Straight-tusked elephant
Straight-tusked elephant Temporal range: Mid-Late Pleistocene ~0.78–0.03 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓ | |
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Skeleton in Rome | |
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Skull in Germany | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Proboscidea |
Family: | Elephantidae |
Genus: | †Palaeoloxodon |
Species: | †P. antiquus |
Binomial name | |
†Palaeoloxodon antiquus | |
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Approximate range of P. antiquus | |
Synonyms | |
Elephas antiquus (Falconer & Cautley, 1847) |
The straight-tusked elephant (Palaeoloxodon antiquus) is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene (around 781,000– 40/30,000 years ago). One of the largest known elephant species, mature fully grown bulls on average had a shoulder height of 4 metres (13 ft) and a weight of 13 tonnes (29,000 lb). 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 eastwards into Russia. Skeletons found in association with stone tools and wooden spears suggest they were scavenged and hunted by early humans, including Neanderthals. It is the ancestor of most dwarf elephants that inhabited islands in the Mediterranean.
Description
![](http://upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Palaeoloxodon_antiquus_size_comparison.png/220px-Palaeoloxodon_antiquus_size_comparison.png)
![](http://upload.wikimedia.org/wikipedia/commons/thumb/2/25/Reproducci%C3%B3n_de_Palaeoloxodon_antiquus.jpg/220px-Reproducci%C3%B3n_de_Palaeoloxodon_antiquus.jpg)
Like many other members of the genus Palaeoloxodon, P. antiquus possesses a well developed parieto-occipital crest at the top of the cranium. The crest functioned to anchor muscle tissue, including the splenius as well as an additional muscle layer called the "extra splenius" (which was likely similar to the "splenius superficialis" found in Asian elephants) which wrapped around the top of the head. 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 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, particularly 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]
![](http://upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Palaeoloxodon_antiquus_size_diagram.svg/220px-Palaeoloxodon_antiquus_size_diagram.svg.png)
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. P. antiquus was on average considerably larger than any living elephant. 90% of mature fully grown straight-tusked elephant bulls are estimated to have had shoulder heights in the region of 3.8–4.2 m (12.5–13.8 ft) (average 4 m (13.1 ft)) and a weight between 10.8–15 tonnes (24,000–33,000 lb) (average 13 tonnes (29,000 lb)) (for comparison, 90% of mature fully grown bulls of the largest living elephant species, the African bush elephant have height between 3.04 to 3.36 metres (10.0 to 11.0 ft) and a mass between 5.2–6.9 tonnes (11,000–15,000 lb)). Extremely large bulls, such those represented by a now lost pelvis and tibia collected from San Isidro del Campo in Spain in the late 19th century, may have reached shoulder heights of 4.6 m (15.1 ft) and body masses of over 19 tonnes (42,000 lb).[3][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 and thick tusk (having a circumference of around 77 centimetres (30 in) where it would have exited the skull) 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 reached shoulder heights exceeding 3 metres (9.8 ft) and weights exceeding 5.5 tonnes (12,000 lb) (for comparison, female African bush elephants reach an average shoulder height of 2.6 metres (8.5 ft) and body mass of 3 tonnes (6,600 lb).[3] Newborn and young calves were likely around the same size as those of modern elephants.[7] 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 (3,200–3,300 lb), which is comparable to a similarly aged African bush elephant.[8]
History of discovery and taxonomy
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 by the Collegium Medicum in the nearby city of Gotha to be purely mineral in nature, Wilhelm Ernst Tentzel, a polymath in the employ of the ducal court of Saxe-Gotha-Altenburg, correctly recognised that they represented the remains of an elephant.[9] The species was named in 1847 by British palaeontologists Hugh Falconer and Proby Cautley for remains found in East Sussex, England as Elephas antiquus.[10] The genus Palaeoloxodon was first named in 1924 by Japanese paleontologist Matsumoto Hikoshichirō as a subgenus of Loxodonta, and E. antiquus subsequently assigned to the genus.[11] Some experts historically regarded the larger Asian species Palaeoloxodon namadicus as a variant or subspecies,[12] but they are now considered distinct.[2] Historically, the genus Palaeoloxodon has at times been regarded as a subgenus of Elephas (which contains the living Asian elephant), but this is no longer supported.[13]
![](http://upload.wikimedia.org/wikipedia/commons/thumb/b/b7/Palaeoloxodon_phylogeny.svg/285px-Palaeoloxodon_phylogeny.svg.png)
