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The straight-tusked elephant ( Palaeoloxodon antiquus) is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene. One of the largest known elephant species, mature fully grown bulls on average had a shoulder height of and a weight of , placing them among the largest land mammals ever. Straight-tusked elephants likely lived very similarly to modern elephants, with herds of adult females and juveniles and solitary adult males. The species was primarily associated with temperate and Mediterranean woodland and forest habitats, flourishing during interglacial periods, when its range would extend across Europe as far north as Great Britain and Denmark and eastwards into Russia, while persisting in southern Europe during , when northern Europe was occupied by Steppe mammoth and later Woolly mammoth. Skeletons found in association with stone tools and in one case, a wooden spear, suggest they were scavenged and hunted by early humans, including Homo heidelbergensis and their Neanderthal successors.
The species is part of the genus Palaeoloxodon (whose other members are also sometimes called straight-tusked elephants), which emerged in Africa during the Early Pleistocene, before dispersing across Eurasia at the beginning of the Middle Pleistocene, with the earliest record of Palaeoloxodon in Europe dated to around 800–700,000 years ago, around the time of the extinction of the previously dominant mammoth species Mammuthus meridionalis. The straight-tusked elephant is the ancestor of over half a dozen named (and several more unnamed) species of that inhabited islands in the Mediterranean, some of which shrunk to only 2% the size of their mainland ancestor. The straight-tusked elephant became extinct during the latter half of the Last Glacial Period, with the youngest remains found in the Iberian Peninsula, dating to around 44,000 years ago. Possible even younger records include a single tooth from the Netherlands that has been dated to around 37,000 years ago, and footprints from the southern part of the Iberian Peninsula dated to 28,000 years ago.
The skull is proportionally both very wide and tall. Like many other members of the genus Palaeoloxodon, P. antiquus possesses a well-developed growth of bone at the top of the cranium above the nasal opening called the parieto-occipital crest, originating from the occipital bone of the skull roof which projects forwards and overhangs the rest of the skull. The crest was probably an anchor for muscles, including the Splenius muscles, as well as an additional muscle layer that wrapped around the top of the head, called the "extra splenius". The latter was likely similar to the "splenius superficialis" found in . The crest likely developed to support the very large size of the head, as the skulls of Palaeoloxodon are the largest proportionally and in absolute size among proboscideans. Two morphs of P. antiquus were previously thought to exist in Europe on the basis of differences in the parieto-occipital crest, one more similar to the South Asian Palaeoloxodon namadicus. These differences were shown to be age-related (Ontogeny variation), with the crest being more pronounced in older individuals, as well as due to distortion during fossilisation (Taphonomy variation). P. antiquus differs from P. namadicus in having a less stout cranium and more robust limb bones, and in lacking a teardrop-shaped indentation behind the eye socket (infraorbital depression). The premaxillary bones (which contain the tusks) are fan-shaped and very broad in front view. The tusks are very long relative to the size of the body and vary from straight to slightly curved. The teeth are high crowned (hypsodont), with each third molar having approximately 16–21 lamellae (ridges).
Females were considerably larger than living female elephants and comparable in size with African bush elephant bulls, with female individuals from the Neumark Nord population in Germany reaching shoulder heights and weights rarely exceeding and respectively (though several relatively young females at the site would likely have exceeded this size when fully grown). A particularly large female known from a pelvis found near Binsfeld in Germany has been estimated to have had a shoulder height of and a weight of . For comparison, 90% of fully grown female African bush elephants reach an shoulder height of and body mass of under optimal growth conditions. Newborn and young calves were likely around the same size as those of modern elephants. A largely complete 5 year old calf from Cova del Rinoceront in Spain was estimated to have a shoulder height of and a body mass of , which is comparable to a similarly aged African bush elephant.
