lagomorph The lagomorphs () are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae (rabbits and hares) and the Ochotonidae ( pikas). There are 110 recent species of lagomorph, of which 109 species in t ...

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

Sardinia Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...

Corsica Corsica ( , , ; ; ) is an island in the Mediterranean Sea and one of the Regions of France, 18 regions of France. It is the List of islands in the Mediterranean#By area, fourth-largest island in the Mediterranean and lies southeast of the Metro ...

Roman times In modern historiography, ancient Rome is the Roman civilisation from the founding of the Italian city of Rome in the 8th century BC to the collapse of the Western Roman Empire in the 5th century AD. It encompasses the Roman Kingd ...

Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Ochotona''), from which it is estimated to have diverged around 30 million years ago.

Anatomy

The full skeletal structure of the Sardinian pika was reconstructed in 1967, thanks to the numerous finds of bones in Corbeddu Cave, which is near Oliena, Sardinia. Some years later, from these remains, the same researchers led by paleontologist Mary R. Dawson from the US were able to create a plaster reconstruction with good accuracy, and provide a thorough description of the skeleton's morphology published in 1969. The Sardinian pika was probably much stockier and more robust than extant species of pikas, and it probably resembled a sort of cross between a large wild rabbit and a pika. The first articulated skeletons of ''P. sardus'' were reported in 2016. ''Prolagus sardus'' weighed about . This is more than its ancestor '' Prolagus figaro'', which is the only other member of ''Prolagus'' that was found in Sardinia and weighed about , and is larger than most mainland species of '' Prolagus.'' Compared to mainland species of ''Prolagus'', ''P. sardus'' had larger and more hypsodont (high crowned) teeth. The Sardinian pika experienced anagenic evolution, with an increasing body size and shifting dental morphology over time.

Ecology

Abundant

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

and

subfossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

remains of ''P. sardus'' are known from several localities across

Corsica Corsica ( , , ; ; ) is an island in the Mediterranean Sea and one of the Regions of France, 18 regions of France. It is the List of islands in the Mediterranean#By area, fourth-largest island in the Mediterranean and lies southeast of the Metro ...

and

Sardinia Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...

hint at the once broad geographical range of this ''Prolagus'' species: it lived from sea level up to at least 800 m (2,624 ft.) in a variety of

habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...

s (grasslands, shrublands). Its morphology suggests that it was capable of traversing rocky terrain, and was probably a proficient jumper and capable of digging, but was not adapted for running. The tooth hypsodonty has been suggested to have been an adaptation to an abrasive diet. The abundance of mass accumulations of broken bones (

bone bed A bone bed is any Geology, geological stratum or deposition (geology), deposit that contains bones of whatever kind. Inevitably, such deposits are Sedimentary rock, sedimentary in nature. Not a formal term, it tends to be used more to describe esp ...

s) suggest that the

population density Population density (in agriculture: Standing stock (disambiguation), standing stock or plant density) is a measurement of population per unit land area. It is mostly applied to humans, but sometimes to other living organisms too. It is a key geog ...

was high. A sample of Late Pleistocene specimens from Medusa Cave, Sardinia found that they had a high incidience rate of

arthritis Arthritis is a general medical term used to describe a disorder that affects joints. Symptoms generally include joint pain and stiffness. Other symptoms may include redness, warmth, Joint effusion, swelling, and decreased range of motion of ...

relative to extant lagomorphs. This is suggested to be the result of ageing due to having a longer lifespan than mainland lagomorphs. Skeletochronology suggests that individuals of ''Prolagus sardus'' reached a lifespan of approximately 8 years, which is longer than mainland lagomorphs of equivalent size. The Sardinian pika was likely preyed on by the two native species of terrestrial carnivores, a canine (the Sardinian dhole), and a

mustelid The Mustelidae (; from Latin , weasel) are a diverse family of carnivoran mammals, including weasels, badgers, otters, polecats, martens, grisons, and wolverines. Otherwise known as mustelids (), they form the largest family in the suborde ...

('' Enhydrictis galictoides'') which were specialized for hunting small prey. Other likely predators include birds of prey such as the endemic owl species '' Bubo insularis''.

Evolution and extinction

The taxonomy of '' Prolagus'' has been the subject of controversy. It is either considered a member of the family Ochotonidae, which includes living

pikas A pika ( , or ) is a small, mountain-dwelling mammal native to Asia and North America. With short limbs, a very round body, an even coat of fur, and no external tail, they resemble their close relative the rabbit, but with short, rounded ears. ...

(which all belong to the genus ''Ochotona''), or the only member of the family Prolagidae. A partial mitochondrial genome from ''Prolagus sardus'' suggests that ''Prolagus'' is more closely related to living pikas than to

Leporidae Leporidae () is the family of rabbits and hares, containing over 70 species of extant mammals in all. The family name comes from "Lepus", hare in Latin. Together with the pikas, the Leporidae constitute the mammalian order Lagomorpha. Leporidae ...

