Greek tortoise

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Greek tortoise
Temporal range: Pliocene–Holocene Possible Late Miocene record
In Greece
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Testudines
Suborder: Cryptodira
Superfamily: Testudinoidea
Family: Testudinidae
Genus: Testudo
T. graeca
Binomial name
Testudo graeca
Note allopatric ranges of "Maghreb" (T. g. graeca) and "Greek" (T. g. ibera) populations
  • T. g. graeca
  • Testudo graeca
    Linnaeus, 1758
  • Testudo pusilla
    Linnaeus, 1758
  • Chersine pusilla
    Merrem, 1820
  • Testudo mauritanica
    A.M.C. Duméril & Bibron, 1835
  • Testudo mauritonica
    Kercado, 1835 (ex errore)
  • Testudo whitei
    Bennett, 1836
  • Peltastes mauritanicus
    Gray, 1873
  • Testudo graeca graeca
    Mertens, 1946
  • Testudo gracea
    Nutaphand, 1979 (ex errore)
  • Testudo whitie
    Highfield & Martin, 1989 (ex errore)
  • Furculachelys whitei
    — Highfield, 1990
  • Testudo graeca sarda
    Ballasina, 1995 (nomen nudum)
  • Testudo graeca whitei
    — Artner, 1996
  • T. g. anamurensis
  • Testudo graeca anamurensis
    Weissinger, 1987
  • Testudo ibera anamurensis
    — Highfield, 1990
  • Testudo terrestris anamurensis
    — David, 1994
  • Testudo anamurensis
    — Vetter, 2002
  • Testudo graeca amurensis
    Ferri, 2002 (ex errore)
  • T. g. antakyensis
  • Testudo antakyensis
    Perälä, 1996
  • Testudo graeca antakyensis
    — Zwartepoorte, 2000
  • Testudo terrestris antakyensis
    — Bour, 2002
  • Testudo ibera antakyensis
    — Artner, 2003
  • T. g. armeniaca
  • Testudo graeca armeniaca
    Chkhikvadze, 1989
    (nomen nudum)
  • Testudo graeca armeniaca
    Chkhikvadze & Bakradze, 1991
  • Testudo graeca armaniaca
    Chkhikvadze & Bakradze, 1991
    (ex errore)
  • Testudo armeniaca
    — Vetter, 2002
  • Testudo terrestris armeniaca
    — Bour, 2002
  • T. g. buxtoni
  • ? Testudo ecaudata
    Pallas, 1814
  • Testudo buxtoni
    Boulenger, 1921
  • Testudo terrestris buxtoni
    — Bour, 2002
  • Testudo ibera buxtoni
    — Artner, 2003
  • ; T. g. cyrenaica
  • Testudo graeca cyrenaica
    Pieh & Perälä, 2002
  • Testudo cyrenaica
    — Vetter, 2002
  • Testudo cyrenaika
    Stettner, 2004 (ex errore)
  • T. g. floweri
  • Testudo floweri
