Marginated Tortoise
Encyclopedia
The Marginated tortoise (Testudo marginata) is a species of tortoise
Tortoise
Tortoises are a family of land-dwelling reptiles of the order of turtles . Like their marine cousins, the sea turtles, tortoises are shielded from predators by a shell. The top part of the shell is the carapace, the underside is the plastron, and the two are connected by the bridge. The tortoise...

 found in Greece, Italy and the Balkans in southern Europe. It is the largest Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

an tortoise
Tortoise
Tortoises are a family of land-dwelling reptiles of the order of turtles . Like their marine cousins, the sea turtles, tortoises are shielded from predators by a shell. The top part of the shell is the carapace, the underside is the plastron, and the two are connected by the bridge. The tortoise...

, reaching a weight of up to 5 kg
Kilogram
The kilogram or kilogramme , also known as the kilo, is the base unit of mass in the International System of Units and is defined as being equal to the mass of the International Prototype Kilogram , which is almost exactly equal to the mass of one liter of water...

 (11 pound
Pound (mass)
The pound or pound-mass is a unit of mass used in the Imperial, United States customary and other systems of measurement...

s) and a length of 35 cm (14 inches). Its shell is oblong and has a notable thickness around the middle of the body. The posterior end of the shell has a saw
Saw
A saw is a tool that uses a hard blade or wire with an abrasive edge to cut through softer materials. The cutting edge of a saw is either a serrated blade or an abrasive...

-like formation, flanged outward like a bell
Bell (instrument)
A bell is a simple sound-making device. The bell is a percussion instrument and an idiophone. Its form is usually a hollow, cup-shaped object, which resonates upon being struck...

. The carapace
Carapace
A carapace is a dorsal section of the exoskeleton or shell in a number of animal groups, including arthropods such as crustaceans and arachnids, as well as vertebrates such as turtles and tortoises. In turtles and tortoises, the underside is called the plastron.-Crustaceans:In crustaceans, the...

 of adult specimens is almost completely black
Black
Black is the color of objects that do not emit or reflect light in any part of the visible spectrum; they absorb all such frequencies of light...

, with yellow
Yellow
Yellow is the color evoked by light that stimulates both the L and M cone cells of the retina about equally, with no significant stimulation of the S cone cells. Light with a wavelength of 570–590 nm is yellow, as is light with a suitable mixture of red and green...

 highlights. The front sides of the limbs are covered with large scales
Scale (zoology)
In most biological nomenclature, a scale is a small rigid plate that grows out of an animal's skin to provide protection. In lepidopteran species, scales are plates on the surface of the insect wing, and provide coloration...

. The tail
Tail
The tail is the section at the rear end of an animal's body; in general, the term refers to a distinct, flexible appendage to the torso. It is the part of the body that corresponds roughly to the sacrum and coccyx in mammals, reptiles, and birds...

 is notable for a lengthwise marking and for an undivided carapace over the tail. The Marginated Tortoise is herbivorous, and hibernates for the winter.

Taxonomy

The marginated tortoise was formally described by German naturalist Johann David Schoepf
Johann David Schoepf
Johann David Schoepff, or Schoepf, or Schöpf, was a German botanist, zoologist, and physician....

 in 1789, its specific epithet marginata a straightforward derivation from the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 term for 'marginated'.

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

The Sardinian marginated tortoise (Testudo marginata sarda) is the name usually used to separate the population
Population
A population is all the organisms that both belong to the same group or species and live in the same geographical area. The area that is used to define a sexual population is such that inter-breeding is possible between any pair within the area and more probable than cross-breeding with individuals...

 on Sardinia. These tortoises have less strongly bent tiles in the posterior of the carapace, and the posterior of the carapace is almost smooth compared with the saw-like Testudo marginata marginata. Clearly distinct according to morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

 and entirely allopatric, it cannot be distinguished by mtDNA cytochrome b
Cytochrome b
Cytochrome b/b6 is the main subunit of transmembrane cytochrome bc1 and b6f complexes. In addition, it commonly refers to a region of mtDNA used for population genetics and phylogenetics.- Function :...

 and nDNA ISSR sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

 analysis. 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. Though it is not clear whether this occurred in prehistoric times or more recently, this might be resolved by dating subfossil
Subfossil
Subfossil refers to remains whose fossilization process is not complete, either for lack of time or because the conditions in which they were buried were not optimal for fossilization....

 remains. Sequence evolution at least in mtDNA is known to proceed much more slowly in some turtles and tortoises than in others; the rate of the mitochondrial 12S rRNA gene in Testudo is probably a rather low 1-1.63% per million years (as this fits best the paleobiogeographical situation), limiting the resolution provided by molecular systematics.

