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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 zarudnyi NIKOLSKY 1896 
Common NamesE: Mediterranean Spur-thighed Tortoise
G: Maurische Landschildkröte
Russian: Средиземноморская черепаха 
SynonymTestudo graeca LINNAEUS 1758: 198
Testudo pusilla LINNAEUS 1758: 198
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 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
Testudo whitei
Tistudo mauritonica — KERCADO 1835 (error typographicus)
Testudo graeca graeca — MERTENS 1946
Testudo graeca graeca — BONIN et al 2006
Testudo graeca graeca — PIEH & PHILIPPEN 2007
Testudo graeca graeca — HERZ 2013
Testudo graeca whitei — HERZ 2013
Testudo (Testudo) graeca graeca — TTWG 2017: 152

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 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 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 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 flavominimaralis HIGHFIELD & MARTIN 1990 (fide KUYL et al. 2002)
Testudo terrestris ibera — DAVID 1994
Testudo graeca ibera — BISCHOFF 2002
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 flavominimaralis — HERZ 2013
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 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 graeca nabeulensis (HIGHFIELD 1990)
Furculachelys nabeulensis HIGHFIELD 1990: 32
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 nabeulensis — HERZ 2013
Testudo (Testudo) graeca — TTWG 2017: 153

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 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 
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

anamurensis: Cape Anamur and SW coast of Turkey.

antakyensis: S Turkey

cyrenaica: Libya (endemic)

flavominimaralis: Tunisia

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

ibera: S Serbia, Greece (Lesbos, Samos), Romania through Iran.

lamberti: NW Morocco

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

nikolskii: Republic of Georgia, Russia (western Caucasus)

pallasi: Russia (Daghestan)

perses: SE Turkey, NE Iraq, SC Iran (Zagros mountains)

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

terrestris: Israel, Lebanon, Jordan, Syria, SE Turkey.

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

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).  
ReproductionT. graeca has been reported to have temperature-dependent sex determination (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] 
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.).

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.

PERÄLÄ (2002) described T. perses as follows: “T. perses is distinguished from all other taxa in the genus Testudo sensu stricto (in the sense of Gmira, 1995; Lapparent de Broin, 2000a, 2000b, 2001; Hervet, 2001; Perälä, 2002 in this volume), by the following combination of characters subject to individual variation and pertaining to adults and both sexes unless stated otherwise: Robustly built tortoise with a xiphiplastral hinge (showing xiphiplastral kinesis), ventrally short diagonal length of first costal, short fifth vertebral, dorsally wide supracaudal, very broad fourth vertebral and, with extremely long and wide anal scutes; anal midline seam (interanal) typically much longer than seam separating femorals; transverse seam separating humerals and femorals typically double-S-shaped, with or without a laterally straight proximal section; broad first vertebral; first vertebral almost always widest and sharp-edged anteriorly and, frequently elongated anteriorly in its proximal parts (modestly infundibular); fifth vertebral ventrally narrow; midbody (at bridge) of medium width relative to length, and rounded in dorsal view; in lateral view, outline of carapace mostly ventrally indented at anterior portion of first vertebral and, posteriormost marginals covering part of supracaudal (lateral outline of semi-adults can resemble that of T. kleinmanni); outline of carapace relatively angulate in anterior view, with perpendicular outermost (upper) surfaces of bridge marginals (i.e. lateral surfaces above the marginal crest which is well developed), edged upper arch (at costals) and relatively flat top with a slightly elevated proximal portion (central vertebral area); in posterior view the lower outline has a bell-shaped curvature due to marginal flaring, especially in males; carapace highest at slightly less than 2/3 length; supracaudal slightly flared to the rear, wide and almost box-shaped (rectangular) in females except for a vertically thin to more pronounced, laterally oriented and spiky offshoots towards the marginals in ventral corners, and with a more or less 90-100 degree inner angle between dorsolateral and dorsal sides; supracaudal of males more massive, usually flared to the rear (only rarely inverted) and also with ventrolateral offshoots, the inner angle between dorsolateral and dorsal sides being in the region of 90-120° and, with proximally elongated, smoothly U-shaped ventral extension becoming less apparent and more V-shaped towards the southeast of the range; outermost lateral extensions of bridge marginals (the crest separating dorsal and ventral portions of the scutes) typically in a 12-17 degree angle with respect to horizontal level, the open angle oriented cranially; inguinals relatively large, usually longer than broad, typically not touching femorals, and associated with smaller accessory scales (which may touch femorals); anterior and posterior free marginals flared to strongly flared, and serrated, convexly-rimmed towards the outside, and typically without much reversion; anteriormost (1st) marginals elongated, often protruding infront of second marginals; posteriormost (11th) marginals often extending posteriorly beyond supracaudal; lateral flaring of posterior marginals begins at, and is inclusive of, 8th marginal; 8th marginal featuring an anteriorly skewed, ventrally oriented half-dome to pyramid-shaped bulge with a more or less rounded apex and outer surface; greatest width at 8th or (more often) 9th marginals; anterior and posterior free marginals may extend beyond bridge marginals in dorsolateral (and ventrolateral) view, more so in males than in females; anterior plastron lobe ventrally convex in lateral view; nuchal scute (cervical) long and relatively narrow, and widest posteriorly or at around 3/4 length; gulars clearly set off from anterior lobe, and relatively long both ventrally and dorsally; length of bridge very variable, and not typically relatively shorter in males than in females (as opposed to most Testudo s. s.); anterior feet relatively massive and thick, featuring relatively long, almost drop-shaped, triangularish to four-edged scales with longest lateral sides facing claws, closely packed against the surface, and with generally not much overlap between scales; fore-feet with five relatively straight claws, the innermost of which are shorter than the rest, posterior extremities with four claws; tail relatively long, longer in males than in females, and very slender towards the tip; tip of tail features mostly a conspicuously enlarged, medium-sized twin-scale (not claw); upper thigh region typically with (at least) one horny spur on each thigh; thigh-spurs very broad at base; outline of open eyes in life more round than oval; bony carapace rim proximally convex towards the anterior; triturating surfaces of maxilla featuring only one longitudinal, modestly elevated central ridge; parietal bones straight or just very slightly notched in lateral view; supraoccipital crest (parietal-supraoccipital complex) long.