    Bodenheimer, 1935
  • Testudo graeca floweri
    — Mertens, 1946
  • Testudo terrestris floweri
    — David, 1994
  • Testudo ibera floweri
    — Artner, 2003
  • T. g. ibera
  • Testudo ibera
    Pallas, 1814
  • Chersus iberus
    — Brandt, 1852
  • Testudo iberia
    Blyth, 1853 (ex errore)
  • Medaestia ibera
    — Wussow, 1916
  • Testudo ibera racovitzai
    Călinescu, 1931
  • Testudo graeca ibera
    — Mertens, 1946
  • Testudo ibera ibera
    — Gmira, 1993
  • Testudo terrestris ibera
    — David, 1994
  • T. g. lamberti
  • Testudo graeca lamberti
    Pieh & Perälä, 2004
  • Testudo lamberti
    — Perälä, 2004
  • T. g. marokkensis
  • Testudo graeca marokkensis
    Pieh & Perälä, 2004
  • Testudo marokkensis
    — Perälä, 2004
  • T. g. nabeulensis
  • ? Testudo flavominimaralis
    Highfield & Martin, 1989
  • Furculachelys nabeulensis
    Highfield, 1990
  • Testudo nabeulensis
    — Welch, 1994
  • ? Testudo graeca flavominimaralis
    — Artner, 1996
  • Testudo graeca nabeulensis
    — Artner, 1996
  • T. g. nikolskii
  • Testudo graeca nikolskii
    Chkhikvadze & Tuniyev, 1986
  • Testudo ibera nikolskii
    — Highfield, 1990
  • Testudo terrestris nikolskii
    — David, 1994
  • Testudo graeca niiolskii
    Paull, 1997 (ex errore)
  • Testudo nikolskii
    — Vetter, 2002
  • T. g. pallasi
  • Testudo graeca pallasi
    Chkhikvadze, 1989
    (nomen nudum)
  • Testudo graeca pallasi
    Chkhikvadze & Bakradze, 2002
  • Testudo pallasi
    — Danilov & Milto, 2004
  • T. g. perses
  • Testudo perses
    Perälä, 2002
  • Testudo ibera perses
    — Artner, 2003
  • T. g. soussensis
  • Testudo graeca soussensis
    Pieh, 2001
  • Testudo soussensis
    — Vetter, 2002
  • T. g. terrestris
  • Testudo terrestris
    Forsskål, 1775
  • ? Testudo zolhafa
    Forsskål, 1831 (nomen nudum)
  • ? Testudo zolkafa
    Forsskål, 1831 (nomen nudum)
  • ? Testudo zohalfa
    Forsskål, 1835 (nomen nudum)
  • Testudo graeca terrestris
    — Wermuth, 1958
  • Testudo terrestris terrestris
    — David, 1994
  • Testudo ibera terrestris
    — Artner, 2003
  • T. g. zarudnyi
  • Testudo zarudnyi
    Nikolsky, 1896
  • Testudo graeca zarudnyi
    — Mertens, 1946
  • Testudo ibera zarudnyi
    — Gmira, 1993
  • Testudo terrestris zarudnyi
    — David, 1994

