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eschweizerbartxxx sng-
© E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), 2007, ISSN 0341–4116
Cour. Forsch.-Inst. Senckenberg | 259 | 287 – 297 | 2 Figs | Frankfurt a. M., 13. 12. 2007
Patterns of Late Quaternary megafaunal extinctions in Europe and northern Asia
With 2 gs
Anthony J. Stuart & Adrian M. LiSter
Abstract
This paper summarizes the results so far of our ‘Late Quaternary Megafaunal Extinctions’ project, focussing
on an assessment of latest available dates for selected target species from Europe and northern Asia. Our
approach is to directly radiocarbon-date material of extinct megafauna to construct their spatio-temporal
histories, and to seek correlations with the environmental and archaeological records with the aim of estab-
lishing the cause or causes of extinction. So far we have focussed on Mammuthus primigenius, Coelodonta
antiquitatis, and Megaloceros giganteus, and are accumulating data on Panthera leo/spelaea, Crocuta
crocuta and Ursus spelaeus. Attempts to date Palaeoloxodon antiquus and Stephanorhinus hemitoechus
(from southern Europe) were largely unsuccessful.
The pattern of inferred terminal dates is staggered, with extinctions occurring over ca. 30 millennia,
with some species previously thought extinct in the Late Pleistocene – M. primigenius and M. giganteus
– surviving well into the Holocene. All species show dramatic range shifts in response to climatic/vegeta-
tional changes, especially the beginning of the Last Glacial Maximum, Late Glacial Interstadial, Allerød,
Younger Dryas and Holocene, and there was a general trend of progressive range reduction and fragmenta-
tion prior to nal extinction. With the possible exceptions of P. antiquus, S. hemitoechus and Homo nean-
derthalensis, extinctions do not correlate with the appearance of modern humans. However, although most
of the observed patterns can be attributed to environmental changes, some features – especially failures to
recolonize – suggest human involvement.
Key words: Late Quaternary, megafauna, extinctions, Northern Eurasia, radiocarbon dating
Authors’ addresses: A. J. Stuart (corresponding author) & A. M. LiSter, Department of Palaeontology, Natural History Museum, Cromwell
Road, London SW7 5BD, UK and A. J. Stuart, University of Durham, School of Biological and Biomedical Sciences, South Road, Durham,
DH1 3LE, UK, <Tony@megafauna.freeserve.co.uk>
Authors’ addresses: A. J. Stuart (corresponding author) & A. M. LiSter, Department of Palaeontology, Natural History Museum, Cromwell
Road, London SW7 5BD, UK and A. J. Stuart, University of Durham, School of Biological and Biomedical Sciences, South Road, Durham,
DH1 3LE, UK, <Tony@megafauna.freeserve.co.uk>
Introduction
We live today in a zoologically impoverished world from
which many of the largest and most spectacular large
vertebrates (megafauna) have disappeared in the recent
geological past. Formerly most of these extinctions were
thought to have occurred before the end of the Pleis-
tocene ca. 10 ka (10,000 radiocarbon years BP, ca. 11,600
calendar years), but it is now clear that some megafaunal
species, such as woolly mammoth and giant deer, became
extinct in the Holocene (Vartanyan et al. 1993, Guthrie
2004, Stuart et al. 2004).
As data continue to accumulate, the patterns and
processes of Late Quaternary extinctions appear increas-
ingly complex (Stuart et al. 2004, BarnoSky et al. 2004,
Stuart 2005), while the crucial question of cause or
causes remains unresolved. Since Paul S. Martin (1967,
1984, Martin & SteadMan 1999) rst championed the
‘overkill’ hypothesis, in which extinctions were attributed
to the impact of human hunters, the subject has gener-
ated strong views as to the nature and extent of human
involvement in these extinctions (Martin & kLein 1984,
MacPhee 1999, GraySon et al. 2001, BarnoSky et al.
2004). If true, ‘overkill’ – in which extinctions are attrib-
uted to predation by Stone Age modern humans Homo
sapiens – can be seen as the beginning of a new episode
of ‘mass extinction’ which continues at an accelerated
pace today – the so-called ‘Sixth Extinction’ (MacPhee
1999, Benton 2003). The alternative hypothesis of en-
vironmental change necessitates that a unique event or
series of climatic events occurred in the Late Pleistocene
(Sher 1997, Guthrie 1984), as no comparable wave of
extinctions had taken place in the previous 0.8 Ma de-
spite a series of major climatic uctuations (Stuart 1991,
BarnoSky et al. 2004). Some, notably Sher (1997), have
argued that the last glacial-interglacial cycle was indeed
signicantly different, because of unprecedented changes
in circulation of the Arctic Ocean, and that the resulting
eschweizerbartxxx sng-
Stuart & LiSter: Late Quaternary megafaunal extinctions
288
replacement of arctic steppe-tundra by modern boggy
tundra and coniferous forest in the early Holocene caused
the extinction of mainland woolly mammoths.
A number of authors (Stuart 1991, 1999, BarnoSky
et al. 2004, hayneS & eiSeLt 1999) have suggested a
combination of ‘overkill’ and environmental change, in
which extinctions resulted from human hunting of mega-
faunal populations subject to habitat fragmentation and
the stress of climatic/vegetational changes.
A further hypothesis, termed ‘hyperdisease’, involv-
ing the spread of diseases to megafaunal species by
immigrating humans (MacPhee & Marx 1997) appears
unlikely to have produced the observed timing of extinc-
tions or the body size spectra of the species affected (aL-
roy 1999, Stuart 1999, LyonS et al. 2004).
