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© Rune Midtgaard


Taxonomy of the family Testudinidae
Bibliography of the genus Testudo
Biodiversity of the family Testudinidae








Genus
Testudo

Palearctic Tortoises, Eurasian Tortoises, Mediterranean Tortoises

Paläarktische Landschildkröten

Palæarktiske Landskildpadder

1758 Testudo Linnaeus (type species: Testudo graeca Linnaeus 1758)
1820 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)
1830 Chersus Wagler (type species: Testudo marginata Schoepff 1792; syn. Fritz & Havas 2006)
1869 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)
1870 Testudinella Gray [not Testudinella Bory 1822 (Rotatoria)] (type species: Testudo horsfieldii Gray 1844; syn. Fritz & Havas 2006)
1872 Peltonia Gray [substitute name for Peltatstes Gray 1869] (syn. Fritz & Havas 2006)
1916 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)
1957 Pseudotestudo Loveridge & Williams (type species: Testudo kleinmanni Lortet 1883; syn. Fritz & Havas 2006)
1966 Agrionemys Khozatsky & Mlynarski (type species: Testudo horsfieldii Gray 1844; syn. Fritz & Havas 2006; Fritz & Bininda-Emonds 2007)
1990 Furculachelys Highfield (type species: Furculachelys nabeulensis Highfield 1990; syn. Fritz, Bischoff, Martens & Schmidtler 1996; Fritz & Havas 2006)
2006 Eurotestudo De Lapparent, Bour, Parham & Perälä (type species: Testudo hermanni Gmelin 1789; syn. Fritz & Bininda-Emonds 2007; Fritz & Kraus 2008; Bour & Ohler 2008)

Contents: 5 species, none of which are endemic.
Endemism: 0% 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.

Testudo graeca

Spur-thighed Tortoise, Mediterranean Spur-thighed Tortoise, Moorish Tortoise

Maurische Landschildkröte

Maurisk Landskildpadde - (i ældre litteratur fejlagtigt kaldt Græsk Landskildpadde)

1758 Testudo graeca Linnaeus
1758 Testudo pusilla Linnaeus (Fritz & Havas 2006)
Chersine pusilla Merrem 1820
1775 Testudo terrestris Forskål (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo graeca terrestris Wermuth 1958
Testudo ibera terrestris Artner 2003
1814 Testudo ibera Pallas (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Medaestia ibera Wussow 1916
Testudo graeca ibera Mertens 1946
Testudo terrestris ibera David 1994
1835 Testudo mauritanica Duméril & Bibron (Fritz & Havas 2006)
Peltastes mauritanicus Gray 1873
1836 Testudo whitei Bennett in White (Iverson 1992, Fritz, Bischoff, Martens & Schmidtler 1996)
Peltastes marginatus var. whitei Gray 1870
Furculachelys whitei Highfield 1990
Testudo graeca whitei Kuyl, Ballasina, Dekker, Maas, Willemsen & Goudsmit 2002
1896 Testudo zarudnyi Nikolski (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo graeca zarudnyi Mertens 1946
Testudo ibera zarudnyi Gmira 1993
Testudo terrestris zarudnyi David 1994
1920 Testudo ibera var. bicaudalis Venzmer (Fritz & Havas 2006)
1921 Testudo buxtoni Boulenger (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo terrestris buxtoni Bour 2002
Testudo ibera buxtoni Artner 2003
Testudo graeca buxtoni Fritz, Hundsdörfer, Siroky, Auer, Kami, Lehmann, Mazanaeva, Turkosan & Wink 2007
1931 Testudo ibera racovitzai Calinescu (Fritz & Havas 2006)
1935 Testudo floweri Bodenheimer (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo graeca floweri Mertens 1946
Testudo terrestris floweri David 1994
Testudo ibera floweri Artner 2003
1986 Testudo graeca nikolskii Chkhikvadze & Tuniev (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo ibera nikolskii Highfield 1990
Testudo terrestris nikolskii David 1994
Testudo nikolskii Vetter 2002
1987 Testudo graeca anamurensis Weissinger (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo ibera anamurensis Highfield 1990
Testudo terrestris anamurensis David 1994
Testudo anamurensis Vetter 2002
1989 Testudo flavominimaralis Highfield & Martin (Fritz & Havas 2006)
Furculachelys minimaralis [lapsus] van der Kuyl, Ballasina, Dekker, Maas, Willemsen & Goudsmit 2002
1990 Furculachelys nabeulensis Highfield
Testudo nabeulensis Welch 1994
Testudo graeca nabeulensis Pieh & Perälä 2002
1991 Testudo graeca armeniaca Tschchikwadse & Bakradse (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo armeniaca Vetter 2002
Testudo terrestris armeniaca Bour 2002
1996 Testudo antakyensis Perälä (Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo graeca antakyensis Zwartepoorte in Artner & Meier 2000
Testudo terrestris antakyensis Bour 2002
Testudo ibera antakyensis Artner 2003
2000 Testudo graeca soussensis Pieh
Testudo soussensis Vetter 2002
2002 Testudo graeca cyrenaica Pieh & Perälä
Testudo cyrenaica Vetter 2002
2002 Testudo graeca pallasi Chkhikvadze & Bakradze
Testudo pallasi Danilov & Milto 2004
2002 Testudo perses Perälä (Fritz & Havas 2006; Parham, Turkosan, Stuart, Arakelyan, Shafei, Macey & Papenfuss 2006)
Testudo ibera perses Artner 2003
2003 Testudo graeca lamberti Pieh & Perälä
Testudo lamberti Perälä in Pieh & Perälä 2004
2003 Testudo graeca marokkensis Pieh & Perälä
Testudo marokkensis Perälä in Pieh & Perälä 2004

