Testudo graeca 

Scope: Europe
Language: English

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Taxonomy [top]

Kingdom Phylum Class Order Family
Animalia Chordata Reptilia Testudines Testudinidae

Scientific Name: Testudo graeca
Species Authority: Linnaeus, 1758
Regional Assessments:
Common Name(s):
English Common Tortoise, Greek Tortoise, Moorish Tortoise, Spur-thighed Tortoise
French Tortue Mauresque
Spanish Tortuga Mora
Furculachelys nabeulensis Highfield, 1990
Testudo anamurensis Weissinger, 1987
Testudo antakyensis Perälä, 1996
Testudo armeniaca Chkhikvadze & Bakradze, 1991
Testudo buxtoni Boulenger, 1921
Testudo flavominimaralis Highfield & Martin, 1989 (nomen dubium
Testudo floweri Bodenheimer, 1935
Testudo graeca ssp. cyrenaica Pieh & Perälä, 2002
Testudo graeca ssp. soussensis Pieh, 2001
Testudo ibera Pallas, 1814
Testudo nikolskii Chkhikvadze & Bakradze, 1991
Testudo pallasi Chkhikvadze & Bakradze, 2002
Testudo perses Perälä, 2002
Testudo terrestris Forsskål, 1775
Testudo whitei Bennett in White, 1836
Testudo zarudnyi Nikolsky, 1896
Taxonomic Notes: This taxon is clearly a complex of several species, but its taxonomy remains very confused without resolution, and so it is treated here as a single species. Several subspecies or possibly synonyms have been treated as full species by some authors. The subspecies cyrenaica and soussensis were elevated to full species by Vetter, (2002) without supporting information, a move supported by J. Perala (pers. comm. 2004). Testudo flavominimaralis is usually treated as a synonym of Testudo graeca graeca, and it was not accepted as valid by Iverson (1992) and Schleich et al. (1996); nor was it discussed in detail by Pieh (2002:110). Its taxonomic status and distribution are uncertain, and it is treated here as a synonym of T. graeca, though it has not been formally synonymised. Testudo nabeulensis from Tunisia and northwestern Libya is usually treated as a synonym of Testudo graeca, but Pieh (2002:110) who took the view that it is a valid taxon within the Testudo graeca complex. Testudo anamurensis, from the southern coastal region of Turkey, was described as a subspecies of Testudo graeca and is treated as such by most recent studies (Perala 2002b:35, Pieh et al. 2002), but it was implied to merit full species status by the phylogenies presented by Perala (2002b:38-39) and is presented as such by Perala (2002c:87) and Iverson (Checklist update MS). It is reported to occur in micro-sympatry with T. ibera at Side by Bruekers (1997). Testudo antakyensis, from the southern Anamos Mountains region of southern Turkey southwards to northern Israel and northwestern Jordan, was described as a separate species but might be a variant of T. graeca based on mitochondrial DNA sequences, and is treated here as a synonym pending further information. Testudo armeniaca, from northeastern Turkey, southern Armenia, northwestern Iran and possibly Azerbaijan, was originally described as a subspecies of Testudo graeca, and has been treated as such by most recent studies (Perala 2002b:35, Pieh et al. 2002), but it was implied to merit full species status by phylogenies presented by Perala (2002b:38-39) and it was presented as such by Iverson (Checklist update MS). Testudo buxtoni, from the southwestern Caspian Sea shore of northern Iran, was previously treated as a synonym within the Testudo graeca complex, specifically of T. ibera, but was considered 'likely to be a valid taxon' by Perala (2002c:84), following a hint by Chkhikvadze and Bakradze (1991). Testudo floweri, from the Levantine coastal plain and inland lowland areas in Israel and Lebanon, is usually considered to be a synonym of T. graeca, but might merit species status. Testudo ibera, from southeastern Europe to Iran (depending on the taxonomy followed), is usually treated as a subspecies of T. graeca, but elevation of the 'ibera-group' to species level was proposed by Gmira (1993) and Perala (2002c). Testudo nikolskii, from the northern Black Sea coast, has recently been considered to merit species rank, but its distinctness from T. ibera was questioned by Leontyeva et al. (2002). Testudo pallasi, from the northern and eastern Caucasus, was described as a subspecies of T. graeca, but was elevated to species level by Iverson, (Checklist update MS). Testudo perses, from southeastern Turkey and much of Iran, was previously would have been considered part of the Testudo graeca complex, specifically related to T. ibera and/or T. zarudnyi, but was described as a separated species by Perala (2002c:81). Testudo terrestris, from the northern Mesopotamian region, has a confused nomenclatural history, but elevation of the 'terrestris-group' to species level was proposed by Gmira (1993) and Perala (2002). Testudo whitei, possibly from Algeria, is generally considered to be a synonym of T. graeca (Buskirk et al. 2001), but it was elevated to species status by Highfield and Martin (1989) and Ballasina et al. (2002); it was considered a nomen dubium by Pieh (2002:110). Testudo zarudnyi, from eastern Iran, was traditionally treated as a subspecies of T. graeca, but was treated as full species by Perala (2002c:83). A full taxonomic review of this species required from across its entire range to identify those lineages that have diverged sufficiently to merit full species status.

