Straight-tusked elephant

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). Recovered individuals have reached up to 4–4.2 metres (13.1–13.8 ft) in height, and an estimated 11.3–15 tonnes (11.1–14.8 long tons; 12.5–16.5 short tons) in weight. The straight-tusked elephant probably lived in small herds, flourishing in interglacial periods, when its range would extend as far north as Great Britain. Isolated tusks are often found while partial or whole skeletons are rare, and there is evidence of predation by early humans. It is the ancestral species of most dwarf elephants that inhabited islands in the Mediterranean.

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 ↓
Skull and model (from Ambrona)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Proboscidea
Family:	Elephantidae
Genus:	†Palaeoloxodon
Species:	†P. antiquus
Binomial name
†Palaeoloxodon antiquus (Falconer & Cautley, 1847)
Approximate range of P. antiquus
Synonyms
Elephas antiquus (Falconer & Cautley, 1847) Loxodonta antiquus^[1]

Description

 

Complete skeleton in Rome.

Palaeoloxodon antiquus was quite large, with individuals reaching 4 metres (13.1 ft) in height. Like other members of Palaeoloxodon, P. antiquus possesses a well developed parieto-occipital crest at the top of the cranium that anchored the splenius as well as possibly the rhomboid muscles to support the large head, 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] One approximately 40-year-old male specimen of P. antiquus measured about 3.81 metres (12.5 ft) tall and weighed about 11.3 tonnes (11.1 long tons; 12.5 short tons), while another from Montreuil weighed about 15 tonnes (14.8 long tons; 16.5 short tons) and was about 4.2 metres (13.8 ft) tall.^[3] and had long, slightly upward-curving tusks.^[4] P. antiquus's legs were slightly longer than those of modern elephants. This elephant is thought, based on the shape of the hyoid bone, to have had an 80-cm-long tongue that could be projected a short distance from the mouth to grasp leaves and grasses. With this tongue and a flexible trunk, straight-tusked elephants could graze or browse on Pleistocene foliage about 8 metres (26 ft) above ground.^[5]

 

Life restoration

Taxonomy

The species was first named in 1847 by Hugh Falconer and Proby Cautley for remains found in East Sussex as Elephas antiquus.^[6] The genus Palaeoloxodon was first named in 1924 by Hikoshichiro Matsumoto as a subgenus of Loxodonta, and E. antiquus subsequently assigned to the genus.^[7] Some experts regarded the larger Asian species Palaeoloxodon namadicus as a variant or subspecies,^[8] but they are now considered distinct.^[2] Historically, the genus Palaeoloxodon has at times been regarded as a subgenus of Elephas, but a 2007 study of hyoid characteristics amongst living and fossil elephants has largely led to an abandonment of this hypothesis.^[9] In 2016, a Mitochondrial DNA sequence analysis of P. antiquus suggested that its closest extant relative was the African forest elephant (L. cyclotis). The paper argues that P. antiquus is closer to L. cyclotis than L. cyclotis is to the African bush elephant, L. africana, thus invalidating the genus Loxodonta as currently recognized.^[10][11] A subsequent study published by Palkopoulou et al. (2018) based on the 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 basal to the common ancestor of forest and bush elephants, which subsequently hybridized with members of the lineage related to extant African forest elephants and with the lineage related to woolly mammoths.^[12]

Evolution

P. antiquus is believed to derive from the African P. recki. P. antiquus first appears during the early Middle Pleistocene, around 0.8–0.6 Ma, appearing around 780 kya in Italy; its earliest known appearance in northern Europe is in Suffolk around 600 kya. 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.^[13]

Behaviour

Straight-tusked elephants probably lived in small herds of about 5 to 15 individuals.^[14] Like its recent relatives, the straight-tusked elephant would have been heavily dependent on fresh water, which greatly influenced its migration.^[15] 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, though microwear only reflects the diet in the last few days or weeks before death, so this may be reflecting seasonal dietary variation. Mesowear suggests a browsing predominant mixed feeding diet, with clear niche separation from the more grazing dominated diet of the sympatric Mammuthus trogontherii.^[16][17]

