Sardinian pika

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Sardinian pika [1]
Temporal range: Middle Pleistocene - Holocene 0.8–0.0002  Ma
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Prolagus3.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Lagomorpha
Family:† Prolagidae
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 the last surviving member of Prolagus, a genus of lagomorph with a fossil record spanning 20 million years once widespread throughout Europe during the Miocene and Pliocene epochs. Its closest living relatives are modern pikas (which all belong to the genus Ochotona ), from which it is estimated to have diverged around 30 million years ago.

Contents

Anatomy

Skeleton of P. sardus Prolagus skeleton SI.jpg
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]

Glires

Rodentia

Lagomorpha

Leporidae (rabbits and hares)

† Prolagus

Ochotona (living pikas)

The earliest species of Prolagus appeared in Europe 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 Sardinia sometime after 810 BC (based on radiocarbon dating), 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). [23] Its extinction was possibly due to agricultural practices, the introduction of predators (dogs, cats and small mustelids) and ecological competitors (rodents, rabbits and hares). [25] 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. [26] 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. [27] [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]

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. [29]

Related Research Articles

<span class="mw-page-title-main">Lagomorpha</span> Order of mammals

The lagomorphs are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae and the Ochotonidae (pikas). There are 110 recent species of lagomorph of which 109 are extant, including 10 genera of rabbits, 1 genus of hare and 1 genus of pika. The name of the order is derived from the Ancient Greek lagos + morphē.

<span class="mw-page-title-main">Pika</span> Genus of mountain-dwelling mammal

A pika 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. The large-eared pika of the Himalayas and nearby mountains lives at elevations of more than 6,000 m (20,000 ft).

<span class="mw-page-title-main">Sardinian dhole</span> Extinct species of carnivore

The Sardinian dhole 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.

<i>Mammuthus lamarmorai</i> Extinct species of mammal

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.

<span class="mw-page-title-main">Gaoligong pika</span> Species of mammal

The Gaoligong pika is a species of mammal in the family Ochotonidae. It is endemic to China. Many of the general physical characteristics of the pika species are shared by Gaoligong pikas. However, the Gaoligong pika is specifically characterized by unique physical characteristics, including a red-brown colored crown around the neck and black behind the ears. They can produce one litter per year and can live up to three years. Their behavior is currently undetermined due to limited information available about the species. This is due to the inaccessibility of their habitat.

<span class="mw-page-title-main">Hoffmann's pika</span> Species of mammal

Hoffmann's pika is a species of mammal in the pika family, Ochotonidae, that is endemic to Mongolia. It is currently listed as endangered by the International Union for Conservation of Nature (IUCN).

<i>Asoriculus</i> Extinct genus of red-toothed shrew

Asoriculus is an extinct genus of terrestrial shrews in the subfamily Soricinae and tribe Nectogalini, native to Europe and North Africa, from the Late Miocene until the late Holocene. The genus is closely related and possibly ancestral to the also recently-extinct Balearic shrews (Nesiotites), with their closest living relative being the Himalayan shrew.

<i>Prolagus</i> Extinct genus of mammals

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.

<i>Megalenhydris</i> Extinct genus of giant otter

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.

<span class="mw-page-title-main">Tsing-ling pika</span> Species of mammal

The Tsing-ling pika is a species of pika endemic to the mountains in Central China. It inhabits mountainous forests and shrublands. It is a poorly known species.

<i>Macaca majori</i> Extinct species of monkey

Macaca majori, commonly known as the dwarf macaque, is a prehistoric macaque from the Early Pleistocene of Sardinia, Italy. It descended from the Barbary macaque. Its temporal range spans from about 2 million to 0.8 million years ago, during the Nesogoral faunal complex, alongside the goat-antelope Nesogoral, the pig Sus sondaari, the hyena Chasmaporthetes, the pika Prolagus, the shrew Asoriculus, the mole Talpa tyrrhenica, the mustelid Pannonictis, and the dormouse Tyrrhenoglis.

<span class="mw-page-title-main">Giant pika</span> Extinct species of mammal

The giant pika or Wharton's pika is an extinct mammal species in the family Ochotonidae. It lived during the Pleistocene and early Holocene in northern parts of North America. Very similar forms have also been found also in Siberia.

<i>Rhagamys</i> Extinct rodent genus

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.

<i>Talpa tyrrhenica</i> Extinct species of mammal

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.

<span class="mw-page-title-main">Corbeddu Cave</span> Cave and archaeological site in Italy

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.

Rhagapodemus is a genus of extinct rodent from the Miocene to Pleistocene periods. Most species are known from European localities, although R. debruijni comes from India.

<i>Enhydrictis</i> Extinct genus of mustelid

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.

References

  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 2024-11-20). ISSN   0375-7633.{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (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...58210645M. doi: 10.1016/j.palaeo.2021.110645 . hdl: 2183/28801 . 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. 33 (2): 504–516. 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): 13635. 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. ↑ 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.
  26. ↑ "Prolagus sardus factsheet". Archived from the original on 10 February 2009. Retrieved 10 January 2017.
  27. ↑ 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.
  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. 1 2 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)
  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.
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