British fossil elephants – Deposits

By Adrian M Lister

Fig. 1. From a sensible scale mannequin on the Pure Historical past Museum, London. Observe the sloping again and the double ‘finger and thumb’ on the finish of the trunk. (© Pure Historical past Museum, London.)

The elephant household (Elephantidae), like that of people, originated in Africa. Finds from the late Miocene of southern and japanese Africa present that, by between seven and 6 million years in the past, true elephants had arisen, most likely from superior mastodonts, that are associated to stegodons. Between these dates and about 4 million years in the past, the earliest representatives of the three nice shares of elephants – the African elephant (Loxodonta), Asian elephant (Elephas) and mammoth (Mammuthus) all make their look within the African fossil document. Loxodonta, in fact, stayed in Africa, whereas Elephas finally migrated north and east into its present vary in south-east Asia.

The primary true elephant fossils in Europe are of the Mammuthus lineage. In Britain, these first make their look within the Pink Crag of Suffolk, now dated to round 2.6 million years previous. The fossils should not frequent, however three well-preserved molars from Rendlesham will be seen in Ipswich Museum. This materials has just lately been attributed to the species Mammuthus rumanus, on the premise of the primitive look of the again molars with solely ten full enamel loops (Lister and van Essen, 2003).

Fig. 2. A molar from one of many earliest mammoths in Europe, *Mammuthus rumanus*, from the Pink Crag of Suffolk, Ipswich Museum. (Photograph by H van Essen.)

Within the succeeding Norwich Crag, the elephant fossils are recognized because the extra superior species Mammuthus meridionalis. This title, actually which means ‘southern mammoth’, was based mostly on sort materials from Italy, therefore ‘southern’ in a European context, though it was one of many northernmost elephants on the earth on the time! I want the casual time period ‘ancestral mammoth’ as a result of this species unfold broadly in Europe and Asia, endured for at the least one million and a half years (about 2.2 to 0.7 million years in the past), and was most likely on the root of all later mammoth species (Lister and Bahn, 2015).

Fig. 3. The pelvic bone of a foetal ancestral mammoth, *Mammuthus meridionalis*, recognized by Dick Mol. It was discovered within the Wroxham Crag Formation (formally Weybourne Crag), simply above the stone bed at Weybourne, Norfolk. (Photo by A.Cruickshanks. Length, 10cm.)

Mammuthus meridionalis, with its massive, thick-enamelled teeth (typically with 12 to 14 enamel loops in the back molars) is particularly common in the lower part of the Cromer Forest Bed formation. In the late nineteenth to early twentieth century heyday of collecting on the Norfolk coast, the deposit at the base of the cliff at Bacton gained the name ‘elephant bed’ because of the abundance of elephantine fossils found there.

Fig. 4. A rare tongue bone (stylohyoid) from *Mammuthus meridionalis* found from the Norwich Crag Formation at Easton Bavents, Suffolk. (Photo by A Cruickshanks. Length, 14cm.)

However, it was fishermen from the oyster beds, three-quarters of a mile out to sea off Happisburgh, Norfolk who dredged the largest numbers of remains. An account given by Woodward (1833) is both amusing and exasperating from the point of view of the modern collector. He describes how, in the 1820s, ‘many hundred specimens of the molar teeth of the elephant were destroyed by the fishermen, who amused themselves by breaking them, their wonder being excited by the grinders separating into laminae’!

Mammuthus meridionalis stood up to four metres high, had robust, spirally twisted but relatively short tusks, and was a browser in temperate woodlands. Its coat was presumably sparse, like that of living elephants. Its successor, the so-called ‘steppe mammoth’, Mammuthus trogontherii, was equally large but shows a marked change in feeding adaptation, with higher-crowned molars comprising 20 or so enamel loops. It had plainly gone over to a diet consisting of a larger proportion of grass, corresponding to the expansion of grassland habitats as the ice ages progressed.

The spectacular discovery of the West Runton mammoth, one of the most important British fossil finds of the twentieth century, has provided us with much new information about this species. It is fortuitous that the almost complete skeleton lay within the celebrated type deposits of the Cromerian interglacial (about 700,000 years ago), and its discovery has been the catalyst for a multidisciplinary re-study of the site (Stuart and Lister, in prep.).

The massive male skeleton had been preserved lying on its right side and had been scavenged by hyenas prior to burial, as shown by chewed mammoth foot bones and perfectly preserved hyena coprolites in place around the skeleton. The most remarkable feature of the skeleton was a severely distorted knee joint, clear evidence that the animal had suffered a major accident during its life that had left one hind leg permanently dislocated, possibly accounting for the animal’s inability to extricate itself from the river deposits at West Runton where it died.

