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Testudo graeca LINNAEUS, 1758

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Higher TaxaTestudinidae, Testudinoidea, Testudines (turtles)
SubspeciesTestudo graeca graeca LINNAEUS 1758
Testudo graeca armeniaca CHKHIKVADZE & BAKRADSE 1991
Testudo graeca buxtoni BOULENGER 1921
Testudo graeca cyrenaica PIEH & PERÄLÄ 2002
Testudo graeca ibera PALLAS 1814
Testudo graeca marokkensis PIEH & PERÄLÄ 2004
Testudo graeca nabeulensis HIGHFIELD 1990
Testudo graeca perses PERÄLÄ 2002
Testudo graeca soussensis PIEH 2000
Testudo graeca terrestris FORSKÅL 1775
Testudo graeca whitei BENNETT in WHITE 1836
Testudo graeca zarudnyi NIKOLSKY 1896 
Common NamesE: Mediterranean Spur-thighed Tortoise
G: Maurische Landschildkröte
Russian: Средиземноморская черепаха
Italian: testuggine moresca 
SynonymTestudo graeca LINNAEUS 1758: 198
Testudo pusilla LINNAEUS 1758: 198
Testudo graeca — SONNINI & LATREILLE 1802: 65
Testudo pusilla — SONNINI & LATREILLE 1802: 92
Testudo graeca — LINK 1807: 53
Testudo ibera PALLAS 1814 (fide KHALIKOV, pers. comm.)
Testudo mauritanica DUMÉRIL & BIBRON 1835: 45
Testudo graeca — DUMÉRIL & BIBRON 1835: 49
Testudo mauritanica — DUMÉRIL & BIBRON 1854: 219
Testudo graeca — DUMÉRIL & BIBRON 1854: 219
Testudo ibera — BOULENGER 1836
Testudo ibera — WERNER 1899: 376
Testudo graeca — ENGELMANN et al. 1993
Testudo graeca — SCHLEICH, KÄSTLE & KABISCH 1996: 148
Testudo graeca — LAPPARENT DE BROIN 2001
Testudo dagestanica CHIKHIKVADZE, MAZANAEVA & SHAMMAKOV 2011
Testudo graeca dagestanica — HERZ 2013
Testudo (Testudo) graeca — TTWG 2017: 151

Testudo graeca graeca LINNAEUS 1758
Testudo pusilla LINNAEUS 1758
Testudo mauritanica DUMÉRIL & BIBRON 1835: 45
Tistudo mauritonica — KERCADO 1835 (error typographicus)
Testudo pusilla — BOETTGER 1879: 81
Testudo graeca graeca — MERTENS 1946
Testudo graeca graeca — BONIN et al 2006
Testudo graeca graeca — PIEH & PHILIPPEN 2007
Testudo graeca graeca — HERZ 2013
Testudo (Testudo) graeca graeca — TTWG 2017: 152
Testudo (Testudo) graeca graeca — TTWG 2021

Testudo graeca antakyensis PERÄLÄ 1996
Testudo antakyensis PERÄLÄ 1996
Testudo (graeca) antakyensis — BONIN et al 2006
Testudo graeca antakyensis — PIEH & PHILIPPEN 2007
Testudo graeca antakyensis — HERZ 2013

Testudo graeca armeniaca CHKHIKVADZE & BAKRADSE 1991
Testudo graeca armeniaca CHKHIKVADZE 1989 (nom. nud.)
Testudo graeca pallasi CHKHIKVADZE & BAKRADZE 2002
Testudo (graeca) pallasi — BONIN et al 2006
Testudo (graeca) armeniaca — BONIN et al 2006
Testudo graeca armeniaca — AUER 2007
Testudo graeca armeniaca — PIEH & PHILIPPEN 2007
Testudo graeca pallasi — PIEH & PHILIPPEN 2007
Testudo graeca pallasi — HERZ 2013
Testudo graeca passasi [sic] — HERZ 2013: 113
Testudo graeca armeniaca — HERZ 2013
Testudo graeca armeniaca — TUNIYEV et al. 2014
Testudo graeca armeniaca — JAVANBAKHT et al. 2017
Testudo (Testudo) graeca armeniaca — TTWG 2017: 152
Testudo (Testudo) graeca armeniaca — TTWG 2021

