Greek tortoise

The Greek tortoise (Testudo graeca), also known commonly as the spur-thighed tortoise or Moorish tortoise, 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. It has the largest known genome of all reptiles.

Greek tortoise Temporal range: Possible Late Miocene record
T. g. nabeulensis male in Tunisia
Conservation status
Vulnerable  (IUCN 3.1)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Testudines
Suborder:	Cryptodira
Superfamily:	Testudinoidea
Family:	Testudinidae
Genus:	Testudo
Species:	T. graeca
Binomial name
Testudo graeca Linnaeus, 1758
Note allopatric ranges of "Maghreb" (T. g. graeca) and "Greek" (T. g. ibera) populations
Synonyms
List 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

Geographic range

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.

Evolution

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

Characteristics

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

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 supracaudal scute over the tail	Supracaudal scute 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

Subspecies

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.

• 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, 1990 – Tunisian tortoise, Tunisia
• T. g. cyrenaica Pieh & Perälä, 2002 – Libya
• T. g. ibera Pallas, 1814 – Turkey
• T. g. armeniaca Chkhikvadze & Bakradse, 1991 – Armenian 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.

• 
  in Greece
• 
  T. g. ibera in Turkey
• 
  T. g. ibera, 4 years
• 
  juvenile T. g. nabeulensis in Tunisia

Sexing

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.

Behavior

Hibernation

Testudo graeca hibernates during cold months, emerging as early as February in hot coastal areas. Individual tortoises may emerge during warm days even during winter.

Mating and reproduction

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.

Trade

The Greek tortoise is commonly traded as a pet in source countries such as Morocco and Spain, despite the illegality of this trade. 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.

Food

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.

See also

• Mediterranean tortoise
• Timothy (tortoise)
• Jackson ratio

External links

• European Tortoises. (in German).
• Broasca Testoasa Greaca - Testudo graeca. (in Romanian).