Straight-tusked elephant

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Straight-tusked elephant
Temporal range:
Ma
↓
Skeleton in Rome
Skull in Germany
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Genus: †Palaeoloxodon
Species:
†P. antiquus
Binomial name
†Palaeoloxodon antiquus
(Falconer & Cautley, 1847)
Approximate range of P. antiquus
Synonyms

Elephas antiquus (Falconer & Cautley, 1847)
Loxodonta antiquus[1]

The straight-tusked elephant (Palaeoloxodon antiquus) is an extinct species of

Mediterranean
.

Description

Size of a 40 year old adult male compared to a human
Model

Like many other members of the genus

proboscideans. Two morphs of P. antiquus were previously suggested to exist in Europe on the basis of parieto-occipital crest variation, one more similar to P. namadicus, but these were shown to be the result of ontogenetic variation and taphonomic distortion. P. antiquus can be distinguished from the Indian P. namadicus based on its less stout cranium and more robust limb bones.[2] The premaxillary bones (which contain the tusks) are fan-shaped and very broad in front-view. The tusks are very long relative to body size and vary from straight to slightly curved.[3] The teeth are high crowned (hypsodont), with each third molar having approximately 16–21 ridges/lamellae.[4] The forelimbs of P. antiquus, particuarly the humerus and scapular, are proportionally longer than those of living elephants, resulting in the high position of the shoulder. The head represents the highest point of the animal, with the back being somewhat sloped though irregular in shape. The spines of the back vertebrae are noticeably elongate. The body (including the pelvis) was broad relative to extant elephants. The tail was relatively long. Although not preserved, the body was probably only sparsely covered in hair, similar to extant elephants, and probably had large ears.[3]

As with modern elephants, the species was sexually dimorphic, with males being substantially larger than females, with this size dimorphism being more pronounced than in living elephants.[3] P. antiquus was on average considerably larger than any living elephant. In a 2015 study, one approximately 40-year-old male specimen of P. antiquus from Viterbo in Italy was estimated at about 3.81 metres (12.5 ft) tall at the shoulders and estimated to weigh about 11.3 tonnes (11.1 long tons; 12.5 short tons), while a fragmentary specimen including the humerus from Montreuil, France, was estimated to weigh about 15 tonnes (14.8 long tons; 16.5 short tons) and be 4.2 metres (13.8 ft) tall at the shoulder. The average male has been estimated to have had a shoulder height of 4 m (13.1 ft) and a weight of 13 tonnes (12.8 long tons; 14.3 short tons) (for comparison, bulls of the largest living elephant species, the African bush elephant on average have a shoulder height of around 3.2 metres (10.5 ft) and a mass of around 6 tonnes (5.9 long tons; 6.6 short tons)).[5] Adult males had tusks typically around 3.5–4 metres (11–13 ft) long, with masses comfortably exceeding 100 kilograms (220 lb), with the preserved portion of one particularly large tusk from Aniene, Italy measured to be 3.9 metres (13 ft) in length, having an estimated mass of over 190 kilograms (420 lb).[6] Females probably only reached a shoulder height of about 3 metres and weight of 5.5 tonnes (5.4 long tons; 6.1 short tons).[3] A largely complete 5 year old calf from Cova del Rinoceront in Spain was estimated to have a shoulder height of 178–187 centimetres (5.8–6.1 ft) and a body mass of 1.45–1.5 tonnes (1.43–1.48 long tons; 1.60–1.65 short tons), which is comparable to a similarly aged African bush elephant.[7]

History of discovery and taxonomy

In 1695, remains of a straight-tusked elephant were collected from

Loxodonta, and E. antiquus subsequently assigned to the genus.[10] Some experts historically regarded the larger Asian species Palaeoloxodon namadicus as a variant or subspecies,[11] but they are now considered distinct.[2] Historically, the genus Palaeoloxodon has at times been regarded as a subgenus of Elephas, but a 2007 study of hyoid characteristics amongst living and fossil elephants has largely led to an abandonment of this hypothesis.[12]

Phylogeny showing the placement of Palaeoloxodon antiquus in relation to other elephantids based on nuclear genomes, after Palkopoulou et al. 2018

In 2016, a

mammoths (~5%). The African forest elephant ancestry was more closely related to modern West African forest elephants than to other African forest elephant populations. This hybridisation likely occurred in Africa, prior to migration of Palaeoloxodon into Eurasia,[15] and appears to be shared with other Palaeoloxodon species.[16]

Evolution

Like other Eurasian Palaeoloxodon species, P. antiquus is believed to derive from the African

Mammuthus trogontherii. There appears to be no overlap between M. meridionalis and P. antiquus, which suggests that the latter might have outcompeted the former. During P. antiquus's hundreds of thousands of years of existence, its morphology remained relatively static, unlike European mammoth populations.[17]

Distribution and habitat

Eemian interglacial in a temperate forest
landscape characterised by grassy areas, open woodland and closed forest

P. antiquus is known from abundant finds across Europe, ranging northwards to Great Britain and as far east as European Russia during interglacial periods. Fossils are also known from Israel, Iran and probably Turkey in West Asia, and also possibly from Kazakhstan and Tajikistan. During glacial periods P. antiquus permanently resided in the Mediterranean region.[18][19] It is primarily associated with temperate and Mediterranean forest and woodland habitats, as opposed to the colder open steppe environments inhabited by contemporary mammoths,[20] though the species is also known to have inhabited open grasslands.[21] Many of the remains attributed to the species from Western Asia are primarily done so for geographical reasons, and it has been suggested that some of these specimens actually belong to P. recki.[22]

