No subspecies are currently recognized. Formerly, T. m. marginata and T. m. sarda were recognised, however, later research (Fritz et al. 2005, Perez at al. 2012) showed that these "subspecies" are genetically inseparable from the species, T. marginata. In this account, we include T. weissingeri within T. marginata following TTWG (2021).

Global and European regional assessment: Least Concern (LC)
EU 27 regional assessment: Least Concern (LC)

This species is endemic to Europe, where it is considered to be native to southern Albania and Greece, with occurrence in Italy (Sardinia and on some satellite islands) the result of historical introduction from Greece. It is assessed as Least Concern both globally and for the majority of EU 27 Member States since, although its currently known Area of Occupancy (AOO) is restricted, at 568 km², it occurs in many more than ten locations and is not severely fragmented, and has an area of extensive, suitable habitat which appears not to be under significant threat. It also has a presumed large population, and it is unlikely to be declining fast enough to qualify for listing in a threatened category. In Italy, where introduced, the current national Red List assessment considers the species NT.

This species is endemic to Europe, where it occurs as native in Greece (the northwestern, central and southern mainland from the Axios (Vardar) River to the Peloponnese) and adjoining extreme southern Albania, including a number of islands (Alonissos, Skiathos, Skyros, Kyra Panagia, Valaxa, Salamina, Sfaktiria, Sapietza, Poros, Dokos, Psili, Romvi, Spetses, Spetsopoula, Trizonia, Kythera, Elafonisos and Euboea; Escoriza et al. 2023). The species occurs across a wide altitudinal range, from 0 to 1,600 m asl (Peloponnese, Greece). The native range area of occupancy (AOO) has not been estimated, but both the AOO and the extent of occurrence (EOO) are considered to exceed the threshold for Near Threatened or a threatened category.

The species has also been recorded from Calabria in southern peninsula Italy (Leonetti et al. 2020). The species is also introduced to several Aegean and Ionian islands (e.g. Amorgos, Andros, Chios, Corfu, Crete, Kefalonia, Lefkada, Lesvos, Milos, Naxos, Paros, Samos, Syros), Cyprus, and in southern Bulgaria (TTWG 2021, Tzoras et al. 2021, Escoriza et al. 2023). Introduced individuals (not mapped) have been reported in mainland Italy. In addition, newly reported cases of released or escaped individuals are cited from southern Catalonia (Spain; Pfau and Budó 2019).

The species also occurs in Sardinia and on some of its satellite islands, where it is fairly common (at least at the local scale), observed also on Isola di San Pietro off southwestern Sardinia. It may occur on Sardinia as a result of human introduction in the Classical era (see Bringsoe et al. 2001: 300); TTWG (2021) consider the Italian subpopulations to be the result of historical introduction, and Leonetti et al. (2020) consider the species to be naturalised in Italy. However, the Sardinian populations have been assigned to a distinct subspecies (T. m. sarda) with slight colour and morphological differences from conspecifics from elsewhere. The species has been reported for western Anatolia, Türkiye (Brinkmeier et al. 1989), however, this report required confirmation (Sindaco et al. 2000) and Türkozan et al. (2001) surveyed the area mentioned by Brinkmeier et al. (1989) and found only T. graeca ssp. ibera.

The species is generally considered to occur at low densities (Stubbs 1989). Willemsen (1995) and Bringsoe et al. (2001) noted that the preferred habitat of the species is very hard to penetrate and survey, thus records of low-density occurrence might pertain to marginal habitats and the true density may be higher than presently recorded. It is a very rare and localised species in Albania (Haxhiu 2004), whereas it is relatively easy to observe in Sardinia (L. Luiselli and M. Capula, unpublished observations; Corti emet al. 2022). The current population trend is considered to be stable.

A subpopulation in Sardinia, which was perceived as intensively collected in the 1950s, persists at 'normal' density (S. Fritz, in Bringsoe et al. 2001: 318). Populations of T. m. weissingeri are apparently dense, as Bour (1996) examined 384 animals and Artner (1996) encountered 30 animals in four hours, and 80 animals in two days. Bour (1996) recorded a male-dominated population structure, and very few juveniles (juveniles of T. marginata are very cryptic) (data summarised in Bringsoe et al. 2001: 317).