At around the same time the steppe mammoth appeared in Europe, another entirely new kind of elephant, the so-called straight-tusked elephant Palaeoloxodon antiquus, also made its appearance. It too was a massive beast, four metres or more at the shoulder and weighing in at around ten tonnes, and distinguished from the mammoths by its double-domed skull and tusk sockets widely-diverging in front view. Its long tusks, while not twisted like those of the mammoths, were not really straight but gently curved.

While the steppe mammoth occupied more of a grazing niche, the straight-tusker was more of a browser. It used to be thought that the ancestral mammoth, Mammuthus meridionalis, had split into these two lineages in Europe. However, newer evidence shows this to be incorrect (Lister et al., 2005). M. trogontherii remains as old as 1.7 million years have been found in China, suggesting that the species arose there (presumably from an eastern population of M. trogontherii), so its appearance in Europe a million or so years later represents a migrational rather than an evolutionary event.

Meanwhile, Palaeoloxodon antiquus is of a completely different lineage and its roots can be traced back to P. recki of the African Plio-Pleistocene, from where it appears to have migrated into Europe around 0.75 million years ago. It first appears in Britain in the early Middle Pleistocene deposits of the Cromer Forest Bed formation, including a couple of molars from the Pakefield deposits that recently provided evidence of early human occupation in Britain (Parfitt et al., 2005).

Fig. 5. The massive skull from P. antiquus in front view. Stuttgart Museum. (Photo A Lister.)

From the late Middle Pleistocene, remains of both lineages are abundant in Britain. A massive mammoth skull from the brickworks at Ilford, Essex, was collected by Sir Antonio Brady as early as 1863, in time for it to be described in detail by Leith Adams in his monograph British Fossil Elephants (from which the title of the present article is respectfully taken). The skull (though quite heavily restored with plaster) is still on display in the main entrance hall of the Natural History Museum in London.

Fig. 6. The magnificent skull and tusks of a mammoth found at Ilford, Essex in 1863. (© Natural History Museum, London.)

More recently, hundreds of mammoth molars and tusks have been recovered from Thames deposits, of similar age to those at Ilford, at Stanton Harcourt in Oxfordshire (Buckingham et al., 1996). This interval (marine isotope stage 7, around 200,000 years ago) is the last time that mammoths are found in a temperate context in Britain. By the time of the penultimate cold stage (MIS 6, around 160,000 years ago), advanced woolly mammoths (Mammuthus primigenius) had taken their place, probably having emigrated from Siberia (Lister et al., 2005).

Important finds of straight-tusked elephant in Britain include the famous ‘Upnor elephant’, a headless but otherwise largely complete skeleton found in clay deposits of the River Medway in 1913 (Andrews and Cooper, 1928), formerly mounted at the Natural History Museum. A remarkable discovery was made in 1964 in deposits at Aveley, not far from Ilford and in the same Thames terrace (Blezard, 1966). The skeleton of a straight-tusked elephant was found directly overlain by that of a mammoth, in deposits with pollen indicating a transition from a more wooded to a more open environment.

Fig. 7. The excavation of the elephant and mammoth skeletons at Aveley, Essex, 1964. (© Natural History Museum, London.)

Partial skeletons of Palaeoloxodon have also been found in interglacial deposits under the beach at Selsey, West Sussex, probably of similar age to those at Ilford and Aveley. These have also been found at Deeping St James near Peterborough, in deposits of the subsequent, last interglacial (MIS 5e, about 120,000 years ago). The species soon after disappears from the British record, although it hung on in southern Europe until close to the start of the last glacial maximum (around 25,000 years ago) and, as dwarf forms, on Mediterranean islands, even later.

Fig. 8. Mounting the Upnor Palaeoloxodon skeleton at the Natural History Museum in the 1920s. (© Natural History Museum, London.)

Woolly mammoth fossils are abundant in many cave and river deposits of the last cold stage (broadly MIS 4-2) in Britain, although the assemblages vary greatly in their composition. While, in river deposits, the remains of adults are commonest (probably due to preservation and collecting bias), the remains are overwhelmingly those of juveniles in caves such as Kent’s Cavern (in Devon) and Pin Hole (in Derbyshire). The explanation probably lies in these having been accumulated by spotted hyenas (Crocuta), whose ability to lug joints of mammoth meat into the cave would have been limited to that from the youngest animals (Lister, 2001).