There is strong evidence that the pattern and timing
of Late Pleistocene and Holocene extinctions was very
different in each zoogeographical region (Stuart 1991,
roBertS et al. 2001, BarnoSky et al. 2004, FLannery
2005) so that, while also maintaining a global perspec-
tive, each region needs to be studied on its own merits.
Because many Late Quaternary Extinctions fall within
the range of radiocarbon (14C) dating there is enormous
potential for resolving the chronology of these extinc-
tions in detail, in marked contrast to the problems of
stratigraphic resolution for older extinction episodes
(haLLaM & WiGnaLL 1997, Benton 2003). Establish-
ing a reliable chronology for the extinct megafauna of
each zoogeographical region is essential for testing these
hypotheses. Radiocarbon dating is an especially useful
tool for Late Quaternary extinctions that occurred within
the reliable chronological range of this method, mainly
post-dating ca 35–30,000 14C years BP, in some cases ex-
tending back to ca. 40–50,000 14C years BP. In this paper
dates are given as uncalibrated radiocarbon dates (ka =
thousands of radiocarbon years BP), or as calendar dates,
including calibrated 14C dates, (cal. ka = thousands of cal-
endar years BP). Many of the radiocarbon determinations
collated for this paper are beyond the current limit of the
INTCAL04 calibration dataset (reiMer et al. 2004), so
here radiocarbon dates have been tentatively compared
using the data published by FairBankS et al. (2005). This
dataset comprises 230Th/234U/238U dates and radiocarbon
AMS dates of coral from Barbadian, central and western
Pacic locations, which extend back to ca. 55 ka BP.
Northern Eurasia is a particularly fruitful region for
the study of Late Quaternary extinctions, not only be-
cause of the wealth of available archaeological, palaeon-
tological and environmental data, but also because in this
region most extinctions occurred well within the range of
radiocarbon dating. Building on the large number of ra-
diocarbon dates already available, it is possible to analyse
the record to much higher standards and resolution than
elsewhere. In contrast to the situation in North America
where the arrival of Clovis technology, major environ-
mental change and megafaunal extinction are thought to
have been closely coincident and therefore difcult to
unravel (Martin & SteadMan 1999), it has been apparent
for some time that in northern Eurasia Late Quaternary
extinctions were staggered over a long period (Stuart
1991), and recent discoveries have served to strengthen
this observation. This situation potentially allows much
better discrimination of possible causes.
This paper draws on the results of our rst research
programme (1999–2002) ‘Late Quaternary Megafaunal
Extinctions Project’ (LQME project), funded by the
UK Natural Environment Research Council (NERC),
together with some preliminary ndings from our second
programme (2006–9). The project used both new dates
from samples we submitted to the Oxford Radiocarbon
Accelerator Unit (ORAU), and dates available in the
literature. The principal species investigated are: Mam-
muthus primigenius – woolly mammoth, Megaloceros
giganteus – giant deer (‘Irish elk’), and Coelodonta an-
tiquitatis – woolly rhinoceros, Crocuta crocuta – spotted
hyaena, Panthera spelaea/leo – lion, and Ursus spelaeus
– cave bear. Attempts were also made to date apparently
late specimens of Palaeoloxodon antiquus – straight-
tusked elephant, and Stephanorhinus hemitoechus – nar-
row-nosed rhino. Radiocarbon dating of these species
met with very limited success, but we have collated
and critically assessed available data relevant to their
extinction chronology. Hippopotamus amphibius and
Crocuta crocuta, although still extant, are considered
part of the Late Quaternary extinctions phenomenon
from a Eurasian point of view, and the demise of Homo
neanderthalensis (Neanderthal man) can also be regarded
as a part of the Late Quaternary extinctions phenomenon
(SteWart 2005); see discussion. Other taxa, including
Equus hydruntinus, which appears to have survived in
the Balkans until ca. 6 cal. ka (SPaSSoV & iLieV 2002) and
Homotherium latidens, recently claimed to have survived
in Europe until ca. 28 ka (ca. 32 cal. ka) (reuMer et al.
2003), are not included in the present study.
Our approach is to obtain radiocarbon dates directly
on megafaunal material, thereby minimising problems
of stratigraphic control and context. All too often mega-
faunal material, when dated, proved to be older than its
apparent context, probably due to such factors as poor ex-
cavation, post-depositional mixing of layers, or the incor-
poration of older material, e.g. mammoth ivory, collected
by humans (Stuart 2005). Some AMS (accelerator mass
spectrometry) radiocarbon dates produced in the earlier
days of the method are now known to be signicantly
in error. A recent study (JacoBi et al. 2006, hiGhaM et al.
2006), re-dating mammalian fossils using modern tech-
niques (notably ultraltration), found that many previous
dates were signicantly too young, presumably because
of inadequate removal of contaminants (see below: Cro-
cuta crocuta, spotted hyaena).
In our research work, whenever possible the validity
of any outstanding or unexpected dates was corroborated
through independent dating by another 14C laboratory.
Samples of tooth, bone or antler were obtained from Eu-
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Cour. Forsch.-Inst. Senckenberg, 259, 2007
289
rope and Siberia, both by visiting relevant museums and
also where appropriate by ‘mail order’ from colleagues,
subject to careful checks on the taxonomic identication
of the material. Care was taken to ensure correct iden-
tication of the dated material by means of diagnostic
characters, e.g. with Megaloceros giganteus (commonly
confused with bovid or red deer material) and Ursus spe-
laeus (confused with brown bear U. arctos). In the case
of ‘mail order’ samples, colleagues were asked to provide
scaled photographs of the sampled material as a further
check on correct identication.