Range

They preferred warm conditions and flourished in the interglacial periods during the current Ice Age, expanding their range from Southern Europe as far North as Great Britain during the warmer interglacials, while permanently residing in Mediterranean Europe during glacial periods. The straight-tusked elephant became extinct in Britain near the beginning of the Weichselian glaciation, about 115,000 years ago.^[18] P. antiquus likely survived until around 28,000 years ago in the southern Iberian Peninsula, based on footprints.^[19]

Excavations

 

Skeleton in Naturkunde Museum, Berlin

 

Illustration from 1916

 

Full-size reconstruction in the Museum of Barnstaple and North Devon

Finds of isolated tusks are relatively common in Great Britain. For example, a tusk of this elephant was found during the construction of the Swan Valley Community School in Swanscombe, Kent. However, finds of whole or partial skeletons of this elephant are very rare.

Skeleton finds in Britain are known from only a few sites. Two sites were found in the Lower Thames basin, one at Upnor, Kent and one at Aveley, Essex. Paleontological and archaeological excavations in advance of High Speed 1 revealed the 400,000-year-old skeleton of a straight-tusked elephant in the Ebbsfleet Valley, near Swanscombe. It was lying at the edge of what would once have been a small lake. Flint tools lay scattered around, suggesting the elephant had been cut up by a group of the early humans around at the time, likely Homo heidelbergensis.^[20]

On the European mainland, many remains of the straight-tusked elephant have been found. In addition to skeletons, some sites contained additional archaeological material, as in the Ebbsfleet site. In Greece, three partial skeletons have been recovered from the province of West Macedonia,^{[21][22][23][24]} and a Palaeoloxodon antiquus butchering site has been excavated near Megalopolis, in the Peloponnese.^[25][26]

Straight-tusked elephant remains have been found with flint tools at a number of other sites, such as Torralba and Aridos in Spain, Notarchirico in Italy, and Gröbern and Ehringsdorf in Germany.

A Palaeolithic scratched figure of an elephant head in the Vermelhosa area, Portugal, near the Côa Valley Park, is reported to be the depiction of an Palaeoloxodon antiquus.^[27] The Iberian peninsula may have served as the last European refuge of the straight-tusked elephant. According to João Luís Cardoso,^[28] the species survived until 30,000 years BP in Portugal.

Elephants and early humans

Both early human beings and the straight-tusked elephant reached the European continent in the late Early Pleistocene. Sites where early humans and the straight-tusked elephant appear together have been frequently documented. One of the earliest finds is in an approximately 600,000-year-old site near Heidelberg, the location of the early human jawbone dubbed Mauer 1. Fossils were also found washed up in a course of the Neckar river. Isernia la Pineta in Italy is likely to be the same age or slightly younger. Remains of lower jaws are well documented in these three locations, alongside stone artefacts and numerous bones of rhinos, wild cattle, hippos and horses. Since the remains of the trunks do not show human markings, extinction by natural causes can not be ruled out.

The clearest proof to date that this animal was among the prey of early humans was provided in 1948 in Lehringen, Germany, near the Aller river. A skeleton of a straight-tusked elephant was found with a yew spear between the ribs, and lithic artifacts around the head. The find dates to the Eemian interglacial period.^[29] A similar find of a carcass exploited by humans from the same time period was found in 1987 in the Gröbern strip mine in Saxony-Anhalt. There were over 20 flint artifacts found, but there was no evidence of a hunting weapon.^[30] A second elephant skeleton found in Gröbern the same year proved to be free of any markings of human use.^[31]

