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Citation: Midtgaard, Rune. RepFocus - A Survey of the Reptiles of the World. (www.repfocus.dk).
Latest update: April 5th, 2026.


Taxonomy of the family Testudinidae
Bibliography of the genus Testudo
Biodiversity of the family Testudinidae
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 Genus
Testudo
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Image		 Palearctic Tortoises, Eurasian Tortoises, Mediterranean Tortoises
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Image		 Paläarktische Landschildkröten
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Image		 Palæarktiske Landskildpadder
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1758Image Testudo Linnaeus (type species: Testudo graeca Linnaeus 1758)
1820Image Chersine Merrem [substitute name for Testudo Linnaeus 1758] (type species: Testudo graeca sensu Merrem 1820 [not Testudo graeca Linnaeus 1758] = Testudo hermanni hermanni Gmelin 1789 or Testudo hermanni boettgeri Mojsisovich 1889; syn. Fritz & Kraus 2008)
1830Image Chersus Wagler (type species: Testudo marginata Schoepff 1792; syn. Fritz & Havas 2006)
1869Image Peltastes Gray [not Peltastes Illiger 1807 (Hymenoptera); not Peltastes Agassiz 1838 (Echinodermata); not Peltastes Fischer-Waldheim 1839 (Orthopthera)] (type species: Peltastes graecus sensu Gray 1869 = Testudo graeca ibera Pallas 1814; syn. Fritz & Havas 2006)
1870Image Testudinella Gray [not Testudinella Bory 1822 (Rotatoria)] (type species: Testudo horsfieldii Gray 1844; syn. Fritz & Havas 2006)
1872Image Peltonia Gray [substitute name for Peltatstes Gray 1869] (syn. Fritz & Havas 2006)
1916Image Medaestia Wussow (type species: Medaestia graeca sensu Wussow 1916 [not Testudo graeca Linnaeus 1758] = Testudo hermanni hermanni Gmelin 1789 or Testudo hermanni boettgeri Mojsisovich 1889; syn. Fritz & Kraus 2008)
1957Image Pseudotestudo Loveridge & Williams (type species: Testudo kleinmanni Lortet 1883; syn. Fritz & Havas 2006)
1966Image Agrionemys Khozatsky & Mlynarski (type species: Testudo horsfieldii Gray 1844; syn. Fritz & Havas 2006; Fritz & Bininda-Emonds 2007)
1990Image Furculachelys Highfield (type species: Furculachelys nabeulensis Highfield 1990; syn. Fritz, Bischoff, Martens & Schmidtler 1996; Fritz & Havas 2006)
2006Image Eurotestudo De Lapparent, Bour, Parham & Perälä (type species: Testudo hermanni Gmelin 1789; syn. Fritz & Bininda-Emonds 2007; Fritz & Kraus 2008; Bour & Ohler 2008)
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Contents: 5 species, none of which are endemic.
Endemism: 0% Image 100%
Remarks: Some authors place horsfieldi in a separate genus, Agrionemys. Gmira (1993) also placed hermanni in Agrionemys. Most recent works retain Agrionemys as a synonym of Testudo (e.g., Squalli-Houssaini & Blanc 1989; Fritz & Havas 2006; Vetter 2011).
Distribution: S. Europe, N. Africa, Middle East, C. Asia.
Reported from: Afghanistan, Albania, Algeria, Armenia, Azerbaijan, Bosnia-Herzegovina, Bulgaria, China (Xinjiang), Croatia, Egypt (incl. Sinai), France (incl. Corsica), Georgia, Greece (incl. Aegean Islands [incl. Sporades Islands]), Iran, Iraq, Israel, Italy (incl. Aeolian Islands, Capraia, Elba, Pantelleria, Pelagie Islands [incl. Lampedusa, Linosa Islands], Sicily), Jordan, Kazakhstan, Kyrgyzstan, Lebanon, Libya, Mongolia, Montenegro, Morocco, North Macedonia, Pakistan, Romania, Russia, Serbia, Slovenia, Spain, Syria, Tajikistan, Tunisia, Turkey (incl. European Turkey), Turkmenistan, Uzbekistan.
