Marginated tortoise

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Marginated tortoise
Left: Greek marginated tortoise (T. m. marginata)
Right: Sardinian marginated tortoise (T. m. sarda)

Least Concern  (IUCN 3.1)[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Testudines
Suborder: Cryptodira
Superfamily: Testudinoidea
Family: Testudinidae
Genus: Testudo
Species:
T. marginata
Binomial name
Testudo marginata
Schoepff, 1789
Subspecies
  • Greek marginated tortoise (T. m. marginata)
    — Schoepff, 1789
  • Sardinian marginated tortoise (T. m. sarda)
    — Mayer, 1992
Synonyms[2]
  • Testudo marginata
    Schoepff, 1793
  • Testudo graja
    Hermann, 1793
  • Chersine marginata
    Merrem, 1820
  • Chersus marginatus
    Wagler, 1830
  • Testudo campanulata
    Gray, 1831 (nomen nudum)
  • Testudo graji
    Gray, 1831 (
    ex errore
    )
  • Testudo campanulata
    Strauch
    , 1862
  • Peltastes marginatus
    — Gray, 1869
  • Peltastes marginata
    — Gray, 1872
  • Testudo nemoralis
    Schreiber, 1875
  • Testudo marginata sarda
    Mayer, 1992
  • Testudo marginata weissingeri
    Trutnau, 1994
  • Testudo weissingeri
    Bour, 1996
  • Testudo marginata marginata
    — Rogner, 1996

The marginated tortoise (Testudo marginata) is a

Testudinidae. The species is endemic to Greece, Italy, and the Balkans in Southern Europe. It is the largest European tortoise. The marginated tortoise is herbivorous, and brumates for the winter.[3]

Taxonomy

Carapace shapes of Greek (left) and Sardinian marginated tortoises

The marginated tortoise was formally described by German naturalist Johann David Schoepff in 1789; its specific epithet marginata is a straightforward derivation from the Latin term for 'marginated'.

The

nominate subspecies is the Greek marginated tortoise, Testudo marginata marginata. Three additional subspecies
of marginated tortoises have been named:

The Sardinian marginated tortoise (T. m. sarda) is the name usually used to separate the

Lineage sorting
has not occurred to a considerable degree; consequently, the more geographically isolated Sardinian population is presumably of quite recent origin.

Indeed, it appears to derive from a deliberate introduction by humans.

An

last ice age, presumably larger than mainland tortoises.[7] However, numerous additional fossils have since refuted the validity of this subspecies.[8]

A population of small and light-colored marginated tortoises exists on the southwestern coast of the Peloponnesus, between

apomorphies seem related to this. Considering ice age-associated climate and sea level changes, this population is probably not older than a few thousand years; as it is not geographically isolated, it should be considered a local form, and not even a subspecies T. m. weissingeri. Notably, a similar situation is found in Hermann's tortoises living in the same region.[9][4]

Testudo marginata is also closely related to the Greek or

common tortoise (Testudo graeca). Both have very similar bodily characteristics—oblong carapaces, large scales on the front legs, large coverings for the head and cone-like scales on the upper thighs, undivided tail coverings, moveable stomach plates, and lack of tail spikes. Presumably, Testudo marginata diverged from ancestral Testudo graeca as a population more suited for life in the mountainous regions. Evidence in favor of this is the wide geographical region and the extremely large number of subspecies of Testudo graeca, including a subspecies in Turkey with strongly bent carapace tiles, like the marginated tortoise. Testudo marginata on the other hand, despite the two subspecies, presents a much more unified appearance, which points toward an earlier appearance in evolutionary
history. In captivity, the two species often cross-breed, but this should be avoided.

According to the 2005 DNA sequence data,

Mediterranean, or the supposed "clade" is invalid and the similarity due to convergent evolution
.

Description

The marginated tortoise is the largest European tortoise, reaching a weight of up to 5 kg (11 lb) and a length of 35 cm (14 in). Its

eggs in the soil
in May and June.

Distribution and habitat

Close-up of fore limbs and head, showing the particularly large scales

The natural range of the marginated tortoise is southern

Peloponnesus to Mount Olympus. They are also found in isolated zones of the Balkans and Italy, and northeastern Sardinia
.

The marginated tortoise lives in more mountainous regions than Hermann's tortoise. It can be found in elevations as high as 1,600 m (5,200 ft). The black color of the carapace is helpful for survival in this environment, as it allows the tortoise to absorb a great deal of heat in a short time, helping it maintain its

body temperature
. Early in the morning, marginated tortoises bask in the sun to raise their body temperature, and then search for food. After feeding, the tortoises return to their shelters in the hot midday hours, leaving them again in the late afternoon.

Behaviour

Marginated tortoises are fairly calm and relaxed, although they can be somewhat territorial in the wild. They have a controlled temper and are generally slow to anger. If they are not given the proper diet in captivity, however, they will become rather aggressive and might mistakenly attack if they feel threatened.