An extinct subspecies described as T. marginata cretensis persisted on Crete
Crete
Crete is the largest and most populous of the Greek islands, the fifth largest island in the Mediterranean Sea, and one of the thirteen administrative regions of Greece. It forms a significant part of the economy and cultural heritage of Greece while retaining its own local cultural traits...

 before the end of the last ice age and perhaps into early historic times

A population of small and light-colored marginated tortoises exists on the southwestern coast of the Peloponnesus, between Kalamata
Kalamata
Kalamata is the second-largest city of the Peloponnese in southern Greece. The capital and chief port of the Messenia prefecture, it lies along the Nedon River at the head of the Messenian Gulf...

 and south of Stoupa
Stoupa
Stoupa is a village on the coast of the southern Peloponnese peninsula in Greece. It is part of the municipal unit of West Mani, in Messenia and the historic region of Mani Peninsula....

. The so-called "Dwarf marginated tortoise" was described as a new species Testudo weissingeri, but it is not recognizably distinct phylogenetically. Unlike the Sardinian population, it occupies an extremely restricted range with very arid habitat, and its apomorphies seem related to this. Considering ice age
Ice age
An ice age or, more precisely, glacial age, is a generic geological period of long-term reduction in the temperature of the Earth's surface and atmosphere, resulting in the presence or expansion of continental ice sheets, polar ice sheets and alpine glaciers...

-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 Testudo marginata weissingeri. It is notable that a similar situation is found in Hermann's Tortoises living in the same region.

Testudo marginata is also closely related to the Greek tortoise
Greek Tortoise
The spur-thighed tortoise or Greek tortoise is one of four European members of the Testudinidae family of tortoises...

 or ppur-thighed tortoise (Testudo graeca). Both have very similar bodily characteristics, an oblong carapace, large scales on the front legs, large covering for the head and cone-like scales on the upper thigh, undivided tail covering, moveable stomach plates, and lack of a tail spike. 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
Turkey
Turkey , known officially as the Republic of Turkey , is a Eurasian country located in Western Asia and in East Thrace in Southeastern Europe...

 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 evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

ary history. In captivity, the two species often cross-breed, but this should be avoided.

According to the 2005 DNA sequence data, these species do not seem to hybridize to a notable extent in the wild, though they are obviously very close relatives, and as evidenced bby morphology some allele
Allele
An allele is one of two or more forms of a gene or a genetic locus . "Allel" is an abbreviation of allelomorph. Sometimes, different alleles can result in different observable phenotypic traits, such as different pigmentation...

 flow still occurs, but slowly. The Egyptian tortoise appears to represent a lineage that diverged from the same ancestral stock southwards into northeastern Africa around the same time as the marginated tortoise's ancestors diverged in Greece. These two are actually more similar to each other than to T. graeca regarding DNA sequence data, but considering biogeography, this is either due to (rather unlikely) dispersal across the Mediterranean, or the supposed "clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

" is invalid and the similarity due to convergent evolution
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...

.

Description

The marginated tortoise is the largest Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

an tortoise
Tortoise
Tortoises are a family of land-dwelling reptiles of the order of turtles . Like their marine cousins, the sea turtles, tortoises are shielded from predators by a shell. The top part of the shell is the carapace, the underside is the plastron, and the two are connected by the bridge. The tortoise...

, reaching a weight of up to 5 kg
Kilogram
The kilogram or kilogramme , also known as the kilo, is the base unit of mass in the International System of Units and is defined as being equal to the mass of the International Prototype Kilogram , which is almost exactly equal to the mass of one liter of water...

 (11 pound
Pound (mass)
The pound or pound-mass is a unit of mass used in the Imperial, United States customary and other systems of measurement...

s) and a length of 35 cm (14 inches). Its shell is oblong and has a notable thickness around the middle of the body. The posterior end of the shell has a saw
Saw
A saw is a tool that uses a hard blade or wire with an abrasive edge to cut through softer materials. The cutting edge of a saw is either a serrated blade or an abrasive...

-like formation, flanged outward like a bell
Bell (instrument)
A bell is a simple sound-making device. The bell is a percussion instrument and an idiophone. Its form is usually a hollow, cup-shaped object, which resonates upon being struck...

. The carapace
Carapace
A carapace is a dorsal section of the exoskeleton or shell in a number of animal groups, including arthropods such as crustaceans and arachnids, as well as vertebrates such as turtles and tortoises. In turtles and tortoises, the underside is called the plastron.-Crustaceans:In crustaceans, the...

 of adult specimens is almost completely black
Black
Black is the color of objects that do not emit or reflect light in any part of the visible spectrum; they absorb all such frequencies of light...

