Greek tortoise

{{Multiple issues|

}}

{{Speciesbox

| fossil_range = {{fossilrange|Pliocene|Holocene|earliest=late miocene}} Possible Late Miocene record

| image = Tunisian tortoise (Testudo graeca nabeulensis) male Cap Bon.jpg

| image_caption = T. g. nabeulensis
male in Tunisia

| status = VU

| status_system = IUCN3.1

| status_ref = {{cite iucn |author=Tortoise. |author2=Freshwater Turtle Specialist Group |date=1996 |title=Testudo graeca |volume=1996 |page=e.T21646A9305693 |doi=10.2305/IUCN.UK.1996.RLTS.T21646A9305693.en |access-date=19 November 2021}}

| genus = Testudo

| species = graeca

| authority = Linnaeus, 1758

| range_map = Areale Testudo graeca.svg

| range_map_caption = Note allopatric ranges of "Maghreb" (T. g. graeca) and "Greek" (T. g. ibera) populations

| synonyms_ref = {{Cite journal|journal=Vertebrate Zoology|title=Checklist of Chelonians of the World|year=2007|last1=Fritz|first1=Uwe|author1-link=species:Uwe Fritz|last2=Havaš|first2=Peter|author2-link=species:Peter Havaš|volume=57|issue=2|pages=296–300|doi=10.3897/vz.57.e30895 |s2cid=87809001 |issn=1864-5755|url=http://www.cnah.org/pdf_files/851.pdf|archive-url=https://web.archive.org/web/20110501060224/http://www.cnah.org/pdf_files/851.pdf|archive-date=1 May 2011|access-date=29 May 2012|url-status=live |doi-access=free }}

| synonyms = {{collapsible list|bullets = true

| T. g. graeca

|Testudo graeca
{{small|Linnaeus, 1758}}

|Testudo pusilla
{{small|Linnaeus, 1758}}

|Chersine pusilla
{{small|— Merrem, 1820}}

|Testudo mauritanica
{{small|A.M.C. Duméril & Bibron, 1835}}

|Testudo mauritonica
{{small|Kercado, 1835}} (ex errore)

|Testudo whitei
{{small|Bennett, 1836}}

|Peltastes mauritanicus
{{small|— Gray, 1873}}

|Testudo graeca graeca
{{small|— Mertens, 1946}}

|Testudo gracea
{{small|Nutaphand, 1979}} (ex errore)

|Testudo whitie
{{small|Highfield & Martin, 1989}} (ex errore)

|Furculachelys whitei
{{small|— Highfield, 1990}}

|Testudo graeca sarda
{{small|Ballasina, 1995}} (nomen nudum)

|Testudo graeca whitei
{{small|— Artner, 1996}}

| T. g. anamurensis

|Testudo graeca anamurensis
{{small|Weissinger, 1987}}

|Testudo ibera anamurensis
{{small|— Highfield, 1990}}

|Testudo terrestris anamurensis
{{small|— David, 1994}}

|Testudo anamurensis
{{small|— Vetter, 2002}}

|Testudo graeca amurensis
{{small|Ferri, 2002}} (ex errore)

| T. g. antakyensis

|Testudo antakyensis
{{small|Perälä, 1996}}

|Testudo graeca antakyensis
{{small|— Zwartepoorte, 2000}}

|Testudo terrestris antakyensis
{{small|— Bour, 2002}}

|Testudo ibera antakyensis
{{small|— Artner, 2003}}

| T. g. armeniaca

|Testudo graeca armeniaca
{{small|Chkhikvadze, 1989}}
(nomen nudum)

|Testudo graeca armeniaca
{{small|Chkhikvadze & Bakradze, 1991}}

|Testudo graeca armaniaca
{{small|Chkhikvadze & Bakradze, 1991}}
(ex errore)

|Testudo armeniaca
{{small|— Vetter, 2002}}

|Testudo terrestris armeniaca
{{small|— Bour, 2002}}

| T. g. buxtoni

|? Testudo ecaudata
{{small|Pallas, 1814}}

|Testudo buxtoni
{{small|Boulenger, 1921}}

|Testudo terrestris buxtoni
{{small|— Bour, 2002}}

|Testudo ibera buxtoni
{{small|— Artner, 2003}}

|; T. g. cyrenaica

|Testudo graeca cyrenaica
{{small|Pieh & Perälä, 2002}}

|Testudo cyrenaica
{{small|— Vetter, 2002}}

|Testudo cyrenaika
{{small|Stettner, 2004}} (ex errore)

| T. g. floweri

|Testudo floweri
{{small|Bodenheimer, 1935}}

|Testudo graeca floweri
{{small|— Mertens, 1946}}

|Testudo terrestris floweri
{{small|— David, 1994}}

|Testudo ibera floweri
{{small|— Artner, 2003}}

| T. g. ibera

|Testudo ibera
{{small|Pallas, 1814}}

|Chersus iberus
{{small|— Brandt, 1852}}

|Testudo iberia
{{small|Blyth, 1853}} (ex errore)

