Greek tortoise

Greek tortoise (Testudo graeca), also known as the spur-thighed tortoise or Moorish tortoise, is a species of tortoise in the family Testudinidae. It is a medium sized herbivorous testudinae, widely distributed in the Mediterranean region.^[3][4]

Greek tortoise Temporal range: Pliocene–Holocene PreꞒ Ꞓ O S D C P T J K Pg N Possible Late Miocene record
T. g. nabeulensis male in Tunisia
Conservation status
Vulnerable  (IUCN 3.1)[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[2]
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

Testudo graeca is recognized for its longevity, with verified lifespans exceeding 100 years and anecdotal reports suggesting ages over 125 years.^[5] Among reptiles, it has one of the largest known genomes.^[6]

Geographic range

The geographic distribution of the Greek tortoise (Testudo graeca) includes North Africa, Southern Europe, and Southwest Asia. It is commonly found along the Black Sea coast of the Caucasus, extending from Anapa, Russia, to Sukhumi, Abkhazia, Georgia. Additional populations are present in parts of Georgia, Armenia, Iran, and Azerbaijan.^[7]

Evolution

The oldest confirmed fossil attributed to Testudo graeca originates from the Early Pliocene of Greece.^[8] However, fossils tentatively identified as Testudo cf. graeca have also been reported from the Middle and Late Miocene of both Greece and Turkey, suggesting a more ancient and geographically diverse origin.^[9][10]

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 classification of the Greek tortoise (Testudo graeca) into subspecies is complex and sometimes inconsistent due to its extensive distribution across North Africa, Southern Europe, and Southwest Asia. Diverse environmental conditions across this range have resulted in a wide array of morphological variations. As of 2023, at least 20 subspecies have been described, with the following 12 currently recognized as valid:^[11]

• 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 – Tunisia
• T. g. cyrenaica Pieh & Perälä, 2002 – Libya
• T. g. ibera Pallas, 1814 – Turkey
• T. g. armeniaca Chkhikvadze & Bakradse, 1991 – Armenia
• T. g. buxtoni Boulenger, 1921 – Caspian Sea region
• 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

The recognition and delimitation of these subspecies are challenging due to overlapping morphological traits such as body size, shell shape, color patterns, and the degree of curvature at the carapace edges. Some populations formerly assigned to T. graeca have since been reassigned to different species or genera.

Genetic diversity within T. graeca is further demonstrated by interbreeding between geographically distinct populations, resulting in variable offspring. For this reason, geographical origin is often considered the most reliable method of identification.

Among the most distinctive subspecies is the Tunisian tortoise (T. g. nabeulensis), noted for its bright coloration and small size. However, it is also one of the most sensitive, poorly suited for outdoor enclosures in temperate climates, and incapable of prolonged hibernation.

Populations from northeastern Turkey are notably robust. Some of the largest individuals, weighing up to 7 kg (15 lb).

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

Sexing

Males of T. graeca exhibit several distinct physical characteristics that differentiate them from females. They are typically smaller in size and possess longer tails that taper evenly to a point. The male's cloacal opening is situated farther from the base of the tail. On the plastron, or underside of the shell, males show a slight concavity, which aids in mounting during mating, whereas females have a flat plastron. Additionally, the posterior portion of a male’s carapace is usually broader than its length, and the rear marginal scutes often curve 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.^[12]

Mating and reproduction

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

Reproductive behavior in T. graeca begins shortly after emerging from hibernation.^[13] Males actively pursue females, displaying courtship behaviors such as circling, biting at the limbs, ramming, and mounting attempts. During copulation, males emit squeaking sounds and display a red tongue by opening their mouths.

Females generally remain still during copulation, bracing with their front legs and moving rhythmically in response to the male’s actions. A single successful mating can result in multiple clutches of eggs. In captivity, males and females are often kept separate to avoid aggression. If multiple males are housed together, dominant behavior may occur, including attempts to mount other males. An imbalanced male-to-female ratio can lead to serious aggression and injury.^[14]

Prior to oviposition, females become noticeably restless, engaging in behaviors such as sniffing and digging to locate suitable nesting sites. In the final days before laying, females may display dominant behavior, such as mock mounting and vocalizations. This behavior may help establish social dominance and ensure minimal disturbance during egg laying. The specifics of oviposition resemble those observed in related species like the marginated tortoise.

Trade

The Greek tortoise (Testudo graeca) is frequently traded as a pet, particularly in source countries such as Morocco and Spain, despite existing legal restrictions on the trade of wild-caught individuals.^[15][16][17]

This practice poses a conservation risk, as it may contribute to unsustainable removal of individuals from wild populations for both local sale and international export. Furthermore, concerns have been raised regarding the welfare conditions under which the tortoises are kept and transported, with reports of inadequate housing and care leading to high mortality rates in captivity.^[18][19]

Food

In captivity, Greek tortoises (Testudo graeca) commonly consume a variety of leafy greens, with a particular preference for dandelion leaves and similar vegetation. While they may readily eat lettuce, it is generally not recommended as a staple food, as it lacks the essential nutrients required to support their long-term health and survival.^[20][21]

See also

• Mediterranean tortoise
• Timothy (tortoise)
• Jackson ratio