This is a list of European species extinct in the Holocene that covers extinctions from the Holocene epoch, a geologic epoch that began about 11,650 years before present (about 9700 BCE)[A] and continues to the present day.[1]
Most recent remains dated to 3040-1840 BCE. A painting on the Ancient Egyptian tomb of Rekhmire (1470-1445 BCE) depicting exotic animals brought to Egypt as tribute by foreign peoples, includes a picture of an animal interpreted as a dwarf elephant by some authors.[4]
Most recent remains dated to 348 BCE - 283 CE.[9] Though hunted by the original human inhabitants of the islands, it likely became extinct due to Roman agricultural practices, the introduction of predators (dogs, cats, and small mustelids) and ecological competitors (rodents, rabbits, and hares).[10] Transmission of pathogens by rabbits and hares could have been another factor.[11] Survival into modern history, even as late as 1774 on the smaller island of Tavolara, has been hypothesised from the description of unknown mammals by later Sardinian authors; however, this interpretation remains dubious owing to anatomical discrepancies.[12]
Most recent remains dated to 9650 BCE in the Ponto-Caspian region, 9550 BCE in Boreal Europe, 9450 BCE in the British Isles, 8850 BCE in Northwestern Germany, 8750 BCE in northern Central Europe, 6050 BCE[2] in the Carpathian Basin, the Middle Holocene in the Middle Urals,[13] and 1220 BCE in the Southern Urals.[2] This species avoids human disturbance strictly and is considered an excellent indicator of the health of steppe ecosystems, as a result.[14]
Most recent remains dated to 9650 BCE in the Ponto-Caspian Region, 9550 BCE in Boreal Europe, 8750 BCE in northern Central Europe, 8250 BCE in the Franco-Cantabrian region, 6050 BCE in Northwestern Germany, 5850 BCE[2] in the Carpathian Basin, and Late Holocene in the Urals.[14]
Most recent remains at Mochlos dated to the Bronze Age. It was outcompeted and replaced by the house mouse accidentally introduced by sailors from the eastern Mediterranean.[15]
Most recent remains at Escorca, Mallorca dated to 4840-4690 BCE, coinding with the period of initial human settlement in the island. It could have succumbed to diseases carried by introduced commensal mammals.[17]
Highly specialised for grazing, with the narrowest range of all Pleistocene ground quirrels. The latest possible date is the Atlantic, and its extinction was probably related to the local collapse of mammoth steppe.[18]
Most recent remains in north Central Europe are dated to 8750 BCE.[2] However, remains have been dated to the late Holocene in Ukraine, and one account describes a large, previously unidentified, "red-cheeked" ground squirrel from the early 20th century of the Dnipropetrovsk area.[20]
Most recent remains at Alcúdia dated to 3030-2690 BCE, coinding with the period of initial human settlement in the island. It could have succumbed to diseases carried by introduced commensal mammals.[17]
Most recent remains in the Franco-Cantabrian region dated to 9350 BCE.[2] Other lion remains from Italy and northern Spain could indicate that a small form survived in mountain areas until the Preboreal and Boreal, respectively.[22]
Remains were found in Shengavit and Urartu, Armenia dating to the 4th-3rd millennium BCE. It is also depicted in rock art of the 4th-1st millennium BCE, where it can be differenciated from the leopard by the shape of its paws and unretracted claws. Possibly survived in Armenia until the Middle Ages before disappearing due to hunting.[23]
According to the alternate hypothesis, the modern lion expanded into southern Europe and replaced the cave lion there already in the Late Glacial, surviving in Italy and northern Spain until the Preboreal or Boreal.[22] A possible second colonization event took place in the Balkans during the Atlantic and Subboreal periods, reaching as far as Hungary, southwestern Ukraine, and Greece. In the Iron Age the lion strongly declined until it disappeared from these regions, possibly because of hunting and habitat loss caused by increasing human population and livestock rearing.[24] In 370 CE the Greco-Roman orator Themistius mentioned that lions had disappeared from Thessaly, their last Balkan stronghold.[C] Lions were also hunted historically across Transcaucasia, and were reportedly common in the ungulate-rich Kura-Araz and Mughan plains, up to the Absheron Peninsula, until 900 CE.[26]
A cold-adapted subspecies of the leopard, Panthera pardus spelaea, was widespread in Europe during the Pleniglacial and Late Glacial.[27] A 8850 BCE record from the Franco-Cantabrian region,[2] another from the Preboreal or Boreal of Greece, and two from the Sub-Atlantic of western and southern Ukraine could indicate that leopards survived or recolonized the continent in the Holocene. However, later remains from Hellenistic and Roman sites are confidently attributed to imports from Asia and Africa.[24]