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Image 		Testudo graeca Image Image
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Image		 Spur-thighed Tortoise, Mediterranean Spur-thighed Tortoise, Moorish Tortoise
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Image		 Maurische Landschildkröte
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Image		 Maurisk Landskildpadde - (i ældre litteratur fejlagtigt kaldt 'Græsk Landskildpadde')
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1758Image Testudo graeca Linnaeus
1758Image Testudo pusilla Linnaeus (syn. Fritz & Havas 2006)
Image Chersine pusilla Merrem 1820
1775Image Testudo terrestris Forskål (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo graeca terrestris Wermuth 1958
Image Testudo ibera terrestris Artner 2003
1814Image Testudo ibera Pallas (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Medaestia ibera Wussow 1916
Image Testudo graeca ibera Mertens 1946
Image Testudo terrestris ibera David 1994
1835Image Testudo mauritanica Duméril & Bibron (syn. Fritz & Havas 2006)
Image Peltastes mauritanicus Gray 1873
1836Image Testudo whitei Bennett in White (syn. Iverson 1992, Fritz, Bischoff, Martens & Schmidtler 1996)
Image Peltastes marginatus var. whitei Gray 1870
Image Furculachelys whitei Highfield 1990
Image Testudo graeca whitei Kuyl, Ballasina, Dekker, Maas, Willemsen & Goudsmit 2002
1896Image Testudo zarudnyi Nikolski (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo graeca zarudnyi Mertens 1946
Image Testudo ibera zarudnyi Gmira 1993
Image Testudo terrestris zarudnyi David 1994
1920Image Testudo ibera var. bicaudalis Venzmer (syn. Fritz & Havas 2006)
1921Image Testudo buxtoni Boulenger (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo terrestris buxtoni Bour 2002
Image Testudo ibera buxtoni Artner 2003
Image Testudo graeca buxtoni Fritz, Hundsdörfer, Siroky, Auer, Kami, Lehmann, Mazanaeva, Turkosan & Wink 2007
1931Image Testudo ibera racovitzai Calinescu (syn. Fritz & Havas 2006)
1935Image Testudo floweri Bodenheimer (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo graeca floweri Mertens 1946
Image Testudo terrestris floweri David 1994
Image Testudo ibera floweri Artner 2003
1986Image Testudo graeca nikolskii Chkhikvadze & Tuniev (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo ibera nikolskii Highfield 1990
Image Testudo terrestris nikolskii David 1994
Image Testudo nikolskii Vetter 2002
1987Image Testudo graeca anamurensis Weissinger (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo ibera anamurensis Highfield 1990
Image Testudo terrestris anamurensis David 1994
Image Testudo anamurensis Vetter 2002
1989Image Testudo flavominimaralis Highfield & Martin (syn. Fritz & Havas 2006)
Image Furculachelys minimaralis [lapsus] van der Kuyl, Ballasina, Dekker, Maas, Willemsen & Goudsmit 2002
1990Image Furculachelys nabeulensis Highfield
Image Testudo nabeulensis Welch 1994
Image Testudo graeca nabeulensis Pieh & Perälä 2002