Assessment Information [top]

Red List Category & Criteria: Vulnerable A2bcde+4bcde (Regional assessment) ver 3.1
Year Published: 2004
Date Assessed: 2004-12-17
Needs updating
Assessor(s): van Dijk, P.P., Corti, C., Mellado, V.P. & Cheylan, M.
Reviewer(s): Cox, N.A. & Temple, H.J.
At the European and EU level this species is assessed as Vulnerable based on population declines exceeding 30% over 3 generations (generation length = 25 years).

This draft regional assessment is based on draft information gathered at the IUCN Mediterranean Red Listing Workshop - Freshwater Fishes, Reptiles and Amphibians (Malaga [Spain] 13-17 December 2004) and remains under current review by the IUCN-SSC Tortoise and Freshwater Turtle Specialist Group - please contact Peter Paul van Dijk with any comments.

Geographic Range [top]

Range Description:This species ranges from the Mediterranean basin, east to Iran, with populations in North Africa, southern Europe and West Asia. It occurs from near sea level to 1,900m altitude (in Morocco - Lambert 1983, in Buskirk et al. 2001:136). In North Africa it ranges from western and northern Morocco, through northern Algeria and northern and central Tunisia to northwestern Libya, with an isolated population in Cyrenaica in northeastern Libya. In southern Europe it is known from isolated populations in southern Spain, Mallorca (Spain), west-central Sardinia (Italy), Mal di Ventre (off western Sardinia, Italy), and from the eastern Balkans where it is distributed more continuously in southeastern Serbia, eastern Macedonia, most of Bulgaria, eastern Romania, northeastern Greece and European Turkey. There is an isolated population on the Black Sea coast of southern Russia, extending into northwestern Georgia. Records from Ukraine require confirmation. Genetic analysis suggests that the Spanish populations are introductions from northern Africa in historical times (Alvarez et al. 2000), and all Italian populations are also presumed to be introduced. There are scattered records (probably of introduced individuals, not established populations, and therefore not mapped - Buskirk et al. 2001:136) from southern France, mainland Italy and the Balkan region (outside the main range). In West Asia it occurs widely in Turkey (except the northeast), south through northern and western Syria, Lebanon, northern and central Israel, western Jordan, east through northern Iraq, much of Iran, and southern Turkmenistan, and north through much of Azerbaijan, Armenia, eastern Georgia, and southern Russia (in Dagestan). It ranges very close to Afghanistan and might occur there. For further information, see Iverson (1992), Buskirk et al. (2001), Perala (2002c), and Pieh (2002). The range of the possible synonym Testudo flavominimaralis is not clear. The type locality is stated as "North Africa. Precise biotope and range not known. Believed to include Libya." It was restricted to "Libya" by Highfield (1990) (see also Buskirk et al. (2001)).
Countries occurrence:
Algeria; Armenia (Armenia); Azerbaijan; Bulgaria; Georgia; Greece; Iran, Islamic Republic of; Iraq; Israel; Jordan; Lebanon; Libya; Macedonia, the former Yugoslav Republic of; Morocco; Romania; Russian Federation; Serbia (Serbia); Syrian Arab Republic; Tunisia; Turkey; Turkmenistan
Present - origin uncertain:
Italy (Italy (mainland) - Introduced, Sardegna); Spain (Baleares, Spain (mainland))
Additional data:
Upper elevation limit (metres):1900
Range Map:Click here to open the map viewer and explore range.