In 2016, a mitochondrial DNA sequence analysis of P. antiquus found their mitochondrial genomes were nested within the mitochondrial genome diversity of the African forest elephant (Loxodonta cyclotis), with analysis of a partial nuclear genome supporting a closer relationship with L. cyclotis than to the African bush elephant (L. africana).[14][15] A subsequent study published in 2018 by the same authors based on the complete nuclear genome indicated a more complicated relationship between straight-tusked elephants and other species of elephants; according to this study, the biggest genetic contribution to straight-tusked elephants comes from a lineage of elephants that was most closely related but basal to the common ancestor of forest and bush elephants (~60% of total genomic contribution), which hybridized with African forest elephants (>33%) and to a lesser extent with 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,[16] and appears to be shared with other Palaeoloxodon species.[17]
Evolution
Like other Eurasian Palaeoloxodon species, P. antiquus is believed to derive from the African Palaeoloxodon recki. P. antiquus first appears during the early Middle Pleistocene, with its first records around 780,000 years ago in Italy and Israel. Its earliest known appearance in northern Europe is in England around 600,000 years ago. Its arrival coincided with the extinction of the European mammoth species Mammuthus meridionalis and the migration of Mammuthus trogontherii (the steppe mammoth) into Europe from Asia. 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.[18]
Distribution and habitat
![](http://upload.wikimedia.org/wikipedia/commons/thumb/b/ba/Straight_tusked_elephant_Eemian_landscape.jpg/270px-Straight_tusked_elephant_Eemian_landscape.jpg)
Palaeoloxodon 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, western Iran[19] and probably Turkey[20] in West Asia. Some remains of the species have also been reported from Central Asia in northeastern Kazakhstan and Tajikistan. During glacial periods P. antiquus permanently resided in the Mediterranean region. Among the northernmost reported records of the species are from the banks of the Kolva river in the Russian Urals at around the 60th parallel north.[19] 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, such as those from Israel, actually belong to P. recki.[21] The straight-tusked elephant is primarily associated with temperate and Mediterranean forest and woodland habitats, as opposed to the colder open steppe environments inhabited by contemporary mammoths,[22] though the species is also known to have inhabited open grasslands, and is thought to have been tolerant of a range of environmental conditions.[23] Straight-tusked elephants rarely coexisted alongside mammoths, although they occasionally did so, like at the Ilford locality in Britain dating to the Marine Isotope Stage 7 interglacial where both steppe mammoths and P. antiquus are found. At this locality, the two species appear to have engaged in dietary niche partitioning.[4]
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), 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),[19][24] 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).[24] The effect of straight tusked elephant and other extinct megafauna on vegetation likely resulted in increased openness of woodland habitats.[25]
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 males leaving these groups to live solitarily upon reaching adolesence around 14-15 years of age,[7] with adult males likely engaging in combat with each other during musth similar to living elephants.[26] 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 likely of a herd of P. antiquus have been found in dune deposits southern Spain, dating to MIS 5 (130-80,000 years ago).[7] Due to their larger size, straight-tusked elephants are suggested to have finished growing 10 to 15 years later than living elephants, continuing to grow after 50 years of age. They may also have lived longer than extant elephants, with lifespans perhaps in excess of 80 years.[3]
Dental microwear studies suggest that the diet of P. antiquus was highly variable according to the local conditions, ranging from almost completely grazing to nearly totally browsing (feeding on leaves, stems and fruits of high-growing plants), though microwear only reflects the diet in the last few days or weeks before death, so this may be reflecting seasonal dietary variation,[27] as is found in living elephants.[23] Isotopic analysis of a specimen from Greece suggests that it alternately consumed more browse during the dry (presumably summer) months and more grass during the wet (presumably winter) months.[23] A dental mesowear study of specimens from Britain found that most examined specimens had a browsing predominant mixed feeding diet, similar to that inferred for modern African bush elephants and Asian elephants.[28]
Potential gnaw marks suggested to have been made by hyenas and 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 hyenas do on elephants in Africa today.[29]
