British fossil elephants

Copyright 2009
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British fossil elephants
By Adrian M Lister
T
he
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
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.
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 widelydiverging 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,
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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
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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 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.
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 massive skull from P. antiquus in front
view. Stuttgart Museum. Photo A. Lister.
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The excavation of the elephant and mammoth skeletons at Aveley, Essex, 1964.
© Natural History Museum, London.
& 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. 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
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Mounting the Upnor Palaeoloxodon skeleton
at the Natural History Museum in the 1920s.
© 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).
The magnificent skull and tusks of a mammoth found at Ilford, Essex in
1863. © Natural History Museum, London.
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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.
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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.
Adrian Lister works in 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.
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
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. Lateglacial 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’, 4752.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.
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