In 1695, remains of a straight-tusked elephant were collected from travertine deposits near Burgtonna in what is now Thuringia, Germany. While these remains were originally declared to be purely mineral in nature by the Collegium Medicum in the nearby city of Gotha, Wilhelm Ernst Tentzel, a polymath in the employ of the ducal court of Saxe-Gotha-Altenburg, correctly identified them as elephant remains. The Burgtonna skeleton was one of the specimens that Johann Friedrich Blumenbach described in his publication naming the woolly mammoth ( Mammuthus primigenius, originally Elephas primigenius) in 1799. The remains of straight-tusked elephants continued to be attributed to woolly mammoths until the 1840s.Davies, Paul; (2002) The straight-tusked elephant ( Palaeoloxodon antiquus) in Pleistocene Europe. Doctoral thesis (Ph.D), UCL (University College London).40
The straight-tusked elephant was scientifically named in 1847 by British palaeontologists Hugh Falconer and Proby Cautley as Elephas ( Euelephas) antiquus.Falconer, H and Cautley, P. T. (1845-1849). Fauna Antiqua Sivalensis, being the fossil zoology of the Sewalik Hills in the north of India. London, Smith, Elder and Co. Note: Part 1 published in 1845, Part 2 in 1846, Parts 3 through 8 in 1847, Part 9 in 1849.40-41 The type specimen is a mandible (lower jaw) with a second molar (M2006). The exact provenance of the specimen is unknown, though it probably originates from Britain, and possibly the site of Grays in Essex, southeast England. This specimen was originally considered to be that of a mammoth, and the attribution to E. antiquus was made in a hand-written correction.40 The common name "straight-tusked elephant" was used for the species as early as 1873 by William Boyd Dawkins. In 1924, the Japanese paleontologist Matsumoto Hikoshichirō assigned E. antiquus to his new taxon Palaeoloxodon, which he classified as a subgenus of Loxodonta (which includes the living African elephants).40 The species has a confused taxonomic history, with at least 21 named synonyms.41 In publications in the 1930s and 1940s (the latter published posthumously), Henry Fairfield Osborn assigned the species to its own genus Hesperoloxodon, which was followed by some later authors, but is now rejected.42 In his widely cited 1973 work, Origin and evolution of the Elephantidae, Vincent J. Maglio sunk P. antiquus into the South Asian P. namadicus, as well as Palaeoloxodon back into Elephas (which contains the living Asian elephant). While the sinking of Palaeoloxodon into Elephas (with Palaeoloxodon sometimes being treated as a subgenus of Elephas) gained considerable traction in the following decades, today both P. antiquus and Palaeoloxodon are considered distinct.
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
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. Fossil tracks of newborns, calves and adults, which are likely of a herd of P. antiquus, have been found in dune deposits in southern Spain, dating to the early Late Pleistocene (Marine Isotope Stage (MIS) 5, around 130–80,000 years ago). Some straight-tusked elephant populations may have engaged in seasonal migrations, as occurs in living elephants. Isotopic analysis of male tooth specimens from Neumark Nord in Germany indicate that over the 8 years preserved by the teeth, these individuals had engaged in (perhaps seasonal) migrational movements. While some remained relatively local to the Neumark Nord area, others had migrated considerable distances to and from the Neumark Nord area, perhaps as far as away from where they eventually died.Armaroli, Elena; Lugli, Federico ; Tacail, Théo ; Kindler, Lutz; Gaudzinski-Windheuser, Sabine; Roebroeks, Wil; Vonhof, Hubert ; Cipriani, Anna ; Tütken, Thomas; Müller, Wolfgang (August 2025) " Combined Time-Resolved Sr-O-C Isotopes and Proteomics Unravel the Ecology of Extinct Straight-Tusked Elephants at Neumark-Nord (~125 ka, Saxony-Anhalt, Germany)" Abstract book of the 11th International Symposium on Biomolecular Archaeology p. 138
Due to their larger size, straight-tusked elephants are thought to have finished growing 10 to 15 years later than living elephants, continuing to grow after 50 years of age in males, and to around 40 years of age in females, the latter comparable to the growth period of African bush elephant bulls. They may also have lived longer than extant elephants, with lifespans perhaps in excess of 80 years.
Dental microwear studies suggest that the diet of P. antiquus was highly variable, ranging from almost completely grazing to almost completely browsing (feeding on leaves, stems and fruits of high-growing plants). However, microwear only reflects the diet in the last few days or weeks before death, so the observed dietary variation may be seasonal, as is the case with living elephants. Isotopic analysis of a specimen from Greece suggests that it was primarily browsing during the dry (presumably summer) months and consumed more grass during the wet (presumably winter) months. Dental mesowear analysis of specimens from Britain and Greece suggests that the diet also varied according to local environmental conditions, with individuals occupying more grass-dominated open environments having a greater grazing-related wear signal, but that overall straight-tusked elephants tended to have browsing-dominated mixed feeding diets. Preserved stomach contents of German specimens found at Neumark Nord suggests that in temperate Europe, its diet included trees such as maple, Tilia, hornbeam, hazel, alder, beech, Fraxinus, oak, elm, spruce and possibly juniper, as well as other plants like ivy, Pyracantha, Artemisia, mistletoe ( Viscum), ( Carduus and Cirsium), grass and sedges ( Carex), as well as members of Apiaceae, Lauraceae, Rosaceae, Caryophyllaceae and Asteraceae (including the subfamily Lactuceae).Grube, R., Palombo, M., Iacumin, P. & Di Matteo, A. What did the fossil elephants from Neumark–Nord eat? In Elefantenreich – Eine Fossilwelt in Europa. (ed. Meller, H.) 253–272 (Landesamt für Denkmalpflege und Archäologie Sachsen-Anhalt, 2010). (p. 258 for relevant content)
Straight-tusked elephants rarely coexisted alongside mammoths, although they occasionally did so, like at the Ilford locality in Britain that dates to the Marine Isotope Stage (MIS) 7 interglacial (~200,000 years ago) and where both and P. antiquus are found. At this locality, the two species appear to have engaged in dietary niche partitioning. During interglacial periods, P. antiquus existed as part of the Palaeoloxodon antiquus large-mammal assemblage, along with other temperate adapted megafauna species, including the hippopotamus ( Hippopotamus amphibius), rhinoceroses belonging to the genus Stephanorhinus (Merck's rhinoceros S. kirchbergensis and the narrow-nosed rhinoceros S. hemitoechus), the European water buffalo ( Bubalus murrensis), bison ( Bison spp.), Irish elk ( Megaloceros giganteus), aurochs ( Bos primigenius), fallow deer ( Dama spp.), roe deer ( Capreolus capreolus), red deer ( Cervus elaphus), moose ( Alces alces), wild horse ( Equus ferus) and wild boar ( Sus scrofa). Carnivores included Eurasian lynx ( Lynx lynx) European leopards ( Panthera pardus spelaea), ( Crocuta spelaea), Panthera spelaea ( Panthera spelaea), Wolf ( Canis lupus) and ( Ursus arctos). Some authors have argued, in accordance with the Vera/wood-pasture hypothesis, that the effects of straight-tusked elephants and other extinct megafauna on vegetation likely resulted in increased openness of woodland habitats, though this conclusion has been disputed by other authors.