|Medaestia ibera
{{small|— Wussow, 1916}}

|Testudo ibera racovitzai
{{small|Călinescu, 1931}}

|Testudo graeca ibera
{{small|— Mertens, 1946}}

|Testudo ibera ibera
{{small|— Gmira, 1993}}

|Testudo terrestris ibera
{{small|— David, 1994}}

| T. g. lamberti

|Testudo graeca lamberti
{{small|Pieh & Perälä, 2004}}

|Testudo lamberti
{{small|— Perälä, 2004}}

| T. g. marokkensis

|Testudo graeca marokkensis
{{small|Pieh & Perälä, 2004}}

|Testudo marokkensis
{{small|— Perälä, 2004}}

| T. g. nabeulensis

|? Testudo flavominimaralis
{{small|Highfield & Martin, 1989}}

|Furculachelys nabeulensis
{{small|Highfield, 1990}}

|Testudo nabeulensis
{{small|— Welch, 1994}}

|? Testudo graeca flavominimaralis
{{small|— Artner, 1996}}

|Testudo graeca nabeulensis
{{small|— Artner, 1996}}

| T. g. nikolskii

|Testudo graeca nikolskii
{{small|Chkhikvadze & Tuniyev, 1986}}

|Testudo ibera nikolskii
{{small|— Highfield, 1990}}

|Testudo terrestris nikolskii
{{small|— David, 1994}}

|Testudo graeca niiolskii
{{small|Paull, 1997}} (ex errore)

|Testudo nikolskii
{{small|— Vetter, 2002}}

| T. g. pallasi

|Testudo graeca pallasi
{{small|Chkhikvadze, 1989}}
(nomen nudum)

|Testudo graeca pallasi
{{small|Chkhikvadze & Bakradze, 2002}}

|Testudo pallasi
{{small|— Danilov & Milto, 2004}}

| T. g. perses

|Testudo perses
{{small|Perälä, 2002}}

|Testudo ibera perses
{{small|— Artner, 2003}}

| T. g. soussensis

|Testudo graeca soussensis
{{small|Pieh, 2001}}

|Testudo soussensis
{{small|— Vetter, 2002}}

| T. g. terrestris

|Testudo terrestris
{{small|Forsskål, 1775}}

|? Testudo zolhafa
{{small|Forsskål, 1831}} (nomen nudum)

|? Testudo zolkafa
{{small|Forsskål, 1831}} (nomen nudum)

|? Testudo zohalfa
{{small|Forsskål, 1835}} (nomen nudum)

|Testudo graeca terrestris
{{small|— Wermuth, 1958}}

|Testudo terrestris terrestris
{{small|— David, 1994}}

|Testudo ibera terrestris
{{small|— Artner, 2003}}

| T. g. zarudnyi

|Testudo zarudnyi
{{small|Nikolsky, 1896}}

|Testudo graeca zarudnyi
{{small|— Mertens, 1946}}

|Testudo ibera zarudnyi
{{small|— Gmira, 1993}}

|Testudo terrestris zarudnyi
{{small|— David, 1994}}

}}

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.{{Cite journal |last1=Alsafy |first1=Mohamed A. M. |last2=El-sharnobey |first2=Nermin K. A. |last3=El-Gendy |first3=Samir A. A. |last4=Abumandour |first4=Mohamed A. |last5=Ez Elarab |first5=Samar M. |last6=Rashwan |first6=Ahmed M. |last7=Hanafy |first7=Basma G. |date=October 2024 |title=Macroscopic, microscopic, and immunofluorescent characterization of the Greek tortoise ( Testudo graeca graeca ) oropharyngeal floor with concern to its feed adaptation as a herbivorous land reptile |url=https://analyticalsciencejournals.onlinelibrary.wiley.com/doi/10.1002/jemt.24619 |journal=Microscopy Research and Technique |language=en |volume=87 |issue=10 |pages=2385–2398 |doi=10.1002/jemt.24619 |pmid=38808586 |issn=1059-910X}}{{Cite journal |last1=Sereau |first1=Matthieu |last2=Lagarde |first2=Frédéric |last3=Bonnet |first3=Xavier |last4=El Mouden |first4=El Hassan |last5=Slimani |first5=Tahar |last6=Dubroca |first6=Laurent |last7=Trouvé |first7=Colette |last8=Dano |first8=Stéphanie |last9=Lacroix |first9=André |date=2010-06-01 |title=Does testosterone influence activity budget in the male Greek tortoise (Testudo graeca graeca)? |url=https://linkinghub.elsevier.com/retrieve/pii/S0016648010000857 |journal=General and Comparative Endocrinology |volume=167 |issue=2 |pages=181–189 |doi=10.1016/j.ygcen.2010.03.002 |pmid=20226191 |issn=0016-6480}}