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).


Testudo graeca
© Rune Midtgaard

Testudo hermanni

Greek Tortoise, (Hermann's Tortoise)

Griechische Landschildkröte, Europäische Landschildkröte

Græsk Landskildpadde, (Hermanns Landskildpadde)

1789 Testudo hermanni Gmelin
Protestudo hermanni Chkhikvadze 1983
Agrionemys hermanni Gmira 1993
Eurotestudo hermanni Lapparent de Broin, Bour, Parham & Perälä 2006
1881 Testudo graeca bettai Lataste (Fritz & Havas 2006)
1889 Testudo graeca var. boettgeri Mojsisovics
Testudo hermanni boettgeri Bour 1987
Testudo boettgeri Vetter 2002
Eurotestudo boettgeri Lapparent de Broin, Bour, Parham & Perälä 2006
1899 Testudo graeca var. hercegovinensis Werner (Fritz & Havas 2006)
Testudo hercegovinensis Perälä 2002
Testudo boettgeri hercegovinensis Artner 2003
Testudo hermanni hercegovinensis Vinke & Vinke 2004
Eurotestudo hercegovinensis Lapparent de Broin, Bour, Parham & Perälä 2006
1932 Testudo enriquesi Parenzan (Fritz & Havas 2006)
1952 Testudo hermanni robertmertensi Wermuth

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).


Testudo hermanni
© Rune Midtgaard

Testudo horsfieldii

Steppe Tortoise, Central Asian Tortoise, Afghan Tortoise, Four-toed Tortoise, (Horsfield's Tortoise)