, with yellow
Yellow
Yellow is the color evoked by light that stimulates both the L and M cone cells of the retina about equally, with no significant stimulation of the S cone cells. Light with a wavelength of 570–590 nm is yellow, as is light with a suitable mixture of red and green...

 highlights. The ventral shell is lighter coloured and has pairs of triangular markings with the points facing the rear of the animal. The front sides of the limbs are covered with large scales
Scale (zoology)
In most biological nomenclature, a scale is a small rigid plate that grows out of an animal's skin to provide protection. In lepidopteran species, scales are plates on the surface of the insect wing, and provide coloration...

. In old female
Female
Female is the sex of an organism, or a part of an organism, which produces non-mobile ova .- Defining characteristics :The ova are defined as the larger gametes in a heterogamous reproduction system, while the smaller, usually motile gamete, the spermatozoon, is produced by the male...

 specimens, the rear flaps of the underside of the plastron are somewhat moveable. The tail
Tail
The tail is the section at the rear end of an animal's body; in general, the term refers to a distinct, flexible appendage to the torso. It is the part of the body that corresponds roughly to the sacrum and coccyx in mammals, reptiles, and birds...

 is notable for a lengthwise marking and for an undivided carapace over the tail. Male
Male
Male refers to the biological sex of an organism, or part of an organism, which produces small mobile gametes, called spermatozoa. Each spermatozoon can fuse with a larger female gamete or ovum, in the process of fertilization...

s have a longer tail, which is thicker at the base than the females. Their underside is more strongly indented. Males are also often larger than the females. The females lay their hard-shelled spherical eggs
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...

 in the soil
Soil
Soil is a natural body consisting of layers of mineral constituents of variable thicknesses, which differ from the parent materials in their morphological, physical, chemical, and mineralogical characteristics...

 in May and June.

Distribution and habitat

The natural range of the marginated tortoise is southern Greece
Greece
Greece , officially the Hellenic Republic , and historically Hellas or the Republic of Greece in English, is a country in southeastern Europe....

, from the Peloponnesus to Mount Olympus
Mount Olympus
Mount Olympus is the highest mountain in Greece, located on the border between Thessaly and Macedonia, about 100 kilometres away from Thessaloniki, Greece's second largest city. Mount Olympus has 52 peaks. The highest peak Mytikas, meaning "nose", rises to 2,917 metres...

. They are also found in isolated zones of the Balkans
Balkans
The Balkans is a geopolitical and cultural region of southeastern Europe...

 and Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

, and northeastern Sardinia
Sardinia
Sardinia is the second-largest island in the Mediterranean Sea . It is an autonomous region of Italy, and the nearest land masses are the French island of Corsica, the Italian Peninsula, Sicily, Tunisia and the Spanish Balearic Islands.The name Sardinia is from the pre-Roman noun *sard[],...

.

The marginated tortoise lives in more mountainous regions than Hermann's Tortoise (Testudo hermanni). It can be found in elevations as high as 1,600 m (5,250 feet). 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.

Diet

Marginated tortoises are herbivorous, their diet consisting primarily of plants from their native Mediterranean region. In captivity, the primary foodstuffs are dandelions, clover
Clover
Clover , or trefoil, is a genus of about 300 species of plants in the leguminous pea family Fabaceae. The genus has a cosmopolitan distribution; the highest diversity is found in the temperate Northern Hemisphere, but many species also occur in South America and Africa, including at high altitudes...

, salad leaves such as rocket or watercress
Watercress
Watercresses are fast-growing, aquatic or semi-aquatic, perennial plants native from Europe to central Asia, and one of the oldest known leaf vegetables consumed by human beings...

, and also leaves from plantain, radish
Radish
The radish is an edible root vegetable of the Brassicaceae family that was domesticated in Europe, in pre-Roman times. They are grown and consumed throughout the world. Radishes have numerous varieties, varying in size, color and duration of required cultivation time...

, and crowfoot plants. Sometimes even grass and hay will be eaten. The feeding of fruit and tomatoes is to be discouraged as they may cause diarrhea. Lettuce will be eagerly devoured, but owing to its low nutrient value should not generally be given. Protein levels in the diet should be kept very low and in no account should they be given sources of animal protein such as meat, snails, worms, and cat, dog or terrapin food.

Additionally they should be given daily calcium supplements that are essential for a correct shell development. A mixture of various calcium salts is best, mainly calcium lactate
Calcium lactate
Calcium lactate is a white crystalline salt made by the action of lactic acid on calcium carbonate. It is used in foods and given medicinally. Its E number is E327.Calcium lactate is often found in aged cheeses...

, and calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...

, with no phosphorus
Phosphorus
Phosphorus is the chemical element that has the symbol P and atomic number 15. A multivalent nonmetal of the nitrogen group, phosphorus as a mineral is almost always present in its maximally oxidized state, as inorganic phosphate rocks...

 content.