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.[30][26] These include the Gesher Benot Ya'akov[31] (c. 780 kya,[32] though other authors attribute the remains to P. recki[2]) and Revadim Quarry[33] (sometime between 780 kya and 300-500 kya[32]) sites in Israel, the Ambrona AS3[34] (MIS 12 c. 478-425 kya) , Aridos 1 (MIS 11-9 424-300 kya) & 2[35] (MIS 11 c. 425-375 kya) sites in Spain, the Notarchirico[36] (c. 670-610 kya,[32] though the evidence for butchery at this site is disputed[37]), Ficoncella[38] (c. 500 kya), Castel di Guido[39][40][41] (c. 400 kya[41]), La Polledrara di Cecanibbio (325–310 kya)[32] and Poggetti Vecchi (c. 171 kya)[42] sites in Italy, the Marathousa 1 site (c. 500-400 kya)[43] in Greece, the Ebbsfleet site (c. 425-375 kya)[44] in England, and the Schöningen (c. 300 kya)[45] Gröbern[30][46] Taubach[46] Lehringen[47][30] and Neumark Nord[30] (all dating to the Eemian interglacial, c. 130-115 kya) sites in Germany.[30][46]
The creation of these sites is likely attributable to Homo heidelbergensis and Neanderthals.[26] Stone tools used at these sites include flakes, choppers, bifacial tools like handaxes, as well as cores.[26] At some sites, the bones of straight-tusked elephants[41] and in some cases their ivory[48] 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.[26][46] 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.[26] 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.[26]
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.[47][30] 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.[30][46] 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.[30]
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.[22][49]
Extinction
Palaeoloxodon antiquus retreated from northern Europe after the end of the Eemian interglacial around 115,000 years ago due to climatic conditions becoming unfavourable, and fossils after that time during the Last Glacial Period are rare. A molar from the cave deposits of Grotta Guattari in central 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),[50] with later dating done in 2023 suggesting an age of deposition in the cave of around 66-65,000 years ago.[51][52] Another late Italian record has been reported from Mousterian layers in Barma Grande cave in northwest Italy, probably dating to around the same time as Grotta Guattari, which has been suggested to display evidence of butchery by Neanderthals.[48] Other late remains have been reported from several sites in the Iberian Peninsula,[50][22] including El Castillo cave in northern Spain, dating to 43,000 years ago (alternatively calibrated to around 49,570–44,250 years Before Present[53]), and Foz do Enxarrique (a sequence of terrace deposits of the Tagus river) in central-eastern Portugal, originally dated to around 34-33,000 years ago,[22] but later revised to around 44,000 years ago.[54] 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,[55] but it has been suggested that this date needs independent confirmation, due to only representing a single sample.[56] It has been suggested that P. antiquus likely survived until around 28,000 years ago in the southern Iberian Peninsula based on footprints found in Southwest Portugal[57] and Gibraltar.[58] While some authors have argued that climate change was primarily responsible for the extinction of the straight-tusked elephant,[57] others have argued that climate change alone cannot account for the species extinction.[50] Human hunting has been suggested to have possibly played a contributory role, but the importance of this is uncertain.[57][50]
The extinction of P. antiquus and other temperate adapted European megafauna has resulted in the severe loss of functional diversity in European ecosystems.[24]
Dwarfed descendants
Dwarf elephants that presumably evolved from the straight-tusked elephant are known from many Mediterranean islands, spanning from Sicily and Malta in the west to Cyprus in the east.[50] The responsible factors for the dwarfing of island mammals are thought to be the reduction in food availability, predation and competition.
References
- ^ Benoit, J., Legendre, L. J., Tabuce, R., Obada, T., Mararescul, V., & Manger, P. (2019). Brain evolution in Proboscidea (Mammalia, Afrotheria) across the Cenozoic. Scientific Reports, 9(1), 9323. https://doi.org/10.1038/s41598-019-45888-4
- ^ a b c Larramendi, Asier; Zhang, Hanwen; Palombo, Maria Rita; Ferretti, Marco P. (February 2020). "The evolution of Palaeoloxodon skull structure: Disentangling phylogenetic, sexually dimorphic, ontogenetic, and allometric morphological signals". Quaternary Science Reviews. 229: 106090. Bibcode:2020QSRv..22906090L. doi:10.1016/j.quascirev.2019.106090. S2CID 213676377.
- ^ a b c d e Larramendi, Asier; Palombo, Maria Rita; Marano, Federica (2017). "Reconstructing the life appearance of a Pleistocene giant: size, shape, sexual dimorphism and ontogeny of Palaeoloxodon antiquus (Proboscidea: Elephantidae) from Neumark-Nord 1 (Germany)" (PDF). Bollettino della Società Paleontologica Italiana (3): 299–317. doi:10.4435/BSPI.2017.29. ISSN 0375-7633. Archived from the original (PDF) on 2023-09-30.
- ^ a b Saarinen, Juha; Lister, Adrian M. (October 2016). "Dental mesowear reflects local vegetation and niche separation in Pleistocene proboscideans from Britain: Dental Mesowear in Pleistocene Proboscideans". Journal of Quaternary Science. 31 (7): 799–808. doi:10.1002/jqs.2906.
- ^ Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61. doi:10.4202/app.00136.2014. S2CID 2092950.
- ^ Larramendi, Asier (2023-12-10). "Estimating tusk masses in proboscideans: a comprehensive analysis and predictive model". Historical Biology: 1–14. doi:10.1080/08912963.2023.2286272. ISSN 0891-2963.