Testudo graeca buxtoni BOULENGER 1921
Testudo buxtoni BOULENGER 1921 (valid fide PIEH et al. 2002)
Testudo buxtoni — PERÄLÄ 2001
Testudo buxtoni — PIEH & PHILIPPEN 2007
Testudo graeca buxtoni — AUER & HERZ 2007
Testudo graeca buxtoni — HERZ 2013
Testudo graeca buxtoni — JAVANBAKHT et al. 2017
Testudo (Testudo) graeca buxtoni — TTWG 2017: 152
Testudo (Testudo) graeca buxtoni — TTWG 2021

Testudo graeca cyrenaica PIEH & PERÄLÄ 2002
Testudo graeca cyrenaica — BONIN et al 2006
Testudo graeca cyrenaica — PIEH & PHILIPPEN 2007
Testudo graeca cyrenaica — HERZ 2013
Testudo (Testudo) graeca cyrenaica — TTWG 2017: 153
Testudo (Testudo) graeca cyrenaica — TTWG 2021

Testudo graeca ibera PALLAS 1814
Testudo ibera PALLAS 1814: 18
Testudo georgicana GÜLDENSTEDT 1814 (nomen nudum)
Testudo ecaudata PALLAS 1814 (nomen dubium fide PIEH et al. 2002)
Testudo ibera var. bicaudalis VENZMER 1918
Testudo ibera racovitzai CALINESCU 1931
Testudo graeca ibera — MERTENS 1946
Testudo graeca nikolskii CHKHIKVADZE & TUNIEV 1986
Testudo graeca nikolskii — TUNIYEV 1990
Testudo terrestris ibera — DAVID 1994
Testudo graeca ibera — STOEV 2000
Testudo graeca ibera — BISCHOFF 2002
Testudo graeca ibera — TÜRKOZAN et al. 2005
Testudo (graeca) ibera — BONIN et al 2006
Testudo (graeca) nikolskii — BONIN et al 2006
Testudo graeca nikolskii — PIEH & PHILIPPEN 2007
Testudo graeca ibera — AUER 2007
Testudo graeca ibera — PIEH & PHILIPPEN 2007
Testudo graeca nikolskii — HERZ 2013
Testudo ibera racovitzai — HERZ 2013
Testudo graeca ibera — HERZ 2013
Testudo graeca ibera — JAVANBAKHT et al. 2017
Testudo (Testudo) graeca ibera — TTWG 2017: 153
Testudo (Testudo) graeca ibera — TTWG 2021

Testudo graeca marokkensis PIEH & PERÄLÄ 2004
Testudo graeca marokkensis — BONIN et al 2006
Testudo graeca lamberti — BONIN et al 2006
Testudo graeca lamberti — PIEH & PHILIPPEN 2007
Testudo graeca marokkensis — PIEH & PHILIPPEN 2007
Testudo graeca lamberti — HERZ 2013
Testudo graeca marokkensis — HERZ 2013
Testudo (Testudo) graeca marokkensis — TTWG 2017: 153
Testudo (Testudo) graeca marokkensis — TTWG 2021

Testudo graeca nabeulensis (HIGHFIELD 1990)
Furculachelys nabeulensis HIGHFIELD 1990: 32
Testudo flavominimaralis HIGHFIELD & MARTIN 1990 (fide KUYL et al. 2002)
Testudo nabeulensis — IVERSON
Testudo nabeulensis — VETTER 2002
Testudo graeca nabeulensis — KUYL et al. 2002
Testudo nabeulensis — ROGNER 2006
Testudo graeca nabeulensis — BONIN et al 2006
Testudo graeca nabeulensis — PIEH & PHILIPPEN 2007
Testudo graeca flavominimaralis — HERZ 2013
Testudo graeca nabeulensis — HERZ 2013
Testudo flavominimaralis — BAUER et al. 2017: 265
Testudo (Testudo) graeca — TTWG 2017: 153
Testudo (Testudo) graeca nabeulensis — TTWG 2021