In this paper we focus on the pattern and interpretation
of the terminal dates for megafaunal species. Full datasets
and discussion of the complex dynamic changes in distri-
butions preceding extinctions will be given elsewhere.
Results
Extinctions before ca. 35 cal. ka
Hippopotamus amphibius, hippopotamus
During the Last Interglacial [Marine Isotope Stage (MIS)
5e] the range of the hippopotamus included Africa and
Mediterranean Europe, extending as far north as Britain,
but not further east than the Rhine Valley (Stuart 1991).
However, its suggested survival in Southern Europe
signicantly post-dating MIS 5e (Stuart 1991) cannot
be substantiated because of stratigraphical uncertainties
and/or misidentications. For example we re-determined
a radio-ulna (in the IPH, Paris), labelled as ‘hippopota-
mus’ from the ‘Mousterian B’ level at Cueva del Castillo,
Northern Spain as a large bovid. No samples of hip-
popotamus were submitted for dating, as none appeared
likely to lie within the range of the radiocarbon method.
The Pleistocene distribution of hippopotamus and its
association at many sites with southern thermophiles
such as Trapa natans (water chestnut) (Stuart 1991)
indicates that it was very sensitive to low temperatures.
Its disappearance from Europe might therefore relate to
the climatic deterioration within MIS 5e ca 122–115 cal.
ka (North Greenland Ice Core Project Members 2004), or
perhaps later in MIS 5.
Palaeoloxodon antiquus, straight-tusked elephant
In marked contrast to woolly mammoth, which had its
maximum range in the cold stages, straight-tusked el-
ephant was widely distributed over most of Western Eu-
rope in interglacials, including the Eemian ca. 130–117
cal. ka (Last Interglacial, Marine Isotope Stage 5e) in
association with regional temperate and Mediterranean
forests (Stuart 1991, 2005). With the onset of the Last
Cold Stage (equivalent to MIS 5d–2) and the spread of
treeless steppe-tundra vegetation over most of Europe, it
apparently withdrew to areas of southern Europe, where
areas of relict temperate woodland persisted (tzedakiS &
Bennett 1996, Van andeL & tzedakiS 1996). Because re-
mains are so few and the age range is close to or beyond
the reliable limits of radiocarbon dating, it is difcult to
reconstruct the detailed history of the latest P. antiquus;
and moreover the stratigraphical context of several nds
is uncertain. The evidence for Last Cold Stage survival is
based mainly on sparse records of Palaeoloxodon mate-
rial from Iberian and Italian cave sequences, in levels
with Mousterian artefacts and/or with associated absolute
dates (Stuart 2005).
At present, the best evidence comes from level 18
at Cueva del Castillo, northern Spain (aLtuna 1972,
BernaLdo de QuiroS 1982, caBrera-VaLdéS et al. 1996)
where P. antiquus molars are recorded in association
with Aurignacian artefacts, overlying a Mousterian level
with ESR dates averaging 70 ± 8 cal. ka. Radiocarbon
dates on two deciduous molars (almost certainly from
the same individual) of ca. 42.9 and > 47.3 ka (LQME
project; Stuart 2005), and charcoal dates of ca. 40 ka
from the same context, are best regarded as minimum
ages. A signicantly later record is claimed from Foz
do Enxarrique, Portugal, where an unworn upper molar
plate from Level C, has associated U-series dates (on
horse teeth) of ca. 33–34 cal. ka (BruGaL & raPoSo 1999,
SouSa & FiGueiredo 2001, Stuart 2005). However, this
record must be regarded with caution because the single
P. antiquus specimen was not dated directly and, as is
common elsewhere, the possibility of derivation from an
older level cannot be ruled out.
Clearly more work and new nds are needed to de-
termine more accurately the extinction chronology of
straight-tusked elephant. For the present we can say that
it survived well beyond the end of the Last Interglacial
in Iberia and Italy, and probably to at least ca. 50 cal. ka
in Iberia.
A date of 32,500 ± 500 ka on a partial molar from
Raalte, the Netherlands (MoL et al. 2005) appears anoma-
lous in that so far there are no other post Eemian records
from Central or Northern Europe, and requires further
investigation.
Stephanorhinus hemitoechus, ‘interglacial’ or ‘narrow-
nosed’ rhinoceros
Like P. antiquus, S. hemitoechus was widespread across
Northern Eurasia in the Last Interglacial and retreated to
southern Europe in the Last Cold Stage (Stuart 1991).
Largely because we are again dealing with events close
to the reliable limit of radiocarbon dating, the extinction
chronology of S. hemitoechus is uncertain at present,
although there are several cave sequences in southern
France, Spain and Portugal where it is recorded, from late
Mousterian levels (aLtuna 1972, BernaLdo de QuiroS
eschweizerbartxxx sng-
Stuart & LiSter: Late Quaternary megafaunal extinctions
290
1982, Stuart 1991), suggesting survival to perhaps 42
cal. ka. Putative later records from Aurignacian levels
(e.g. Lezetxiki Cave, Northern Spain; aLtuna 1972) need
careful checking as to stratigraphic context, and ideally di-
rect dating. A single tooth is recorded from an Aurignacian
level (level 11) in Bacho Kiro Cave, Bulgaria (KozłowsKi
1982) in broad association with a 14C date on charcoal of
37,650 ± 1450 (OxA–3183), (cal. 42,542 ± 1068).
Samples of S. hemitoechus from Cueva del Castillo
(see above, P. antiquus) submitted for radiocarbon dating
(LQME project) gave dates of 42–45 ka – probably mini-
mum ages. Other samples from Spain (Lezetxiki Cave)
and Italy (Grotta della Cala, Castelcivita Cave, Sora
Cave) contained insufcient collagen for dating.