- ^ a b c Neto de Carvalho, Carlos; Belaústegui, Zain; Toscano, Antonio; Muñiz, Fernando; Belo, João; Galán, Jose María; Gómez, Paula; Cáceres, Luis M.; Rodríguez-Vidal, Joaquín; Cunha, Pedro Proença; Cachão, Mario; Ruiz, Francisco; Ramirez-Cruzado, Samuel; Giles-Guzmán, Francisco; Finlayson, Geraldine (2021-09-16). "First tracks of newborn straight-tusked elephants (Palaeoloxodon antiquus)". Scientific Reports. 11 (1): 17311. Bibcode:2021NatSR..1117311N. doi:10.1038/s41598-021-96754-1. ISSN 2045-2322. PMC 8445925. PMID 34531420.
- ^ Palombo, Maria Rita; Sanz, Montserrat; Daura, Joan (December 2021). "The complete skeleton of a straight-tusked elephant calf from Cova del Rinoceront (Late Pleistocene, NE Iberian Peninsula): New insights into ontogenetic growth in Palaeoloxodon antiquus". Quaternary Science Reviews. 274: 107257. Bibcode:2021QSRv..27407257P. doi:10.1016/j.quascirev.2021.107257. S2CID 244088519.
- ^ Rieppel, Olivier (2022-05-04). "The first ever described dinosaur bone fragment in Robinet's philosophy of nature (1768)". Historical Biology. 34 (5): 940–946. Bibcode:2022HBio...34..940R. doi:10.1080/08912963.2021.1954176. ISSN 0891-2963.
- ^ Hugh Falconer and Proby Thomas Cautley: Fauna antiqua Sivalensis
- ^ 松本彦七郎 (1924). 日本産化石象の種類(略報) [Types of fossil elephants from Japan]. 地質学雑誌 (in Japanese). 31 (371): 255–272. doi:10.5575/geosoc.31.371_255.
- ^ Mol, D., De Vos, J., Reumer, J.W.F., 1999. Praeovibos priscus (Bovidae, Artiodactyla, mammalia) from the North Sea and aspects of its paleoecology. In: Reumer, J.W.F., De Vos, J. (Eds.), 1999. Elephants have a snorkel! Papers in Honour of Paul Y. Sondaar. Deinsea 7, 223–232.
- ^ Shoshani, J.; Ferretti, M. P.; Lister, A. M.; Agenbroad, L. D.; Saegusa, H.; Mol, D.; Takahashi, K. (2007). "Relationships within the Elephantinae using hyoid characters". Quaternary International. 169–170: 174–185. Bibcode:2007QuInt.169..174S. doi:10.1016/j.quaint.2007.02.003.
- ^ Callaway, E. (2016-09-16). "Elephant history rewritten by ancient genomes". Nature. doi:10.1038/nature.2016.20622. S2CID 89500906.
- ^ Meyer, Matthias; Palkopoulou, Eleftheria; Baleka, Sina; Stiller, Mathias; Penkman, Kirsty E H; Alt, Kurt W; Ishida, Yasuko; Mania, Dietrich; Mallick, Swapan; Meijer, Tom; Meller, Harald; Nagel, Sarah; Nickel, Birgit; Ostritz, Sven; Rohland, Nadin; Schauer, Karol; Schüler, Tim; Roca, Alfred L; Reich, David; Shapiro, Beth; Hofreiter, Michael (6 June 2017). "Palaeogenomes of Eurasian straight-tusked elephants challenge the current view of elephant evolution". eLife. 6: e25413. doi:10.7554/eLife.25413. PMC 5461109. PMID 28585920.
- ^ Eleftheria Palkopoulou; Mark Lipson; Swapan Mallick; Svend Nielsen; Nadin Rohland; Sina Baleka; Emil Karpinski; Atma M. Ivancevic; Thu-Hien To; R. Daniel Kortschak; Joy M. Raison; Zhipeng Qu; Tat-Jun Chin; Kurt W. Alt; Stefan Claesson; Love Dalén; Ross D. E. MacPhee; Harald Meller; Alfred L. Roca; Oliver A. Ryder; David Heiman; Sarah Young; Matthew Breen; Christina Williams; Bronwen L. Aken; Magali Ruffier; Elinor Karlsson; Jeremy Johnson; Federica Di Palma; Jessica Alfoldi; David L. Adelson; Thomas Mailund; Kasper Munch; Kerstin Lindblad-Toh; Michael Hofreiter; Hendrik Poinar; David Reich (2018). "A comprehensive genomic history of extinct and living elephants". Proceedings of the National Academy of Sciences of the United States of America. 115 (11): E2566–E2574. Bibcode:2018PNAS..115E2566P. doi:10.1073/pnas.1720554115. PMC 5856550. PMID 29483247.
- ^ Lin, Haifeng; Hu, Jiaming; Baleka, Sina; Yuan, Junxia; Chen, Xi; Xiao, Bo; Song, Shiwen; Du, Zhicheng; Lai, Xulong; Hofreiter, Michael; Sheng, Guilian (July 2023). "A genetic glimpse of the Chinese straight-tusked elephants". Biology Letters. 19 (7). doi:10.1098/rsbl.2023.0078. ISSN 1744-957X. PMC 10353889. PMID 37463654.
