Testudo marginata

Schoepff, 1792
Marginated tortoise

Recognition
The elongated carapace (to 38 cm) is domed and highest just behind the center with abruptly descending sides, a weak cervical notch, and the serrated posterior rim greatly flared. A narrow cervical scute is present. Vertebrals are wider than long; the 5th is expanded. Areolae of the vertebral and pleural scutes may be raised slightly and are surrounded by growth annuli. Eleven marginals lie on each side; the undivided supracaudal is very flared, as also are the posterior marginals. The carapace is black to dark brown with yellow vertebral and pleural areolae, and a yellow bar on each marginal. The plastron is well-developed, and its forelobe upturned, tapered toward the front, and shorter and narrower than the hindlobe, which bears a posterior notch. The plastral formula is: abd > an > hum >< pect >< fem > gul; the paired gulars are thickened and may project slightly beyond the carapacial rim. A single axillary and one or two moderately sized inguinals which may touch the femoral scute occur on the broad bridge. Plastron and bridge are yellow, with some dusky pigment on the bridge and two rows of large black triangular blotches along the plastron. The head is of moderate size with a nonprotruding snout and a hooked upper jaw. Its large prefrontal and frontal scales may be longitudinally divided or subdivided; other head scales are small. The head is black, the chin yellow or gray, and the jaws black or brown. Limbs are brown to yellowish brown. Four or five longitudinal rows of large overlapping scales lie on the anterior surface of each foreleg; each forefoot has five claws. Spurlike scales lie on each heel, and weak tubercles may occur on the thighs. The tail may lack a terminal claw.
In wild populations, males grow slightly larger than females; however, the largest known captives are females. Males also have broader heads, more strongly flared rear marginals, and longer, thicker tails with the vent near the tip.

Distribution
Testudo marginata ranges from southwestern Albania through Greece and on the Aegean Islands of Skyros and Poros; introduced populations have been established in Sardinia and Tuscany, Italy (Iverson, 1992).

Geographic Variation
The Sardinian population was described as a separate subspecies (Testudo marginata sarda Mayer, 1992), but Fritz et al. (1995) regarded the characters of adults as persisting juvenile characters and synonymized this form with T. marginata.

Habitat
Today T. marginata is found most often in rocky, dry scrub (phrygana, garrigue) hillsides close to cultivated areas (Stubbs, 1989b); it sometimes enters olive groves. After emergence from hibernation in the spring, the tortoises often move down into the taller, grassy vegetation bordering cultivated fields where there is more food and possibly better nesting sites.

Natural History
Little is known of this tortoise, and most data come from captive individuals. Mating usually occurs in the spring (April to early June), but Clark (1963) observed a wild pair in courtship on the island of Spetsai in the fall (mistakenly reported as Testudo graeca ibera, R. J. Clark, pers. comm.). Courtship is aggressive and consists of chasing, butting of the female's shell, and biting of her head and limbs (Hine, 1982). Hine (1982) reported that females respond to the male's guttural utterances with deliberate head movements from side to side.
Eggs are laid during June and July in excavations about 10 cm deep. In captivity, some females fail to dig a nest and instead deposit the eggs at random about their enclosures. Clutches range from 3 to 11 eggs; hatching occurs from late August into October, depending on soil temperatures. Eggs are nearly spherical to ellipsoidal (31-37 x 27-36 mm) with brittle shells.
Hatchlings have rounded carapaces (about 35 mm) that lack posterior flaring. The plastral hinge is also nonfunctional. Each carapacial scute has a yellow areolus surrounded by a dark border.
Testudo marginata is herbivorous and will accept a wide variety of grasses, flowers, and fruits in captivity.

IUCN Red List Status (1996)
Not listed.

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