The extinction of S. hemitoechus appears to have
occurred well before the Last Glacial Maximum, but a
detailed chronology is lacking at present.
Extinctions close to the onset of the Last Glacial
Maximum, ca. 26 cal. ka
Crocuta crocuta, spotted hyaena
Spotted hyaena, a carnivore and scavenger now conned
to Sub-Saharan Africa, was widespread across mid lati-
tude northern Eurasia during the Last Cold Stage (kahL-
ke 1994), but did not reach Beringia or the Americas. The
limited data available so far from Western Europe and
the Urals indicate possible extinction in Europe by ca. 32
cal. ka, broadly corresponding with the onset of the Last
Glacial Maximum (g. 1).
A number of hyaena remains from Britain (JacoBi et al.
2006, hiGhaM et al. 2006), dated some years previously by
the Oxford Radiocarbon Accelerator Unit, have been re-dat-
ed recently by the same unit using improved pre-treatment
methods, especially ultraltration. They have demonstrated
that several of the previous hyaena dates are erroneously
young, due to incomplete removal of consolidants. For
example, an earlier date of 22,880 ± 240 (OxA–6115) on
a tooth from Robin Hood Cave, Derbyshire, England, has
been shown to be much too young. Re-dating after prepa-
ration using ultraltration produced a date of > 52,800.
JacoBi et al. (2006) regard as suspect even the new date of
23,120 ± 130 (OxA–13659) on the re-sampled mandible
from Goat’s Hole (Paviland), South Wales. In view of these
results, the date of 24,000 ± 300 (OxA–4234) from Castle-
pook Cave, Ireland, will be checked by further dating. All
of the British hyaena dates appear older than ca. 27 ka.
The youngest reliable dates available so far from con-
tinental Europe (LQME project, using ultraltration) are
from Grotta Paglicci, Italy, 26,120 ± 330 (OxA–10523)
(cal. 30,841 ± 153); and “a cave in the Balkan Range”,
Bulgaria, 26,600 ± 170 (OxA–11551) (cal. 31,053 ± 86).
Claims of survival into the late Glacial and Holocene,
e.g. in southern Europe (carrion et al. 2001) and China
(tonG 2004) are the subject of current investigation.
Ursus spelaeus, cave bear
Cave bear is the only extinct megafaunal species that was
probably conned to Europe (kahLke 1994). It is recorded
from both cave and open sites, although most of the remains
come from the former – largely representing animals that
died in hibernation. In contrast to the omnivorous surviving
brown bear Ursus arctos, Ursus spelaeus appears to have
been essentially herbivorous, as shown by cranial and dental
morphology and low δ15N isotope values of bone collagen
(raBeder et al. 2000, BocherenS et al. 1994, Pacher & Stu-
art submitted). The diet very probably included a substantial
percentage of high quality herbaceous vegetation, implying
that such vegetation was available in the Alps and karst areas
of Central Europe, at least in some phases, during the Last
Cold Stage before the onset of the Last Glacial Maximum.
U. spelaeus probably disappeared from the Alps and
adjacent areas – currently the only region for which there
is good evidence – by ca. 24,000 14C years BP (ca. 28
cal. ka) (Pacher & Stuart submitted). Climatic cooling
and inferred decreased vegetational productivity around
the onset of the Last Glacial Maximum were very prob-
ably major contributors to its disappearance from this
region. There is little evidence so far of direct interaction
between cave bears and humans (Pacher 1997, 2002).
In collaboration with M. Pacher we are investigating
the possibility that cave bear survived signicantly later
elsewhere, especially in Southern Europe.
Extinctions in the Late Glacial and Holocene, ca.
15–4 cal. ka
Mammuthus primigenius, woolly mammoth
The woolly mammoth has by far the best coverage in terms
of 14C dates (SuLerzhitSky 1997, VaSiL’chuk et al. 1997,
kuzMin et al. 2001, MacPhee et al. 2002, Stuart et al. 2002,
Sher et al. 2005, Stuart 2005). In the middle of the Last
Cold Stage, MIS 3 ca. 44–24 cal. ka, woolly mammoths
occurred more or less continuously from Western Europe,
including Ireland, Britain, Spain and Italy, across Eastern
Europe and Siberia to China and Japan, and via Beringia to
the northern half of North America (kahLke 1994, Stuart et
al. 2002, aGenBroad 2005, Sher et al. 2005).
Shortly before ca. 13.8 cal. ka (ca.12 ka) mammoth
disappeared entirely and rather suddenly from Europe
and most of northern Asia (g. 2) (Stuart 1991, Sher
1997, Stuart et al. 2002, 2004). Signicantly, this dra-
matic event does not correlate with the marked warming
and spread of shrub-grassland vegetation over much of
Europe, which occurred at the beginning of the Late Gla-
cial Interstadial ca. 15.5–15.0 cal. ka, but does correlate
with the major loss of open biomes at the onset of the
Allerød (the rather cooler later part of the Late Glacial In-
terstadial) when boreal birch and pine woodland became
widely established (hoek 2001, Litt et al. 2003).
eschweizerbartxxx sng-
Cour. Forsch.-Inst. Senckenberg, 259, 2007
291
There is strong evidence (several radiocarbon dates from
more than one laboratory) that woolly mammoth popula-
tions continued to live in the far north of mainland Siberia
on the Taimyr Peninsula for a further two millennia (Sher
1997, SuLerzhitSky 1997, VaSiL’chuk et al. 1997, MacPhee
et al. 2002), in association with persistent open steppe-tun-
dra vegetation (Sher 1997). The latest known dates suggest
survival into the earliest Holocene: 9,670 ± 60 (GIN–1828)
(cal. 11,109 ± 94); 9,780 ± 40 (GIN–8256) (cal. 11,256 ±
72); 9,860 ± 50 (GIN–1495) (cal. 11,385 ± 102), and 9,920
± 60 (GrA–17350) (cal. 11,503 ± 128) (SuLerzhitSky 1997,
kuzMin et al. 2001, MacPhee et al. 2002, Stuart et al.
