Sardinian pika[1]
Temporal range: Middle Pleistocene - Holocene
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Lagomorpha
Family: Ochotonidae
Genus: †Prolagus
Species:
†P. sardus
Binomial name
†Prolagus sardus
(Wagner, 1829)
Synonyms

Prolagus corsicanus

The Sardinian pika (Prolagus sardus) 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 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.

Anatomy

Skeleton of P. sardus

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.[3] 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.[3] The first articulated skeletons of P. sardus were reported in 2016.[4]

Prolagus sardus weighed about 504–525 g (17.8–18.5 oz). This is more than its ancestor Prolagus figaro, which is the only other member of Prolagus that was found in Sardinia and weighed about 398–436 g (14.0–15.4 oz), and is larger than most mainland species of Prolagus.[5]

Compared to mainland species of Prolagus, P. sardus had larger and more hypsodont (high crowned) teeth.[6][7] The Sardinian pika experienced anagenic evolution, with an increasing body size and shifting dental morphology over time.[8]

Ecology

Abundant fossil and subfossil remains of P. sardus are known from several localities across Corsica and Sardinia 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.)[9] in a variety of habitats (grasslands, shrublands).[10] 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.[3][11] The tooth hypsodonty has been suggested to have been an adaptation to an abrasive diet.[7] The abundance of mass accumulations of broken bones (bone beds) suggest that the population density was high.[11][12][9] A sample of Late Pleistocene specimens from Medusa Cave, Sardinia found that they had a high incidience rate of arthritis relative to extant lagomorphs. This is suggested to be the result of ageing due to having a longer lifespan than mainland lagomorphs.[13] Skeletochronology suggests that individuals of Prolagus sardus reached a lifespan of approximately 8 years, which is longer than mainland lagomorphs of equivalent size.[14]

The Sardinian pika was likely preyed on by the two native species of terrestrial carnivores, a canine (the Sardinian dhole), and a mustelid (Enhydrictis galictoides) which were specialized for hunting small prey.[15][16] Other likely predators include birds of prey such as the endemic owl species Bubo insularis.[17]

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 (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, which contains rabbits and hares, with an estimated divergence between living pikas and Prolagus about 30 million years ago.[18] The earliest species of Prolagus appeared during the Early Miocene, around 20 million years ago.[19]

The ancestor of the Sardinian pika, Prolagus figaro, arrived in the Corsican-Sardinian microcontinent at the early-late Pliocene boundary around 3.6 million years ago, likely due to an emergent land connection with Italy caused by a sea level drop.[20][21] 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 last stage of the Miocene, with well known remains from central Italy from the latest Miocene and early Pliocene.[21] The oldest unambiguous remains of Prolagus sardus date back from the Middle Pleistocene,[12] a time when both islands were periodically connected due to sea level changes. Reassessment of palaeontological data has shown that the distinction made by early authors between two contemporaneous taxa (P. sardus and P. corsicanus) is probably unfounded,[22][9] as the Sardinian pika exhibits only subtle anagenetic evolution of its anatomy and body size through time.[8]

Humans first arrived in Corsica-Sardinia around 10,000 years Before Present (BP).[23] 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 colonists of Corsica.[24]

The Sardinian pika became extinct in Corsica and Sardinia sometime after 810 BC,[23] probably during Roman times[25] due to agricultural practices, the introduction of predators (dogs, cats and small mustelids) and ecological competitors (rodents, rabbits and hares).[26] 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.[27] 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.[25][23]

Historical references

The 2nd century BCE Greek historian Polybius 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.[28] The most recent zooarchaeological remains of pikas in Corsica were dated to either late pre-Roman or Roman times, between 348 BCE and 283 CE.[25]

Survival of the Sardinian pika up into modern history has been hypothesised from the description of unknown mammals by later Sardinian authors; however, this interpretation remains dubious owing to anatomical discrepancies.[29] 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').[30] 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.[31][32] 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.[33]

