EIGHT

British fossil elephantsBy Adrian M Lister

A realistic scale model of a woolly mammoth © Natural History Museum, London. Note the sloping back and the double ‘finger and thumb’ at the end of the trunk.

The elephant family (Elephantidae), like that of humans, originated in Africa.

Finds from the late Miocene of southern and eastern Africa show that, by between seven and six million years ago, true elephants had arisen, probably from stegodons, which are distant relatives of mastodonts. Between those dates and about four million years ago, the earliest representatives of the three great stocks of elephants – the African elephant (Loxodonta), Asian elephant (Elephas) and mammoth (Mammuthus) all make their appearance in the African fossil record. Loxodonta, of course, stayed in Africa, while Elephas eventually migrated north and east into its current range in south-east Asia. The first true elephant fossils in Europe are of the Mammuthus lineage. In Britain, these first make their appearance in the Red Crag of Suffolk, now dated to around 2.6

million years old. The fossils are not common, but three well-preserved molars from Rendlesham can be seen in Ipswich Museum. This material has recently been attributed to the species Mammuthus rumanus, on the basis of the primitive appearance of the back molars with only ten complete enamel loops (Lister & van Essen, 2003).In the succeeding Norwich and Wroxham Crags, the elephant fossils are identified as the more advanced species Mammuthus meridionalis. This name, literally meaning ‘southern mammoth’, was based on type material from Italy, hence ‘southern’ in a European context, although it was one of the northernmost elephants in the world at the time! I prefer the informal term ‘ancestral mammoth’ because this species spread widely in Europe and Asia, endured for at least a million and a half years (ca. 2.2 to 0.7mya), and was probably at the root of

all later mammoth species (Lister & Bahn, 2007).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 19th to early 20th 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. But the largest number of remains were dredged by fishermen from the oyster beds three-quarters of a mile out to sea off Happisburgh, Norfolk. 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

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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 of 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 1995 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 & Lister, in prep.). The massive male skeleton had been preserved lying on its right side and had been scavenged by hyaenas prior to burial, as shown by chewed mammoth foot bones and perfectly preserved hyaena 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 kind of elephant entirely, 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, more recent 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. meridionalis), 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 0.75mya. It first appears in the British fossil record 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). From the late Middle Pleistocene, remains of both lineages are abundant in Britain. A massive mammoth skull from the brickearths 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. More recently, hundreds of mammoth molars and tusks have been recovered from Thames deposits, of a 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 migrated 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

A molar from one of the earliest mammoths in Europe, Mammuthus rumanus. From the Red Crag of Suffolk, Ipswich Museum. Photo H. van Essen.

The massive skull from P. antiquus in front view. Stuttgart Museum. Photo A. Lister.

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

The pelvic bone of a foetal ancestral mammoth, Mammuthus meridionalis, identified by Dick Mol. It was found in situ in the Wroxham Crag Formation (formally Weybourne Crag), just above the stone bed at Weybourne, Norfolk. Photo by A. Cruickshanks. Length, 10cm.

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

accumulated by spotted hyaenas (Crocuta), whose ability to lug joints of mammoth meat into the cave would have been limited to those 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 remains 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

& Lister 1987).

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

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

a transition from a more wooded to a more open environment. A partial skeleton of Palaeoloxodon has also been found in interglacial deposits under the beach at Selsey, West Sussex, probably of a similar age to those at Ilford and Aveley. Another was found at Deeping St James near Peterborough, in deposits of the subsequent, last interglacial (MIS 5e, ca. 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. 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

& 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

m a m m o t h , in deposits with pollen

indicating

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

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, England. 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. 2007. 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.Stuart AJ, Kosintsev PA, Higham TFG & 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) in prep. The West Runton Mammoth and its Cromerian Environment. Quaternary International. Woodward, S. 1833. An Outline of the Geology of Norfolk. London: Longman.

ELEVEN

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 & Bahn, 2007). 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 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.

References

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

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.

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www.ukge.co.uk Taken from Issue 13 www.ukfossils.co.uk

Copyright 2009 Deposits Magazine www.depositsmag.biz

www.ukge.co.uk Taken from Issue 13 www.ukfossils.co.uk