Greek tortoise

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Greek tortoise
Temporal range: Pliocene–Holocene
O
S
D
C
P
T
J
K
Pg
N
Possible Late Miocene record
Tunisian tortoise (Testudo graeca nabeulensis) male Cap Bon.jpg
T. g. nabeulensis
male in Tunisia
Scientific classification OOjs UI icon edit-ltr.svg
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
Areale Testudo graeca.svg
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

The Greek tortoise (Testudo graeca), also known commonly as the spur-thighed tortoise [1] or Moorish tortoise, [3] 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. [4]

Contents

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, [5] but specimens referred to as Testudo cf. graeca are known from the Late and Middle Miocene in Greece and Turkey. [6] [7]

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 tortoiseHermann's tortoise
Large symmetrical markings on the top of the headOnly small scales on the head
Large scales on the front legsSmall scales on the front legs
Undivided carapace over the tailTail carapace almost always divided
Notable spurs on each thighNo spurs
Isolated flecks on the spine and rib platesIsolated flecks only on the spinal plates
Dark central fleck on the undersideTwo black bands on the underside
Shell somewhat oblong rectangularOval shell shape
Widely stretched spinal platesSmall spinal plates
Movable posterior plates on undersideFixed plates on underside
No tail spurTail 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. [8]

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.

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. [3]

Mating and reproduction

A pair of Testudo graeca mating in Mountain Yamanlar Nature Park, Izmir Province, Turkey A pair of Testudo graeca mating.jpg
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. [9] [10] [11] 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. [12]

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. [13]

See also

Related Research Articles

Researched and devised by Dr. Oliphant Jackson, the Jackson ratio is a method of determining whether a member of the tortoise species Testudo graeca or Testudo hermanni is maintaining its optimum bodyweight, which is necessary for a successful hibernation.

<span class="mw-page-title-main">Tortoise</span> Family of turtles

Tortoises are reptiles of the family Testudinidae of the order Testudines. Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like other members of the suborder Cryptodira, they retract their necks and heads directly backward into the shell to protect them.

<span class="mw-page-title-main">Marginated tortoise</span> Species of tortoise

The marginated tortoise is a species of tortoise in the family Testudinidae. The species is endemic to Greece, Italy, and the Balkans in Southern Europe. It is the largest European tortoise. The marginated tortoise is herbivorous, and brumates for the winter.

<span class="mw-page-title-main">Hermann's tortoise</span> Species of tortoise

Hermann's tortoise is a species of tortoise native to Europe.

<span class="mw-page-title-main">Aldabra giant tortoise</span> Species of tortoise

The Aldabra giant tortoise is a species of tortoise in the family Testudinidae. The species is endemic to the Seychelles, with the nominate subspecies, A. g. gigantea native to Aldabra atoll. It is one of the largest tortoises in the world. Historically, giant tortoises were found on many of the western Indian Ocean islands, as well as Madagascar, and the fossil record indicates giant tortoises once occurred on every continent and many islands with the exception of Australia and Antarctica.

<span class="mw-page-title-main">African spurred tortoise</span> Species of tortoise

The African spurred tortoise, also called the sulcata tortoise, is an endangered species of tortoise inhabiting the southern edge of the Sahara Desert, the Sahel, in Africa. It is the largest mainland species of tortoise in Africa, and the third-largest in the world, after the Galapagos tortoise and Aldabra giant tortoise. It is the only living species in its genus, Centrochelys, with the five other species in the family already extinct.

<span class="mw-page-title-main">Russian tortoise</span> Species of tortoise

The Russian tortoise, also commonly known as the Afghan tortoise, the Central Asian tortoise, the four-clawed tortoise, the four-toed tortoise, Horsfield's tortoise, the Russian steppe tortoise, the Soviet Tortoise, and the steppe tortoise, is a threatened species of tortoise in the family Testudinidae. The species is endemic to Central Asia from the Caspian Sea south through Iran, Pakistan and Afghanistan, and east across Kazakhstan to Xinjiang, China. Human activities in its native habitat contribute to its threatened status.

<span class="mw-page-title-main">Leopard tortoise</span> Species of tortoise

The leopard tortoise is a large and attractively marked tortoise found in the savannas of eastern and southern Africa, from Sudan to the southern Cape Province. It is the only extant member of the genus Stigmochelys, although in the past, it was commonly placed in Geochelone. This tortoise is a grazing species that favors semiarid, thorny to grassland habitats. In both very hot and very cold weather, it may dwell in abandoned fox, jackal, or aardvark burrows. The leopard tortoise does not dig other than to make nests in which to lay eggs. Given its propensity for grassland habitats, it grazes extensively upon mixed grasses. It also favors succulents and thistles.

<span class="mw-page-title-main">Red-footed tortoise</span> Species of tortoise

The red-footed tortoise is a species of tortoise from northern South America. These medium-sized tortoises generally average 30 cm (12 in) as adults, but can reach over 40 cm (16 in). They have dark-colored, “loaf”-shaped carapaces with a lighter patch in the middle of each scute, and a somewhat lighter-colored plastron (underbelly). They also have dark limbs dotted with brightly-colored scales, from which they get their name, that range from pale yellow to vivid or dark red. Visible differences are noted between red-footed tortoises from varying eco-regions. They are closely related to the more easterly-distributed yellow-footed tortoise of the Amazon Basin.

<span class="mw-page-title-main">Indian star tortoise</span> Species of tortoise

The Indian star tortoise is a threatened tortoise species native to India, Pakistan and Sri Lanka where it inhabits dry areas and scrub forest. It has been listed as Vulnerable on the IUCN Red List since 2016, as the population is thought to comprise more than 10,000 individuals, but with a declining trend. It is threatened by habitat loss and poaching for the illegal wildlife trade. It was upgraded to CITES Appendix I in 2019 by full consensus among all member states, giving it the highest level of international protection from commercial trade. Conservation group TRAFFIC found 6,040 were seized globally that were intended to be sold in the pet trade.Currently they are commonly bred in many countries to be sold as pets.