Population [top]

Population:The abundance of this species varies greatly across its range. In Algeria it was historically very common in the coastal region, and surveys during the period 1976-1981 encountered tortoises at rates varying between 0.1 to 10 tortoises per man-hour searching (summary in Lambert 1995). In Libya, tortoises were considered to be quite common in vegetated areas of Cyrenaica, but there was no status data from Tripolitania (Lambert 1995: 11). Morocco traditionaly supplied the European pet trade, peaking at several 100,000 animals annually in the 1970s. It was reported common in 1958, but populations were reported as depleted in northwestern and central Morocco by 1969 and 1981. Based on encounter rates, populations in some areas of Morocco were estimated to have suffered over 90% decline over the period 1900-1984 (review in Lambert 1995). The Souss Valley population is declining as result of past pet trade and ongoing habitat degradation (Bayley and Highfield 1996). It was historically common on the Tunisian plains, but some collection for international and regional trade has been documented; encounter rates suggest that populations have been depleted somewhat (review by Lambert 1995). There is a good healthy population of the species on Sardinia. In Spain the population protected in the Donana region is stable, but the populations in southeastern Spain and Mallorca continue to decline (Lambert 1995, Andreu 2003). The taxon Testudo anamurensis was apparently fairly easily encountered during a visit to southern Turkey by Bruekers (1997), suggesting that it is not particularly rare. Populations of in Israel were reported to be healthy and substantial, with no indication of decline (Lambert 1995:13). The taxon Testudo ibera has disappeared from land brought under irrigated agriculture in Azerbaijan, and there has been an overall in decline in former Soviet Union (Kuzmin 2002:93). The taxon Testudo nikolskii reaches densities of an average of 0.5 individuals per hectare in optimal habitat; in more densely forested areas the density declines to 0.2-0.4 indiv/ha (Inozemtsev and Pereshkolnik 1985). Populations are dominated by mature adult animals, with apparently low recruitment, but population structure remained stable between 1985-1999 (Inozemtsev and Pereshkolnik 1985, Leontyeva et al. 2002). At a range of 8250 km2, this suggests a maximum population of 40,000 animals. For the taxon Testudo pallasi, Mazanaeva (2002:63) documented declining population densities and loss of range during the period 1991-2001, particularly in the northern part of historical distribution, noting that no tortoises could be found now where previously they were commonly seen. Comparison with range maps from 1977 and before indicate up to 80% loss of area of occurrence in Dagestan (data in Mazanaeva 2002:60-62). Most surveyed localities had densities of less than one individual tortoise per 5 ha, although a few localities had significantly higher densities and two localities had densities of about 1.5 tortoises per ha (Mazanaeva 2002:62). Historical observations from the 1930s-1950s (reviewed by Mazanaeva 2002:60) suggest high densities (on the order of a dozen or more animals/ha). No data are available on range or population density trends of this taxon outside Dagestan.
Current Population Trend:Decreasing
Additional data:
Population severely fragmented:No

Habitat and Ecology [top]

Habitat and Ecology:This species inhabits a variety of dry, open scrubby habitats, meadows and pastures, sand dunes, forest, heathlands, and open habitats through its wide range, generally on a sandy-calcareous substrate (Bayley and Highfield 1996, Buskirk et al. 2001). Areas of strongly salty substrate and sparse vegetation, as well as accumulations of large rocks and steep slopes, tend to be avoided (Alekperov 1978, Kuzmin 2002:90). Testudo graeca is a generalist vegetarian, feeding on a wide variety of leaves, buds, flowers, seeds and fruits of grasses, herbs and shrubs, as well as small invertebrates such as snails, arthropods, and carrion. It has been recorded as dispersing plant seeds. Mature females usually produce two or three clutches (extremes one to four) of about two to four eggs (extremes one to seven) (Buskirk et al. 2001:153). The taxon Testudo nikolskii inhabits undisturbed deciduous forests, dispersing into more open areas in summer, including the vicinity of human habitation. Preferred areas are open forest of mixed low-growing Oaks (Quercus), Juniperus, Artemisia, Rhus, Pistacia and Jasminum with a well-developed herbaceous layer, as well as more open patches. Inhabited areas are below 600, rarely up to 800m alt, and preferably between 50 and 300m, with a preference for south-facing gentle middle and lower slopes. It feeds on a variety of herbs and shrub leaves, and small items of animal origin (Buskirk et al. 2001: 150-152; Leontyeva et al. 2002). The taxon Testudo pallasi inhabitsed a wide variety of habitats, including open lowland forests, hill and mountain slopes and a variety of other open habitats like steppe and semi-desert and some anthropogenic landscapes (Bannikow 1951, in Buskirk et al. 2001:150-155, Mazanaeva 2002). As a result of loss of prime habitat to agriculture and infrastructure development, tortoise populations are now focused on marginal habitats such as anthropogenic habitats (orchards, vegetable gardens), riverine floodplain areas and coastal sand dunes (Mazanaeva, 2002:63). The taxon feeds on herbs, predominantly legumes, with some compositae, fruits and berries, and snails and insects also taken (Bannikov, in Buskirk et al. 2001:151). Females mature at a CL over 16 cm, at an estimated age of 12-14 years, and can produce up to 3 clutches of about 5 (range 2-8) eggs per year (Bannikov, Nadjafov et al. in Buskirk et al. 2001:155).

