Taxonomic Notes
This taxon is clearly a complex of several species, and 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. Perälä (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 Türkiye, was described as a subspecies of Testudo graeca and is treated as such by most recent studies (Perälä 2002b:35, Pieh et al. 2002), but it was implied to merit full species status by the phylogenies presented by Perälä (2002b:38-39) and is presented as such by Perälä (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 Türkiye 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 Türkiye, 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 (Perälä 2002b:35, Pieh et al. 2002), but it was implied to merit full species status by phylogenies presented by Perälä (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 Perälä (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 the 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 elevated to species level by Iverson (TTWG checklist update manuscript), but is currently retained as a synonym (TTWG 2021).
Testudo perses, from southeastern Türkiye 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 separate species by Perälä (2002c:81). Testudo terrestris, from the northern Mesopotamian region, has a confused nomenclatural history, but the elevation of the 'terrestris-group' to species level was proposed by Gmira (1993) and Perälä (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 a full species by Perälä (2002c:83). A full taxonomic review of this species is required across its entire range to identify those lineages that have diverged sufficiently to merit full species status.
Justification
European regional assessment: Near Threatened (NT)
EU 27 regional assessment: Near Threatened (NT)
In the European region, the Common Tortoise occurs as native in the eastern Balkans (from Serbia and Kosovo, south to Greece, and east to Romania, Bulgaria and European Türkiye). The species also occurs in two small subpopulations in southern Spain (origin uncertain but included here as native), and most likely also occurs in the Spanish North African territories. Records from Crimea require confirmation and are excluded, and subpopulations on the Balearic Islands, Italy (the peninsula and Sardinia), southern France, and the Balkan region outside of the presumed native range are excluded.
The species faces a wide range of anthropogenic threats that result in habitat conversion, degradation and loss. Human commensal predators (mainly dogs but also crows and other species) predate tortoises and destroy nests; this is one of the main threats in the Balkan area. The removal of individuals from the wild for the pet trade continues.
However, the population trend across the native range of the species in Europe is not known. At the European and EU levels, this species is assessed as Near Threatened based on suspected population declines that approach but do not exceed 30% over both the past three generation lengths (A2bcde) and an ongoing decline that encompasses the past two generation lengths and the future one generation length (generation length = 25 years) (4bcde).
Geographic Range Information
In the European region, the Common Tortoise (Testudo graeca) occurs as native in southwestern (southern mainland Spain; but see discussion below) and southeastern Europe; the eastern Balkans, where it is distributed more continuously in southeastern and southern Serbia and Kosovo (Tomović et al. 2004; L. Tomović pers. comm. January 2023), eastern North Macedonia, most of Bulgaria, eastern Romania (Cogălniceanu et al. 2013), northeastern and Aegean Greece, and European Türkiye. The species most likely also occurs in the Spanish North African territories. Records from Ukraine (mountainous southern mountains of Crimea) require confirmation and are excluded here.
Genetic analysis suggests that the mainland Spanish subpopulations are introductions from northern Africa in historical times (Alvarez et al. 2000), however, TTWG (2021) considers the species to be native in mainland Spain. Archaeological evidence (from the first and second centuries AD) of its presence in the Iberian Peninsula was confirmed by Boneta Jiménez et al. (2023), but no there is as yet no evidence in the paleontological record of the presence of Testudo graeca in the Iberian Peninsula (Pérez-García 2017). The origin of the species in Spain is still to be confirmed, and Spanish subpopulations are included in this European regional assessment.
The Common Tortoise also occurs as isolated introduced subpopulations on the Balearic Islands, Italy (occasional records from the peninsula and in west-central and western (Mal di Ventre) Sardinia. The Sardinian subpopulation is derived from North African Common Tortoise (Vamberger et al. 2010, 2011); it was not introduced from the Balkans. Individuals randomly found in continental Italy (not mapped) perhaps could originate from the Balkans, and as so far known there are no viable populations in mainland Italy. There are scattered records of probably introduced individuals with no established populations (not mapped) from southern France (Gmira 1993, Buskirk et al. 2001:136), mainland Italy and the Balkan region (outside the main range).