In the Caucasus, the leopard was hunted to extinction from most of the region by the 1950s or 1960s,[28] but still survives in small areas of the North Caucasus, southern Armenia, and Azerbaijan.[29] These leopards belong to the Persian subspecies Panthera pardus tulliana, which also occurs in Anatolia.[30] In 1889 an Anatolian leopard was killed in the Greek island of Samos after swimming from Asia. Local folklore suggests that similar events have happened in the island at different times in history.[31]
Present permanently in the Caucasus region and along the Caspian and eastern Azov coasts, the Terek and Kuban rivers, and the estuary of the Don river during the 10th-12th centuries CE, with vagrants recorded as far as Chernihiv, Ukraine.[28] Last recorded in Mingrelia and Imeretia at the beginning of the 17th century, Armenia in the early 19th century, eastern Georgia in 1936,[26] and Azerbaijan's Talysh Mountains in 1966. Last three were all vagrants intruding after tigers stopped breeding in the respective area.[28]
Exterminated by livestock farmers. The last confirmed individual was killed in 1924 near Bellolampo; unconfirmed killings near Palermo were reported between 1935 and 1938, and unconfirmed sightings between 1960 and 1970.[32]
Most recent remains dated to 7050-6550 BCE in Riparo Fredian, Italy (with doubts)[33] and Les Coves de Santa Maira, Spain.[34] Claims of 21st century presence of dhole in the Caucasus are erroneous.[35]
Most recent remains in Corsica dated to 9910-9710 BCE and Sardinia to 9531-9196 BCE, roughly coinciding with modern human colonization of the islands.[36]
Martens, polecats, otters, badgers, and weasels (family Mustelidae)
Known from a single skeleton found in a cave with no stratigraphical context but estimated to be Late Pleistocene or early Holocene,[37] 68050-8050 BCE.[38]
Western Europe to western Siberia,[39] Anatolia?[40]
Historical sources record wild horses living until the 12th century in Denmark, 13th in Germany,[41] 14th in Portugal, 16th in Spain,[42] the Vosges, East Prussia, and Lithuania; 18th in the northern Carpathians[41] and southern Urals,[43] and 19th in Poland and Ukraine.[44] The last in the wild was killed in Askania-Nova in 1879, and the last in captivity died in the Moscow Zoo in 1887.[41] Some sources treat them as wild, untameable animals of different nature to horses, and others as feral horses or hybrids, casting doubt on the moment when pure wild horses became extinct in the continent. Despite this, the IUCN considers the subspecies E. f. ferus valid. The Tatar-Cossack word "tarpan" was popularized for European wild horses in the 19th century, though today is sometimes limited to horses from central and eastern Europe.[44]
Paleogenomics suggest that horses were domesticated independently in the Ponto-Caspian steppe and expanded to the rest of Europe by the Bronze Age. Early nomadic pastoralists likely released their horses to graze freely at night, resulting in feral populations and hybridization with wild horses. Wild mares were also captured to replenish domestic herds, breaking down the social order of wild herds and diminishing their reproduction. Around 600-1100 CE, the original high genetic diversity of domestic horses dropped to current levels.[44] In historical times European wild horses were hunted for their meat, hide, traditional medicine, sport, and to protect crops and livestock hay deposits during the winter.[42][41] Several horse breeds have been claimed to have recent tarpan ancestry including the Konik, Sorraia, Exmoor pony, Hucul pony, Bosnian Mountain Horse, Estonian Native, and Gotland pony. However, genetic and historical evidence indicate that all are typical domestic horses.[44]
Remains dated to 8050 BCE in Western Europe, 3550 BCE in Italy,[2] 3300-2700 BCE in Karanovo, Bulgaria; 3200-2500 BCE in Los Millares, Spain; 2050 BCE in southern Central Europe,[2] and 1500-500 BCE in Keti, Armenia. Questionable remains in Didi-gora, Georgia dated to 1075 BCE. The hydruntine inhabited open steppe habitat that became rarer and fragmented in the Holocene, making it more vulnerable to human exploitation.[45]
Probably present in the deserts between the Volga and Ural rivers until the 18th or 19th century, when it was extirpated due to increasing hunting with firearms and seizure of waterholes for livestock use. 18th century records from Voronezh, Russia are considered unreliable.[47] It was first reintroduced to Askania-Nova, Ukraine in 1950.[48] In 2020 Rewilding Europe released kulan in the Tarutyns'kyj steppe near the Danube Delta.[49] It has also announced plans to release kulan in Spain as proxy for the hydruntine.[50]
Possibly calved in the Mediterranean in ancient times.[51] All few confirmed individuals in Europe since 1999 were identified as vagrants from the North American population, and known calving areas in Africa appear to be depleted.[52]
Most recent remains at Ijmuiden, Netherlands were dated to 550 CE.[54] A vagrant from the Pacific population dispersed over the Arctic Ocean and was seen in Europe in 2010.[55][56]