Diet

Marginated tortoises are herbivorous, their diets consisting primarily of plants from their native Mediterranean region.

Reproduction

Immediately after waking from

copulation
, the male opens his mouth, showing his red tongue and making loud cries. The tone of the copulation cry is almost sobbing with long, deep tones, in contrast to T. hermanni, which uses a much higher-toned, peeping noise.

During mating, the female stands still and holds her head to the side, looking up to see the opened mouth of the male. The red tongue apparently serves a signalling function. The female moves her head from left to right in the same rhythm as the male's cries.

Afterwards, the female seeks out an adequate location to lay her eggs. Once such a place is found, the female stands still, propping both front legs firmly against the ground. Then she digs out a hole with her hind legs, alternating between left and right, beginning with simply scratching the ground but eventually moving large quantities of soil which are piled up beside the hole. The depth of the hole is determined by the length of her hind legs. If the ground is too hard to dig, the female releases water from her anal gland to soften it.

Once the hole is dug, egg-laying begins. Each egg is gently rolled back into the hole. After the last egg, the female immediately begins refilling the hole, again using her hind legs. Finally, she stamps the opening closed with her feet so that the ground regains its natural hardness. Larger animals may lay eggs as many as three times per summer, with about 15 eggs per clutch.

A marginated tortoise emerges into the world.

The incubation period averages about 100 days under natural conditions, which is relatively short among tortoises. Many tropical tortoises have incubation periods of up to 200 days. The relatively short time is an adaptation to the subtropical Mediterranean climate, where the summers are not as long. In an incubator, this time is notably shorter: with an incubation temperature of 31.5 °C (88.7 °F) the eggs will begin hatching after 60 days.

Unlike bird eggs, the

albumen of reptile eggs are not separated by a membrane. After a few days, the heavy yolk components sink to the bottom of the egg. On top of this floats the embryonal
disk, surrounded by albumen, so the tortoise eggs cannot be turned after the yolk settles without damaging or killing the embryo.

It is possible to see with the naked eye if the eggs are developing healthily. Freshly laid eggs have a gray-white color. Shortly thereafter, a bright white spot forms on the uppermost point of the egg. This spot gradually grows until the entire egg is bright white.

After the embryo has developed fully in the egg, the young animal breaks the shell with its egg tooth from inside, creates a small opening, and for the first time fills its lungs with air. Afterwards, it pulls back into the egg and works on the shell with its beak until it opens completely. In nature, the animal remains below ground for the first two weeks, where it is safe from predators, yet is still able to grow, as it is nourished by the yolk sac. The young animals lead cautious and secretive lives, normally remaining in the shade. They avoid full sunlight because of the great danger of overheating.

Marginated tortoises grow very rapidly. In an ideal biotope, or with good handling, they gain 100–500 g (3.5–17.6 oz) yearly. This quick rate of growth lasts throughout their youth. After the 20th year of life, further growth is minimal. They may live between 100 and 140 years, according to the best estimates of scientists.

  • Two marginated tortoises mating
    Two marginated tortoises mating
  • Building a nest burrow
    Building a nest burrow
  • Finished burrow
    Finished burrow
  • Laying eggs
    Laying eggs
  • Free at last
    Free at last
  • Egg shells with skins
    Egg shells with skins
  • T. marginata young
    T. marginata young
  • Terrarium for raising young
    Terrarium for raising young

In captivity

The caudal plate (above the tail) is not divided as in Hermann's tortoise.

Gallery

  • A male T. marginata marginata, identified by the long tail with broad base
    A male T. marginata marginata, identified by the long tail with broad base
  • A female T. marginata with a broad-edged carapace: The cloacal opening is visible on the tail.
    A female T. marginata with a broad-edged carapace: The cloacal opening is visible on the tail.
  • A young T. marginata on its back
    A young T. marginata on its back
  • Old T. m. sarda
    Old T. m. sarda
  • T. marginata
    T. marginata
  • Greek tortoise, T. graeca
    Greek tortoise, T. graeca
  • Hybrid, T. marginata × T. graeca (father × mother)
    Hybrid, T. marginata × T. graeca (father × mother)

See also

References

  1. . Retrieved 12 November 2021.
  2. .
  3. .
  4. ^ a b c d Fritz et al. (2005)
  5. ^ Avise et al. (1992)
  6. ^ van der Kuyl et al. (2002)
  7. ^ Georgalis et al. (2013)
  8. ^ "The fossil record of turtles from the Pleistocene of Crete (Greece)". Comptes Rendus Palevol. 21 (35): 771–799. 2022.
  9. ^ Fritz et al. (2006)
  10. ^ van der Kuyl et al. (2002), Fritz et al. (2005)

Bibliography

External links