Use and Trade [top]

Use and Trade: Historically an important pet animal, recently less so; some use for tourist souvenir manufacture.

Threats [top]

Major Threat(s): Habitat degradation and loss is widely cited as a significant threat (Lambert 1995, Bayley and Highfield 1996 - Morocco).
Past pet trade collection has involved very large numbers of animals and has been indicated as a major factor in population depletion, particularly in Morocco and Algeria (Lambert 1995). Recent collection for pet trade, largely domestic but including offering animals to European tourists, has been recorded from Tunisia (Highfield and Bayley 1996). In Morocco, tortoise shells were used in the manufacture of tourist souvenirs (Highfield and Bayley 1996). Release of captive tortoises from different populations into T. graeca habitat represents dangers of genetic pollution as well as introduction of pathogens (Andreu 2003). For the taxon Testudo nikolskii, concern has been expressed that the population structure dominated by large mature animals indicates compromised recruitment and therefore future stability of the taxon, but this appears to be a survey artefact as a monitored population proved stable over a decade (Inozemtsev and Pereshkolnik 1985, Leontyeva et al. 2002). Historically, the taxon Testudo pallasi in Dagestan was exploited commercialy, their meat and eggs being used for human consumption as well as feed for pigs and to supply factories producing food for salmon farms. While no quantitative data are available on this exploitation, historical data indicate that harvest, especially of females, was intensive and and led to near-total extirpation in some regions (Bannikov 1951, in Mazanaeva 2002:60). Tortoise habitat in Dagestan is focused on the Primorskaya Lowland and adjacent foothill region; this region was used for human resettlement during the 1960-1980s, followed by expansion of residential infrastructure. Deforestation, intensive land use, use of inorganic fertilisers and pesticides, overgrazing by cattle, extensive plantations, and sand mining, have been factors in loss of tortoise habitat (Mazanaeva 2002:63). Road killing, human commensal predators [mainly dogs but also crows and others] killing tortoises and destroying nests, and collection of tortoises as living souvenirs by tourists, have further been implicated as significant treats to Dagestan tortoises (Mazanaeva 2002:63-64). Natural predators, a significant cause of mortality of juveniles and adults (Buskirk et al. 2001:150-155), have probably have been 'subsidised' by anthropogenic changes to tortoise habitat, allowing higher predator density and predator impact on tortoises.

Conservation Actions [top]

Conservation Actions: It occurs in many protected areas. A variety of non-governmental organisations and private individuals work on in-situ conservation efforts (species and habitat), and ex-situ animal rescue and captive breeding. The species is included in CITES Appendix II and EU regulation EEC No.338/97 prohibits import into the EU unless captive-bred. The species is protected under domestic legislation in Algeria, Tunisia, and many other countries. Populations of the varying taxa, subspecies and evolutionarily significant units need to be confirmed as occurring in adequately protected areas.
Population monitoring of threatened populations would be appropriate, as would further taxonomic, ecological and conservation-focused research. Assessment of taxonomic status is first priority for the taxon flavominimaralis, and many other subspecies. For the taxon Testudo pallasi, improved environmental management and awareness has reduced habitat degradation pressures somewhat in the coastal habitats in recent years (Mazanaeva 2002:63). Mazanaeva (2002:65) called for the establishment of several modest-sized protected areas, particularly for the sand dune areas which still hold relatively good populations and offer good conditions for reproduction; she also recommended population and natural history studies and monitoring, as well as ex-situ conservation breeding and re-introduction, and creating awareness among local human population.

Citation: van Dijk, P.P., Corti, C., Mellado, V.P. & Cheylan, M. 2004. Testudo graeca. The IUCN Red List of Threatened Species 2004: e.T21646A9305080. . Downloaded on 24 March 2017.
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