Phosphorus inhibits absorption of calcium and so a high phosphorus content in the diet can cause abnormal development ("shell pyramiding") even though calcium supplements are given on a daily basis.

Reproduction

Immediately after waking from hibernation
Hibernation
Hibernation is a state of inactivity and metabolic depression in animals, characterized by lower body temperature, slower breathing, and lower metabolic rate. Hibernating animals conserve food, especially during winter when food supplies are limited, tapping energy reserves, body fat, at a slow rate...

 the mating instinct starts up. The males follow the females with great interest, encircling them, biting them on the limbs, ramming them, and trying to mount them. During 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 Testudo hermanni, in which there is 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. It appears that the red tongue 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
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...

. 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.
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
Celsius
Celsius is a scale and unit of measurement for temperature. It is named after the Swedish astronomer Anders Celsius , who developed a similar temperature scale two years before his death...

 (89 °F
Fahrenheit
Fahrenheit is the temperature scale proposed in 1724 by, and named after, the German physicist Daniel Gabriel Fahrenheit . Within this scale, the freezing of water into ice is defined at 32 degrees, while the boiling point of water is defined to be 212 degrees...

) the eggs will begin hatching after 60 days.

Unlike bird eggs, the yolk and albumen of reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...

 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 embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...

nal disk, surrounded by albumen. For this reason 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
Egg tooth
In some egg-laying animals, the egg tooth is a small, sharp, cranial protuberance used by offspring to break or tear through the egg's surface during hatching...

 from inside, creates a small opening, and for the first time fill its lung
Lung
The lung is the essential respiration organ in many air-breathing animals, including most tetrapods, a few fish and a few snails. In mammals and the more complex life forms, the two lungs are located near the backbone on either side of the heart...

s 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. Here 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
Biotope
Biotope is an area of uniform environmental conditions providing a living place for a specific assemblage of plants and animals. Biotope is almost synonymous with the term habitat, but while the subject of a habitat is a species or a population, the subject of a biotope is a biological community.It...

, or with good handling, they gain 100–500 g
Gram
The gram is a metric system unit of mass....

 (4–17 oz
Ounce
The ounce is a unit of mass with several definitions, the most commonly used of which are equal to approximately 28 grams. The ounce is used in a number of different systems, including various systems of mass that form part of the imperial and United States customary systems...

) yearly. This quick rate of growth lasts throughout their youth. After the twentieth year of life, further growth is minimal. They may live to between 100 and 140 years, according to the best estimates of scientists.

In captivity

In a terrarium

Tortoises can thrive in captivity if the owner understands their needs well. A heat lamp is attached to the tortoise's table, directed in such a way that the 60-watt
Watt
The watt is a derived unit of power in the International System of Units , named after the Scottish engineer James Watt . The unit, defined as one joule per second, measures the rate of energy conversion.-Definition:...

 reflector bulb is some 15–20 cm (6–8 in) from the floor of the table, which is covered with about 5 cm of substraite, loam based soil and play sand. When the lamp is turned on in the morning, the animals emerge from the ground, bask in the light to warm themselves, and begin to eat. They should be fed several times a day with clover, dandelions and garden weeds (check first as a few are harmful. Some owners give pellet food occasionally, these should be avoided, only feed your tortoise what they would naturally eat in the wild, their digestive systems do not cope well with protein, of which there is a lot in pellet foods. The animals also require ultraviolet light, thus they should be allowed to bask in the sun
Sun
The Sun is the star at the center of the Solar System. It is almost perfectly spherical and consists of hot plasma interwoven with magnetic fields...

light daily, but for only a short period of time. In the summer, they can be taken outdoors for this purpose, in the winter if not hybernated they need access to UVB light every day. Care must be taken to avoid allowing the tortoise to overheat to help, use water in a shallow dish or(use a digital thermometer to keep a check).

Outdoors

In temperate zones, marginated tortoises can be kept outside from approximately mid-March to October. Their pen should be in the sunniest place in the garden, preferably close to the house. It is important to provide a wooden house where they can get into the shade. They should be provided enough gravel
Gravel
Gravel is composed of unconsolidated rock fragments that have a general particle size range and include size classes from granule- to boulder-sized fragments. Gravel can be sub-categorized into granule and cobble...

that they can completely bury themselves when needed.

The animals will leave the house in the morning, warm themselves, and eat. Afterwards, they return to the house. In the late afternoon, they will reemerge from their shelter. Tortoises do not need as much sunlight as many assume is required for a reptile. By October, they will take longer and longer pauses during the day. At this point, they should be transferred into a roomy crate filled with dry leaves and kept in a cool room. With a temperature under 10 °C (50 °F), they will hibernate until mid-March, though younger animals will awaken much earlier.

External links

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