North Caucasus and the Transcaucasian coast of the Black Sea
Hunted to extinction by the beginning of the 20th century. The subspecies' validity is questioned because moose from Russia later colonized the North Caucasus naturally over the 20th century.[57]
Most recently dated to 8718 BCE in Teppa u Lupinu, Corsica and 5641–5075 BCE in Grotta Juntu, Sardinia. It survived the first human colonization of the islands, but became extinct when Neolithic peoples arrived.[36]
Survived into the early Holocene of Scania and (as the subspecies C. c. palmidactyloceros) in northern Italy, Switzerland, and possibly the French Alps while the temperate forest-adapted red deer replaced it in the rest of Europe. The dwarf subspecies C. c. tyrrhenicus existed in Capri after the post-glacial sea level rise.[60]
Cattle, goats, antelopes, and others (family Bovidae)
Declined after the Russian conquest of the Caucasus as a result of increased hunting, deforestation, and domestic cattle rearing. The subspecies was protected in the 1890s when it was limited to 442 animals in the area between the Belaya and Laba rivers. However an epizootic outbreak in 1919 reduced the animals to just 50, and the last individuals were poached in 1927.[61] The only captive animal, a male, lived in Germany between 1908 and 1925 and bred with females of the lowland wisent subspecies. As a result, several wisent populations carry its genes today.[62]
Most recent remains dated to 1130-1060 BCE near the Oyat river in Western Russia. However this date was not calibrated and the remains could be older.[64] Recent calibrated dates include 9450 BCE in the Southern Urals, 8650 BCE in the Middle Urals, and 7550 BCE in Boreal Europe.[2]
Declined as a result of hunting, deforestation for agriculture, competition with livestock for pastures, and diseases transmitted by domestic cattle. The last individual in the Jaktorow forest of Mazovia, Poland died in 1627,[65] and the last in Sofia, Bulgaria in the late 17th or early 18th century.[66][67] There are different active projects to breed aurochs-like cattle and release them in the wild as proxy for the aurochs.
Most recent confirmed remains in Kolomna, Russia dated to 10811 BCE, during the Last Glacial Period.[68] However, unique genetic introgression into local domestic water buffaloes and possible remains from the Neolithic of southeastern Europe (9000-7000 BCE) and Atlantic of Austria (7000-4000 BCE) suggest that the native European species of water buffalo survived into the Holocene.[69] In 2019, Rewilding Europe released domestic buffaloes in the Danube Delta as proxy for the European water buffalo.[70]
Hunted to extinction around 1890. A different subspecies of Spanish ibex naturally colonized the Peneda-Gerês National Park in the Portuguese ibex's former range during the 21st century.[71][72]
The last individual, a female, died at Ordesa National Park in 2000. A single cloned individual was born on July 30, 2003, but died several minutes later,[74] making this the first case of biological taxonde-extinction and a taxon becoming extinct twice. In 2014, Spanish ibexes from the Guadarrama Mountains were released in the French Pyrenees as proxy for the Pyrenean ibex.[72]
Most recent remains dated to 3969-3759 BCE in Menorca, 3649-3379 BCE in Cabrera,[75] and 2830-2470 BCE in Mallorca. The timeframe allows to confidently exclude climate change as a reason for the extinction and blame it solely on the first human settlers to the islands.[76]
The last wild population in Poland's Białowieża Forest was hunted to extinction during World War I. A captive herd was returned to Bialowieza in 1929; it was made of zoo animals, some of which were hybridized with other subspecies or species of bison. Individuals with American bison ancestry were removed from Bialowieza in 1936, and with Caucasian wisent ancestry in 1950. The Bialowieza herd was fully returned to the wild in 1952 and subsequently used as stock for pure lowland herds in Poland, Lithuania, and Belarus.[77] The Caucasian-lowland hybrid line was introduced to the Kavkazsky Nature Reserve in 1940, in the Caucasian wisent's former range, and allowed to roam free from 1946.[78] Other hybrid wisent herds were later established in the Carpathians, Ukraine, and Russia.[77]
Most recent remains in Sweden were dated to 7050 BCE.[81] The first reintroduction attempt was made at Gurskøya, Norway in 1925, but all animals died because of the unfavorable climate or poaching. Another herd was released at Hjerkinn in the Dovre mountains in 1932. These animals are presumed to have been exterminated during World War II, though there were unconfirmed sightings of muskoxen at Tafjord in 1942 and 1951. The definitive successful reintroduction in Dovre was made in 1947.[82] In 1971 a herd left Dovre after being harassed by tourists and established itself in Harjedalen, Sweden. Norwegians also introduced muskoxen to Svalbard in 1929, outside of the muskox's natural range, but this population died out by the 1970s.[81]