2002). Possible evidence of mammoth survival post 14 cal.
ka in southwest Siberia (orLoVa et al. 2004) requires further
investigation.
Although the data are limited at present, there are
strong indications that there was a modest re-expansion
of mammoth range ca. 12.6–11.7 cal. ka (ca. 10.5–10.0
ka) from Taimyr into the Yamal/Gydan Peninsulas (north-
west Siberia) and thence into northeast Europe (Stuart
et al. 2002, Stuart 2005; g. 2). This re-immigration to
Europe, after an absence of about 1.5 millennia, can be
plausibly linked to the renewed cold and open vegeta-
tional conditions of the Younger Dryas. The latest records
for Europe are from: Zhidikhovo peat bog, Cherepovets,
north of Moscow (rib from a partial skeleton), 9,760 ±
40 (GIN–8885c), 9,810 ± 100 (GIN–8676a) (cal. 11,315
± 168), 9,840 ± 50 (GIN–8885b) (cal. 11,350 ± 97); Pu-
urmani, Estonia (molar), 10,100 ± 100 (Hela–423) (cal.
11,847 ± 219), 10,200 ± 100 (Hela–425) (cal. 12,061 ±
232) (LõuGaS et al. 2002, Stuart et al. 2002). Since these
are similar dates by different laboratories from two dis-
tinct sites, the results are probably reliable, but conrma-
tion of each by an independent laboratory is desirable.
The nal extinction of mammoth therefore seems to
have occurred in mainland Eurasia (both in northeast
Europe and northern Siberia) in the very early Holocene
(kuzMin et al. 2001, MacPhee et al. 2002, Stuart et al.
2002). This event occurred soon after, but not coinciden-
tally, with the rapid warming that marks the beginning of
the Holocene, and can be plausibly correlated with the
loss of the steppe-tundra biome and widespread establish-
ment of temperate and boreal forests in mid latitudes and
boggy tundra in the far north (Sher 1997).
However, it seems improbable that habitats suitable
for woolly mammoth were entirely eliminated throughout
mainland northern Eurasia at this time, especially in view
of the Holocene presence of ‘steppe-tundra’ on Wrangel
Island, where an isolated population continued to ca. 4.1
cal. ka (LonG et al. 1994, Vartanyan et al. 1993, 1995).
Recently a mammoth molar from St Paul Island in the
Bering Sea off Alaska (Pribilof Islands) has been 14C dated
by the Arizona AMS laboratory to 7,908 ± 100 (AA26010)
(cal. 8,895 ± 124) and 8,015 ± 85 (AA34501) (cal. 9,013 ±
89) (Guthrie 2004). These Arizona dates have been inde-
pendently corroborated (LQME project) by an ORAU date
of 8,010 ± 40 (OxA–13027) (cal. 9,018 ± 45) on the same
specimen. Moreover, three specimens (probably from one
individual mammoth) from Qagnax Cave on the same is-
land have produced even younger dates ranging from 5800
± 80 (Beta 190142) (cal. 6601 ± 97) to 5630 ± 40 (Beta
190141B) (cal. 6404 ± 42) (yeSner et al. 2005).
Coelodonta antiquitatis, woolly rhinoceros
Although widely regarded as a ‘fellow traveller’ of the
mammoth and similarly adapted to the ‘steppe-tundra’ bi-
ome, judging by the relative scarcity of its remains woolly
rhinoceros was less common, and its range was less exten-
sive. It was absent from Ireland and north-central Siberia
(kahLke 1994) and did not reach North America.
After a contraction of range during part of the Last
Glacial Maximum, woolly rhinoceros returned to most ar-
eas in the Late Glacial, but failed to recolonize Britain and
southern Europe. In collaboration with E. WiLLerSer and
J. BinLaden, many more dates on woolly rhinoceros are
being done to get a full picture of its latest history, but the
youngest available records from Western Europe suggest
they had disappeared from this region by ca. 16–14 cal. ka
(ca. 13–12 ka), approximately coincident with either the
warming at the onset of the Late Glacial Interstadial (ca.
14.5 cal. ka), or with the marked reduction in open habi-
tats at the beginning of the Allerød (ca. 14 cal. ka). So far
there are no later dates from Siberia (Garutt & BoeSkoroV
2001, BinLaden et al. 2007).
The latest available dates include: Lobva Cave, Urals
Russia 12,275 ± 55 (KIA–5670) (cal. 14,233 ± 173); Grotto
Kotel, Urals Russia 13,245 ± 65 (OxA–10921) (cal. 15,802
± 149); Grotto Pershinsky 1, Urals, Russia 13,575 ± 65
(OxA–10928) (cal. 16,262 ± 159); Gönnersdorf, Germany
13,610 ± 80 (OxA–10201) (cal. 16,311 ± 173); and Vau-
marcus, Neuchatel, Switzerland 13,980 ± 140 (ETH–8777)
(cal. 16,827 ± 242). The suggestion that woolly rhinoceros
survived into the early Holocene in the Urals is based on
a single conventional 14C date of 9,510 ± 260 (IPAE–93,
former 14C laboratory, Ekaterinburg) (koSintSeV 1999).
