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Straight-tusked elephant

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Straight-tusked elephant
Temporal range: Mid-Late Pleistocene
~0.78–0.03 Ma
↓
Skull and model (from Ambrona)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Genus: †Palaeoloxodon
Species:
†P. antiquus
Binomial name
†Palaeoloxodon antiquus
(Falconer & Cautley, 1847)
Approximate range of P. antiquus
Synonyms

Elephas antiquus (Falconer & Cautley, 1847)

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 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.

Description

Complete skeleton in Rome.

Palaeoloxodon antiquus was quite large, with individuals reaching 4 metres (13.1 ft) in height. Like other members of Paleoloxodon, P. antiquus possesses a well developed parieto-occipital crest (POC) at the top of the cranium, for anchoring 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 in Europe were previously suggested to exist in Europe based on POC variation, one more similar to P. namadicus, but these were shown to be the result of ontogenetic variation and taphonomic distortion.[1] One approximately 40-year-old male 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.[2] and had long, slightly upward-curving tusks.[3] P. antiquus's legs were slightly longer than those of modern elephants. This elephant is thought to have had an 80-cm-long tongue that could be projected a short distance from the mouth to grasp leaves and grasses.[4] 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 subsequently assigned E. antiquus to the genus.[7] Some experts regarded the larger Asian species Palaeoloxodon namadicus as a variant or subspecies[8] but they are currently considered distinct.[1] 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 DNA sequence analysis of P. antiquus suggested that its closest extant relative may be 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) 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

Palaoloxodon antiquus is believed to derive from the African P. recki. Palaeoloxodon antiquus first appears during the early Middle Pleistocene, around 0.8-0.6 Ma, with an early occurrence of around 780 kya in Italy, its earliest appearance in northern Europe is in Suffolk around 600 kya. Its arrival is co-incides 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 ' 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 five to 15 individuals.[5] Like its recent relatives, the straight-tusked elephant would have been heavily dependent on fresh water, which greatly influenced its migration.[14] Its dietary habits changed depending on food supply across the seasons, browsing in spring and summer and grazing in autumn and winter. Maple, hornbeam, hazelnut, alder, ash, beech and ivy were consumed, based on found gastric contents and food residues adhering to tooth remains. Microscopic examination of signs of wear on the surfaces of back teeth also show the use of different food resources.

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 periods, while permantently residing in Mediterranean Europe during glacial periods. It is assumed that they preferred wooded environments. The straight-tusked elephant became extinct in Britain near the beginning of the Weichselian glaciation, about 115,000 years ago.

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.[15]

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,[16][17][18][19] and a Palaeoloxodon antiquus butchering site has been excavated near Megalopolis, in the Peloponnese.[20][21]

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.[22] The Iberian peninsula may have served as the last European refuge of the straight-tusked elephant. According to João Luís Cardoso,[23] the species survived until 30,000 years BP in Portugal.

Germany

Finds located particularly far to the north are documented not only from individual remains from Mecklenburg-Western Pomerania and Lower Saxony.[24] Fossils of at least ten P. antiquus have been found in the Schöningen opencast mine since the excavations began in the 1980s. These were individual finds of ribs, tusks or vertebrae that had to be hurriedly recovered from the approaching bucket wheel excavator during rescue excavations.[25] The P. antiquus discovered in 2017 is the only almost completely preserved skeleton. Since the site is in the open-cast mining area on the preserved spear base, the recovery took place without time pressure in close consultation with the conservation workshop of the Lower Saxony State Office for Monument Preservation.[26] Further analyzes of the environmental and climatic conditions at the time of death of the animal, as well as the sedimentation processes of the former lake, are carried out by the Technical University of Braunschweig, the University of Lüneburg and the University of Leiden in the Netherlands. These sediment samples were analyzed for micro-fauna, micro-morphology, Limnology and Paleobotany.[27] The 300.000 year-old nearly entire remains of a female straight-tusked elephant were revealed by University of Tübingen researchers and the Senckenberg Centre for Human Evolution in May in 2020 in Schöningen in Germany. According to the archaeozoologist Ivo Verheijen, 6.8 tones older skeleton with battered teeth had a shoulder height of about 3.2 metres. Researchers also uncovered two long bones and 30 small flint flakes were used as tools for knapping among the elephant bones. “We found both 2.3-m-long tusks, the complete lower jaw, numerous vertebrae and ribs as well as large bones belonging to three of the legs and even all five delicate hyoid bones,” said an archaeologist Dr. Jordi Serangeli.[28][29][30] The P. antiquus is probably a female with a shoulder height of approximately 3.2 m (10 ft) and weight around 6.8 tons. During the excavation, 300 bones and a total of 700 bone parts were recovered, spread over an area of around 64 m2 (690 sq ft). The finds include the 2.3 meter long tusks, the lower jaw, vertebrae and ribs, as well as the bones of three legs and the entire hyoid bone.[31] The skeletal parts were largely in an anatomically correct arrangement. Various parts were missing, such as the pelvis, the left front leg with the shoulder blade and the foot bones of two legs. The preservation of the bones is mostly very good, only the more than one meter high skull of the individual had disintegrated into hundreds of small parts. This was caused by the comparatively light construction of the skull, which, like all elephants, consists of numerous air-filled and honeycomb-like hollow chambers. Overall, the animal lay parallel to the former lake shore with the head in the north and the rear in the south. The age of the animal is estimated to be around 50 years due to the worn teeth and osteoarthritis of the bones.[32]