Potential gnaw marks suggested to have been made by cave hyenas and cave lions on the bones of straight-tusked elephants have been reported at some localities, which suggests that these species likely at least scavenged on the remains of straight-tusked elephants like lions and Spotted hyena do on elephants in Africa today. Remains of juvenile straight-tusked elephants are known from Kirkdale Cave in northern England and Grotta Guattari in central Italy, which are well known cave hyena dens and may suggest predation upon juvenile straight-tusked elephants by cave hyenas, as is exhibited by closely related living spotted hyenas on African elephant calves.
These sites are likely attributable to Homo heidelbergensis and Neanderthals. Stone tools used at these sites include Lithic flake, choppers, bifacial tools like Hand axe, as well as Lithic core . At some sites, the bones of straight-tusked elephants and in at least one case their ivory were used to make tools. 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. 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. 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 injuries caused by combat with other male elephants during musth. At the Lehringen site in north Germany, dating to the Eemian/Last Interglacial (around 130–115,000 years ago) a skeleton of a mature adult P. antiquus, around 45 years of age, was found with a complete (though currently fractured) spear/lance between its ribs, with flint artifacts found close by, providing unequivocal evidence that this specimen was hunted,H. Thieme, S. Veil, Neue Untersuchungen zum eemzeitlichen Elefanten-Jagdplatz Lehringen, Ldkr. Verden. Kunde 36, 11–58 (1985). though it has been suggested the elephant may have already been mired prior to being killed. The spear/lance, which is around long, is made of Taxus baccata ( Taxus baccata) , which has both a durable and elastic wood, properties that may have been deliberately selected for.Milks, A. (2020) Yew wood, would you? An exploration of the selection of wood for Pleistocene spears. In: Berihuete-Azorin, M., Martin Seijo, M., Lopez-Bulto, O. and Pique, R. (eds.) The Missing Woodland Resources: Archaeobotanical studies of the use of plant raw materials. Advances in Archaeobotany, 6 (6). Barkhuis Publishing, Groningen, pp. 5-22. The Lehringen spear/lance is one of the oldest known wooden weapons after the Clacton Spear (also made of yew wood) and the Schöningen spears, has been suggested to have served as a handheld thrusting spear rather than as a throwing weapon. The current c-curved bent shape of the spear suggests that the spear was thrust upwards into the elephants abdomen, and may have been deformed by the elephant falling on it (the current fractured state of the spear is thought to have been due to much later sediment compaction).
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. The remains of at least 57 elephants were found at Neumark Nord; the study authors estimated that they accumulated over a time span of around 300 years and that one elephant was hunted once every 5–6 years at the site. in Germany]]At the Lower Palaeolithic Bilzingsleben site in Germany and Stránská Skála 1 site in the Czech Republic, bones of straight-tusked elephants have been found engraved with multiple nearly straight lines, either parallel or converging, of unclear purpose.
There are no that unambiguously depict P. antiquus. 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 these could also potentially depict . ARCA A., 2014 - Elephas antiquus depicted at Vermelhosa rock art? TRACCE Online Rock Art Bulletin, 31, online
Some island dwarf elephant descendants survived considerably later than the youngest confirmed straight tusked elephant records, with the Sicilian Palaeoloxodon cf. mnaidriensis surviving until sometime after 32,000 years ago, with one record perhaps as late as 20,000-19,000 years ago, while Palaeoloxodon cypriotes on Cyprus survived until at least around 12–11,000 years ago.
The extinction was part of the Late Pleistocene megafauna extinctions, which resulted in the extinction of most large terrestrial mammals globally. The extinction of P. antiquus and other temperate adapted European megafauna has resulted in a severe loss of functional diversity in European ecosystems.
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