma RUMMLER & FRITZ, 1991 from Assam, central Thailand and further west are recognized. The populations in between these two areas are more clinal variants than subspecies.

(46) Cuora galbinifrons: STUART & PARHAM (2004) elevated the subspecies Cuora galbinifrons bourreti OBST & REIMANN, 1994 and Cuora galbinifrons picturata LEHR, FRITZ & OBST, 1998 to species rank. These forms, however, are quite similar to the nominate form in external morphology, especially C. g. bourreti. FRITZ et al. (2006) showed that remarkable osteological similarities are present in C. bourreti and C. picturata (however, some—if not most—of their sample material probably consisted of specimens originating from the turtle trade without confirmed locality data). In the same paper FRITZ et al. (2006) argued for conspecifity of C. galbinifrons and C. bourreti due to intergradation on Hainan Island, as previously described by DE BRUIN & ARTNER (1999)—a clear sign for subspecies status.

(47) Cuora pani: ARTNER (2003) first argued for conspecifity of Cuora aurocapitata LUO & ZONG, 1988 and Cuora pani SONG, 1984. Morphological similarities are overwhelming and intergradation obviously occurs according to own observations.

(48) Cyclemys: another "highlight" of modern taxonomy. No less than three revisions have been published during the last two decades, all of them with FRITZ as senior (twice) or junior (once) author (FRITZ et al. 1997, 2008, GUICKING et al. 2002). In their most recent study, FRITZ et al. (2008) oversplit the genus by far. They mainly rely on genetics neglecting even obvious morphological similarities (up to the point of indistinctiveness). Their newly described dark-bellied species Cyclemys enigmatica, Cyclemys fusca, and Cyclemys gemeli are probably only clinal variants of the very variable Cyclemys oldhamii, but as the authors state themselves: "gross morphology of the dark-bellied species is so similar that reliable species determination is only possible when genetic markers are used". So their proposed species concept of Cyclemys is not followed here until enough specimens including samples from zones of possible intergradation will be investigated.

(49) Cyclemys atripons: includes Cyclemys pulchristriata FRITZ, GAULKE & LEHR, 1997. The two forms are morphologically undistinguishable, but FRITZ et al. (2008) claim them to be two separate species.

(50) Cyclemys oldhamii: includes Cyclemys enigmatica FRITZ, GUICKING, AUER, SOMMER, WINK & HUNDSDOERFER, 2008, Cyclemys fusca FRITZ, GUICKING, AUER, SOMMER, WINK & HUNDSDOERFER, 2008, Cyclemys gemeli FRITZ, GUICKING, AUER, SOMMER, WINK & HUNDSDOERFER, 2008, and Cyclemys tcheponensis (BOURRET, 1939). These forms are clinal variations within a very variable species and consequently not recognized.

(51) Malayemys subtrijuga: includes Malayemys macrocephala (GRAY, 1859) as subspecies due to close resemblance in external morphology and unlikeliness of reproductive incompatibility.

(52) Mauremys caspica: includes Mauremys rivulata (VALENCIENNES, 1833) as subspecies. M. rivulata intergrades with M. c. caspica in southeastern Turkey (FRITZ et al. 2008) and obviously also in Jordan and Syria (ARTNER pers. obs.). VAMBERGER et al. (2012) recently proved Mauremys caspica siebenrocki WISCHUF & FRITZ, 1997 and Mauremys caspica ventrimaculata WISCHUF & FRITZ, 1996 invalid.

(53) Mauremys leprosa vanmeerhaeghei: FRITZ et al. (2006) suppressed M. leprosa vanmeerhaeghei as a synonym of M. l. saharica. Both taxa (living in distinct drainages) are easily separated by morphology and thus both kept as subspecies.

(54) Melanochelys trijuga: it seems that there is some clinal variation within this species rather than subspecific differentiation. While M. t. coronata is clearly distinguishable by morphological means, M. t. indopeninsularis shares characteristics with both M. t. trijuga and M. t. edeniana, and is thus considered not a subspecies but rather a part of a clinal variation zone between the latter two subspecies. A comprehensive study of the subspecies of M. trijuga including morphometric and genetic data is needed.

(55) Melanochelys trijuga thermalis: includes M. t. parkeri. This taxon is undistinguishable from M. t. thermalis (own observations).

(56) Morenia: both forms are morphologically similar but not to the extent to synonymize them.

(57) Notochelys: includes Leucocephalon MCCORD, IVERSON, SPINKS & SHAFFER, 2000. This is supported by similar morphology and by the molecular results of SPINKS et al. (2004).

(58) Rhinoclemmys punctularia: includes Rhinoclemmys diademata (MERTENS, 1954) and R. melanosterna (GRAY, 1861). Both forms are very similar to the nominate one in external morphology and readily interbreed with R. p. punctularia in captivity (FRITZ 1995).

(59) Sacalia quadriocellata: as already mentioned by DE BRUIN & ARTNER (1999), Hainan specimens are morphologically distinguishable from mainland ones. In a more recent publication, SHI et al. (2008) have confirmed these results using genetic data. Thus the Hainan form is clearly a subspecies for which the name Sacalia quadriocellata insulensis ADLER, 1962 is available.

(60) Kinosternon flavescens: includes Kinosternon arizonense GILMORE, 1922 and K. durangoense IVERSON, 1979 as subspecies. Both forms have been elevated to species status by SERB et al. (2001). However, this elevation is unwarranted as gross morphology is very similar (pers. observations). I have also bred K. f. flavescens and K. f. arizonense in captivity and the hatchlings are actually indistinguishable from each other (ARTNER unpubl.).

(61) Kinosternon hirtipes: for decades, no specimens of Kinosternon hirtipes megacephalum IVERSON, 1981 have been found in the wild and there are no specimens present in captivity either, so that this subspecies must be considered extinct and is thus not included in here. A genetic survey of all other currently recognized subspecies is in dire need.

(62) Kinosternon integrum: includes Kinosternon chimalhuaca BERRY, SEIDEL & IVERSON, 1996 and Kinosternon oaxacae BERRY & IVERSON, 1980 as subspecies. Both forms are very similar to the nominate one in external morphology and reproductive incompatibility seems unlikely.

(63) Sternotherus minor: includes Sternotherus depressus TINKLE & WEBB, 1955 as a subspecies due to striking similarities in gross morphology and reproductive incompatibility being highly unlikely.

(64) Platysternon megacephalum: own personal observations and published data (DE BRUIN & ARTNER 1999) in Southeast Asia do not support any subspecies recognition. The data presented by ERNST & LAEMMERZAHL (2002) are not reliable in separating different forms of Platysternon megacephalum. Specimens with a color pattern claimed typical for P. m. shiui ERNST & MCCORD, 1987 can be found almost everywhere in Southeast Asia and are very common on Hainan Island (DE BRUIN & ARTNER, 1999). A combined morphological and genetic analysis would be rewarding.

(65) Chelonoidis nigra: after the death of "Lonesome George" on June 24th, 2012, Chelonoidis nigra abingdonii is extinct. The nominate subspecies Chelonoidis nigra nigra (QUOY & GAIMARD, 1824) and the subspecies C. n. phantastica (VAN DENBURGH, 1907) are extinct (PRITCHARD 1996) and therefore not included in this list. After many years of wise hesitation, the TURTLE TAXONOMY WORKING GROUP in its 2011 and 2012 issues considers the remaining 7 subspecies to be full species although the subspecies interbreed readily (PRITCHARD 1996) and their morphological similarities also clearly mark them as subspecies.