Solitudo Temporal range: | |
---|---|
Material of Solitudo robusta | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | Testudines |
Suborder: | Cryptodira |
Superfamily: | Testudinoidea |
Family: | Testudinidae |
Subfamily: | Testudininae |
Genus: | † Solitudo Valenti et al. 2022 |
Type species | |
†Solitudo robusta (Leith-Adams, 1877) [1] | |
Other species | |
|
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).
The oldest discoveries of fossil turtles now included under Solitudo were made in the 19th century, with Leith-Adams describing remains from Zebbug Cave (Malta) as Testudo robusta. In 1914 Testudo gymnesica was described based on Pliocene material found on Menorca. [2] Younger material was later discovered in the Zubbio di Cozzo San Pietro cave in northern Sicily. These remains, which include a femur, phalanx, pubis and an ischium, were found in what is thought to be a prehistoric funeral site, however repeated human or animal activity changed the stratigraphy of the site. Due to this the turtle remains, although suspected to have belonged to a single individual, were dispersed across the site. Additionally, these finds have been found to be unrelated to the human activity at the site, having been deposited there independently of the human remains. These remains were described in 2022 by Valenti and colleagues, who found sufficient anatomical evidence to establish a new genus they dubbed Solitudo, with "Testudo" robusta serving as the type species. The remains from Sicily were used to erect a new species, Solitudo sicula and Solitudo gymnesica became a new name for "Testudo" gymnesica from Menorca. [3]
The name Solitudo derives from the Latin word for loneliness or solitude, which reflects the isolation of islands where members of this genus have been found. The name ending with "tudo" furthermore serves as an allusion to the genus Testudo with which the genus shares characteristics with. The species name of the youngest member, Solitudo sicula, is the female form of the Latin word for "Sicilian". [3]
The different species of Solitudo differ from the tortoises of the Testudo genus mostly in the anatomy of the trochanters. In Testudo, the trochanters are fused at the proximal end of the bone through the presence of a rounded ridge which is absent in Solitudo, leaving the trochanters only incompletely fused. The femur is generally slender and its head narrower than the width between the two trochanters. Additionally, the femoral head is oriented in a 25-45° angle when viewed from above, while other genera of tortoise have femoral heads with more anteroposterior orientation. [3]
Solitudo sicula is distinguished by the other species of the genus by having trochanters that are at the same height as the femoral head, additionally, the femoral head is also more elliptical than in more derived Solitudo species. The entire femur of S. sicula measures 104 mm (4.1 in) long, which is notably larger than that of the biggest extant tortoise of Europe, the Marginated tortoise. Assuming similar proportions, this would mean that Solitudo sicula may have reached a straight carapace length of 50–60 cm (20–24 in). [3]
Larger sizes have been estimated for the older species. Solitudo robusta may have reached a carapace length of up to 85 cm (33 in), while some estimates suggest that Solitudo gymnesica could have reached a carapace length of 1.1–1.3 m (3.6–4.3 ft). [4]
Although repeated attempts at extracting genetic material were conducted, no methods yielded DNA which rendered molecular phylogenetic analysis impossible. Due to this restriction, the relationship between Solitudo and other circum-Mediterranean tortoises was established on the basis of femur morphology, which is well known in the relevant taxa. Morphological analysis concluded that among the tortoises found around the Mediterranean, three separate femur morphotypes could be found. The "Testudinini morphotype", which is generally slender with a more narrow and oblique head, the "‘Geochelonini morphotype" which was found to be more robust and with a less oblique head and the "Insular morphotype" which typically shows a non-oblique head, slender diaphysis and trochanters that unlike in the other groups are not fully joined at the proximal end of the bone. The clade that contains all Solitudo species is based on a single synapomorphy, the incompletely fused trochanter, supported by the geographical proximity. [3]
The Solitudo sicula is recovered in a basal position within the genus, its femur still retaining some similarities to the "Testudinini morphotype". Besides the three named species of Solitudo, a fourth, unnamed taxon from Monte Pellegrino was also found to fall within the "Insular morphotype", sharing the unfused trochanter that defines the clade. The idea that this specific morphotype is the result of the tortoise's insular habitat is dismissed on account of other insular species retaining their ancestral morphology, such as the Galápagos tortoise and Gran Canaria giant tortoise. The exact relationship between Solitudo and the other tortoises surrounding the Mediterranean remains unknown until better material, in particular fossils of the head and shell, is found. [3]
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Remains of Solitudo are exclusively known from Mediterranean islands. The type species, Solitudo robusta, was found on the island of Malta and has been dated to the Middle Pleistocene. Solitudo gymnesica is known from Pliocene Menorca and Solitudo sicula from Sicily. Dating the pubis of Solitudo sicula revealed that the animal lived approximately 12.5 ± 0.5 thousand years BP. Solitudo sicula would have shared its home island with the extant Hermann's Tortoise, which has been continuously present on Sicily since at least the Middle Pleistocene. There are also remains of unnamed giant tortoises found on Sicily, they were however not contemporary with Solitudo and instead correlate with the Middle Pleistocene. [3]
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.