Globally, this species ranges from the Mediterranean basin, east through West Asia and the Caucuses (Chkhikvadze and Bakradze 1991) to Iraq, Iran and, possibly, western Afghanistan and through North Africa (TTWG 2021). It occurs from near sea level to 1,900 m altitude (in Morocco; Lambert, 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 (Lambert 1995). There is an isolated population on the Black Sea coast of southern Russia, extending into northwestern Georgia. In West Asia, it occurs widely in Türkiye (except the northeast; Sindaco et al. 2000), south through northern and western Syria (Lymberakis and Kalionzopoulou 2003), Lebanon (Hraoui-Bloquet et al. 2002), northern and central Israel, western Jordan (Sindaco et al. 2000), east through northern Iraq, much of Iran, and southern Turkmenistan, and north through much of Azerbaijan, Armenia, eastern Georgia, and southern Russia (in Dagestan; Chkhikvadze and Bakradze 1991). It ranges very close to Afghanistan and might occur there.
Population Information
In the European region, this is a species with a wide distribution that inhabits both heavily developed sites (including urban areas, intensive agriculture and touristic areas) and remote sites (shrublands and maquis far from human intervention), and the abundance of this species varies greatly across its range.
In Serbia, there are only a small number of records in the past ten years; survey efforts mainly in the south of Serbia reported <10 individuals in the last 10-15 years; it is a marginal species, with reproduction only known in one locality in southern Serbia, but there is no evidence of decline (L. Tomović pers. comm. January 2023).
In North Macedonia, a population decline is inferred as a result of agricultural development and the suspected illegal collection from the wild for farms (L. Tomović pers. comm. January 2023). Preliminary studies suggest that its subpopulations are much less dense than those of Hermann’s Tortoise with which it shares its habitats almost throughout its entire national range (D. Arsovski pers. comm. January 2023); a subpopulation that has been studied for four years in an urban park in Skopje revealed a population density of only 2.2 ind./ha opposed to ~11 ind./ha for a syntopic population of Hermann’s Tortoises (Frcovki et al. 2022). Nevertheless, observations in the southeastern corner of North Macedonia reveal a possible opposite trend, but studies are needed to confirm this trend (D. Arsovski pers. comm. January 2023).
In Greece, the population trend is not known (C. Papoutsakis, draft Greece national Red List assessment, October 2023).
The species is widespread in eastern Romania, except for the flood-prone areas of the Danube Delta. It can reach high densities in favourable habitats, with some tens of adults/ha being observed (Buică et al. 2013).
In Spain, the subpopulation protected in the Doñana National Park is stable, but subpopulations in southeastern mainland Spain continue to decline.
In its non-native European range, there are two healthy subpopulations of the species on Sardinia. Out of the European region, the species underwent significant declines across large parts of North Africa as a result of over-collection and habitat conversion, but the current situation is not known. The population trend across the rest of its range outside of Europe is more mixed, stable in some areas, but with declines reported in others.
Habitat and Ecology Information
This species inhabits a variety of dry, open scrubby habitats, meadows and pastures, sand dunes, forests, heathlands, and open habitats through its wide range, generally on a sandy-calcareous substrate (Buskirk et al. 2001).
It is also found in more open, rocky areas (L. Tomović pers. comm. 2023). Areas of strongly salty substrate and sparse vegetation, as well as accumulations of large rocks and steep slopes, tend to be avoided (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. More specifically, this species feeds on predominantly legumes, with some compositae, fruits and berries, and snails and insects are also taken (Bannikov, cited in Buskirk et al. 2001:151). It has also been recorded to feed on carrion, and, for instance, two adult individuals from one of the two Sardinian subpopulations were observed while feeding on the corpse of a dead bird (L. Luiselli and L. Rugiero, unpublished data).
It is a well-known disperser of 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 m, rarely up to 800 m asl, and preferably between 50 and 300 m, 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). The former subspecies pallasi (now placed as a synonym) inhabited 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 (Bannikov, cited in Buskirk et al. 2001:150-155, Mazanaeva 2002). Females mature at a CL over 16 cm, at an estimated age of 12-14 years, and can produce up to three clutches of about five (range two to eight) eggs per year (Bannikov et al. in Buskirk et al. 2001:155).
The introduced subpopulations on Sardina occur in Mediterranean evergreen maquis habitats, often in rocky or sandy soils (Corti et al. 2007). Testudo graeca mainly inhabits coastal zones characterized by open uncultivated areas, dune vegetation and Mediterranean maquis. T. graeca seems to be active throughout the year but with decreased or no activity on the hottest summer days. In the autumn, characterised by relatively mild temperatures, intense activity was recorded. Courtship and mating were observed more frequently in the autumn (September-November) than in spring (February-March). On the Sardinian island of Mal di Ventre Island, females are larger than males and individuals of both sexes are larger than those of the main island.