Testudo graeca perses PERÄLÄ 2002
Testudo perses PERÄLÄ 2002
Testudo perses — TÜRKOZAN et al. 2004
Testudo (graeca) perses — BONIN et al 2006
Testudo graeca perses — PIEH & PHILIPPEN 2007
Testudo graeca perses — HERZ 2013

Testudo graeca soussensis PIEH 2000
Testudo graeca soussensis — BONIN et al 2006
Testudo graeca soussensis — PIEH & PHILIPPEN 2007
Testudo graeca soussensis — LEPTIEN 2009
Testudo graeca soussensis — HERZ 2013
Testudo (Testudo) graeca soussensis — TTWG 2017: 153

Testudo graeca terrestris FORSKÅL 1775
Testudo terrestris FORSKÅL 1775 (non T. t. FERMIN 1765)
Testudo zolkafa FORSKÅL 1831 (nomen nudum)
Testudo zohalfa FORSKÅL 1835 (nomen nudum)
Testudo floweri BODENHEIMER 1935
Testudo graeca floweri — MERTENS 1946
Testudo graeca terrestris — WERMUTH 1958
Testudo graeca terrestris — ESTERBAUER 1985
Testudo graeca anamurensis WEISSINGER 1987
Testudo terrestris terrestris — DAVID 1994
Testudo terrestris — PERÄLÄ 2001
Testudo floweri — PERÄLÄ 2001
Testudo floweri — LAHONY et al. 2002
Testudo graeca terrestris — HRAOUI-BLOQUET et al. 2002
Testudo terrestris — PERÄLÄ & BOUR 2004
Testudo (graeca) terrestris — BONIN et al 2006
Testudo (graeca) anamurensis — BONIN et al 2006
Testudo graeca floweri — BONIN et al 2006
Testudo graeca floweri — PIEH & PHILIPPEN 2007
Testudo graeca anamurensis — PIEH & PHILIPPEN 2007
Testudo graeca terrestris — PIEH & PHILIPPEN 2007
Testudo graeca anamurensis — HERZ 2013
Testudo graeca floweri — HERZ 2013
Testudo graeca terrestris — HERZ 2013
Testudo graeca terrestris — JAVANBAKHT et al. 2017
Testudo (Testudo) graeca terrestris — TTWG 2017: 154
Testudo (Testudo) graeca terrestris — TTWG 2021

Testudo graeca whitei BENNETT in WHITE 1836
Testudo whitei BENNETT in WHITE 1836: 361
Peltastes marginatus whitei — GRAY 1870: 12
Testudo marginata whitei
Furculachelys whitei — HIGHFIELD 1990
Testudo graeca whitei — HERZ 2013
Testudo (Testudo) graeca whitei — TTWG 2021

Testudo graeca zarudnyi NIKOLSKY 1896
Testudo zarudnyi NIKOLSKY 1896
Testudo zarudnyi — SCHMIDT 1939
Testudo graeca zarudnyi — MERTENS 1946
Testudo terrestris zarudnyi — DAVID 1994
Testudo zarudnyi — PERÄLÄ 2001
Testudo (graeca) zarudnyi — BONIN et al 2006
Testudo graeca zarudnyi — PIEH & PHILIPPEN 2007
Testudo graeca zarudnyi — HERZ 2013
Testudo graeca zarudnyi — JAVANBAKHT et al. 2017
Testudo (Testudo) graeca zarudnyi — TTWG 2017: 154
Testudo graeca zarudnyi — ANANJEVA et al. 2020
Testudo (Testudo) graeca zarudnyi — TTWG 2021 
DistributionS Spain (introduced to Mallorca, Formentera)
Italy incl. Sardinia and Sicily (introduced), France (introduced),
N Greece (including Aegean islands, e.g. Limnos, Lesbos, Chios, Samos, Samothraki),
Serbia, Montenegro, Albania, Romania, Bulgaria, Turkey,
Morocco, Algeria, Tunisia, Libya, Israel, Syria, Jordan, Iran, N Iraq
Russia (eastwards to Transcaucasus; Dagestan),