This date must be regarded with considerable caution in the
absence of details of the pre-treatment methods used, or cor-
roborative evidence from this or other sites in the region.
Megaloceros giganteus, giant deer
Formerly widespread across the mid latitudes of Northern
Eurasia (kahLke 1994, Stuart et al. 2004), giant deer
apparently vacated most of Europe for much of MIS 2
including the Last Glacial Maximum, possibly surviving
in refugia north of the Black Sea and/or in Siberia (Stuart
et al. 2004). It reappeared, in Northwest Europe only, from
ca. 14.7 cal. ka, corresponding to the Late Glacial Inter-
stadial warming and a more productive vegetational pulse
(Bølling/Allerød), but this population was subsequently
extirpated in the low-productivity Younger Dryas (Moen et
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292
al. 1999, Stuart et al. 2004). The demise of giant deer in
Ireland at this time is unequivocally linked to environmen-
tal change alone, not overkill, as humans did not colonize
this area for another 2.5 millennia (WoodMan 1985).
Radiocarbon dating at Oxford (LQME project) and
Kiel has revealed previously unsuspected survival of
giant deer well into the Holocene (Stuart et al. 2004).
The youngest dates for giant deer known so far are from
open sites in Western Siberia, immediately east of the
Urals: Kamyshlov Mire, Western Siberia 6,816 ± 35
(KIA–5669 – on a rib) (cal. 7,655 ± 68), and 6,881 ±
3 8 (OxA–13015 – skull) (cal. 7,728 ± 72) from an as-
sociated skeleton; Redut, Miass River, Western Siberia
6,968 ± 33 (KIA–5668) (cal. 7,826 ± 70) and 7,034 ± 34
(OxA–13014) (cal. 7,900 ± 72), both on a cervical verte-
bra from an associated skull and vertebrae.
At present the timing of the nal extinction of giant
deer is uncertain, and is the subject of further work. Its
late survival in the Urals/Western Siberia region can be
attributed to the persistence there of mixed woodland-
grassland environments through the Younger Dryas. Its
extirpation from this region, although this might have
occurred signicantly later than 7.8 cal. ka, could plau-
sibly relate to vegetational changes ca. 9–7.8 cal. ka, in
which closed forest occupied the hills, and dry grassland
the plains – neither suitable for giant deer (PanoVa 1996,
Stuart et al. 2004). However, its demise might also relate
to the appearance of the Neolithic in this region ca. 9 cal.
ka (SerikoV 2000) with probably increased human popu-
lations and environmental modication.
Panthera leo spelaea, cave lion
Today lion is conned to Sub-Saharan Africa and one small
area in Northwest India (Gir Forest), but until ca. 200 years
ago it occurred also in North Africa, and through Turkey
and the Levant to Iran and northern India. Archaeological
nds from Neolithic, Bronze Age and Iron Age levels in
Bulgaria and Greece indicate Holocene presence of lion in
the Balkans until ca. 3 cal. ka (ninoV 1999).
The lion (‘cave lion’) that was widespread in Eurasia
through the Late Pleistocene is now generally regarded as an
extinct subspecies P. leo spelaea, or even species Panthera
spelaea (turner & antón 1997, SotnikoVa & nikoLSkiy
2006) and so has the status of an additional extinct megafau-
nal taxon in the Late Quaternary of Northern Eurasia. The
distinctiveness of the Late Pleistocene lion from modern Af-
rican populations is shown by ancient DNA studies (BurGer
et al. 2004) and analysis of cave art (turner & antón
1997). On morphological grounds SPaSSoV & iLieV (1994)
regard the Holocene lion material from the Balkans as P. leo,
which they infer immigrated from Asia Minor early in the
Holocene, after the extinction of P. spelaea.
From our accumulating dataset it is clear that, unlike
spotted hyaena (see above), cave lion survived into the
Late Glacial across wide areas of Europe. The latest dates
(LQME project) are: Zigeuenerfels, Sigmaringen, Germa-
ny 12375 ± 50 (OxA–17268) (cal. 14219 ± 112); Abri des
Cabones, Ranchot, France 12,565 ± 50 (OxA–12021) (cal.
14,853 ± 101); Podsemnich Ochotnikov, Urals 13,500 ±
65 (OxA–11349) (cal. 16,158 ± 157); Grotto Verhnegu-
bahinsky, Urals 13,560 ± 70 (OxA–10909) (cal. 16,241
± 163); Grotto Viasher, Urals 13,570 ± 70 (OxA–10908)
(cal. 16,255 ± 163); and Urtiaga, Spain, 13,770 ± 120
(OxA–10121) (cal. 16,534 ± 216). The published date
of 10,670 ± 160 (OxA–729) (cal. 12,678 ± 119) from
Lathum, Netherlands was done early in the life of the
Oxford laboratory, and improved pre-treatments are now
routinely applied by this laboratory to material in this age
range (Stuart et al. 2004, hiGhaM et al. 2006, JacoBi et al.
2006). A new determination 44,850 ± 650 (OxA–16715)
(cal. 48,792 ± 992) shows that the specimen is very much
older than previously thought.
Discussion
The pattern of Eurasian extinctions, summarised in g. 1,
is based on the evidence currently available, in particular
a substantial dataset of radiocarbon dates made directly
on megafaunal material. It emphasises the complex, rag-
ged or staggered pattern of extinctions in Northern Eura-
sian megafauna (Stuart 1991, Martin & Stuart 1995).
This pattern contrasts with a claimed much shorter time
range for extinctions in North America (Martin 1984,
Martin & SteadMan 1999).