Threats Information
Habitat conversion, degradation and loss is widely cited as a significant threat. As a result of loss of prime habitat to agriculture and infrastructure development, tortoise populations are now focused in marginal habitats such as anthropogenic habitats (orchards, vegetable gardens), riverine floodplain areas and coastal sand dunes, and this is currently the case in both Serbia and in North Macedonia. Mechanisation of agriculture is also a threat through direct mortality (P.P. van Dijk pers. comm. January 2023).
Past pet trade collection has involved very large numbers of animals and has been indicated as a major factor in population depletion in some parts of the global range of the species, and the removal of individuals from the wild for the pet trade continues. The release of tortoises of captive origin from different sources into T. graeca habitat represents the dangers of genetic pollution as well as the introduction of pathogens (Andreu 2003).
Road mortality, human commensal predators (mainly dogs but also crows and others) killing tortoises and destroying nests; this is one of the main threats in the Balkan area, and collection of tortoises as living souvenirs by tourists, have further been implicated as significant threats. Natural predators, a significant cause of mortality of juveniles and adults (Buskirk et al. 2001:150-155), have probably been 'subsidised' by anthropogenic changes to tortoise habitat, allowing higher predator density and predator impact on tortoises.
For the introduced Sardinian subpopulations, illegal harvesting, fires, fragmentation and loss of habitat due to the expansion of intensive agriculture as well as the expansion of tourist and urban infrastructures. The disappearance of open or semi-open areas, optimal for foraging and spawning may cause a decline. In Romania, animals are occasionally killed in agriculture fields and vegetation mowing (Vlad et al. 2020), and the frequency of scars on the carapace is higher in populations from areas with elevated human activities (Buică et al. 2014). Vegetation fires can cause high mortality and injuries.
Historically, the species was exploited commercially in parts of its range out of Europe (Dagestan); their meat and eggs used for human consumption as well as feed for pigs and to supply factories producing food for salmon farms. The use for these purposes in Europe is not known.
Use and Trade Information
This species was historically an important pet animal, recently less so. There is some use for tourist souvenir manufacture in North Africa; this requires confirmation for Europe. There is still substantial trade, but this is primarily sourced from captive-bred legal stock, but with some ongoing illegal trade from wild sources (P.P. van Dijk pers. comm. January 2023). Historically it was also overhunted for food consumption in various European countries, as this meat of this species was allowed by the Catholic religion even during the days/periods in which other types of meat were not allowed; this use is probably now insignificant.
Three tortoise farms in North Macedonia breed this species (D. Arsovski pers. comm. 2023). The Macedonian Red Data Book of Fauna states: “National reports from the Ministry of Environment and Physical Planning report 150 and 850 exported individuals in 2016 and 2017, respectively. Since farm-born individuals cannot be distinguished from free-ranging animals, and since adult individuals (>10 cm long, at least eight years old) get a much higher price or smaller individuals are not imported at all (Canada and USA, personal communications from farmers), there is a high incentive to collect from the wild and use farms as platforms for “legal export” of tortoises (Arsovski and Sterijovski 2023).
Conservation Actions Information
The species occurs in many protected areas across its European range. The species is included in CITES Appendix II and EU regulation EEC No.338/97 which prohibits the import into the EU unless captive-bred. The species is protected under domestic legislation in Italy and many other countries.
The conservation status varies from country to country, and in general it is less threatened in the eastern countries of its range. Population monitoring of threatened subpopulations would be appropriate, as would further taxonomic, ecological, and conservation-focused research. In Italy, where it is introduced, it was considered Near Threatened in the Italian Red List because it has a limited area of occupancy of less than 2,000 km
2 (Ballasina 1995). Currently, it is known in this country in only two locations, although there is no evidence that either of these remnant subpopulations is in decline. However, potentially these localities are threatened by alteration and degradation of the habitat and possibly by poaching.
It is assessed as Vulnerable in North Macedonia (
Arsovski and Sterijovski 2019) and as Critically Endangered in Serbia. Eastern Hermann's Tortoise (
Testudo hermanni boettgeri) is assessed as Endangered (A3ac) in Bulgaria (
Beshkov 2011).
A number of non-governmental organisations and private individuals work on
in situ conservation efforts (species and habitat), and
ex situ animal rescue and captive breeding.
Reintroductions and habitat management would benefit this species, along with further research into its taxonomy and population trends.