The species bred in Kazakhstan and southern Siberia and wintered in western Morocco and Tunisia, being present in Europe during migration or as a vagrant. It likely disappeared as a result of habitat alteration in Asia and overhunting in Africa. The last confirmed record worldwide was in Hungary, in 2001.[84]
Originally hunted for its feathers, meat, fat, and oil; as it grew rare, also to supply collectionists. The last pair on the eastern Atlantic was killed on Eldey Island, off Iceland in 1844.[85]
Extirpated from Europe before 1650 as a result of habitat loss, climate change, and direct persecution.[87] A 1738 painting made in England by Eleazar Albin was based on a stuffed specimen or an older depiction.[88] In 1991 a gradual reintroduction project using handreared chicks began at Alpenzoo Innsbruck in Austria, and in 2011 a migratory population was established between southern Germany, Austria, and Tuscany. A second reintroduction project started in southern Spain in 2004.[87]
Described as different separated species including Bubo insularis, before being recognized as a subspecies of the Asian brown fish owl.[91] The most recent remains in Corsica date to 7433-7035 BCE. In Corsica-Sardinia it could have been locally adapted to prey on the Sardinian pika, disappearing after human arrival with it.[9]
Occasional winter visitor to southwest Andalusia until the end of the 19th century, with a single later record of a bird shot in Jerez de la Frontera in 1998.[92]
Most recent remains dated to 5295 BCE. The causes of extinction are presumed human-induced due to the lack of climatic changes at the time, such as the introduction of exotic predators like feral dogs, pigs, and garden dormice by the first human settlers.[97]
Eastern coast of North America and the Baltic region
Last known Baltic specimen was caught in 1996 near Muhumaa, Estonia.[98] It was reintroduced to the Oder river in 2009,[99] and to the Narva in 2013.[100]
Caspian Sea, Volga, Ural and Terek river drainages
Last recorded in the Ural in the 1960s.[109] All spawning grounds were lost after dams were built in the Volga, Ural, and Terek river drainages. The species continues to exist in captivity, from which it is released periodically in its native range. However, illegal fishing and hybridization with the introduced nelma remain threats to its survival.[109]
Disappeared around 1940 as a result of water pollution.[110] Though treated as a different species since about 1700, a genetic study in 2023 found the houting indistinguishable from the lavaret (Coregonus lavaretus) still extant in Great Britain, the Alpine area, and waterways it was introduced to.[111][112]
Last recorded in the 1960s. Extinct as a result of hybridization with the three-spined stickleback; the springs it inhabited were separated from the latter's habitat by a hypersaline lake acting as barrier between the species, until irrigation works transformed the lake into a brackish one that was invaded by migratory three-spined sticklebacks.[113]
Only known from a male adult and a nymph found on a dead Iberian lynx in 1997, itself a critically endangered species with low population density and disjunct distribution at the time. Besides difficulties in mixing and exchanging populations, the lice was threatened by the fact that lynxes taken to captive breeding centers were systematically deloused.[118][119]
Last collected in 1938. Both the Main and the Rhine were heavily polluted around that time and all local caddisfly species disappeared. Although other caddisflies returned after water quality improved, this species has not been recorded since.[126]
Last recorded in 1987 and deemed extinct as a result of water substraction, which peaked in 1988. However, fresh shells collected in 2009 may hint to its continued survival.[128]
Islets of Dyo Adelfoi, Megali Zafrano, Karavonisi, and Divounia, inbetween Astypalaia and Karpathos, Greece
Known only from subfossil shells in three islets and last recorded in the fourth in 1985. Likely declined due to habitat alteration caused by fire, tourism, and military construction.[134]
Not seen since its description in 1874. The species has been suggested to be the same as, or related to Drusia deshayesii from northern Morocco and Algeria, as well as an introduced species.[135]
Sea anemones, corals, and zoanthids (class Hexacorallia)
^The source gives "11,700 calendar yr b2k (before CE 2000)". But "BP" means "before CE 1950". Therefore, the Holocene began 11,650 BP. Doing the math, that is c. 9700 BCE.
^A. corsicanus was originally applied to remains from Corsica and A. similis to Sardinia. It was later recognized that A. corsicanus existed in the early Pleistocene of both islands, and A. similis in the late Pleistocene-Holocene, as seen in Moncunill-Sole et al. (2016).
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