<i>Testudo</i> (genus) Genus of tortoises

Testudo, the Mediterranean tortoises, are a genus of tortoises found in North Africa, Western Asia, and Europe. Several species are under threat in the wild, mainly from habitat destruction.

<span class="mw-page-title-main">Wood turtle</span> Species of turtle

The wood turtle is a species of turtle endemic to North America. It is in the genus Glyptemys, a genus which contains only one other species of turtle: the bog turtle. The wood turtle reaches a straight carapace length of 14 to 20 centimeters, its defining characteristic being the pyramidal shape of the scutes on its upper shell. Morphologically, it is similar to the bog turtle, spotted turtle, and Blanding's turtle. The wood turtle exists in a broad geographic range extending from Nova Scotia in the north to Minnesota in the west and Virginia in the south. In the past, it was forced south by encroaching glaciers: skeletal remains have been found as far south as Georgia.

<span class="mw-page-title-main">Tunisian tortoise</span> Subspecies of tortoise

The Tunisian tortoise or Nabeul tortoise is a subspecies of Greek tortoises. It was originally described as a new species in 1990, and even placed in a distinct genus. The spur-thighed or "Greek" tortoises are usually collectively referred to as Testudo graeca, but this covers a wide variety of subspecies that have very different ecological and morphological characteristics and appear to comprise at least three phylogenetic lineages. As its name implies, it is found in Tunisia and nearby Algeria.

<span class="mw-page-title-main">Galápagos tortoise</span> Species of reptile

The Galápagos tortoise or Galápagos giant tortoise is a very large species of tortoise in the genus Chelonoidis. The species comprises 15 subspecies. It is the largest living species of tortoise, and can weigh up to 417 kg (919 lb). They are also the largest extant terrestrial cold-blooded animals (ectotherms).

<span class="mw-page-title-main">Kleinmann's tortoise</span> Species of tortoise

Kleinmann's tortoise, also called commonly the Egyptian tortoise, Leith's tortoise, and the Negev tortoise, is a critically endangered species of cryptodire turtle in the family Testudinidae. The species is native to Egypt, Libya, and Israel. The species was once more widespread, but its numbers are now dwindling. The species is nearly extinct in Egypt, and complete extinction in the wild is a looming threat unless more actions are taken to protect this species.

<span class="mw-page-title-main">Yellow-footed tortoise</span> Species of reptile

The yellow-footed tortoise, also known as the Brazilian giant tortoise, is a species of tortoise in the family Testudinidae and is closely related to the red-footed tortoise. It is found in the Amazon Basin of South America. The species name has often been misspelled as denticulata, an error introduced in the 1980s when Chelonoidis was elevated to genus and mistakenly treated as feminine, an error recognized and fixed in 2017.

<i>Discoglossus pictus</i> Species of amphibian

Discoglossus pictus, the Mediterranean painted frog or simply painted frog, is a species of frog in the family Alytidae.

<span class="mw-page-title-main">Polyandry in animals</span> Class of mating system in non-human species

In behavioral ecology, polyandry is a class of mating system where one female mates with several males in a breeding season. Polyandry is often compared to the polygyny system based on the cost and benefits incurred by members of each sex. Polygyny is where one male mates with several females in a breeding season . A common example of polyandrous mating can be found in the field cricket of the invertebrate order Orthoptera. Polyandrous behavior is also prominent in many other insect species, including the red flour beetle, the adzuki bean weevil, and the species of spider Stegodyphus lineatus. Polyandry also occurs in some primates such as marmosets, mammal groups, the marsupial genus' Antechinus and bandicoots, around 1% of all bird species, such as jacanas and dunnocks, insects such as honeybees, and fish such as pipefish.

<i>Chelonoidis niger guentheri</i> Subspecies of giant tortoise

Chelonoidis niger guentheri, commonly known as the Sierra Negra giant tortoise or Günther's giant tortoise, is a subspecies of Galápagos tortoise endemic to the Galápagos archipelago in the eastern Pacific Ocean. The specific epithet guentheri honours zoologist Albert Günther.

<i>Chelonoidis niger vandenburghi</i> Subspecies of giant tortoise

Chelonoidis niger vandenburghi, also known as the Volcán Alcedo giant tortoise, the Alcedo Volcano giant tortoise or the Alcedo giant tortoise, is a subspecies of Galápagos tortoise endemic to the Galápagos archipelago in the equatorial eastern Pacific Ocean. The specific epithet vandenburghi honours American herpetologist John Van Denburgh.

References

  1. 1 2 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. 1 2 Pritchard, Dr. Peter C. H. (1979). Encyclopedia of Turtles. Neptune, New Jersey: T. F. H. Publications. ISBN   0-87666-918-6.
  4. "Testudo graeca". The Moirai – Aging Research. 12 September 2016. Archived from the original on 13 September 2018. Retrieved 13 December 2016.
  5. 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.
  6. 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 .
  7. 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)
  8. Genus Testudo at The Reptile Database www.reptile-database.org.
  9. 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. Bibcode:2012EcMod.245...19P. doi:10.1016/j.ecolmodel.2012.03.016. hdl: 10261/67281 .
  10. Bergin, Daniel; Nijman, Vincent (2014). "Open, Unregulated Trade in Wildlife in Morocco's Markets, TRAFFIC Bulletin" . Retrieved 23 March 2015.
  11. 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.
  12. 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.
  13. "Helpfull [sic] advice for your tortoise diet". www.tortoisecentre.co.uk. Retrieved 29 January 2018.