acording to the 1994 IUCN Red List of Threatened Animals:
Albania, Algeria, Armenia, Azerbaijan, Bulgaria, Egypt, France, E Republic of Georgia, Greece, Iran, Iraq, Israel, Italy, Jordan, Lebanon, Libya, Macedonia, Moldova, Morocco, Romania, Russia, Spain, Syria, Tunisia, Turkey, Turkmenistan

armeniaca: Trans-Caucasus, Iran; Type locality: Megri, southeastern Armenia.

anamurensis: Cape Anamur and SW coast of Turkey. Type locality: Anamurum, southern Turkey.

antakyensis: S Turkey; Type locality: Antakya, Turkey.

buxtoni: N Iran; Type locality: Manjil, between Resht and Kaswin, Iran.

cyrenaica: Libya (endemic); Type locality: Derna (= Darna or Darnah, 32°46'N 22°39'E), Cyrenaica, eastern Libya.

flavominimaralis: Tunisia

floweri: Negev (S arid Israel); type locality: “Negev”.

ibera: S Serbia, Greece (Lesbos, Samos), Romania through Iran. Type locality: Tbilisi, Georgia (designated by Bour 1987).

lamberti: NW Morocco; Type locality: 22 km north of Tétouan (35°34'N 5°22'W), Morocco.

marokkensis: NW Morocco; Type locality: Tarmilete (33°23'N 6°04'W), Morocco.

nabeulensis: Tunisia, NW Libya; Type locality: Nabeul, N Tunisia

nikolskii: Republic of Georgia, Russia (western Caucasus); Type locality: Nebug Settlement, Tuapse County, Krasnodar District, Russia.

pallasi: Russia (Daghestan); Type locality: vicinity of Gilyary-Dag Settlement, Dagestan.

perses: SE Turkey, NE Iraq, SC Iran (Zagros mountains); Type locality: 3 miles W of Lālābād Village and 25 miles NW Kermānshāh (34°27'N 46°50'E), Kermānshāhān Province, west-central Iran.

soussensis: Souss valley, Morocco; Type locality: Vincity of Agadir, 30°28'N, 9°55'W.

terrestris: Israel, Lebanon, Jordan, Syria, SE Turkey. Designated type locality (by neotype designation; Perälä & Bour 2004): Aleppo, Syria.

zarudnyi: E Iran; Type locality: “Birdschan” [Birjand, South Khorasan Province, Iran, 32.87 N 59.22 E].

whitei: Algeria; Type locality: Algiers and its environs, Algeria (Highfield & Martin 1989)