Testudo graeca is recognized for its longevity, with verified lifespans exceeding 100 years and anecdotal reports suggesting ages over 125 years.{{Cite journal |last1=Fritz |first1=Uwe |last2=Havaš |first2=Peter |date=2007-10-31 |title=Checklist of Chelonians of the World |url=https://vertebrate-zoology.arphahub.com/article/30895/ |journal=Vertebrate Zoology |language=en |volume=57 |issue=2 |pages=149–368 |doi=10.3897/vz.57.e30895 |doi-access=free |issn=2625-8498}} Among reptiles, it has one of the largest known genomes.{{cite book |last=Pritchard |first=Peter C. H. |url=https://archive.org/details/completeencyclopediaofturtles |title=Encyclopedia of Turtles |publisher=T.F.H. Publications |year=1979 |isbn=0-87666-918-6 |location=Neptune, NJ |access-date=2024-04-03}}

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.{{cite journal |author=Tortoise & Freshwater Turtle Specialist Group |year=1996 |title=Testudo graeca (Greek Tortoise) |url=https://www.iucnredlist.org/species/21646/9305693 |access-date=2024-04-03 |journal=The IUCN Red List of Threatened Species |doi=10.2305/IUCN.UK.1996.RLTS.T21646A9305693.en}}

Evolution

The oldest confirmed fossil attributed to Testudo graeca originates from the Early Pliocene of Greece.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. 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.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.Staesche K, Karl HV, Staesche U (2007). "Fossile Schildkröten aus der Türkei". In: Fossile Schildkröten aus Drei Kontinenten. 98: 91–149.

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
class="wikitable"
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:Genus Testudo at The Reptile Database. www.reptile-database.org.

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. 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

Estimated by counting growth rings, testudo graeca was always reached earlier in males than in females. Post-maturity females continued to grow slowly in size but with 30–90% greater growth on average compared to males.{{Cite journal |last1=Tiar-Saadi |first1=Manel |last2=Tiar |first2=Ghoulem |last3=Bouslama |first3=Zihad |last4=Široký |first4=Pavel |date=2022-05-23 |title=Mechanisms Determining Body Size and Shape Difference in Algerian Spur-Thighed Tortoises (Testudo graeca) |journal=Animals |language=en |volume=12 |issue=10 |pages=1330 |doi=10.3390/ani12101330 |doi-access=free |issn=2076-2615 |pmc=9137665 |pmid=35625176}}

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.{{cite book |last1=Pritchard |first1=Dr. Peter C. H. |title=Encyclopedia of Turtles |date=1979 |publisher=T. F. H. Publications |isbn=0-87666-918-6 |location=Neptune, New Jersey}}

=Mating and reproduction=

File:A pair of Testudo graeca mating.jpg, İzmir Province, Turkey]]

Reproductive behavior in T. graeca begins shortly after emerging from hibernation.{{Cite web |title=ClinicalKey |url=https://www.clinicalkey.com.au/#!/content/journal/1-s2.0-S0016648010000857 |access-date=2025-04-03 |website=www.clinicalkey.com.au}} 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.{{cite book |last=Highfield |first=Andy |title=Practical Encyclopedia of Keeping and Breeding Tortoises and Freshwater Turtles |publisher=Carapace Press |year=1996 |isbn=1873943067}}

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.Pérez, Irene et al. (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.Bergin, Daniel & Nijman, Vincent (2014). "Open, Unregulated Trade in Wildlife in Morocco's Markets, TRAFFIC Bulletin". Retrieved 23 March 2015.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.

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.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.{{cite journal |last1=Nijman |first1=Vincent |last2=Bergin |first2=Daniel |year=2017 |title=Trade in spur-thighed tortoises Testudo graeca in Morocco: Volumes, value and variation between markets |journal=Amphibia-Reptilia |volume=38 |issue=3 |pages=275–287 |doi=10.1163/15685381-00003109}}

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."Helpful advice for your tortoise diet". www.tortoisecentre.co.uk. Archived from the original on 29 January 2018. Retrieved 29 January 2018.{{cite web |title=Greek Tortoise Diet |url=https://www.tortoisetrust.org/articles/greekdiet.html |access-date=2024-04-03 |website=Tortoise Trust}}

References

External links

[http://www.mayer-richard.de/ European Tortoises. (in German).]
[https://archive.today/20130416152136/http://www.animalutul.ro/reptile/specii-de-reptile/broasca-testoasa-greaca-testudo-graeca/208/ Broasca Testoasa Greaca - Testudo graeca. (in Romanian).]

Category:Testudo (genus)

Category:Reptiles described in 1758

Tortoise

Category:Fauna of Spain

Category:Reptiles of Armenia

Category:Reptiles of Azerbaijan

Category:Turtles of Europe

Category:Turtles of Asia

Category:Species that are or were threatened by the pet trade

Category:Taxa named by Carl Linnaeus

Category:Reptiles of Russia

Category:Reptiles as pets