The Greek tortoise (Testudo graeca), also known commonly as the spur-thighed tortoise,[1] is a species of tortoise in the family Testudinidae. Testudo graeca is one of five species of Mediterranean tortoises (genera Testudo and Agrionemys). The other four species are Hermann's tortoise (T. hermanni), the Egyptian tortoise (T. kleinmanni), the marginated tortoise (T. marginata), and the Russian tortoise (A. horsfieldii). The Greek tortoise is a very long-lived animal, achieving a lifespan upwards of 125 years, with some unverified reports up to 200 years.[3]

Geographic range[edit]

The Greek tortoise's geographic range includes North Africa, Southern Europe, and Southwest Asia. It is prevalent in the Black Sea coast of the Caucasus (from Anapa, Russia, to Sukhumi, Abkhazia, Georgia, to the south), as well as in other regions of Georgia, Armenia, Iran, and Azerbaijan.


The oldest known definitive fossil is from the Early Pliocene of Greece,[4] but specimens referred to as Testudo cf. graeca are known from the Late and Middle Miocene in Greece and Turkey.[5][6]


The Greek tortoise (T. g. ibera) is often confused with Hermann's tortoise (T. hermanni ). However, notable differences enable them to be distinguished.

T. g. ibera
T. graeca, male
Greek tortoise Hermann's tortoise
Large symmetrical markings on the top of the head Only small scales on the head
Large scales on the front legs Small scales on the front legs
Undivided carapace over the tail Tail carapace almost always divided
Notable spurs on each thigh No spurs
Isolated flecks on the spine and rib plates Isolated flecks only on the spinal plates
Dark central fleck on the underside Two black bands on the underside
Shell somewhat oblong rectangular Oval shell shape
Widely stretched spinal plates Small spinal plates
Movable posterior plates on underside Fixed plates on underside
No tail spur Tail bears a spur at the tip


Testudo graeca, 4 years

The division of the Greek tortoise into subspecies is difficult and confusing. Given its huge range over three continents, the various terrains, climates, and biotopes have produced a huge number of varieties, with new subspecies constantly being discovered. As of 2023, at least 20 subspecies have been published, of which the following 12 are recognized as being valid.[7]

  • T. g. graeca Linnaeus, 1758 – northern Africa, southern Spain
  • T. g. soussensis Pieh, 2000 – southern Morocco
  • T. g. marokkensis Pieh & Perälä, 2004 – northern Morocco
  • T. g. nabeulensis Highfield, 1990Tunisian tortoise, Tunisia
  • T. g. cyrenaica Pieh & Perälä, 2002 – Libya
  • T. g. ibera Pallas, 1814 – Turkey
  • T. g. armeniaca Chkhikvadze & Bakradse, 1991Armenian tortoise, Armenia
  • T. g. buxtoni Boulenger, 1921 – Caspian Sea
  • T. g. terrestris Forskål, 1775 – Israel, Jordan, Lebanon
  • T. g. zarudnyi Nikolsky, 1896 – Azerbaijan, Iran
  • T. g. whitei Bennett in White, 1836 – Algeria
  • T. g. perses Perälä, 2002 – Turkey, Iran, Iraq

This incomplete listing shows the problems in the division of the species into subspecies. The differences in form are primarily in size and weight, as well as coloration, which ranges from dark brown to bright yellow, and the types of flecks, ranging from solid colors to many spots. Also, the bending-up of the edges of their carapaces ranges from minimal to pronounced. So as not to become lost in the number of subspecies, recently, a few tortoises previously classified as T. graeca have been assigned to different species, or even different genera.

The genetic richness of T. graeca is also shown in its crossbreeding. Tortoises of different form groups often mate, producing offspring with widely differing shapes and color. Perhaps the best means of identification for the future is simply the place of origin.

The smallest, and perhaps the prettiest, of the subspecies, is the Tunisian tortoise. It has a particularly bright and striking coloration. However, these are also the most sensitive tortoises of the species, so they cannot be kept outdoors in temperate climates, as cold and rainy summers quickly cause the animals to become ill. They are also incapable of long hibernation.

At the other extreme, animals from northeastern Turkey are very robust, such as Hermann's tortoise. The largest specimens come from Bulgaria. Specimens of 7 kg (15 lb) have been reported. In comparison, the Tunisian tortoise has a maximum weight of 0.7 kg (1.5 lb). T. graeca is also closely related to the marginated tortoise (T. marginata). The two species can interbreed, producing offspring capable of reproduction.


Males of T. graeca differ from females in six main points. Firstly, they are generally smaller. Their tails are longer than females and taper to a point evenly, and the cloacal opening is farther from the base of the tail. The underside is somewhat curved, while females have a flat shell on the underside. The rear portion of a male's carapace is wider than it is long. Finally, the posterior plates of the carapace often flange outward.

Mating and reproduction[edit]

A pair of Testudo graeca mating in Mountain Yamanlar Nature Park, İzmir Province, Turkey

In T. graeca, immediately after waking from hibernation, the mating instinct starts up. The males follow the females with great interest, encircling them, biting them in the limbs, ramming them, and trying to mount them. During copulation, the male opens his mouth, showing his red tongue and making squeaking sounds.

During mating, the female stays still, bracing herself with her front legs, moving the front part of her body to the left and right in the same rhythm as the male's cries. One successful mating will allow the female to lay eggs multiple times. When breeding in captivity, the pairs of females and males must be kept separate. If multiple males are in a pen, one takes on a dominant role and will try to mate with the other males in the pen. If more males than females are in a pen, the males might kill each other to mate with the females.

