Clockwise from left:
''[[Greek tortoise|Testudo graeca ibera]]''
''[[Herman's tortoise|Testudo hermanni boettgeri]]''
''[[Herman's tortoise|Testudo hermanni hermanni]]''
''[[Marginated tortoise|Testudo marginata sarda]]''."},"taxon":{"wt":"Testudo"},"authority":{"wt":"[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]"},"type_species":{"wt":"''[[Testudo graeca]]''"},"type_species_authority":{"wt":"[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]"},"subdivision_ranks":{"wt":"Species"},"subdivision":{"wt":""},"range_map":{"wt":"Testudo tortoises - distribution map.png"},"range_map_caption":{"wt":"Distribution of the genus ''Testudo'', showing overlaps and subspecies"},"synonyms":{"wt":"*''Furculachelys'' Highfield, 1990{{cite journal |url=http://images.turtleconservancy.org/documents/2017/crm-7-checklist-atlas-v8-2017.pdf |title=Turtles of the world, 2017 update: Annotated checklist and atlas of taxonomy, synonymy, distribution, and conservation status(8th Ed.)|journal=Chelonian Research Monographs |volume=7 |access-date=October 4, 2019|date=August 3, 2017 |last1=Rhodin | first1=Anders G.J. |last2=Inverson |first2=John B. |last3=Roger |first3=Bour |last4=Fritz |first4=Uwe |last5=Georges |first5=Arthur |last6=Shaffer |first6=H. Bradley |last7=van Dijk |first7=Peter Paul |isbn=978-1-5323-5026-9}}\n*''Chersus'' Gmira, 1993\n:and see text."}},"i":0}}]}" id="mwBA">
Testudo Temporal range: | |
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Four tortoises of the genus Testudo. Clockwise from left: Testudo graeca ibera Testudo hermanni boettgeri Testudo hermanni hermanni Testudo marginata sarda . | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | Testudines |
Suborder: | Cryptodira |
Superfamily: | Testudinoidea |
Family: | Testudinidae |
Genus: | Testudo Linnaeus, 1758 |
Type species | |
Testudo graeca | |
Distribution of the genus Testudo, showing overlaps and subspecies | |
Synonyms | |
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Testudo, the Mediterranean tortoises, are a genus of tortoises found in North Africa, Western Asia, and Europe. Several species are under threat in the wild, mainly from habitat destruction.
They are small tortoises, ranging in length from 7.0 to 35 cm and in weight from 0.7 to 7.0 kg.
The systematics and taxonomy of Testudo is notoriously problematic. Highfield and Martin commented:
Synonymies on Testudo are notoriously difficult to compile with any degree of accuracy. The status of species referred has undergone a great many changes, each change introducing an additional level of complexity and making bibliographic research on the taxa extremely difficult. Most early and not a few later checklists contain a very high proportion of entirely spurious entries, and a considerable number of described species are now considered invalid – either because they are homonyms, non-binomial or for some other reason. [2]
Since then, DNA sequence data have increasingly been used in systematics, but in Testudines (turtles and tortoises), its usefulness is limited: In some of these, at least mtDNA is known to evolve more slowly in these than in most other animals. [3] Paleobiogeographical considerations suggest the rate of evolution of the mitochondrial 12S rRNA gene is 1.0-1.6% per million years for the last dozen million years or so in the present genus [4] and ntDNA evolution rate has been shown to vary strongly even between different population of T. hermanni; [5] this restricts sequence choice for molecular systematics and makes the use of molecular clocks questionable.
The following extant species in the following subgenera are placed here:
The first two are more distinct and ancient lineages than the closely related latter three species. Arguably, T. horsfieldii belongs in a new genus (Agrionemys) on the basis of the shape of its carapace and plastron, [6] and its distinctness is supported by DNA sequence analysis. [7] Likewise, a separate genus Eurotestudo has recently been proposed for T. hermanni; these three lineages were distinct by the Late Miocene as evidenced by the fossil record. [8] Whether these splits will eventually be accepted remains to be seen. The genus Chersus has been proposed to unite the Egyptian and marginated tortoises which have certain DNA sequence similarities, [4] but their ranges are (and apparently always were) separated by their closest relative T. graeca and the open sea and thus, chance convergent haplotype sorting would better explain the biogeographical discrepancy.