Basic coloration (in alcohol): Ground colour of carapace and plastron from pale greenish-yellow (USNM 154515; subadult), to light or very dark-brown colour with a reddish or orange tinge (typical), or even completely grey or black. Plastron is usually not much darker than carapace. The yellowish variety “Golden Greek” is T. g. terrestris FORSKÅL 1775.

Markings: Small juveniles (post-hatchlings) with a transverse dark band located centrally on 1st vertebral; 2nd to 5th vertebrals with small, more or less round central blotches with thin lateral offshoots of dark pigment not connected with the blotch. In lateral view, 1st costal features a horizontal dark stripe located at 1/3 height and running parallel to horizontal level; in 2nd and 3rd costals this band is also at 1/3 height, but it runs in a slight angle so that the posterior end is located highest. 4th costal has a central blotch. 2nd to 4th costals feature additionally centrally located vertical stripes at the anteriormost seams. Plastral markings may be completely absent, or there are tiny black blotches on the lower central region of the abdominals. Anterior marginals may feature dark semi-triangular markings. Bridge marginals with dark bands along anterior vertical borders. Posterior marginals with anterior and dorsal bands connected dorsolaterally. Larger juveniles and semi-adults: The central blotch on 1st vertebral divides laterally into two sister flecks. Costals may retain vertical bands near the anteriormost side of areolae, horizontal bands fade away or develope into larger, irregular blotches. Separately from these, extremely thin centrally oriented stripes appear at anterior and lateral, and anterior and ventrolateral borders of vertebrals and costals. These may develope into dark bands. The supracaudal may feature triangular lateral markings. These may later fade away, especially in the dorsal border region. Markings on marginals become darker and more defined. The plastron is irregularly pigmented with dark grey to blackish elements, darkest regions being located posterolaterally. Gulars and anals may be lighter coloured than other scutes. In adults, regions on and adjacent to the areolae seem to feature most pigmentation, dark blue-grey to black blotches with irregular borders. Markings around scute edges diffuse into centrally oriented rays which may or may not be connected to the central flecks. Dark blotches may appear on posterior ventral corners of (especially) the bridge marginals, or around the marginal crest. Many adult specimens are completely dark: grey, intensively dark-brown or black, with hardly any markings visible. Basic coloration of head and leg scutes varies from greenish-grey and grey to (as typical) light or deepish-dark chocolate-brown. The ground colour on most parts of the head is either uniform or it may feature blackish regions especially in the snout area, and darker brown to black, or, occasionally found yellowish scales towards the temple. Completely black heads are a rarity. The jaw region features often a dark grey to blackish triangular pattern, or a "Schnautzendreieck" sensu Eiselt & Spitzenberger (1967). Leg scutes have mostly a dark-brown, dark-violet to blackish tip, also on the hind legs. The base of leg scutes may be pale brown, pale grey or yellowish. Claws are light or dark chocolate-brown to dark violet and dark grey. Skin in soft parts is pale greyish, light brown or reddish-brown coloured. Eyes in life are very dark brown.”

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). 
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