Potentially signicant impacting factors in Europe
(see g. 1) are:
• Cooling trend (with many oscillations) from MIS 5e
into the Last Cold Stage.
• Appearance of modern humans in Europe, ca. 40 cal.
ka (possibly a little earlier).
• Onset of the Last Glacial Maximum, broadly 26 cal. ka
• Recolonization of Central and Northern Europe by
humans, ca. 16 cal. ka (GaMBLe et al. 2004)
• Onset of Late Glacial Interstadial, ca.14.5 cal. ka
• Spread of trees at onset of Allerød Interstadial, ca. 14
cal. ka.
• Onset of renewed cold of Younger Dryas, ca. 12.7 cal. ka
• Holocene warming and spread of forest, ca. 11.7 cal. ka.
The detailed pattern of megafaunal radiocarbon dates
(g. 2) strongly suggests a close relationship between
vegetational changes and the distributional shifts prior
to extinction. There are clear correlations between cli-
matic/vegetational changes and major range contrac-
tions/expansions in both extinct and extant species. Some
of these events may have led directly to extinctions, but
certain pieces of evidence suggest that humans could also
have been involved, perhaps in more subtle ways than
previously recognized (Stuart et al. 2004; and below).
eschweizerbartxxx sng-
Cour. Forsch.-Inst. Senckenberg, 259, 2007
293
The extinction of Palaeoloxodon antiquus and Steph-
anorhinus hemitoechus in their southern refugia could be
attributed to the overall cooling, and consequent decline of
woodland habitats, in the early and middle part of the Last
Cold Stage, broadly 75–30 cal. ka. Alternatively, their de-
mise could be related to the appearance of modern humans
Homo sapiens, now thought to have immigrated to Europe
ca. 40 cal. ka or possibly a little earlier (StrinGer 2006)
(g. 1). On the basis of the unsatisfactory dating evidence
currently available for these species (see above), we can-
not distinguish between these possibilities. The extinction
of Neanderthals Homo neanderthalensis ca. 35 cal. ka
has also been blamed on climatic deterioration (SteWart
2005), or alternatively on competition from, or even active
persecution by, modern humans (StrauS 2005). The avail-
able evidence suggests that there was an overlap between
the two species of perhaps 5 millennia in Europe (StrinGer
2006) (g.1). Others estimate a longer overlap, e.g. ca. 12
ka (FinLaySon 2005). On present data, the disappearance
of Neanderthals does not obviously correspond with any
major climatic event, but follows the arrival of modern hu-
mans, with a signicant lag between the appearance of one
species of Homo and the demise of the other. However, it
has also been argued that the combination of rapid climatic
uctuations and the arrival of modern humans were factors
in Neanderthal extinction (StrinGer et al. 2003)
The Last Glacial Maximum saw the disappearance
of most megafauna (extinct and extant) from much of
Europe, for variable durations probably related to the
ecology of each species (Stuart et al. 2004). Humans also
vacated most of northern and Central Europe for much of
this time (GaMBLe et al. 2004). The limited data available
so far indicate probable extinction of spotted hyaena (in
Northern Eurasia) and cave bear at the onset of the Last
Glacial Maximum before ca. 25 cal. ka. In contrast, cave
lion was widespread in the Late Glacial and survived until
at least ca. 14.5 cal. ka. Further work is needed, including
many more dates, to investigate possible explanations for
Fig. 1: Summary chart showing current interpretation of the chronology of extinction for megafaunal species in Europe and
Northern Asia. Solid lines: species present on the basis of direct radiocarbon dates. Grey lines: last occurrences in limited regions
(refugia). Dashed lines: occurrence inferred from limited absolute dating, archaeological association or stratigraphy. Question
marks: dates that are poorly evidenced. Note that lines implying continuous presence summarise data for the entire species’ range,
within which there were complex patterns of regional appearances and disappearances (Stuart et al. 2004, Stuart & LiSter in
preparation.). NB. P. antiquus and S. hemitoechus probably occurred only in Southern Europe over the time range shown. LGM:
Last Glacial Maximum; LGI: Late Glacial Interstadial; YD: Younger Dryas.
eschweizerbartxxx sng-
Stuart & LiSter: Late Quaternary megafaunal extinctions
294
these patterns. Intriguingly, on available evidence, mam-
moth, woolly rhinoceros and giant deer failed to return to
Southern Europe after the Last Glacial Maximum, perhaps
because humans continued to occupy these regions in
relatively high population densities through this period,
thus inhibiting recolonization by these megafaunal species
(Stuart et al. 2004). If such a relationship can be demon-
strated by further work, it would have important implica-
tions for understanding the role of humans in megafaunal
extinction.
The extinction of woolly rhinoceros may relate to
the warming at the onset of the Late Glacial Interstadial
ca. 14.5 cal. ka, although the timing of the extinction is
unclear (see above). Another possibility is that it may
have become extinct ca. 14 cal. ka, coincident with ma-
jor loss of the steppe-tundra biome with the onset of the
Allerød (see below). The Late Glacial Interstadial also
broadly correlates with the return of both giant deer
and humans to Northwest Europe. Interestingly, the hu-
man recolonization occurred a little in advance of the
climatic amelioration (GaMBLe et al. 2004). The onset
of the Allerød phase of the Late Glacial Interstadial (ca.
14 cal. ka), accompanied by widespread replacement of
former open habitats by forests, had a profound effect on
the megafauna. Mammoth withdrew apparently rapidly
from most of its range, and was then largely restricted
to north-central Siberia (Taimyr Peninsula) where open
steppe-tundra persisted. In contrast giant deer evidently
ourished (in North West Europe) during the Allerød (g.