References

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  1. ↑ Hoffman, R.S.; Smith, A.T. (2005). "Order Lagomorpha". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 193–194. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. ↑ Smith, A.T.; Lanier, H.C. (2019). "Prolagus sardus". IUCN Red List of Threatened Species. 2019: e.T18338A1737167. doi:10.2305/IUCN.UK.2019-2.RLTS.T18338A1737167.en. Retrieved 17 February 2022.
  3. 1 2 3 Dawson, Mary R. (1969). "Osteology of Prolagus sardus : a Quaternary Ochotonid (Mammalia, Lagomorpha)" (PDF). Palaeovertebrata. 2 (4): 157–190. doi:10.18563/pv.2.4.157-190. Retrieved February 14, 2017.
  4. ↑ Zoboli, Daniel; Caddeo, Guglielmo Angelo (2016). "Articulated skeletons of Prolagus sardus (Mammalia, Lagomorpha) from the Quaternary of Grotta del Campanaccio (Santadi, south-western Sardinia)". Bollettino della Società Paleontologica Italiana (1): 81–83. doi:10.4435/BSPI.2016.08 (inactive 1 August 2023). ISSN 0375-7633.{{cite journal}}: CS1 maint: DOI inactive as of August 2023 (link)
  5. ↑ Moncunill-Solé, Blanca; Tuveri, Caterinella; Arca, Marisa & Angelone, Chiara (2016). "Comparing the body mass variations in endemic insular species of the genus Prolagus (Ochotonidae, Lagomorpha) in the Pleistocene of Sardinia (Italy)". Rivista Italiana di Paleontologia e Stratigrafia. 122 (1): 25–36. doi:10.13130/2039-4942/6905.
  6. ↑ Angelone, Chiara (2005). "Evolutionary trends in dental morphology of the genus Prolagus (Ochotonidae, Lagomorpha) in the Mediterranean islands". Proceedings of the International Symposium "Insular Vertebrate Evolution: The Palaeontological Approach": September, 16–19 Mallorca, 2005, ISBN 84-609-6472-8, págs. 17–26. Societat d'Història Natural de les Balears: 17–26. ISBN 978-84-609-6472-8.
  7. 1 2 Moncunill-Solé, Blanca; Tuveri, Caterinella; Arca, Marisa; Angelone, Chiara (November 2021). "Tooth and long bone scaling in Sardinian ochotonids (Early Pleistocene-Holocene): Evidence for megalodontia and its palaeoecological implications". Palaeogeography, Palaeoclimatology, Palaeoecology. 582: 110645. Bibcode:2021PPP...582k0645M. doi:10.1016/j.palaeo.2021.110645. S2CID 239224107.
  8. 1 2 Angelone, Chiara; Tuveri, Caterinella; Arca, Marisa; López Martínez, Nieves & Kotsakis, Tassos (2008). "Evolution of Prolagus sardus (Ochotonidae, Lagomorpha) in the Quaternary of Sardinia Island (Italy)". Quaternary International. 182 (1): 109–115. Bibcode:2008QuInt.182..109A. doi:10.1016/j.quaint.2007.08.020.
  9. 1 2 3 Vigne, Jean-Denis (July 10, 2003). "Le Lapin rat : Prolagus sardus (Wagner, 1829)". In Pascal, M.; Lorvelec, O.; Vigne, J.-D.; Keith, P.; Clergeau, P. (eds.). Évolution holocène de la faune de Vertébrés de France : invasions et disparitions (Report) (in French). Institut National de la Recherche Agronomique, Centre National de la Recherche Scientifique, Muséum National d'Histoire Naturelle. pp. 108–110.
  10. ↑ Lanfranchi, François de; Weiss, Michel-Claude (1977). "Araguina-Sennola : dix années de fouilles préhistoriques à Bonifacio". Archeologia Corsa (in French). 2: 1–167. ISSN 0154-2656.
