Sardinian dhole Temporal range: Middle Pleistocene-Holocene | |
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Drawing of the skeleton of C. sardous scale bar = 20 cm | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Family: | Canidae |
Subfamily: | Caninae |
Tribe: | Canini |
Genus: | † Cynotherium Studiati, 1857 |
Type species | |
Cynotherium sardous Studiati, 1857 | |
Other species | |
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Former range (in red) |
The Sardinian dhole (genus Cynotherium especially C. sardous) is an extinct insular canid which was endemic to what is now the Mediterranean islands of Sardinia and Corsica during the Middle-Late Pleistocene. It went extinct at the end of the Pleistocene around the time of human settlement of the islands.
Its scientific name means "dog-beast of Sardinia", the genus name from the Ancient Greek : θήρ, romanized: thḗr, lit. 'beast', 'wild animal' and κύων, kyōn, 'dog' and the specific name from the Latin : Sardous, lit. 'of Sardinia', alt. form of Latin : Sardus.
Around the size of a jackal, morphological studies indicate it was a specialised stalking predator of small mammals and birds, with a probable preference for the endemic Sardinian pika.
The oldest remains of Cynotherium on Sardinia date to around the early-Middle Pleistocene transition around 800,000 years ago, associated with a faunal turnover event on Sardinia likely caused low sea levels allowing dispersal to Sardinia-Corsica from mainland Italy. Cynotherium is suggested to have originated from the species Xenocyon lycaonoides of mainland Europe. Some of the older Cynotherium remains are assigned to the separate species Cynotherium malatestai, which is larger than C. sardous and shows intermediate features between it and X. lycaonoides. [1] A 2021 genetic study found that its closest living relative is the dhole, from which it diverged approximately 885,000 years ago. The study found that the lineage of the Sardinan dhole and dhole emerged from the hybridisation between a lineage closely related to the genus Canis and a lineage related to the African wild dog (Lycaon pictus) in proportions of roughly 65% and 35% respectively. The modern dhole was also found to have additional African wild dog related ancestry (making up around 25% of its genome) not found in the Sardinan dhole. [2]
C. sardous was relatively small in size, with a weight around 10 kilograms (22 lb), comparable to a jackal. [3] The skull is slender and wedge-shaped in lateral view. The snout is narrower than in extant dholes, but broader than those of foxes. The postorbital region of the skull is broad, and the zygomatic arches only modestly project outwards. The mastoid is very enlarged and projects outwards. The sagittal crest is weakly developed. The mandible is slender. [4] The attachment sites for the triceps on the scapula, ulna and humerus are large, indicating the muscle, along with the anconeus, was well developed. The scar for the deltoid muscle at the posterior of the deltoid ridge on the humerus is high, large and rugose, indicating this muscle was also well developed. [3] Analysis of its ear morpology, including the significant reduction in the number of cochlear turns, suggests a specialisation towards the hearing of high-frequency sounds, but suggests it was poor at detecting low frequency sounds, and completely unable to detect sounds lower than 250 Hz. [5]
When the ancestor of this canid became confined to the island, its diet became limited to small prey. Cynotherium as a predator specializing in small, fast moving prey is supported by an examination of the animal's anatomy. The evolution of short, powerful limbs, a low neck carriage and increased head and neck mobility suggests an animal specialised for stalking low to the ground, and then quickly pouncing on or running down prey. [4] [3] A suggested preferred prey item of the Sardinian dhole is the Sardinian pika (Prolagus sardus), a large lagomorph that was also endemic and abundant on the island, with birds also being likely targets. [4]
During the Middle and Late Pleistocene Corsica and Sardinia had their own highly endemic depauperate terrestrial mammal fauna which besides the Sardinian dhole and Sardinian pika, included the Tyrrhenian field rat, ( Rhagamys orthodon ) the Tyrrhenian vole ( Microtus henseli ), a shrew ( Asoriculus similis ), a mole ( Talpa tyrrhenica ), a dwarf mammoth (Mammuthuslamarmorai), a galictine mustelid ( Enhydrictis galictoides ), three species of otter ( Algarolutra majori, Sardolutra ichnusae, Megalenhydris barbaricina ) and a deer ( Praemegaceros cazioti ). [6]
The youngest radiocarbon dates for the Sardinan dhole are around 11,500 years BC/13,500 years Before Present (BP), only a few thousand years before the first confirmed human presence on Sardian-Corsica around 8,000 BC/10,000 years BP, and it is assumed that the species was alive when humans arrived on the island. The causes of its extinction are uncertain. [7] The low genetic diversity of sampled individuals suggests that the population size had been small but stable for a long period of time prior to extinction. [2]
Canidae is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid. The family includes three subfamilies: the Caninae, and the extinct Borophaginae and Hesperocyoninae. The Caninae are known as canines, and include domestic dogs, wolves, coyotes, foxes, jackals and other species.
The dhole is a canid native to South, East and Southeast Asia. It is anatomically distinguished from members of the genus Canis in several aspects: its skull is convex rather than concave in profile, it lacks a third lower molar and the upper molars possess only a single cusp as opposed to between two and four. During the Pleistocene, the dhole ranged throughout Asia, with its range also extending into Europe but became restricted to its historical range 12,000–18,000 years ago. It is now extinct in Central Asia, parts of Southeast Asia, and possibly the Korean peninsula and Russia.
Canis is a genus of the Caninae which includes multiple extant species, such as wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and dentition, long legs, and comparatively short ears and tails.