Type locality: "Africa," according to Linnaeus 1758:352; however, the caption of the holotype, Plate 204 of Edwards 1751 reads "Loc. Santa Cruz in West Barbary" (=old fort Santa Cruz near Oran, Algeria, according to Strauch (1862:67). Mertens and Müller (1928:22), designated it as "Santa Cruz, in der Westberberei, Nordafrika" (after KING & BURKE 1989).  
ReproductionOviparous. Temperature-dependent sex determination (TSD) (Pieau 1998). 
TypesIconotype: The life-sized colored figure in Plate 204 of Edwards 1751 according to Loveridge and Williams 1957:264-265.
Holotype: BNHS fide Boulenger 1921, collected by Captain P.A. Buxton, BMNH 1947.3.5.16 (1920.8.6.1) Presented: Bombay Nat. Hist. Soc. Type of Testudo buxtoni, listed as Testudo graeca ibera) [buxtoni]
Holotype: MTD (= MTKD) 31880, adult female, don. Detlef MENDAU VII. 1991 [cyrenaica]
Lectotype: Iconotype (Pallas. Plate II. fig. 2- 3, originally unpublished but reproduced by Strauch 1862:69: Darevsky and Mertens 1973: 100: and Bour 1986: 113). Designation by Bour (1986: 113) [ibera]
Holotype: SMNS 7602 [marokkensis]
Neotype: MNHN 0.1937, designated by Schweiger & Gemel 2020. Previously, USNM 10980 was considered as (questionable) syntype [Testudo mauritanica]
Holotype: unknown, "adult male", without further collection information [nabeulensis]
Holotype: FMNH 130820 [perses]
Paratype: SMNS 4647.1 [soussensis]
Hololype: lost (apparently fide R. G. Webb. pers. comm., cited in Iverson 1992: 286) [terrestris]
Hololype: ZISP 8738 (also given as ZIL) [zarudnyi] 
DiagnosisAdditional details, e.g. a detailed description or comparisons (12019 characters), are available for collaborators and contributors. Please contact us for details. 
CommentDistribution: Erroneously reported from Egypt (SCHLEICH et al. 1996), but does not occur there (BUSKIRK 1996). Testudo graeca has been reported from the Canary Islands but these specimens are most likely escaped pets that do not form viable populations (A. VILADO, pers. comm.). For a map see Escoriza & Hassine 2022: Fig. 1 and Türkozan et al. 2023: Fig. 11.

Hybridization: Testudo graeca ibera has been reported to produce viable hybrids with Testudo horsfieldii, T. hermanni boettgeri, and T. marginata (KABISCH 2001, STEMMLER-GYGER 1963, 1964, HEIMANN 1986, MERTENS 1968, Rogner 2013).

Taxonomy: Testudo buxtoni has been considered as valid species by PIEH et al. (2002). Oğuz et al. (2010) found that “antakyensis”, “anamurensis”, and “terrestris”) are not Tortoise populations from northern and southern Turkey, which may correspond to the ibera and terrestris mt clades, also appear to be morphometrically distinct. Türkozan et al. (2010) found that “antakyensis”, “anamurensis”, and “terrestris” are not morphometrically distinct, but “armeniaca” and “perses” are.

Synonymy: IVERSON 1992 and SCHLEICH et al. 1996 don’t accept T. nabeulensis, which might be a subspecies or variant of Testudo graeca. Testudo dagestanica has been tentatively placed in the synonymy of T. graeca (PP van Dijk, pers. comm. 18 July 2012).

Subspecies: after TTWG 2014. According to Kuyl et al. (2002) only 4 subspecies are recognized: T. g. graeca of North Africa; T. g. ibera of the Balkans, Greece, Turkey, Iran, and Russia; and T. g. terrestris of Libya, Israel, Egypt, and Syria. Little is known about the fourth subspecies, T. g. zarudnyi, which is restricted to the Central Iranian Plateau and Afghanistan. A morphological study suggested that the first three subspecies should be elevated to full species level (Gmira, 1993). More recent genetic analyses by Kuyl et al. (2005) indicated that only 2 subspecies are clearly distinguishable, T. g. graeca and T. g. ibera. The variety “Golden Greek” is actually Testudo graeca terrestris FORSKÅL 1775.

Note that the paper by KUYL et al. (2002) was later criticized by PERÄLÄ (2004).

Type species: Testudo graeca LINNAEUS 1758: 198 is the type species of the genus Testudo LINNAEUS 1758. 
EtymologyT. graeca has been named after its distribution in Greece (”graeca”). However, it has been also suggested that the species has been named after its markings which resemble Greek letters (Y.L. Werner, pers. comm., 5 Jan 2015).

T. g. lamberti has been named after Michael Lambert (1941-2004), British zoologist and herpetologist. See obituary in Hailey (2005).

The genus is named after Latin testūdō,-inis (feminine) = tortoise 
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