1991Image Testudo graeca armeniaca Tschchikwadse & Bakradse (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo armeniaca Vetter 2002
Image Testudo terrestris armeniaca Bour 2002
1996Image Testudo antakyensis Perälä (syn. Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo graeca antakyensis Zwartepoorte in Artner & Meier 2000
Image Testudo terrestris antakyensis Bour 2002
Image Testudo ibera antakyensis Artner 2003
2000Image Testudo graeca soussensis Pieh
Image Testudo soussensis Vetter 2002
2002Image Testudo graeca cyrenaica Pieh & Perälä
Image Testudo cyrenaica Vetter 2002
2002Image Testudo graeca pallasi Chkhikvadze & Bakradze
Image Testudo pallasi Danilov & Milto 2004
2002Image Testudo perses Perälä (syn. Fritz & Havas 2006; Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Image Testudo ibera perses Artner 2003
2003Image Testudo graeca lamberti Pieh & Perälä
Image Testudo lamberti Perälä in Pieh & Perälä 2004
2003Image Testudo graeca marokkensis Pieh & Perälä
Image Testudo marokkensis Perälä in Pieh & Perälä 2004
2024Image Testudo graeca tripolitania Poterala, Rattay, Johnson, Buijlayyil, Kiraz & Ajaj
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Other common names:
anamurensis: Anamurum Spur-thighed Tortoise, Anamur Tortoise
antakyensis: Antakya Tortoise
armeniaca: Armenian Tortoise, Araxes Tortoise
buxtoni: Buxton's Tortoise
cyrenaica: Cyrenaican Spur-thighed Tortoise
floweri: Flower's Dwarf Tortoise, Flower's Tortoise
graeca: Mediterranean Spur-thighed Tortoise
ibera: Eastern Spur-thighed Tortoise, Asia Minor Spur-thighed Tortoise
lamberti: Lambert's Tortoise
marokkensis: Moroccan Tortoise
nabeulensis: Tunisian Tortoise, Nabeul Tortoise
nikolskii: Nikolsky's Spur-thighed Tortoise, Nikolsky's Tortoise
pallasi: Daghestan Tortoise
perses: Zagros Mountain Tortoise
soussensis: Souss Valley Tortoise
terrestris: Middle Eastern Spur-thighed Tortoise, Mesopotamian Tortoise
zarudnyi: Iranian Spur-thighed Tortoise, Central Iranian Tortoise
Remarks: It has been suggested that most subspecies or synonyms (anamurensis, antakyensis, armeniaca, buxtoni, cyrenaica, flavominimaralis, floweri, ibera, lamberti, marokkensis, nabeulensis, nikolskii, pallasi, perses, soussensis, terrestris, whitei, zarudnyi) deserve full species status (e.g., Gmira 1983; Highfield 1990, Highfield & Martin 1989a, 1989b; Vetter 2002). This has not been generally accepted, however, and most authors regard the validity of these forms as separate species as insufficiently documented (e.g., Fritz, Bischoff, Martens & Schmidtler 1996; Iverson 1992; Schleich, Kästle & Kabisch 1996; Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006; Vetter 2011).
Erroneously reported from Egypt (Buskirk 1996; Baha el Din 2006). Although recorded from Albania, there are no recent records from the country (Mizsei, Jablonski, Vegvari, Lengyel & Szabolcs 2017).
Introduced to: Cyprus, France, Gibraltar, Greece (Rhodes, Crete), Italy (Sardinia, Mal di Ventre Island, Sicily), Spain (Balearic Islands [Mallorca, Pityusic Islands (Formentera, Ibiza)], Canary Islands).