The woolly mammoth vacated Britain (and indeed much of western and central Europe) for a few thousand years during the last glacial maximum (LGM, around 25,000 to 20,000 years ago) (Stuart et al 2004). It was previously thought not to have subsequently returned to Britain, until the discovery of the Condover mammoths in 1986 demonstrated their presence between 15,000 to 14,000 years ago. These were found in kettle-hole deposits demonstrably overlying the till of the LGM. The Condover discovery at a working gravel pit a few miles south of Shrewsbury was, like that at West Runton, due to the chance spotting of a few exposed bones, and like it, led to the excavation of an almost complete adult male skeleton, this time of woolly mammoth M. primigenius. The remains of three juvenile mammoths were also found in the same horizon (Coope and Lister, 1987).

Fig. 9. The first discovery of the Condover mammoth at a gravel pit near Shrewsbury in 1986. These limb bones proved to be part of an almost completely preserved skeleton of a wooly mammoth.

The Condover discoveries, plus a few worked ivory fragments from the late Upper Palaeolithic horizons of Kent’s Cavern and Pin Hole, mark the last appearance of wild elephants in Britain. Mammoths persisted in northern Siberia into the early part of the Holocene epoch, until their final extinction several thousand years later on arctic islands such as Wrangel Island and the Pribilof Group (Lister and Bahn, 2015). Advances in molecular biology have allowed substantial portions of the mammoth genome to be recovered from permafrost-preserved remains, and comparison of DNA sequences has shown that the mammoth was more closely related to the Asian than the African elephant (Krause et al. 2006). The phylogenetic position of Palaeoloxodon is still uncertain, although it was clearly closer to the Mammuthus/Elephas than to Loxodonta.

Predictions of the re-creation of a living mammoth, by cloning or other techniques are, in the opinion of the author, unlikely to be realised in the foreseeable future because of the fragmentary state of preservation of all ancient DNA recovered to date.

Fig. 10. The author with the partial skull of the West Runton Mammoth, Mammuthus trogontherii. The skull shows the right tusk and the massive last molar behind. (Photo A J Stuart.)

Professor Adrian Lister works at the Department of Palaeontology at the Natural History Museum in London.

References

Adams, A. L. 1877-81. Monograph on the British Fossil Elephants. London: Palaeontographical Society.

Andrews, C.W. & Cooper, C.F. 1928. On a specimen of Elephas antiquus from Upnor, with further notes on the teeth and skeleton. London: Clowes.

Blezard, R.G. 1966. Field meeting at Aveley and West Thurrock. Proc. Geol. Ass. 77, 273-6.

Buckingham, C.M., Roe, D.A. & Scott, K. 1996. A preliminary report on the Stanton Harcourt Channel deposits (Oxfordshire, England): geological context, vertebrate remains and Palaeolithic stone artefacts. J. Quat. Sci. 11, 397-415.

Coope, G.R. & Lister, A.M. 1987. Late-glacial mammoth skeletons from Condover, Shropshire, Enland. Nature 330, 472-4.

Krause, J., Dear, P.H., Pollack, J.L., Slatkin, M., Spriggs, H., Barnes, I., Lister, A.M., Ebersberger, I., Pääbo, S., & Hofreiter, M. 2006. Multiplex amplification of the mammoth mitochondrial genome and the evolution of Elephantidae. Nature 439, 724-727.

Lister, A.M., 2001. Age profile of mammoths in a Late Pleistocene hyaena den at Kent’s Cavern, Devon, England. Anthrop. Pap. Univ. Kansas. 22, 35-43.

Lister, A. & Bahn, P. (2015). Mammoths: Giants of the Ice Age. Third revised edition. London: Frances Lincoln.

Lister A.M., Sher, A.V., van Essen, H. & Guangbiao Wei, 2005. The pattern and process of mammoth evolution in Eurasia. Quaternary International 126-128, 49-64.

Lister, A.M. & van Essen, H. 2003. Mammuthus rumanus (Stefanescu), the earliest mammoth in Europe. In (Petulescu, a. & Stiuca, E., eds) Advances in Palaeontology ‘Hen to Panta’, 47-52.Romanian Academy, ‘Emil Racovita’ Inst. of Speleology, Bucharest.

Parfitt, S.A., Barendregt, R.W., Breda, M., Candy, I., Collins, M.J., Coope, G.R., Durbidge, P., Field, M.H., Lee, J.R. Lister, A.M., Mutch, R., Penkman, K.E.H., Preece, R.C., Rose, J., Stringer, C.B., Symmons, R., Whittaker, J.E., Wymer, J.J. & Stuart, A.J. 2005. The earliest record of human activity in northern Europe Nature 438, 1008-1012.

StuartAJ, Kosintsev PA, HighamTFG & Lister AM., 2004. Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth. Nature 431, 684-9.

Stuart. A.J. & Lister, A.M. (eds), 2011. The West Runton Mammoth and its Cromerian Environment. Quaternary International.

Woodward, S. 1833. An Outline of the Geology of Norfolk. London: Longman.

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