- ^ Lister, Adrian M. (2004), "Ecological Interactions of Elephantids in Pleistocene Eurasia", Human Paleoecology in the Levantine Corridor, Oxbow Books, pp. 53–60, ISBN 978-1-78570-965-4, retrieved 2020-04-14
- ^ a b c Pushkina, Diana (July 2007). "The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian Palaeoloxodon antiquus assemblage". Mammal Review. 37 (3): 224–245. doi:10.1111/j.1365-2907.2007.00109.x. ISSN 0305-1838.
- ^ Albayrak, Ebru; Lister, Adrian M. (October 2012). "Dental remains of fossil elephants from Turkey". Quaternary International. 276–277: 198–211. Bibcode:2012QuInt.276..198A. doi:10.1016/j.quaint.2011.05.042.
- ^ Pokines, James T.; Lister, Adrian M.; Ames, Christopher J. H.; Nowell, April; Cordova, Carlos E. (March 2019). "Faunal remains from recent excavations at Shishan Marsh 1 (SM1), a Late Lower Paleolithic open-air site in the Azraq Basin, Jordan". Quaternary Research. 91 (2): 768–791. Bibcode:2019QuRes..91..768P. doi:10.1017/qua.2018.113. ISSN 0033-5894.
- ^ a b c d Braun, Ingmar M.; Palombo, Maria Rita (October 2012). "Mammuthus primigenius in the cave and portable art: An overview with a short account on the elephant fossil record in Southern Europe during the last glacial". Quaternary International. 276–277: 61–76. Bibcode:2012QuInt.276...61B. doi:10.1016/j.quaint.2012.07.010.
- ^ a b c Roditi, Effrosyni; Bocherens, Hervé; Konidaris, George E.; Athanassiou, Athanassios; Tourloukis, Vangelis; Karkanas, Panagiotis; Panagopoulou, Eleni; Harvati, Katerina (2024-01-16). "Life-history of Palaeoloxodon antiquus reveals Middle Pleistocene glacial refugium in the Megalopolis basin, Greece". Scientific Reports. 14 (1): 1390. Bibcode:2024NatSR..14.1390R. doi:10.1038/s41598-024-51592-9. ISSN 2045-2322. PMC 10791645. PMID 38228659.
- ^ a b c Davoli, Marco; Monsarrat, Sophie; Pedersen, Rasmus Østergaard; Scussolini, Paolo; Karger, Dirk Nikolaus; Normand, Signe; Svenning, Jens-Christian (January 2024). "Megafauna diversity and functional declines in Europe from the Last Interglacial to the present". Global Ecology and Biogeography. 33 (1): 34–47. Bibcode:2024GloEB..33...34D. doi:10.1111/geb.13778. hdl:11573/1714498. ISSN 1466-822X.
- ^ Pearce, Elena A.; Mazier, Florence; Normand, Signe; Fyfe, Ralph; Andrieu, Valérie; Bakels, Corrie; Balwierz, Zofia; Bińka, Krzysztof; Boreham, Steve; Borisova, Olga K.; Brostrom, Anna; de Beaulieu, Jacques-Louis; Gao, Cunhai; González-Sampériz, Penélope; Granoszewski, Wojciech (2023-11-10). "Substantial light woodland and open vegetation characterized the temperate forest biome before Homo sapiens". Science Advances. 9 (45): eadi9135. Bibcode:2023SciA....9I9135P. doi:10.1126/sciadv.adi9135. ISSN 2375-2548. PMC 10637746. PMID 37948521.
- ^ a b c d e f g Konidaris, George E.; Tourloukis, Vangelis (2021-04-14). "Proboscidea-Homo interactions in open-air localities during the Early and Middle Pleistocene of western Eurasia: a palaeontological and archaeolocigal perspective". Human-Elephant Interactions: From Past to Present. doi:10.15496/publikation-55599.
- ^ Rivals, Florent; Semprebon, Gina M.; Lister, Adrian M. (September 2019). "Feeding traits and dietary variation in Pleistocene proboscideans: A tooth microwear review". Quaternary Science Reviews. 219: 145–153. Bibcode:2019QSRv..219..145R. doi:10.1016/j.quascirev.2019.06.027. S2CID 200073388.
- ^ Saarinen, Juha; Lister, Adrian M. (October 2016). "Dental mesowear reflects local vegetation and niche separation in Pleistocene proboscideans from Britain: Dental Mesowear in Pleistocene Proboscideans". Journal of Quaternary Science. 31 (7): 799–808. doi:10.1002/jqs.2906. S2CID 132421364.