, which contains rabbits and hares, with an estimated divergence between living pikas and ''Prolagus'' about 30 million years ago.The earliest species of ''Prolagus'' appeared in Europe during the Early Miocene, around 20 million years ago. N. López-Martínezbr>Paleobiogeographical history of ''Prolagus'', a European ochotonid (Lagomorpha)
Lynx, 32 (2001), pp. 215-231 The ancestor of the Sardinian pika, ''Prolagus figaro'', arrived in the

Corsica Corsica ( , , ; ; ) is an island in the Mediterranean Sea and one of the Regions of France, 18 regions of France. It is the List of islands in the Mediterranean#By area, fourth-largest island in the Mediterranean and lies southeast of the Metro ...

n-

Sardinia Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...

n

microcontinent Continental crustal fragments, partly synonymous with microcontinents, are pieces of continents that have broken off from main continental masses to form distinct islands that are often several hundred kilometers from their place of origin. Caus ...

at the early-late

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58 Amongst mainland species, the ''P. figaro-P. sardus'' lineage was previously thought to be most closely related to the species ''P. depereti'' known from the Pliocene of France, which was originally described as a subspecies of ''P. figaro''. However, the oldest known remains of ''Prolagus'' from Sardinia, referred to as ''P. aff. figaro'', show closer affinities to the species ''P. sorbinii'', a species of Eastern European origin, which expanded westwards during the

Messinian The Messinian is in the geologic timescale the last age or uppermost stage of the Miocene. It spans the time between 7.246 ± 0.005 Ma and 5.333 ± 0.005 Ma (million years ago). It follows the Tortonian and is followed by the Zanclean, the fir ...

, the last stage of the

Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...

, with well known remains from central Italy from the latest Miocene and early Pliocene. The oldest unambiguous remains of ''Prolagus sardus'' date back from the

Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...

, a time when both islands were periodically connected due to

sea level Mean sea level (MSL, often shortened to sea level) is an mean, average surface level of one or more among Earth's coastal Body of water, bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical ...

changes. Reassessment of palaeontological data has shown that the distinction made by early authors between two contemporaneous

taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...

(''P. sardus'' and ''P. corsicanus'') is probably unfounded, as the Sardinian pika exhibits only subtle anagenetic evolution of its anatomy and body size through time. Humans first arrived in Corsica-Sardinia around 10,000 years

Before Present Before Present (BP) or "years before present (YBP)" is a time scale used mainly in archaeology, geology, and other scientific disciplines to specify when events occurred relative to the origin of practical radiocarbon dating in the 1950s. Because ...

(BP). The presence of ''Prolagus'' facilitated the establishment of the first human communities of the islands. Jean-Denis Vigne found clear evidence that the Sardinian pika was hunted and eaten by people. He found that many of the Sardinian pikas' limb bones were broken and burnt at one end, suggesting that this animal had been roasted and eaten by the

Neolithic The Neolithic or New Stone Age (from Ancient Greek, Greek 'new' and 'stone') is an archaeological period, the final division of the Stone Age in Mesopotamia, Asia, Europe and Africa (c. 10,000 BCE to c. 2,000 BCE). It saw the Neolithic Revo ...

colonists of Corsica. The Sardinian pika became extinct in Sardinia sometime after 810 BC (based on

radiocarbon dating Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for Chronological dating, determining the age of an object containing organic material by using the properties of carbon-14, radiocarbon, a radioactive Isotop ...

), with records from Corsica suggesting that species survived there until sometime between 393 BC and the 6th century AD (overlapping with the period of Carthaginian and Roman Corsica). Its extinction was possibly due to agricultural practices, the introduction of predators (dogs, cats and small mustelids) and ecological competitors (rodents, rabbits and hares). Transmission of pathogens by rabbits and hares introduced to Sardinia and Corsica by the Romans may have also played a role in the species's extinction. Other endemic small mammals like the shrew '' Asoriculus similis'', the Tyrrhenian field rat, and the Tyrrhenian vole, probably also disappeared from Corsica and Sardinia around the same time.

Historical references

The 2nd century BCE Greek historian

Polybius Polybius (; , ; ) was a Greek historian of the middle Hellenistic period. He is noted for his work , a universal history documenting the rise of Rome in the Mediterranean in the third and second centuries BC. It covered the period of 264–146 ...

described in ''The Histories'' the presence of an animal in Corsica locally called the ''kyniklos'' which "when seen from a distance looks like a small hare, but when captured it differs much from a hare in appearance and taste" and which "lives for the most part under the ground". This animal may have been the Sardinian pika, because Corsica at that time was not characterized by the occurrence of any species of hare. Survival of the Sardinian pika up into

modern history The modern era or the modern period is considered the current historical period of human history. It was originally applied to the history of Europe and Western history for events that came after the Middle Ages, often from around the year 1500, ...

has been hypothesised from the description of unknown mammals by later Sardinian authors; however, this interpretation remains dubious owing to anatomical discrepancies. The Medieval Italian poet Fazio Degli Uberti mentioned "a small animal" in Sardinia which was very timid and was called "Solifughi", which means "hiding from the sun", in his 1360 poem ''Dittamondo'' ('Song of the World'). In 1774, Francesco Cetti wrote that the island of Tavolara off the coast of Sardinia had "giant rats whose burrows are so abundant that one might think the surface of the soil had been recently turned over by pigs", which has often been taken as a reference to the Sardinian pika.Kurtén, Björn (1968) Pleistocene Mammals of Europe. Weidenfeld and Nicolson, London However, this was questioned by Barbara Wilkens in a 2000 publication, who suggested that it was more likely that the animals mentioned by Cetti were brown rats.Wilkens B. (2000)
Osservazioni sulla presenza in epoca recente del prolago sardo a Tavolara secondo le notizie di Francesco Cetti
bservations on the recent presence of the Sardinian pika in Tavolara according to the reports of Francesco CettiAtti 3° Convegno Nazionale di Archeozoologia (Siracusa, 2000), 217 -222. (in Italian)

References

{{Taxonbar, from=Q1048895 Lagomorphs Extinct mammals of Europe Mammal extinctions since 1500 Fauna of Sardinia Fauna of Corsica Mammals described in 1829 Taxa named by Johann Andreas Wagner