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Greek tortoise (''Testudo graeca''), also known as the spur-thighed tortoise or Moorish tortoise, is a species of tortoise in the family
Testudinidae Tortoises ( ) are reptiles of the family Testudinidae of the order Testudines (Latin for "tortoise"). Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like ot ...
. It is a medium sized herbivorous testudinae, widely distributed in the Mediterranean region. ''Testudo graeca'' is recognized for its longevity, with verified lifespans exceeding 100 years and anecdotal reports suggesting ages over 125 years. Among reptiles, it has one of the largest known genomes.


Geographic range

The geographic distribution of the Greek tortoise (''Testudo graeca'') includes
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
,
Southern Europe Southern Europe is also known as Mediterranean Europe, as its geography is marked by the Mediterranean Sea. Definitions of southern Europe include some or all of these countries and regions: Albania, Andorra, Bosnia and Herzegovina, Bulgaria, C ...
, and
Southwest Asia West Asia (also called Western Asia or Southwest Asia) is the westernmost region of Asia. As defined by most academics, UN bodies and other institutions, the subregion consists of Anatolia, the Arabian Peninsula, Iran, Mesopotamia, the Armenia ...
. It is commonly found along the
Black Sea The Black Sea is a marginal sea, marginal Mediterranean sea (oceanography), mediterranean sea lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bound ...
coast of the Caucasus, extending from
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, to
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Georgia Georgia most commonly refers to: * Georgia (country), a country in the South Caucasus * Georgia (U.S. state), a state in the southeastern United States Georgia may also refer to: People and fictional characters * Georgia (name), a list of pe ...
. Additional populations are present in parts of
Georgia Georgia most commonly refers to: * Georgia (country), a country in the South Caucasus * Georgia (U.S. state), a state in the southeastern United States Georgia may also refer to: People and fictional characters * Georgia (name), a list of pe ...
,
Armenia Armenia, officially the Republic of Armenia, is a landlocked country in the Armenian Highlands of West Asia. It is a part of the Caucasus region and is bordered by Turkey to the west, Georgia (country), Georgia to the north and Azerbaijan to ...
,
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, and
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.


Evolution

The oldest confirmed fossil attributed to ''Testudo graeca'' originates from the Early Pliocene of Greece. 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.


Characteristics

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


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: * ''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). Testudo graeca CBNestos.JPG, in Greece Graeca 005.jpg, ''T. g. ibera'' in Turkey Graeca0005.JPG, ''T. g. ibera'', 4 years Tunisian tortoise (Testudo graeca nabeulensis) juvenile Cap Bon.jpg, 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.


Mating and reproduction

Reproductive behavior in ''T. graeca'' begins shortly after emerging from hibernation. 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. 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. 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.


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.


See also

* Mediterranean tortoise *
Timothy (tortoise) Timothy (c. 1844 – 3 April 2004) was a Mediterranean spur-thighed tortoise, estimated to be about 160 years old at the time of her death. This made her the UK's oldest known animal resident. In spite of her name, Timothy was female; gende ...
* Jackson ratio


References


External links


European Tortoises. (in German).''Broasca Testoasa Greaca - Testudo graeca''. (in Romanian).
{{Taxonbar, from=Q504549 Testudo (genus) Reptiles described in 1758
Tortoise Tortoises ( ) are reptiles of the family Testudinidae of the order Testudines (Latin for "tortoise"). Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like o ...
Fauna of Spain Reptiles of Armenia Reptiles of Azerbaijan Turtles of Europe Turtles of Asia Species that are or were threatened by the pet trade Taxa named by Carl Linnaeus Reptiles of Russia Reptiles as pets Habitats Directive species