Conversely, the Greek tortoise is widespread and highly diverse. In this and other species, a high number of subspecies has been described, but not all generally accepted, and several (such as the "Negev tortoise" and the "dwarf marginated tortoise") are now considered to be local morphs. Some, such as the Tunisian tortoise, have even been separated as a separate genus Furculachelys, but this is not supported by more recent studies. [9]
Testudo spp. are promiscuous creatures and they follow a polyandrous mating system. [10] Mating involves a courtship ritual of mechanical, olfactory and auditory displays elicited from the male to coerce a female into accepting copulation. [11] Courtship displays are very energetically costly for males, especially because females tend to run away from courting males. [12] The male will chase her, exerting more energy and elaborate displays of ramming and biting. Females are able to judge a male's genetic quality through these displays; only healthy males are able to perform costly courting rituals, suggesting endurance rivalry. [11] These are considered honest signals that are then used to influence pre- and post-copulatory choice, as females are the choosy sex. [10]
Female mate choice offers no direct benefits (such as access to food or territory or parental care). [13] There are, however, indirect benefits of mating with multiple males. Engaging in a polyandrous mating system offers a female guaranteed fertilization, higher offspring diversity and sperm competition to ensure that eggs are fertilized by a high quality male. This is in respect to the "good genes" hypothesis that females receive indirect benefits through her offspring by mating with a quality male, "a male's contribution to a female's fitness is restricted to [his] genes" (Cutuli, G. et al., 2014).
Mating order has no influence on paternity of a clutch so a female's inclination to mate with multiple males and her ability to store sperm allows for sperm competition and suggests cryptic female choice. [14] However, some species do show size-assortative, T. marginata, for example, where large males breed with large females and small males breed with small females. [11] Other species form hierarchies; during male-to-male competition the more aggressive male is considered alpha. [10] Alpha males are more aggressive with their courting as well and have higher mounting success rate than beta males.
A female's reproductive tract contains sperm storage tubules and she is capable of storing sperm for up to four years. [15] This sperm remains viable and when she goes a breeding season without encountering a male she is able to fertilize her eggs with the stored sperm. Storing sperm can also result in multiple paternity clutches; It is quite common among Testudo spp. females to lay a clutch that has been sired by multiple males. And females can lay one to four clutches a breeding season. Sexual dimorphism, promiscuity, long term sperm storage and elaborate courting rituals are factors that effect mate preference, sperm competition and cryptic female choice in genus Testudo. [10]
Tortoises are reptiles of the family Testudinidae of the order Testudines. Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like other members of the suborder Cryptodira, they retract their necks and heads directly backward into the shell to protect them.
The marginated tortoise is a species of tortoise in the family 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.
The Greek tortoise, also known commonly as the spur-thighed tortoise or Moorish tortoise, is a species of tortoise in the family Testudinidae. Testudo graeca is one of five species of Mediterranean tortoises. The other four species are Hermann's tortoise, the Egyptian tortoise, the marginated tortoise, and the Russian tortoise. The Greek tortoise is a very long-lived animal, achieving a lifespan upwards of 125 years, with some unverified reports up to 200 years. It has the largest known genome of all reptiles.
Hermann's tortoise is a species of tortoise native to Europe.
The Russian tortoise, also commonly known as the Afghan tortoise, the Central Asian tortoise, the four-clawed tortoise, the four-toed tortoise, Horsfield's tortoise, the Russian steppe tortoise, the Soviet Tortoise, and the steppe tortoise, is a threatened species of tortoise in the family Testudinidae. The species is endemic to Central Asia from the Caspian Sea south through Iran, Pakistan and Afghanistan, and east across Kazakhstan to Xinjiang, China. Human activities in its native habitat contribute to its threatened status.
The Asian forest tortoise, also known commonly as the Mountain tortoise or Burmese Brown Mountain tortoise, is a species of tortoise in the family Testudinidae. The species is endemic to Southeast Asia. It is believed to be among the most primitive of living tortoises, based on molecular and morphological studies.