  11. 1 2 Zoboli, Daniel; Zedda, Marco; Festa, Vincenzo; Pillola, Gian Luigi; Palombo, Maria Rita (December 2018). "Does a relationship exist between palaeopathologies and insularity ? a case study of some bones of Prolagus Sardus (Wagner, 1829) from Sardinia (Italy)". Alpine and Mediterranean Quaternary. 31 (1): 75–86. doi:10.26382/AMQ.2018.12.
  12. 1 2 Pereira, Elisabeth; Ottaviani-Spella, Marie-Madeleine & Salotti, Michelle (2001). "Nouvelle datation (Pléistocène moyen) du gisement de Punta di Calcina (Conca, Corse du Sud) par la découverte de Talpa tyrrhenica Bate, 1945 et d'une forme primitive de Microtus (Tyrrhenicola) henseli Forsyth-Major, 1882". Geobios (in French). 34 (6): 697–705. Bibcode:2001Geobi..34..697P. doi:10.1016/S0016-6995(01)80031-0.
  13. ↑ Moncunill-Solé, Blanca; Arzi, Boaz; Filliquist, Barbro; Vapniarsky, Natalia; Zavodovskaya, Regina; Angelone, Chiara (2023-03-14). "The extinct osteoarthritic lagomorphs (Prolagus sardus) from Sardinia (Italy) reveal further evidence of life history evolution in insular domains". Palaeoworld. doi:10.1016/j.palwor.2023.03.004. ISSN 1871-174X. S2CID 257548108.
  14. ↑ Fernández-Bejarano, E., Blanco, A., Angelone, C., Zhang, Z., Moncunill-Solé, B., 2022. First approach to life history of the islander Prolagus sardus (Lagomorpha) by studying femoral bone histology. In: Belvedere, M., Mecozzi, B., Amore, O., Sardella, R. (Eds.), Abstract Book of the XIX Annual Conference of the European Association of Vertebrate Palaeontologists, Benvento/Pietraroja, Italy 27th June-2nd July 2002. PalaeoVertebrata, Special Volume 1-2022, pp. 58–59.
  15. ↑ Lyras, George; van der Geer, Alexandra (2006-01-01). "Adaptations of the Pleistocene island canid Cynotherium sardous (Sardinia, Italy) for hunting small prey". Cranium. 23 (1): 51–60. ISSN 0923-5647.
  16. ↑ van der Geer, Alexandra A.; Lyras, Georgios A.; Lomolino, Mark V.; Palombo, Maria Rita; Sax, Dov F. (2013-04-22). "Body size evolution of palaeo-insular mammals: temporal variations and interspecific interactions". Journal of Biogeography. 40 (8): 1440–1450. doi:10.1111/jbi.12119. ISSN 0305-0270. S2CID 37706170.
  17. ↑ Pavia, Marco (2008-05-01). "The evolution dynamics of the Strigiformes in the Mediterranean islands with the description of Aegolius martae n. sp. (Aves, Strigidae)". Quaternary International. Insularity and its Effects. 182 (1): 80–89. Bibcode:2008QuInt.182...80P. doi:10.1016/j.quaint.2007.05.018. hdl:2318/79470. ISSN 1040-6182. S2CID 53347144.
  18. ↑ Utzeri, Valerio Joe; Cilli, Elisabetta; Fontani, Francesco; Zoboli, Daniel; Orsini, Massimiliano; Ribani, Anisa; Latorre, Adriana; Lissovsky, Andrey A.; Pillola, Gian Luigi; Bovo, Samuele; Gruppioni, Giorgio; Luiselli, Donata; Fontanesi, Luca (2023-08-21). "Ancient DNA re-opens the question of the phylogenetic position of the Sardinian pika Prolagus sardus (Wagner, 1829), an extinct lagomorph". Scientific Reports. 13 (1). doi:10.1038/s41598-023-40746-w. ISSN 2045-2322. PMC 10442435. PMID 37604894.