Giant tortoises are any of several species of various large land tortoises, which include a number of extinct species, as well as two extant species with multiple subspecies formerly common on the islands of the western Indian Ocean and on the Galápagos Islands.
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.
Hermann's tortoise is a species of tortoise native to Europe.
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.
Nuralagus is an extinct genus of leporid, with a single species, Nuralagus rex, described in 2011. It lived on Menorca, one of the Balearic Islands in the western Mediterranean during the Pliocene epoch. It is the largest known lagomorph to have ever existed, with an estimated weight of 8–12 kilograms (18–26 lb), nearly double the weight of the average Flemish Giant rabbit. It likely went extinct at the Pliocene-Pleistocene transition when Mallorca and Menorca were united as one island, letting the mammalian fauna of Mallorca, including the goat-like ungulate Myotragus, colonize Nuralagus's habitat.
Palaeoloxodon falconeri is an extinct species of dwarf elephant from the Middle Pleistocene of Sicily and Malta. It is amongst the smallest of all dwarf elephants at under 1 metre (3.3 ft) in height. A member of the genus Palaeoloxodon, it derived from a population of the mainland European straight-tusked elephant.
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.
The Sicilian shrew is a species of mammal in the family Soricidae. It is found in Sicily (Italy) and Gozo (Malta). Its natural habitat is temperate shrubland.
Asoriculus is an extinct genus of terrestrial shrews in the subfamily Soricinae and tribe Nectogalini, native to Europe and North Africa.
Hesperotestudo is an extinct genus of tortoise native to North and Central America from the Early Miocene to the Late Pleistocene. Species of Hesperotestudo varied widely in size, with a large undescribed specimen from the Late Pleistocene of El Salvador reaching
Chelonoidis is a genus of turtles in the tortoise family erected by Leopold Fitzinger in 1835. They are found in South America and the Galápagos Islands, and formerly had a wide distribution in the West Indies.
Hypnomys, otherwise known as Balearic giant dormice, is an extinct genus of dormouse (Gliridae) in the subfamily Leithiinae. Its species are considered examples of insular gigantism. They were endemic to the Balearic Islands in the western Mediterranean from the Early Pliocene until their extinction around 4,000 years ago. They first appeared in the fossil record on Mallorca during the Early Pliocene, presumably due to the Messinian salinity crisis causing a connection with mainland Europe. They later spread to Menorca, and a possible molar is also known from Ibiza. Hypnomys became extinct during the Holocene after human arrival on the Balearics. They were one of only three native land mammals to the islands at the time of human arrival, alongside the shrew Nesiotites and goat-antelope Myotragus.
Pterodromoides is an extinct genus of fulmarine petrel dating from the Late Miocene. It contains a single species, P. minoricensis. Its fossil remains were first discovered at the Punta Nati palaeontological site on the island of Menorca in the Balearic archipelago of the western Mediterranean. An additional specimen from North Carolina, USA has also been referred to this species, suggesting it lived across the North Atlantic. It was described in 2001, with the authors justifying the creation of a new genus by the large orbitonasal opening and characters of the postcranial skeleton, despite the similarity of the cranial osteology to that of Pagodroma.
Megalochelys is an extinct genus of tortoises that lived from the Miocene to Pleistocene. They are noted for their giant size, the largest known for any tortoise, with a maximum carapace length of over 2 m (6.5 ft) in M. atlas. The genus ranged from western India and Pakistan to as far east as Sulawesi and Timor in Indonesia, though the island specimens likely represent distinct species.
The Tenerife giant tortoise is an extinct species of cryptodire turtle in the family Testudinidae endemic to the island of Tenerife, in the Canary Islands during the Middle Pleistocene.
Centrochelys is a genus of tortoise. It contains one living species, the African spurred tortoise, native to the Sahel and adjacent areas. A number of fossil species have been attributed to this genus, but their placement in the genus is considered equivocal.
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.
Chelonoidis monensis, also known as the Mona tortoise, is an extinct species of land tortoise that lived on the island of Mona from the Late Pleistocene to around 1000 BCE. Evidence for the latter date includes cave drawings. All fossil remains have been found either in or near Liro Cave on the east side of Mona. It had a carapace length of around 50 cm (20 in).