- ^ Diedrich, Cajus G. (September 2014). "Late Pleistocene Eemian hyena and steppe lion feeding strategies on their largest prey—Palaeoloxodon antiquus Falconer and Cautley 1845 at the straight-tusked elephant graveyard and Neanderthal site Neumark-Nord Lake 1, Central Germany". Archaeological and Anthropological Sciences. 6 (3): 271–291. Bibcode:2014ArAnS...6..271D. doi:10.1007/s12520-013-0150-7. ISSN 1866-9557.
- ^ a b c d e f g h Gaudzinski-Windheuser, Sabine; Kindler, Lutz; MacDonald, Katharine; Roebroeks, Wil (2023). "Hunting and processing of straight-tusked elephants 125.000 years ago: Implications for Neanderthal behavior". Science Advances. 9 (5): eadd8186. Bibcode:2023SciA....9D8186G. doi:10.1126/sciadv.add8186. PMC 9891704. PMID 36724231.
- ^ N. Goren-Inbar, A. Lister, E. Werker, M.A. Chech A butchered elephant skull and associated artifacts from the Acheulian site of Gesher Benot Ya'aqov Israel Paléorient, 20 (1994), pp. 99-112
- ^ a b c d Santucci, Ernesto; Marano, Federica; Cerilli, Eugenio; Fiore, Ivana; Lemorini, Cristina; Palombo, Maria Rita; Anzidei, Anna Paola; Bulgarelli, Grazia Maria (2016-06-25). "Palaeoloxodon exploitation at the Middle Pleistocene site of La Polledrara di Cecanibbio (Rome, Italy)". Quaternary International. VIth International Conference on Mammoths and their Relatives, Part 2. 406: 169–182. Bibcode:2016QuInt.406..169S. doi:10.1016/j.quaint.2015.08.042. ISSN 1040-6182.
- ^ Rabinovich, R.; Ackermann, O.; Aladjem, E.; Barkai, R.; Biton, R.; Milevski, I.; Solodenko, N.; Marder, O. (October 2012). "Elephants at the Middle Pleistocene Acheulian open-air site of Revadim Quarry, Israel". Quaternary International. 276–277: 183–197. Bibcode:2012QuInt.276..183R. doi:10.1016/j.quaint.2012.05.009.
- ^ Villa, Paola; Soto, Enrique; Santonja, Manuel; Pérez-González, Alfredo; Mora, Rafael; Parcerisas, Joaquim; Sesé, Carmen (2005-01-01). "New data from Ambrona: closing the hunting versus scavenging debate". Quaternary International. Studying Proboscideans: knowledge, Problems and Perspectives. Selected papers from "The world of Elephants" Congress, Rome. 126–128: 223–250. Bibcode:2005QuInt.126..223V. doi:10.1016/j.quaint.2004.03.001. ISSN 1040-6182.
- ^ Yravedra, J.; Domínguez-Rodrigo, M.; Santonja, M.; Pérez-González, A.; Panera, J.; Rubio-Jara, S.; Baquedano, E. (October 2010). "Cut marks on the Middle Pleistocene elephant carcass of Áridos 2 (Madrid, Spain)". Journal of Archaeological Science. 37 (10): 2469–2476. Bibcode:2010JArSc..37.2469Y. doi:10.1016/j.jas.2010.05.007.
- ^ The elephant butchery area at the middle Pleistocene site of Notarchirico (Venosa, Basilicata, Italy) G. Cavarretta, P. Gioia, M. Mussi, M.R. Palombo (Eds.), The World of Elephants. Proceedings of the First International Congress, Consiglio Nazionale delle Ricerche, Roma (2001), pp. 230-236
- ^ Pineda, Antonio; Mecozzi, Beniamino; Iannucci, Alessio; Carpentieri, Marco; Sardella, Raffaele; Rabinovich, Rivka; Moncel, Marie-Hélène (May 2024). "Reevaluating the "elephant butchery area" at the Middle Pleistocene site of Notarchirico (MIS 16) (Venosa Basin, Basilicata, Italy)". Quaternary Science Reviews. 331: 108603. Bibcode:2024QSRv..33108603P. doi:10.1016/j.quascirev.2024.108603.
- ^ Aureli, Daniele; Contardi, Antonio; Giaccio, Biagio; Jicha, Brian; Lemorini, Cristina; Madonna, Sergio; Magri, Donatella; Marano, Federica; Milli, Salvatore; Modesti, Valerio; Palombo, Maria Rita; Rocca, Roxane (2015-04-21). "Palaeoloxodon and Human Interaction: Depositional Setting, Chronology and Archaeology at the Middle Pleistocene Ficoncella Site (Tarquinia, Italy)". PLOS ONE. 10 (4): e0124498. Bibcode:2015PLoSO..1024498A. doi:10.1371/journal.pone.0124498. ISSN 1932-6203. PMC 4405345. PMID 25898322.