Remains have also been recovered from another Eemian site, the former lake basin Neumark-Nord 1 on the northeast edge of the Geisel valley.^[32] Over 1,350 bone finds from straight-tusked elephants have been found from the basin since 1985, including some skeletons that were almost completely preserved. These remains belonged to about 70 individuals. They were found along with other animals such as rhinoceros (Stephanorhinus spp.), aurochs, horse, red deer and fallow deer. A large amount of old or sick elephants died there.^[33] None of the animals showed any traces of deliberate killing by early humans; however, flint artifacts also occurred, including one containing an oak bark extract that was apparently used for tanning.^[34]

Various archaeologists in Germany have dealt with the question of elephant hunting in the Paleolithic. Thorsten Uthmeier from FAU Erlangen-Nürnberg thinks regular elephant hunts are unlikely. With the assumed clan size of five to ten people and a shelf life of 30 days for meat, only animals with a weight of up to one ton, such as cattle, deer or horses, would be considered as game. Elephants would provide up to ten times more meat than the group could consume over the period. However, elephants are still hunted with spears by pygmies in the central African rainforest. Michael Baales from the Ruhr University considers the role of elephants to be important at some European sites, even if, despite the presence of cut marks, it cannot be clearly decided whether the animals were hunted or dead animals were eviscerated. After examining elephant remains, Sabine Gaudzinski-Windheuser from Johannes Gutenberg University. Mainz came to the conclusion that Stone Age material can prove the presence of humans in the vicinity of the sites. Nicholas J. Conard from the Eberhard Karls, University of Tübingen considers the role of elephants in the Paleolithic economy to be elusive.^[35]

In 2021 it was found that 400,000 years ago at Castel di Guido, Italy, a large number of bone tools, made from elephant bone were created. The tools are more sophisticated than tools found at other sites dating from the same period. They appear to have been made using production line assembly. Paola Villa, a University of Colorado Boulder archaeologist, suspects the tools were made by early Neanderthals. Techniques used to make the tools don't appear to have been used at other sites for another 100,000 years.^[36]

Dwarfed descendants

 

Models

Main article: Dwarf elephant § The Mediterranean islands

Elephants that presumably evolved from the straight-tusked elephant are described from many Mediterranean islands, where they evolved into dwarfed elephants.^[37] The responsible factors for the dwarfing of island mammals are thought to be the reduction in food availability, predation and competition.

References

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15. Meller, Harald (2010). Elefantenreich : eine Fossilwelt in Europa; Begleitband zur Sonderausstellung im Landesmuseum für Vorgeschichte Halle 26.03.-03.10.2010 (in German). Halle (Saale: Landesamt für Denkmalpflege und Archäologie Sachsen-Anhalt, Landesmuseum für Vorgeschichte. pp. 314–321. ISBN 978-3-939414-48-3. OCLC 642989683.
16. 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.
17. 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. doi:10.1016/j.quascirev.2019.06.027. S2CID 200073388.
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19. 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. doi:10.1080/10420940.2020.1744586. ISSN 1042-0940. S2CID 216504699.
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30. Altermann, Manfred (1990). Neumark-Gröbern : Beiträge zur Jagd des mittelpaläolithischen Menschen (in German). Berlin: Dt. Verl. d. Wiss. ISBN 3-326-00571-7. OCLC 246658230.
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Further reading

• "Stone Age elephant remains found". BBC News. 2004. Retrieved 2 July 2006.
• Masseti, M. (1994). (PDF). Hystni. 5: 101–105. Archived from the original (PDF) on September 30, 2007.
• Wenban-Smith, F. F.; Bridgland, D. R. (1997). "Newly discovered Pleistocene deposits at Swanscombe: an interim report". Lithics. 17/18: 3–8.
• Wenban-Smith, F. F.; Bridgland, D. R. (2001). "Palaeolithic archaeology at the Swan Valley Community School, Swanscombe, Kent". Proceedings of the Prehistoric Society. 67: 219–259. doi:10.1017/S0079497X00001675. S2CID 128713311.
• Wenban-Smith, F. F.; Allen, P.; Bates, M. R.; et al. (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.

External links

•  Paleontology portal

• TRACCE Online Rack Art Bulletin paper about an elephant head Palaeolithic depiction in Portugal