Bite marks on the bones indicate that predators fed on the P. antiquus after death. Although no traces of human processing were found on the bones, there are indications of the presence of humans on the carcass.[33] So were between the bone about 30 haircuts of flints that the sharpening stone tools emerge. In addition, two bone artifacts were found that were used as tools by humans and had impact marks. These are the foot bones of a red deer and another approximately 12 cm (4.7 in) long piece of bone from an unidentified animal.[34]

Replica of a P. antiquus on the edge of the Elm forest at Schöningen

After the discovery of the P. antiquus skeleton at Schöningen in September 2017, the exposure took almost two years. It was carried out by archaeologists from the Senckenberg Center for Human Evolution and Paleoenvironment at the University of Tübingen in cooperation with the Lower Saxony State Office for Monument Preservation. The scientists announced the discovery publicly in May 2020 at a press conference with the Lower Saxony Minister of Science Björn Thümler in Schöningen.[35] At that point in time, the examination and conservation of the bones had not yet been completed. That took place in the Schöningen Research Museum, which was directly connected to the excavations and is about 300 meters away from the site.[36]

Due to the findings of elephant bones in the open pit, the life-size replica of a P. antiquus was erected above Schöningen in 2018. It stands on the edge of the Elm forest next to an excursion restaurant and is also used for tourist purposes.[37][38] The reconstruction was made on the basis of a skeleton found at the Neumark-Nord site in Geisel valley, Saxony-Anhalt.[39]

For the year 2022, the President of the Lower Saxony State Office for the Preservation of Monuments, Christina Krafczyk, announced an exhibition with a focus on P. antiquus at the Schöningen Research Museum.[40]

Complete skeletons of P. antiquus are relatively rare, especially in Central Europe, the animals usually appear at the various sites in the form of remains of teeth and teeth or individual bones. One of the best-known finds and the next youngest evidence of P. antiquus in Lower Saxony is also offered by the lance of Lehringen, near Verden. The animal perished in a hunting event in the last warm period (Eem warm period) about 120,000 years ago. During this time of the Middle Paleolithic, the Neanderthals lived in Europe. In addition to the lance, which was stuck under or between the ribs of the animal, there were also 27 sharp-edged flint fragments in the vicinity, which were apparently produced on site at short notice. Since the animal had collapsed in the water, the people of that time could neither dismantle the entire carcass, nor recover the lance mentioned above.[41][42]

From a supra-regional perspective, the old find of P. antiquus from Gröbern in Saxony-Anhalt, which is comparable to Lehringen, is significant. There, too, the bank of a former lake hid a complete skeleton, which reconstructs a fully grown male with a shoulder height of around 4.2 meters represents. The almost 200 bones of the elephant were distributed over an area of 20 square meters and were largely still in the anatomical association. There were only major remnants on the skull, chest and hind extremities. In direct connection, a good two dozen flint fragments were found in Gröbern, mostly between the bones. Similar to Lehringen, it is assumed that the people of that time were only able to recover parts of the carcass[43] A second skeleton of P. antiquus found in Gröbern in the same year showed no anthropogenic influence.[44]

Significant finds also came to light in the Geisel valley, Saxony-Anhalt. There, on the shores of a former lake, the Neumark-Nord 1 lake basin, skeletons of around 70 individuals of P. antiquus were found, some of them largely complete; they spread around the lake. The age of the lake basin is discussed and varies, depending on the view, from the penultimate warm period around 200,000 years ago to the last warm period. The numerous skeletal finds and good conservation conditions made it possible to pass on rarely proven bones such as the tongue and sternum.[45] Individual animals showed pathological symptoms. Changes, for example in the form of hip dysplasia, stunted tusks or perforations, which could possibly be traced back to rival fights.[46][47][48] The animals died a natural death, but were later tattered by large predators such as the also documented cave lion or the cave hyena.[49] Individual skeletons were connected with large flint debris, which indicate activities of early humans. One of these artifacts, known as the “carving knife”, still had an organic residue that turned out to be oak bark extract.[44][50]

Several partial skeletons of P. antiquus from the travertines of Bad Cannstatt in Baden-Württemberg, which probably belong to the penultimate warm period, have been documented from southern Germany. Among them is a fully grown bull, the skeleton of which was spread over an area of 25 square meters and which, due to the 122 centimeter long upper arm and 144 centimeter long thigh bone, had a shoulder height of around 4 meters. For most elephant remains, exposure to early humans can be ruled out. Rather, the animals died due to illness or of old age in the vicinity of the spring water.[51]

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.[52] 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.[53] A second elephant skeleton found in Gröbern the same year proved to be free of any markings of human use.[54]

Remains have also been recovered from another Eemian site, the former lake basin Neumark-Nord 1 on the northeast edge of the Geisel valley.[55] 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.[56] 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.[57]

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.[58]

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.[59] 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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Further reading

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