- ^ Saccà, Daniela (October 2012). "Taphonomy of Palaeloxodon antiquus at Castel di Guido (Rome, Italy): Proboscidean carcass exploitation in the Lower Palaeolithic". Quaternary International. 276–277: 27–41. Bibcode:2012QuInt.276...27S. doi:10.1016/j.quaint.2012.03.055.
- ^ Boschian, Giovanni; Saccà, Daniela (March 2015). "In the elephant, everything is good: Carcass use and re-use at Castel di Guido (Italy)". Quaternary International. 361: 288–296. Bibcode:2015QuInt.361..288B. doi:10.1016/j.quaint.2014.04.030.
- ^ a b c Villa, Paola; Boschian, Giovanni; Pollarolo, Luca; Saccà, Daniela; Marra, Fabrizio; Nomade, Sebastien; Pereira, Alison (2021-08-26). Peresani, Marco (ed.). "Elephant bones for the Middle Pleistocene toolmaker". PLOS ONE. 16 (8): e0256090. Bibcode:2021PLoSO..1656090V. doi:10.1371/journal.pone.0256090. ISSN 1932-6203. PMC 8389514. PMID 34437571.
- ^ Aranguren, Biancamaria; Grimaldi, Stefano; Benvenuti, Marco; Capalbo, Chiara; Cavanna, Floriano; Cavulli, Fabio; Ciani, Francesco; Comencini, Giacomo; Giuliani, Claudia; Grandinetti, Giuditta; Mariotti Lippi, Marta; Masini, Federico; Mazza, Paul Peter Anthony; Pallecchi, Pasquino; Santaniello, Fabio (August 2019). "Poggetti Vecchi (Tuscany, Italy): A late Middle Pleistocene case of human–elephant interaction". Journal of Human Evolution. 133: 32–60. Bibcode:2019JHumE.133...32A. doi:10.1016/j.jhevol.2019.05.013. PMID 31358183. S2CID 196643105.
- ^ Panagopoulou, Eleni; Tourloukis, Vangelis; Thompson, Nicholas; Konidaris, George; Athanassiou, Athanassios; Giusti, Domenico; Tsartsidou, Georgia; Karkanas, Panagiotis; Harvati, Katerina (2018-12-20). "The Lower Palaeolithic site of Marathousa 1, Megalopolis, Greece: Overview of the evidence". Quaternary International. The Gates of Europe. 497: 33–46. Bibcode:2018QuInt.497...33P. doi:10.1016/j.quaint.2018.06.031. ISSN 1040-6182. S2CID 133849514.
- ^ Wenban-Smith, F. F.; Allen, P.; Bates, M. R.; Parfitt, S. A.; Preece, R. C.; Stewart, J. R.; Turner, C.; Whittaker, J. E. (July 2006). "The Clactonian elephant butchery site at Southfleet Road, Ebbsfleet, UK". Journal of Quaternary Science. 21 (5): 471–483. Bibcode:2006JQS....21..471W. doi:10.1002/jqs.1033. ISSN 0267-8179. S2CID 55705671.
- ^ Venditti, Flavia; Rodríguez-Álvarez, Bárbara; Serangeli, Jordi; Cesaro, Stella Nunziante; Walter, Rudolf; Conard, Nicholas J. (2022-12-15). "Using microartifacts to infer Middle Pleistocene lifeways at Schöningen, Germany". Scientific Reports. 12 (1): 21148. Bibcode:2022NatSR..1221148V. doi:10.1038/s41598-022-24769-3. ISSN 2045-2322. PMC 9755147. PMID 36522355.
- ^ a b c d e Gaudzinski-Windheuser, Sabine; Kindler, Lutz; Roebroeks, Wil (2023-12-12). "Widespread evidence for elephant exploitation by Last Interglacial Neanderthals on the North European plain". Proceedings of the National Academy of Sciences. 120 (50): e2309427120. Bibcode:2023PNAS..12009427G. doi:10.1073/pnas.2309427120. ISSN 0027-8424. PMC 10723128. PMID 38048457.
- ^ a b H. Thieme, S. Veil, Neue Untersuchungen zum eemzeitlichen Elefanten-Jagdplatz Lehringen, Ldkr. Verden. Kunde 36, 11–58 (1985).
- ^ a b Onoratini, Gérard; Arellano, Almudena; Del Lucchese, Angiolo; Moullé, Pierre Elie; Serre, Frédéric (March 2012). "The Barma Grande cave (Grimaldi, Vintimiglia, Italy): From Neandertal, hunter of "Elephas antiquus", to Sapiens with ornaments of mammoth ivory". Quaternary International. 255: 141–157. Bibcode:2012QuInt.255..141O. doi:10.1016/j.quaint.2011.05.015.