Gopherus is a genus of fossorial tortoises commonly referred to as gopher tortoises. The gopher tortoise is grouped with land tortoises that originated 60 million years ago, in North America. A genetic study has shown that their closest relatives are in the Asian genus Manouria. The gopher tortoises live in the southern United States from California's Mojave Desert across to Florida, and in parts of northern Mexico. Gopher tortoises are so named because of some species' habit of digging large, deep burrows. Most notably, Gopherus polyphemus digs burrows which can be up to 40 feet (12 m) in length and 10 feet (3.0 m) in depth. These burrows are used by a variety of other species, including mammals, other reptiles, amphibians, and birds. Gopher tortoises are 20–50 cm (7.9–19.7 in) in length, depending on the species. All six species are found in xeric habitats. Numerous extinct species are known, the oldest dating to the Priabonian stage of the Late Eocene of the United States.
Geochelone is a genus of tortoises.
The pancake tortoise is a species of flat-shelled tortoise in the family Testudinidae. The species is native to Tanzania and Kenya. There are also small populations in northern Zambia. Its common name refers to the flat shape of its shell.
Aldabrachelys is genus of giant tortoises, including the Aldabra giant tortoise native to the Seychelles, as well as two extinct species, Aldabrachelys abrupta and Aldabrachelys grandidieri known from Madagascar. The genus name derives from Aldabra, a coral atoll in the Seychelles, plus chelys, from the Greek, χέλυς, for "tortoise" or "lyre", since Hermes was the first to invent the lyre from a hollowed out tortoise shell.
The Tunisian tortoise or Nabeul tortoise is a subspecies of Greek tortoises. It was originally described as a new species in 1990, and even placed in a distinct genus. The spur-thighed or "Greek" tortoises are usually collectively referred to as Testudo graeca, but this covers a wide variety of subspecies that have very different ecological and morphological characteristics and appear to comprise at least three phylogenetic lineages. As its name implies, it is found in Tunisia and nearby Algeria.
Kleinmann's tortoise, also called commonly the Egyptian tortoise, Leith's tortoise, and the Negev tortoise, is a critically endangered species of cryptodire turtle in the family Testudinidae. The species is native to Libya and possibly extinct in Egypt. The species was once more widespread, but its numbers are now dwindling, and complete extinction in the wild is a looming threat unless more actions are taken to protect this species.
The yellow-footed tortoise, also known as the Brazilian giant tortoise, is a species of tortoise in the family Testudinidae and is closely related to the red-footed tortoise. It is found in the Amazon Basin of South America. The species name has often been misspelled as denticulata, an error introduced in the 1980s when Chelonoidis was elevated to genus and mistakenly treated as feminine, an error recognized and fixed in 2017.
A spur is an outgrowth of bone covered in a sheath of horn found in various anatomical locations in some animals. Unlike claws or nails, which grow from the tip of the toes, spurs form from other parts of the foot, usually in connection with joints where the toes meet the foot or the foot meets the long bones. Spurs are most commonly found on the hindfeet, though some birds possess spurs at the leading edge of the wings.
In behavioral ecology, polyandry is a class of mating system where one female mates with several males in a breeding season. Polyandry is often compared to the polygyny system based on the cost and benefits incurred by members of each sex. Polygyny is where one male mates with several females in a breeding season . A common example of polyandrous mating can be found in the field cricket of the invertebrate order Orthoptera. Polyandrous behavior is also prominent in many other insect species, including the red flour beetle, the adzuki bean weevil, and the species of spider Stegodyphus lineatus. Polyandry also occurs in some primates such as marmosets, mammal groups, the marsupial genus' Antechinus and bandicoots, around 1% of all bird species, such as jacanas and dunnocks, insects such as honeybees, and fish such as pipefish.
Titanochelon is an extinct genus of giant tortoises known from the Early Miocene to the beginning of the Pleistocene in Europe, extending from the Iberian Peninsula to Anatolia. Some members of the genus were larger than extant giant tortoises, with a shell length of up to 2 m.
Solitudo is an extinct genus of tortoise that was found during the Pliocene and Pleistocene on the Mediterranean islands of Menorca, Malta and Sicily. The genus includes three described species, Solitudo robusta, Solitudo gymnesica and Solitudo sicula as well as a likely fourth, undescribed species from Monte Pellegrino in Sicily. Solitudo sicula, the youngest of the species, died out approximately 12.5 thousand years BP. The largest species, Solitudo gymnesica, has been estimated to have reached a carapace length of 1.1–1.3 m (3.6–4.3 ft).
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