Steppenschildkröte, Vierzehenschildkröte

Russisk Landskildpadde, Firtået Landskildpadde

1844 Testudo horsfieldii Gray
Testudinella horsfieldii Gray 1870
Homopus horsfieldii Theobald 1876
Medaestia horsfieldi Wussow 1916
Agrionemys horsfieldi Mlynarski 1966
1854 Homopus burnesii Blyth (Fritz & Havas 2006)
1906 Testudo baluchiorum Annandale (Fritz & Havas 2006)
Agrionemys baluchiorum Vetter 2002
Agrionemys horsfieldii baluchiorum Artner 2003
1988 Agrionemys horsfieldi kazachstanica Chkhikvadze
Testudo horsfieldii kazachstanica David 1994
Agrionemys kazachstanica Perälä 2002
Agrionemys horsfieldii kazakhstanica [lapsus] Bonin, Devaux & Dupré 2006
Testudo kazachstanica Bickham, Iverson, Parham, Philippen, Rhodin, Shaffer, Spinks & van Dijk 2007
1990 Agrionemys horsfieldi rustamowi Chkhikvadze, Amiranashvili & Ataev
Testudo horsfieldii rustamovi David 1994
Agrionemys rustamovi Perälä 2002
Testudo rustamovi Bickham, Iverson, Parham, Philippen, Rhodin, Shaffer, Spinks & van Dijk 2007
2009 Agrionemys bogdanovi Chkhikvadze, Brushko & Kubykin
Testudo horsfieldii bogdanovi Dijk, Iverson, Shaffer, Bour & Rhodin 2012
2009 Agrionemys kazachstanica kuznetzovi Chkhikvadze, Ataev, Shammakov & Zatoka in Chkhikvadze, Ataev & Shammakov
Testudo horsfieldii kuznetzovi Dijk, Iverson, Shaffer, Bour & Rhodin 2012
2009 Agrionemys kazachstanica terbishi Chkhikvadze in Chkhikvadze, Ataev & Shammakov
Testudo horsfieldii terbishi Dijk, Iverson, Shaffer, Bour & Rhodin 2012

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 (Dijk, Iverson, Shaffer, Bour & Rhodin 2012)
Introduced to: Latvia, USA (Florida).
Distribution: Afghanistan, China (Xinjiang), Iran, Kazakhstan, Kyrgyzstan, Mongolia, Pakistan, Tajikistan, Turkmenistan, Uzbekistan.


Testudo horsfieldii
© Rune Midtgaard

Testudo kleinmanni

Egyptian Tortoise, (Kleinmann's Tortoise)

Ägyptische Landschildkröte

Ægyptisk Landskildpadde

1869 Testudo leithii Günther [not Testudo leithi Carter 1852]
Peltastes leithii Gray 1870
Medaestia leithi Wussow 1916
1883 Testudo kleinmanni Lortet
Pseudotestudo kleinmanni Obst 1978
2001 Testudo werneri Perälä (Fritz & Havas 2006; Attum, Baha el Din, Carranza, Earley, Arnold & Kingsbury 2007)

Other common names:
werneri: Negev Tortoise, (Werner's Tortoise)
Distribution: Egypt (incl. Sinai), Israel, Libya.


Testudo kleinmanni
© Rune Midtgaard

Testudo marginata

Marginated Tortoise

Breitrandschildkröte

Bredrandet Landskildpadde

1792 Testudo marginata Schoepff
Chersine marginata Merrem 1820
Chersus marginatus Wagler 1830
Peltastes marginatus Gray 1869
1782 Testudo tabulata var. campanulata Walbaum [nomen illegitimum] (Fritz & Havas 2006)
1792 Testudo graja Hermann in Schoepff (Fritz & Havas 2006)
1870 Peltastes marginatus var. melas Gray (Fritz & Havas 2006)
1875 Testudo nemoralis Schreiber (Fritz & Havas 2006)
1992 Testudo marginata sarda Mayer (Fritz, Petters, Matzanke & Matzanke 1995)
1994 Testudo marginata weissingeri Trutnau
Testudo weissingeri Bour 1996

Other common names:
weissingeri: Peloponnese Dwarf Tortoise
Introduced to: Cyprus, Greece (Kimolos, Lesbos), Italy (Sardinia), Malta, Turkey.
Distribution: Albania, Greece (incl. Aegean Islands [incl. Sporades Islands]).


Testudo marginata
© Henrik Bringsøe