Distribution: Albania, Algeria, Armenia, Azerbaijan, Bulgaria, Georgia, Greece (incl. Aegean Islands), Iran, Iraq, Israel, Jordan, Lebanon, Libya, Moldova, Montenegro, Morocco, North Macedonia, Romania, Russia, Serbia, Spain (mainland), Syria, Tunisia, Turkey (incl. European Turkey). 		Image
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Testudo graeca
© Rune Midtgaard
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Image 		Testudo hermanni Image Image
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Image		 Greek Tortoise, (Hermann's Tortoise)
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Image		 Griechische Landschildkröte, Europäische Landschildkröte
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Image		 Græsk Landskildpadde, (Hermanns Landskildpadde)
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1789Image Testudo hermanni Gmelin
Image Protestudo hermanni Chkhikvadze 1983
Image Agrionemys hermanni Gmira 1993
Image Eurotestudo hermanni Lapparent de Broin, Bour, Parham & Perälä 2006
1881Image Testudo graeca bettai Lataste (syn. Fritz & Havas 2006)
1889Image Testudo graeca var. boettgeri Mojsisovics
Image Testudo hermanni boettgeri Bour 1987
Image Testudo boettgeri Vetter 2002
Image Eurotestudo boettgeri Lapparent de Broin, Bour, Parham & Perälä 2006
1899Image Testudo graeca var. hercegovinensis Werner (syn. Fritz & Havas 2006)
Image Testudo hercegovinensis Perälä 2002
Image Testudo boettgeri hercegovinensis Artner 2003
Image Testudo hermanni hercegovinensis Vinke & Vinke 2004
Image Eurotestudo hercegovinensis Lapparent de Broin, Bour, Parham & Perälä 2006
1932Image Testudo enriquesi Parenzan (syn. Fritz & Havas 2006)
1952Image Testudo hermanni robertmertensi Wermuth
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Other common names:
boettgeri: Eastern Greek Tortoise, Eastern Hermann's Tortoise
hercegovinensis: Herzegovina Tortoise
hermanni: Western Greek Tortoise, Western Hermann's Tortoise
Remarks: See under genus. The western subspecies, boettgeri, was known for many years as robertmertensi, however, after the type of robertmertensi was shown to be identical with the nominal, eastern subspecies (hermanni), the western subspecies has been referred to as boettgeri, the next available name.
Introduced to: Cyprus, Greece (Rhodes), Italy (Sardinia), Spain (Balearic Islands [Mallorca, Minorca]).
Distribution: Albania, Bosnia-Herzegovina, Bulgaria, Croatia, France (incl. Corsica), Greece (incl. Aegean Islands), Italy (incl. Aeolian Islands, Capraia, Elba, Pantelleria, Pelagie Islands [Lampedusa, Linosa Islands], Sicily), Montenegro, North Macedonia, Romania, Serbia, Slovenia, Spain, Turkey (incl. European Turkey). 		Image
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Testudo hermanni
© Rune Midtgaard
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Image 		Testudo horsfieldii Image Image
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Image		 Steppe Tortoise, Central Asian Tortoise, Afghan Tortoise, Four-toed Tortoise, (Horsfield's Tortoise)
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Image		 Steppenschildkröte, Vierzehenschildkröte
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Image		 Russisk Landskildpadde, Firtået Landskildpadde
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1844Image Testudo horsfieldii Gray
Image Testudinella horsfieldii Gray 1870
Image Homopus horsfieldii Theobald 1876
Image Medaestia horsfieldi Wussow 1916
Image Agrionemys horsfieldi Mlynarski 1966
1854Image Homopus burnesii Blyth '1853' (syn. Fritz & Havas 2006)
1906Image Testudo baluchiorum Annandale (syn. Fritz & Havas 2006)
Image Agrionemys baluchiorum Vetter 2002
Image Agrionemys horsfieldii baluchiorum Artner 2003
1988Image Agrionemys horsfieldi kazachstanica Chkhikvadze
Image Testudo horsfieldii kazachstanica David 1994
Image Agrionemys kazachstanica Perälä 2002
Image Agrionemys horsfieldii kazakhstanica [lapsus] Bonin, Devaux & Dupré 2006
Image Testudo kazachstanica Bickham, Iverson, Parham, Philippen, Rhodin, Shaffer, Spinks & van Dijk 2007