One or two weeks before egg laying, the animals become notably agitated, moving around to smell and dig in the soil, even tasting it, before choosing the ideal spot to lay the eggs. One or two days before egg laying, the female takes on an aggressive, dominant behavior, mounting another animal as for copulation and making the same squeaking sound the male produces during copulation. The purpose of this behavior is to produce respect in the tortoise community so that the female will not be disturbed by the others during egg laying. Further details of egg-laying behavior are the same as those detailed for the marginated tortoise.


The Greek tortoise is commonly traded as a pet in source countries such as Morocco and Spain, despite the illegality of this trade.[8][9][10] This can lead to an unsustainable removal of wild individuals for the local pet trade and for export. Also, welfare concerns exist with this trade, as the animals are not properly housed when being sold, causing a high rate of mortality in captivity.[11]


In captivity, the Greek tortoise loves dandelion leaves and other leafy plants. However, although they also enjoy eating lettuce, it is not recommended to them due to having a lack of nutrients that the tortoises need to survive.[12]

See also[edit]


  1. ^ a b Tortoise.; Freshwater Turtle Specialist Group (1996). "Testudo graeca". IUCN Red List of Threatened Species. 1996: e.T21646A9305693. doi:10.2305/IUCN.UK.1996.RLTS.T21646A9305693.en. Retrieved 19 November 2021.
  2. ^ Fritz, Uwe; Havaš, Peter (2007). "Checklist of Chelonians of the World" (PDF). Vertebrate Zoology. 57 (2): 296–300. doi:10.3897/vz.57.e30895. ISSN 1864-5755. S2CID 87809001. Archived (PDF) from the original on 1 May 2011. Retrieved 29 May 2012.
  3. ^ "Testudo graeca". The Moirai – Aging Research. 12 September 2016. Archived from the original on 13 September 2018. Retrieved 13 December 2016.
  4. ^ Vlachos E (2015). "The Fossil Chelonians of Greece. Systematics – Evolution – Stratigraphy – Palaeoecology". Scientific Annals of the School of Geology, Aristotle University of Thessaloniki, Greece. 173: 1–479.
  5. ^ Vlachos E, Tsoukala E (2014). "Testudo cf. graeca from the new Late Miocene locality of Platania (Drama basin, N. Greece) and a reappraisal of previously published specimens". Bulletin of the Geological Society of Greece. 48: 27–40. doi:10.12681/bgsg.11046.
  6. ^ Staesche K, Karl HV, Staesche U (2007). "Fossile Schildkroten aus der Turkei". Fossile Schildkroten aus Drei Kontinenten. 98: 91–149.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  7. ^ Genus Testudo at The Reptile Database
  8. ^ Pérez, Irene; Tenza, Alicia; Anadón, José Daniel; Martínez-Fernández, Julia; Pedreño, Andrés; Giménez, Andrés (2012). "Exurban sprawl increases the extinction probability of a threatened tortoise due to pet collections". Ecological Modelling. 245: 19–30. doi:10.1016/j.ecolmodel.2012.03.016. hdl:10261/67281.
  9. ^ Bergin, Daniel; Nijman, Vincent (2014). "Open, Unregulated Trade in Wildlife in Morocco's Markets, TRAFFIC Bulletin". Retrieved 23 March 2015.
  10. ^ Nijman, V; Bergin, D (2017). "Trade in spur-Thighed tortoises Testudo graeca in Morocco: Volumes, value and variation between markets". Amphibia-Reptilia. 38 (3): 275–287. doi:10.1163/15685381-00003109.
  11. ^ Bergin, D.; Nijman, V. (2018). "An Assessment of Welfare Conditions in Wildlife Markets across Morocco". Journal of Applied Animal Welfare Science. 22 (3): 279–288. doi:10.1080/10888705.2018.1492408. PMID 30102072. S2CID 51967901.
  12. ^ "Helpfull [sic] advice for your tortoise diet". Retrieved 29 January 2018.

External links[edit]