The species characteristically inhabits 'phrygana' and macchia scrub, dense thorny scrub among rocky outcrops, but also a range of other vegetation types, from coastal dunes to olive groves, small-scale agricultural landscapes and damp areas. It ranges up to 1,600 m (in the Peloponnese) but mostly occurs below 800 m. It generally prefers drier, hotter areas than the sympatric T. hermanni boettgeri (Willemsen 1995, Bringsoe et al. 2001). This taxon ranges from near sea level to about 1,300 m altitude (Mueller 1908, Bour 1996, in Bringsoe et al. 2001), and feeds on a variety of herbs (Bour 1996, in Bringsoe et al. 2001). The weissingeri ecotype (southern Peloponnese) inhabits both phrygana (Spartan shrubby rocky areas) as well as more lush habitats such as olive groves with dense herbaceous ground cover (Artner 2000, Bringsoe et al. 2001). In Sardinia, where it is considered a historical introduction, it often occurs in very dense Macchia scrub (L. Luiselli and M. Capula, unpublished observations).

The diet of this species in the wild has not been studied in detail, but observations on captive individuals suggest that it prefers herbaceous species, including Taraxacus, Bellis, Trifolium, Eruca, Brassica, etc.

There are two mating periods per year: in April-May and in September-October. Clutches consist of 2-17 eggs, but more often between two to seven eggs (Bour 1996, Artner 2000, Bringsoe et al. 2001). The females produce one to three clutches per year (review by Bringsoe et al. 2001), but more frequently twice per year (at least in Sardinia: L. Luiselli and M. Capula, unpublished data). The longevity in the wild of this species has not been carefully studied so far, but anecdotal evidence from captivity would suggest that it may live up to 140 years. Animals require about eight to 14 years to reach maturity (review by Bringsoe et al. 2001).

The main threat is wildfires. It is less impacted by agricultural land conversion and practices than the sympatric T. hermanni boettgeri due to T. marginata's preference for stony habitats (Bringsoe et al. 2001). Tourist infrastructure development eliminates some habitat, especially along coastal sites in Greece and in Sardinia. There is some mortality recorded from fires and trampling of juveniles by livestock, and destruction by farmers as perceived crop pests (Bringsoe et al. 2001).

In Greece, the Marginated Tortoise is mainly threatened by wildfires, as well as habitat modifications and agricultural activities, and it is also illegally collected and traded (Escoriza et al. 2023; C. Papoutsakis pers. comm. 2023).

There is some illegal collection for the pet trade or merely for placing these animals in own gardens as a recreative activity, might still take place. The death on roads of juveniles migrating from their birth sites to their feeding areas has been observed in Sardinia (L. Luiselli and M. Capula, unpublished observations), but it is not only juveniles but also adults for which the current road network may represent a serious global threat.

In northern Sardinia, a huge quantity of individuals are killed by road traffic particularly during the tourist seasons (SHI-Societas Herpetologica Italica, Sardinian Section and local Veterinary authority). In addition, the mechanical maintenance of road embankments and watercourses negatively affects tortoise populations, but it is unlikely that the current road network may represent a serious global threat for this species.

The species is often found in the pet trade, but as it can be easily bred in captivity, the great majority of the traded individuals are certainly captive-bred. Wild individuals are occasionally and opportunistically captured to be kept inside private gardens in Sardinia and Greece (L. Luiselli, M. Capula and E. Tzoras unpublished observations Corti et al. 2022) as well as many are captured to be taken to the Mainland) and very likely also in other areas of its geographic range.

Up to the beginning of the last century, the meat of this tortoise was certainly consumed by humans in several parts of its range. For instance, during the Middle Ages, these tortoises were kept alive in the gardens of monasteries in order to provide meat also during the periods in which, for religious reasons, the consumption of other types of meat was not allowed (De Grossi et al. 1999). This practice has however been terminated for many decades.

The species is included in CITES Appendix II, Annex A of EU Wildlife Trade Regulation 338/97, in Annex II and in Annex IV of the EU Habitats Directive, and in Annex I and Annex II of the Bern Convention (EEA 2022). The species is included in Greek domestic legislation and is protected in the whole Italian territory, although considered introduced there. The species is considered Least Concern in the draft National Red List assessment of the species (C. Papoutsakis pers. comm. 2023).

74 Natura 2000 sites include this species in their designations (EEA 2022).

Illegal collection, fires, road traffic and the use of machinery for the management of slopes and river banks are a serious problem for the Sardinian tortoises, and the same probably applies elsewhere. Hobbyist captive breeding appears to provide sufficient offspring to meet much of the demand (as this species is very easy to keep and breed in captivity), and the collection of wild specimens for the pet trade, although certainly occurring, is likely less strong nowadays compared to a few decades ago. However, at least in Sardinia, the local forest police at the ferry stations continue to seize individuals, often in large numbers, taken illegally from the wild. Establishment of effectively protected areas containing viable populations of the species would be desirable.

Further research is needed on taxonomy (especially with regard to the status of the weissingeri ecotype (Fritz et al. 2005, Escoriza et al. 2023), population dynamics and conservation biology. In particular, long-term studies on single populations are urgently needed.