2), and human populations in Europe also appear to have
increased (GaMBLe et al. 2004).
The renewed cold of the Younger Dryas ca. 12.7 cal.
ka saw the extirpation of giant deer from Western Europe.
Data from Eastern Europe and most of Siberia are lack-
ing at present, but we now know that it survived through
the Younger Dryas and into the Holocene in the southern
Urals and adjacent west Siberia (Stuart et al. 2004). In
marked contrast to the retreat of giant deer, at the same
Fig. 2: Chart of radiocarbon dates less than 16 ka (ca. 19 cal. ka) made directly on material of woolly mammoth Mammuthus
primigenius (black discs) and giant deer Megaloceros giganteus (grey discs) for Europe and selected areas of Siberia (modied
from Stuart et al. 2004). (For clarity northern Germany is grouped with southern Scandinavia). LGI: Late Glacial Interstadial; YD:
Younger Dryas. Note contrasting patterns related to differences in ecology between these two species (see text). Mammoth dates for
most of Europe and Siberia terminate ca. 12 ka (ca. 14 cal. ka) (Allerød), but mammoths survived into the early Holocene in main-
land North Siberia (Taimyr etc), and much later on Wrangel Island (MacPhee et al. 2002, Stuart et al. 2002, Stuart 2005, Sher
et al. 2005, LonG et al. 1994). Outlying mammoth dates ca.10 ka (ca. 11.7 cal. ka) in Northeast Europe, probably represent a brief
Younger Dryas repopulation from Northern Siberia. After a long absence during the Last Glacial Maximum, giant deer reappeared
(in Northwest Europe only) ca. 12.5 ka (ca. 14.7 cal. ka) (Britain) with most dates ca. 12–10.6 ka (ca. 14–12.6 cal. ka). Giant deer
disappeared from most of Europe at the Younger Dryas, but survived into the Holocene in the Urals/West Siberia region.
eschweizerbartxxx sng-
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295
time mammoth underwent limited re-expansion from its
North Siberian refugium into North East Europe, presum-
ably in response to the renewed spread of open herb-rich
vegetation (g. 2; Stuart et al. 2002, Stuart 2005).
The nal major climatic and vegetational event, the
Holocene warming ca. 11.7 cal. ka and extensive spread
of broad-leafed and conifer forests across northern Eura-
sia, broadly coincides with the disappearance of the
last known mainland mammoths (Taimyr) (gs 1, 2),
although several dates suggest that there was a lag of
several hundred years, perhaps reecting a lag between
climate change and vegetational response from dry steppe
tundra to boggy modern tundra (Sher 1997, MacPhee et
al. 2002). At the same time many extant species formerly
co-existing and widespread in the Last Cold Stage, such
as reindeer, arctic fox, horses and lemmings, retreated
either to the tundra or to the steppe as the steppe tundra
biome almost disappeared.
Most of those megafaunal species that did survive
into the Holocene seem to have undergone range frag-
mentation and reduction, before nally becoming extinct
at different times (gs 1, 2). Such a pattern, not correlated
with any marked climatic event (in contrast to the Last
Cold Stage), implies anthropogenic involvement.
There remain many unanswered questions on the
pattern and causes of the range shifts, and their signi-
cance for the nal extinction of megafaunal species. For
example, why did giant deer recolonize only northwest
Europe during the Late Glacial Interstadial? Why did it
subsequently survive only in limited refugia during the
Holocene? The identication of refugia, and of events
within them, is clearly critical for determining causes of
extinction. The possible role of humans in limiting the
natural expansion of these species, and in precipitating
the demise of refugial populations, also requires further
investigation, both in Europe and northern Asia.
Acknowledgements
Our research on megafaunal extinction in Europe and
northern Asia was supported by the Natural Environ-
ment Research Council (Grant # GR3/12599 and NE/
D003105). We are grateful to many individuals who have
contributed expertise and/or samples for 14C dating, but
especially to: Kim aariS-SørenSen, Judy aLLan, Jesus
aLtuna, Ian BarneS, Federico BernaLdo de QuiroS,
Marzia Breda, the late Victoria caBrera-VaLdéS, Ni-
cholas conard, Andrew currant, William daVieS, Linas
dauGnora, Julia FahLke, Irina ForonoVa, Clive GaMBLe,
Sabine GaudzinSki, Aurora GrandaL d’anGLade, Stephen
Green, R. Dale Guthrie, Brian huntLey, Roger Jaco-
Bi, Laura kaaGan, Ralf-Dietrich kahLke, Wighart von
koeniGSWaLd, Pavel V. koSintSeV, Yaroslav V. kuzMin,
Lembi LõuGaS, Jan Van der Made, Anastasia MarkoVa,
Federico MaSini, Lutz Christian MauL, Dick MoL, Lau-
ra niVen, Hannah o’reGan, Marylène Patou, Martina
Pacher, Maria Rita PaLoMBo, Ana Pinto, Benedetto SaLa,
Andrei V. Sher, Nikolai SPaSSoV, John SteWart, Martin
Street, Chris StrinGer, Elaine turner, Pirkko ukkonen,
Valentin ViLLaVerde, Alex VoroBieV, Piotr WoJtaL and
João ziLhão. We are grateful to A. ManGione, B. SaLa,
and Superintendenza per i Beni Archeologici dell’Umbria
for permission to use illustrations of megafauna in Figs 1
& 2. We thank Dick MoL and Jelle reuMer for valuable
comments on the manuscript. Especial thanks go to Tom
hiGhaM and the staff of the Oxford Radiocarbon Accel-
erator Unit.
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Manuscript submitted 2006–05–11
Revised manuscript accepted 2007–08–28
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