Straight-tusked elephant

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Straight-tusked elephant
Temporal range: Mid-Late Pleistocene
~0.78–0.03  Ma
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Elephas antiquus Rom (cropped).JPG
Skeleton in Rome
Elefantenreich 03.jpg
Skull in Germany
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Genus:† Palaeoloxodon
Species:
†P. antiquus
Binomial name
†Palaeoloxodon antiquus
(Falconer & Cautley, 1847)
Palaeoloxodon antiquus map.png
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 elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene (781,000–30,000 years Before Present). It was larger than any living elephant, with adult males suggested to reach 3.81–4.2 metres (12.5–13.8 ft) in shoulder height, and 11.3–15 tonnes (12.5–16.5 short tons) in weight. Like modern elephants, the straight-tusked elephant lived in herds, flourishing during interglacial periods, when its range would extend as far north as Great Britain. Skeletons found in association with stone tools and wooden spears suggest they were scavenged and hunted by early humans, including Neanderthals. It is the ancestral species of most dwarf elephants that inhabited islands in the Mediterranean.

Description

Size of a 40 year old adult male compared to a human Palaeoloxodon antiquus size comparison.png
Size of a 40 year old adult male compared to a human
Model Reproduccion de Palaeoloxodon antiquus.jpg
Model

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 muscule 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, 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 (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 (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 (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 (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 (1.60–1.65 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 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. [8] The species was named in 1847 by Hugh Falconer and Proby Cautley for remains found in East Sussex as Elephas antiquus. [9] The genus Palaeoloxodon was first named in 1924 by Matsumoto Hikoshichirō as a subgenus of 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 (which contains the living Asian elephant), but this is no longer supported. [12]

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

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). [13] [14] 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 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, [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 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 replacement of Mammuthus meridionalis by 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

Life restoration of two straight-tusked elephants during the Eemian interglacial in a temperate forest landscape characterised by grassy areas, open woodland and closed forest Straight tusked elephant Eemian landscape.jpg
Life restoration of two straight-tusked elephants during the 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 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), [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]

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]

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, [26] as is found in living elephants. [21] Isotopic analysis of a specimen from Greece suggests that it alternately consumed more browse during the dry months and more grass during the wet months. [21] A 2016 dental mesowear study of specimens from Britain found that most examined specimens had a browsing predominant mixed feeding diet, with clear 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. [28] [29] These include the Gesher Benot Ya'akov [30] (c. 780 kya, [31] though other authors attribute the remains to P. recki [2] ) and Revadim Quarry [32] (sometime between 780 kya and 300-500 kya [31] ) sites in Israel, the Ambrona AS3 [33] (MIS 12 c. 478-425 kya) , Aridos 1 (MIS 11-9 424-300 kya) & 2 [34] (MIS 11 c. 425-375 kya) sites in Spain, the Notarchirico [35] (c. 670-610 kya, [31] though the evidence for butchery at this site is disputed [36] ), Ficoncella [37] (c. 500 kya), Castel di Guido [38] [39] [40] (c. 400 kya [40] ), La Polledrara di Cecanibbio (325–310 kya) [31] and Poggetti Vecchi (c. 171 kya) [41] sites in Italy, the Marathousa 1 site (c. 500-400 kya) [42] in Greece, the Ebbsfleet site (c. 425-375 kya) [43] in England, and the Schöningen (c. 300 kya) [44] Gröbern [28] [45] Taubach [45] Lehringen [46] [28] and Neumark Nord [28] (all dating to the Eemian interglacial, c. 130-115 kya) sites in Germany. [28] [45]

The creation of these sites is likely attributable to Homo heidelbergensis and 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

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. [48] The responsible factors for the dwarfing of island mammals are thought to be the reduction in food availability, predation and competition.

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<span class="mw-page-title-main">Elephantidae</span> Family of mammals

Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are large terrestrial mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the family are extinct. Only two genera, Loxodonta and Elephas, are living.

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References

  1. ↑ 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
  2. 1 2 3 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.
  3. 1 2 3 4 5 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.
  4. ↑ 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.
  5. ↑ Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61. doi:10.4202/app.00136.2014. S2CID   2092950.
  6. ↑ 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.
  7. ↑ 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.
  8. ↑ 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.
  9. ↑ Hugh Falconer and Proby Thomas Cautley: Fauna antiqua Sivalensis
  10. ↑ 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 .
  11. ↑ 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. Elephantshave a snorkel! Papers in Honour of Paul Y. Sondaar. Deinsea 7, 223–232.
  12. ↑ 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.
  13. ↑ Callaway, E. (2016-09-16). "Elephant history rewritten by ancient genomes". Nature. doi:10.1038/nature.2016.20622. S2CID   89500906.
  14. ↑ 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.
  15. ↑ 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.
  16. ↑ 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.
  17. ↑ 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
  18. 1 2 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.
  19. ↑ 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.
  20. 1 2 3 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.
  21. 1 2 3 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.
  22. ↑ 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.
  23. 1 2 3 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. ISSN   1466-822X.
  24. ↑ 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.
  25. ↑ 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.
  26. 1 2 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.
  27. ↑ 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.
  28. 1 2 3 4 5 6 7 8 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.
  29. 1 2 3 4 5 6 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.
  30. ↑ 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
  31. 1 2 3 4 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.
  32. ↑ 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.
  33. ↑ 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.
  34. ↑ 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.
  35. ↑ 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
  36. ↑ 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. doi: 10.1016/j.quascirev.2024.108603 .
  37. ↑ 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.
  38. ↑ 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.
  39. ↑ 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.
  40. 1 2 3 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.
  41. ↑ 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. doi:10.1016/j.jhevol.2019.05.013. PMID   31358183. S2CID   196643105.
  42. ↑ 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.
  43. ↑ 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.
  44. ↑ 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.
  45. 1 2 3 4 5 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.
  46. 1 2 H. Thieme, S. Veil, Neue Untersuchungen zum eemzeitlichen Elefanten-Jagdplatz Lehringen, Ldkr. Verden. Kunde36, 11–58 (1985).
  47. ↑ ARCA A., 2014 - Elephas antiquus depicted at Vermelhosa rock art? TRACCE Online Rock Art Bulletin, 31, online
  48. 1 2 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.
  49. ↑ 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.
  50. ↑ 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.
  51. ↑ 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.
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