During interglacial periods, P. antiquus existed as part of the Palaeoloxodon antiquus large-mammal assemblage, along with other temperate adapted megafauna species, including the

European water buffalo (Bubalus murrensis), Irish elk (Megaloceros giganteus), aurochs (Bos primigenius), European fallow deer (Dama dama), roe deer (Capreolus capreolus), red deer (Cervus elaphus), moose (Alces alces) and wild boar (Sus scrofa),[18][23] with carnivores including European leopards (Panthera pardus spelaea), cave hyenas (Crocuta spelaea) cave/steppe lions (Panthera spelaea), wolves (Canis lupus) and brown bears (Ursus arctos).[23] The effect of straight tusked elephant and other extinct megafauna on vegetation likely resulted in increased openness of woodland habitats.[24]

See also: Wood-pasture hypothesis § The prehistoric baseline

Behaviour and paleoecology

As with modern elephants, female and juvenile straight-tusked elephants are thought to have lived in matriarchal herds of related individuals, with the adult males living solitary lives. Like modern elephants, the herds would have been restricted to areas with available fresh water due to the greater hydration needs and lower mobility of the juveniles.[25] Due to their larger size straight-tusked elephants are suggested to have finished growing later than living elephants (perhaps more than a decade longer, up to 50 years of age in males), and may also have lived longer than extant elephants, with lifespans perhaps in excess of 80 years.[3]

niche separation from the more grazing dominated diet of the contemporary Mammuthus trogontherii.[27][26]

Relationship with humans

Remains of straight-tusked elephants at numerous sites are associated with stone tools and/or bear cut and percussion marks indicative of butchery. In the case of most sites, it is unclear whether the elephants were hunted or were scavenged, though scavenging of already dead elephants as well as active hunting are likely to have occurred.

Eemian interglacial, c. 130-115 kya) sites in Germany.[28][45]

The creation of these sites is likely attributable to

Neanderthals.[29] Stone tools used at these sites include flakes, choppers, bifacial tools like handaxes, as well as cores.[29] At some sites, the bones of straight-tusked elephants were used to make tools.[40] There is evidence that exploitation of straight-tusked elephants in Europe increased and became more systematic from the mid-Middle Pleistocene (around 500,000 years ago) onwards.[29][45] Based on analysis of sites of straight-tusked elephants with cut marks and/or artifacts, it has been argued that there is little evidence that straight-tusked elephants were targeted preferentially over smaller animals.[29] Most individuals at these sites are subadult to adult and primarily male in sex. The male sex bias likely both represents the fact that adult males, despite their larger size, were more vulnerable targets due to their solitary nature, as well as the tendency of adult male elephants to engage in risky behavior causing them to more frequently die in natural traps, as well as being weakened or killed by injures caused by combat with other male elephants during musth.[29]

At the Lehringen site in north Germany, a skeleton of P. antiquus was found with a spear made of yew wood between its ribs, with flint artifacts found close by, providing clear evidence that this specimen was hunted.[46][28] Studies in 2023 proposed that in addition to Lehringen, the Neumark Nord, Taubach and Gröbern sites, which show evidence of systematic butchery, provided evidence of widespread hunting of straight-tusked elephants by Neanderthals during the Eemian in Germany.[28][45] The remains of at least 57 elephants were found at Neumark Nord, which the authors suggested accumulated over around 300 years, with them estimating that one elephant was hunted around once every 5–6 years at the site.[28]

There are no cave paintings that unambiguously depicts P. antiquus, though an outline drawing of an elephant in El Castillo cave in Cantabria, Spain, as well as a drawing from Vermelhosa in Portugal have been suggested to possibly depict it, but could also potentially depict woolly mammoths.[20][47]

Extinction

P. antiquus retreated from northern Europe at the end of the Eemian interglacial due to climatic conditions becoming unfavourable, and fossils after that time are rare. A molar from Grotta Guattari in Italy has been suggested to date to around 57,000 years ago (though other studies have found it to have an older early Late Pleistocene age), with other remains from the Iberian Peninsula dating from 50-40,000 years ago, and possibly as late as 34,000 years ago based on a tooth found at Foz do Enxarrique in Portugal.[48][20] A late date of around 37,400 years ago has been reported from a single molar found in the North Sea off the coast of the Netherlands,[49] but it has been suggested that this date needs independent confirmation, due to only representing a single sample.[50] A 2020 study suggested that P. antiquus likely survived until around 28,000 years ago in the southern Iberian Peninsula, based on footprints found in Southwest Portugal and Gibraltar. Its extinction is likely to be due to climatic deterioration, possibly in association with hunting.[51]

The extinction of P. antiquus and other temperate adapted European megafauna has resulted in the severe loss of

functional diversity in European ecosystems.[23]

Dwarfed descendants

Main article: Dwarf elephant § The Mediterranean islands

Mediterranean islands, spanning from Sicily and Malta in the west to Cyprus in the east.[48] The responsible factors for the dwarfing of island mammals
are thought to be the reduction in food availability, predation and competition.

References

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