  19. ↑ N. López-Martínez Paleobiogeographical history of Prolagus, a European ochotonid (Lagomorpha) Lynx, 32 (2001), pp. 215-231
  20. ↑ Palombo, Maria Rita (2006). "Biochronology of the Plio-Pleistocene Terrestrial mammals of Sardinia: The state of the art" (PDF). Hellenic Journal of Geosciences. 41: 47–66. ISSN 1105-0004. Retrieved February 14, 2017.
  21. 1 2 Angelone, Chiara; Čermák, Stanislav & Kotsakis, Tassos (2015). "The most ancient lagomorphs of Sardinia: An overview". Geobios. 48 (4): 287–296. Bibcode:2015Geobi..48..287A. doi:10.1016/j.geobios.2015.06.002.
  22. ↑ Pereira, Elisabeth (2001). Le peuplement mammalien quaternaire de Corse (Post-Glaciaire exclu), son contexte environnemental, biologique et physique (Ph.D.) (in French). Université de Corse.
  23. 1 2 3 Valenzuela, Alejandro; Torres-Roig, Enric; Zoboli, Daniel; Pillola, Gian Luigi; Alcover, Josep Antoni (2021-11-29). "Asynchronous ecological upheavals on the Western Mediterranean islands: New insights on the extinction of their autochthonous small mammals". The Holocene. 32 (3): 137–146. doi:10.1177/09596836211060491. ISSN 0959-6836. S2CID 244763779.
  24. ↑ The Archaeology of Animals. Routledge, 2012. 2012-11-12. p. 124. ISBN 9781135106591.
  25. 1 2 3 Vigne, Jean-Denis; Bailon, Salvador & Cuisin, Jacques (1997). "Biostratigraphy of amphibians, reptiles, birds and mammals in Corsica and the role of man in the Holocene faunal turnover" (PDF). Anthropozoologica. 25–26: 587–604. Retrieved February 14, 2017.
  26. ↑ Vigne, Jean-Denis & Valladas, Hélène (1996). "Small Mammal Fossil Assemblages as Indicators of Environmental Change in Northern Corsica during the Last 2500 Years". Journal of Archaeological Science. 23 (2): 199–215. Bibcode:1996JArSc..23..199V. doi:10.1006/jasc.1996.0018.
  27. ↑ "Prolagus sardus factsheet". Archived from the original on 10 February 2009. Retrieved 10 January 2017.
  28. ↑ Alves C., Paulo; Ferrand, Nuno; Hackländer, Klaus (2007-12-29). Lagomorph Biology: Evolution, Ecology, and Conservation. Springer Science & Business Media, 2007. p. 14. ISBN 9783540724469.
  29. ↑ Wilkens, Barbara (2000). Osservazioni sulla presenza in epoca recente del Prolago sardo a Tavolara secondo le notizie di Francesco Cetti. 3° Convegno Nazionale di Archeozoologia (in Italian). Siracusa. pp. 217–222.
  30. ↑ "Il Dittamondo - di: Fazio degli Uberti". bepi1949.altervista.org (in Italian). Retrieved 2017-01-10.
  31. ↑ Kurtén, Björn (1968) Pleistocene Mammals of Europe. Weidenfeld and Nicolson, London
  32. ↑ Storia Naturale di Sardegna (PDF) (in Italian). Ilisso. 2000. p. 190. ISBN 978-88-87825-07-7. Retrieved 10 July 2016.
  33. ↑ Wilkens B. (2000) - Osservazioni sulla presenza in epoca recente del prolago sardo a Tavolara secondo le notizie di Francesco Cetti. [Observations on the recent presence of the Sardinian pika in Tavolara according to the reports of Francesco Cetti] Atti 3° Convegno Nazionale di Archeozoologia (Siracusa, 2000), 217 -222. (in Italian)
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