The Sardinian pika is an extinct species of lagomorph that was endemic to the islands of Sardinia, Corsica and neighbouring Mediterranean islands until its extinction likely in Roman times. It was the last surviving member of Prolagus, a genus of lagomorph once widespread throughout Europe during the Miocene and Pliocene epochs, whose closest living relatives are pikas of the genus Ochotona.
Dwarf elephants are prehistoric members of the order Proboscidea which, through the process of allopatric speciation on islands, evolved much smaller body sizes in comparison with their immediate ancestors. Dwarf elephants are an example of insular dwarfism, the phenomenon whereby large terrestrial vertebrates that colonize islands evolve dwarf forms, a phenomenon attributed to adaptation to resource-poor environments and lack of predation and competition.
Mammuthus lamarmorai is a species of dwarf mammoth which lived during the late Middle and Late Pleistocene on the island of Sardinia in the Mediterranean. It has been estimated to have had a shoulder height of around 1.4 metres (4.6 ft). Remains have been found across the western part of the island.
Caninae is the only living subfamily within Canidae, alongside the extinct Borophaginae and Hesperocyoninae. They first appeared in North America, during the Oligocene around 35 million years ago, subsequently spreading to Asia and elsewhere in the Old World at the end of the Miocene, some 7 million to 8 million years ago.
Palaeoloxodon cypriotes is an extinct species of dwarf elephant that inhabited the island of Cyprus during the Late Pleistocene. A probable descendant of the large straight-tusked elephant of mainland Europe and West Asia, the species is among the smallest known dwarf elephants, with fully grown individuals having an estimated shoulder height of only 1 metre (3.3 ft). It represented only one of two large animal species on the island alongside the Cypriot pygmy hippopotamus. The species became extinct around 12,000 years ago, around the time humans first colonised Cyprus, and potential evidence of human hunting has been found.
Asoriculus is an extinct genus of terrestrial shrews in the subfamily Soricinae and tribe Nectogalini, native to Europe and North Africa.
Prolagus is an extinct genus of lagomorph. Over 20 species have been named, and the genus was abundant and widespread in Europe during the Neogene. However, by the end of the Middle Pleistocene, it was confined to a single species, the Sardinian pika, on the Corsica, Sardinia, and surrounding islands, where it survived into historical times. In North Africa and Western Asia, the genus is known from the Miocene and Pliocene. The scientific name may mean "before hares" or "primitive hares". Its taxonomy is disputed, with it either being considered a member of the family Ochotonidae, which includes living pikas, or the only member of the family Prolagidae.
Xenocyon is an extinct group of canids, either considered a distinct genus or a subgenus of Canis. The group includes Canis (Xenocyon) africanus, Canis (Xenocyon) antonii and Canis (Xenocyon) falconeri that gave rise to Canis (Xenocyon) lycanoides. The hypercarnivorous Xenocyon is thought to be closely related and possibly ancestral to modern dhole and the African wild dog, as well as the insular Sardinian dhole.
Megalenhydris barbaricina is an extinct species of giant otter from the Late Pleistocene of Sardinia. It is known from a single partial skeleton, discovered in the Grotta di Ispinigoli near Dorgali, and was described in 1987. It was larger than any living otter, exceeding the size of South American giant otters (Petrolutra), which can reach two meters in length. The species is one of four extinct otter species from Sardinia and Corsica. The others are Algarolutra majori, Lutra castiglionis and Sardolutra ichnusae. It is suggested to have ultimately originated from the much smaller European mainland species "Lutra" simplicidens, which may be more closely related to Lutrogale than to modern Lutra species. The structure of the teeth points to a diet of bottom dwelling fish and crustaceans. A special characteristic of the species is the flattening of the first few caudal vertebrae. This might point to a slightly flattened tail.
The Corsican red deer, also known simply as the Corsican deer or Sardinian deer, is a subspecies of red deer found on the Mediterranean islands of Sardinia (Italy) and Corsica (France).
Rhagamys is an extinct genus of rodents in the subfamily Murinae, the Old World mice and rats. The genus was established by the Swiss zoologist Charles Immanuel Forsyth Major to accommodate Rhagamys orthodon, which is the only species in the genus. It was endemic to the Mediterranean islands of Corsica and Sardinia, descending from Rhagapodemus, which had colonised the islands around 3.6 million years ago. Its closest living relatives are of the genus Apodemus, which includes the field and wood mice.
Praemegaceros is an extinct genus of deer, known from the Pleistocene and Holocene of Western Eurasia. Praemegaceros is considered to be a genus of "giant deer", with many species having an estimated body mass of around 400 kilograms (880 lb), considerably larger than most living deer.
Talpa tyrrhenica, also known as the Tyrrhenian mole, is an extinct species of mole belonging to the genus Talpa. It was endemic to the Mediterranean islands of Corsica and Sardinia during the Pleistocene epoch.
Microtus (Tyrrhenicola) henseli is an extinct species of vole belonging to the genus Microtus that was endemic to Sardinia and Corsica during the Pleistocene and Holocene.
Mececyon trinilensis, the Trinil dog, is an extinct canid species that lived on the island of Java in Indonesia during the Pleistocene.
The Corbeddu cave is located in the territory of Oliena, municipality of the province of Nuoro, in Sardinia. In this cave found refuge the famous Sardinian bandit Giovanni Corbeddu Salis (1844-1898), from which it took its name.
Enhydrictis is a genus of extinct mustelid, belonging to the subfamily Galictinae. The type species, and best known, is Enhydrictis galictoides from the Pleistocene of Sardinia and Corsica. Some authors attribute species from mainland Eurasia to the genus, but this is disputed, with others considering the genus endemic to Sardinia-Corsica.