- ^ ARCA A., 2014 - Elephas antiquus depicted at Vermelhosa rock art? TRACCE Online Rock Art Bulletin, 31, online
- ^ a b c d e Stuart, Anthony J. (2005). "The extinction of woolly mammoth (Mammuthus primigenius) and straight-tusked elephant (Palaeoloxodon antiquus) in Europe" (PDF). Quaternary International. 126–128. Elsevier BV: 171–177. Bibcode:2005QuInt.126..171S. doi:10.1016/j.quaint.2004.04.021. ISSN 1040-6182.
- ^ Petronio, Carmelo; Rolfo, Mario; Di Mario, Francesco; Ceruleo, Piero; Ferracci, Angelica; Fiore, Ivana; Gatta, Maurizio; Salari, Leonardo (2022-01-26). "Preliminary report on the new faunal remains from Grotta Guattari (Late Pleistocene, San Felice Circeo, Latium)". Bulletin of Regional Natural History. 1 (4): 1–10 Paginazione. doi:10.6093/2724-4393/8927.
- ^ Rolfo, Mario Federico; Bini, Monica; Di Mario, Francesco; Ferracci, Angelica; Giaccio, Biagio; Hsun-Ming, Hu; Isola, Ilaria; Sadori, Laura; Shen, Chuan-Chou; Vignola, Cristiano; Zanchetta, Giovanni (July 2023). "Neanderthal bones collected by hyena at Grotta Guattari, central Italy, 66–65 ka: U/Th chronology and paleoenvironmental setting". Quaternary Science Reviews. 311: 108132. Bibcode:2023QSRv..31108132R. doi:10.1016/j.quascirev.2023.108132.
- ^ Wood, Rachel; Bernaldo de Quirós, Federico; Maíllo-Fernández, José-Manuel; Tejero, José-Miguel; Neira, Ana; Higham, Thomas (April 2018). "El Castillo (Cantabria, northern Iberia) and the Transitional Aurignacian: Using radiocarbon dating to assess site taphonomy". Quaternary International. 474: 56–70. Bibcode:2018QuInt.474...56W. doi:10.1016/j.quaint.2016.03.005.
- ^ Cunha, Pedro P.; Martins, António A.; Buylaert, Jan-Pieter; Murray, Andrew S.; Gouveia, Maria P.; Font, Eric; Pereira, Telmo; Figueiredo, Silvério; Ferreira, Cristiana; Bridgland, David R.; Yang, Pu; Stevaux, José C.; Mota, Rui (2019-01-18). "The Lowermost Tejo River Terrace at Foz do Enxarrique, Portugal: A Palaeoenvironmental Archive from c. 60–35 ka and Its Implications for the Last Neanderthals in Westernmost Iberia". Quaternary. 2 (1): 3. doi:10.3390/quat2010003. hdl:10174/24689. ISSN 2571-550X.
- ^ Mol, Dick; de Vos, John; van der Plicht, Johannes (July 2007). "The presence and extinction of Elephas antiquus Falconer and Cautley, 1847, in Europe". Quaternary International. 169–170: 149–153. Bibcode:2007QuInt.169..149M. doi:10.1016/j.quaint.2006.06.002.
- ^ van der Plicht, J.; Palstra, S.W.L. (June 2016). "Radiocarbon and mammoth bones: What's in a date". Quaternary International. 406: 246–251. Bibcode:2016QuInt.406..246V. doi:10.1016/j.quaint.2014.11.027.
- ^ a b c de Carvalho, Carlos Neto; Figueiredo, Silvério; Muniz, Fernando; Belo, João; Cunha, Pedro P.; Baucon, Andrea; Cáceres, Luis M.; Rodriguez-Vidal, Joaquín (2020-07-02). "Tracking the last elephants in Europe during the Würm Pleniglacial: the importance of the Late Pleistocene aeolianite record in SW Iberia". Ichnos. 27 (3): 352–360. Bibcode:2020Ichno..27..352D. doi:10.1080/10420940.2020.1744586. ISSN 1042-0940. S2CID 216504699.
- ^ Muñiz, Fernando; Cáceres, Luis M.; Rodríguez-Vidal, Joaquín; Neto de Carvalho, Carlos; Belo, João; Finlayson, Clive; Finlayson, Geraldine; Finlayson, Stewart; Izquierdo, Tatiana; Abad, Manuel; Jiménez-Espejo, Francisco J.; Sugisaki, Saiko; Gómez, Paula; Ruiz, Francisco (August 2019). "Following the last Neanderthals: Mammal tracks in Late Pleistocene coastal dunes of Gibraltar (S Iberian Peninsula)". Quaternary Science Reviews. 217: 297–309. Bibcode:2019QSRv..217..297M. doi:10.1016/j.quascirev.2019.01.013.