1990Image Agrionemys horsfieldi rustamowi Chkhikvadze, Amiranashvili & Ataev
Image Testudo horsfieldii rustamovi David 1994
Image Agrionemys rustamovi Perälä 2002
Image Testudo rustamovi Bickham, Iverson, Parham, Philippen, Rhodin, Shaffer, Spinks & van Dijk 2007
2009Image Agrionemys bogdanovi Chkhikvadze, Brushko & Kubykin
Image Testudo horsfieldii bogdanovi Dijk, Iverson, Shaffer, Bour & Rhodin 2012
2009Image Agrionemys kazachstanica kuznetzovi Chkhikvadze, Ataev, Shammakov & Zatoka in Chkhikvadze, Ataev & Shammakov
Image Testudo horsfieldii kuznetzovi Dijk, Iverson, Shaffer, Bour & Rhodin 2012
2009Image Agrionemys kazachstanica terbishi Chkhikvadze in Chkhikvadze, Ataev & Shammakov
Image Testudo horsfieldii terbishi Dijk, Iverson, Shaffer, Bour & Rhodin 2012
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Other common names:
bogdanovi: Fergana Valley Steppe Tortoise
kazachstanica: Kazakhstan Steppe Tortoise
kuznetzovi: Turkmenistan Steppe Tortoise
rustamovi: Kopet-Dag Steppe Tortoise
terbishi: Mongolian Steppe Tortoise
Remarks: See under genus. The Mongolian population may have been introduced (syn. Dijk, Iverson, Shaffer, Bour & Rhodin 2012)
Introduced to: Latvia, USA (syn. Florida).
Distribution: Afghanistan, China (syn. Xinjiang), Iran, Kazakhstan, Kyrgyzstan, Mongolia, Pakistan, Tajikistan, Turkmenistan, Uzbekistan. 		Image
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Testudo horsfieldii
© Rune Midtgaard
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Image 		Testudo kleinmanni Image Image
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Image		 Egyptian Tortoise, (syn. Kleinmann's Tortoise)
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Image		 Ägyptische Landschildkröte
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Image		 Ægyptisk Landskildpadde
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1869Image Testudo leithii Günther [not Testudo leithi Carter 1852]
Image Peltastes leithii Gray 1870
Image Medaestia leithi Wussow 1916
1883Image Testudo kleinmanni Lortet
Image Pseudotestudo kleinmanni Obst 1978
2001Image Testudo werneri Perälä (syn. Fritz & Havas 2006; Attum, Baha el Din, Carranza, Earley, Arnold & Kingsbury 2007)
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Other common names:
werneri: Negev Tortoise, (Werner's Tortoise)
Distribution: Egypt (incl. Sinai), Israel, Libya. 		Image
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Testudo kleinmanni
© Rune Midtgaard
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Image 		Testudo marginata Image Image
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Image		 Marginated Tortoise
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Image		 Breitrandschildkröte
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Image		 Bredrandet Landskildpadde
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1792Image Testudo marginata Schoepff
Image Chersine marginata Merrem 1820
Image Chersus marginatus Wagler 1830
Image Peltastes marginatus Gray 1869
1782Image Testudo tabulata var. campanulata Walbaum [nomen illegitimum] (syn. Fritz & Havas 2006)
1792Image Testudo graja Hermann in Schoepff (syn. Fritz & Havas 2006)
1870Image Peltastes marginatus var. melas Gray (syn. Fritz & Havas 2006)
1875Image Testudo nemoralis Schreiber (syn. Fritz & Havas 2006)
1992Image Testudo marginata sarda Mayer (syn. Fritz, Petters, Matzanke & Matzanke 1995)
1994Image Testudo marginata weissingeri Trutnau
Image Testudo weissingeri Bour 1996
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Other common names:
weissingeri: Peloponnese Dwarf Tortoise
Introduced to: Cyprus, Greece (Aegean Islands [Cyclades Islands (Kimolos), North Aegean Islands (Lesbos)], Ionian Islands [Cephalonia, Corfu, Lefkada, Sapientza, Sphacteria]), Italy (Sardinia), Malta, Turkey.
Distribution: Albania, Greece (incl. Aegean Islands [incl. Sporades Islands]). 		Image
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Testudo marginata
© Henrik Bringsøe