How do you kill 86 mammoths?

Taphonomic investigations of
mammoth megasites
Pat Shipman
*
Department of Anthropology, The Pennsylvania State University, University Park, PA 16802, USA
a r t i c l e i n f o
Article history:
Available online xxx
Keywords:
Mammoths
Dogs
Wolves
Hunting
Age proles
Taphonomy
a b s t r a c t
A series of Eurasian archaeological sites formed between about 40 e 15 ka feature unusually large
numbers of mammoth remains with abundant artefacts and, often, mammoth bone dwellings. None of
these mammoth megasites is dated prior to the appearance of modern humans in Eurasia. This unusual
type of site begs for taphonomic explanation. The large number of individual mammoths and the scarcity
of carnivore toothmarks and gnawing suggest a new ability to retain kill mammoths and control of
carcasses. Age proles of such mammoth-dominated sites with a large minimum number of individuals
differ statistically at the p < 0.01 level from age proles of Loxodonta africana populations that died of
either attritional or catastrophic causes. However, age proles from some mammoth sites exhibit a chain
of linked resemblances with each other through time and space, suggesting the transmission of
behavioral or technological innovation. I hypothesize that this innovation may have been facilitated by an
early attempted domestication of dogs, as indicated by a group of genetically and morphologically
distinct large canids which rst appear in archaeological sites at about 32 ka B.P. (uncal). Testable pre-
dictions of this hypothesis are generated based on ethnographic data.
2014 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
Since the late nineteenth century, archaeological sites domi-
nated by mammoth remains have been an important focus of
research. A key question is how the bones of large numbers of
mammoths, ranging from a minimum number of individuals (MNI)
of about ve to hundreds, were deposited in one place. Despite
much investigation, the cause of death of mammoths in these sites
has remained controversial (e.g., Klima, 1954, 1963; Koz1owski
et al., 1974; Klein, 1982; Vereschagin and Baryshnikov, 1984;
Soffer, 1985, 2003; Haynes, 1991; Fladerer, 2003; Wojtal and
Sobczyk, 2003; Svoboda et al., 2005; Wojtal, 2007; Brugre and
Fontana, 2009; Wojtal et al., 2012). Two dominant hypotheses are
that these sites represent natural deaths which were scavenged by
humans; or that specialized human hunting resulted in such as-
semblages. Here, for convenience, I refer to such sites as mammoth
megasites. An argument could be made that large sites heavily
dominated by other particular species should also be termed
megasites.
Many ambiguities and inconsistencies in excavation and
collection techniques used by different scholars have complicated
attempts to construct an overall consensus on the purpose, for-
mation, taphonomy, and use of these megasites, which span a large
geographic and temporal range. Questions about the accuracy of
radiocarbon dates on many mammoth megasites are unresolved
because recent advances in decontamination, ultraltration, and
the use of strict criteria to eliminate samples with poor preserva-
tion show that many dates in the literature calculated decades ago
are inaccurate (Bronk et al., 2007; Higham et al., 2012; Wood et al.,
2013). Nonetheless, the wealth of material and the inherent fasci-
nation of sites with large numbers of mammoth bones call scholars
to attempt to understand these sites.
Fortunately, many recent researchers have compiled basic in-
formation from zooarchaeological analyses of the sites, including
number of non-mammoth species represented, mammoth MNIs,
mammoth age at death, and mammoth age proles fromindividual
sites. Demographic studies have gained in importance as indicators
of taphonomic processes due to research by several workers (e.g.
Voorhies, 1969; Saunders, 1977; Klein, 1982; Klein and Cruz-Uribe,
1984; Haynes, 1991; Stiner, 1991) comparing age proles of fossil
assemblages to those of animals of known causes of death. In
particular, proles of the age at death of mammoths represented at
various sites have been published and used to formulate
* Permanent address: 3140 Chatham Church Road, Moncure, NC 27559, USA.
E-mail addresses: pat.shipman9@gmail.com, pls10@psu.edu.
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Quaternary International xxx (2014) 1e9
Please cite this article in press as: Shipman, P., How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites, Quaternary
International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048
interpretations about site formation processes. In an attempt to
contribute to progress in deducing the taphonomic histories of
sites, I have used these published age proles for further analysis.
The researchers who have published the age proles on these
sites have also commented on additional indicators of taphonomic
history, such as cutmarks, carnivore toothmarks, evidence of
burning, and skeletal representation. Virtually all sites discussed
here include lithic remains from the Gravettian or Epigravettian
industries, with the exception of Molodova I, level 4, which is a
Mousterian site. No lithic assemblage includes novel tool types that
have been suggested to be designed for killing mammoths. Bones
show scarce signs of carnivore activity and human modication
(usually <5% of the mammoth bones have such evidence). The
human modications take the form of cutmarks, lleting marks,
skinning marks, disarticulation marks, fracturing of long bones for
marrow, and/or shaping of bones as tools. Some sites also include
architectural structures such as mammoth bone huts, refuse dumps
or pits, postholes, and hearths. There is no evidence of pit traps or
drive lanes.
1.1. Regional, temporal, and geographic distribution of mammoth
megasites
Dozens of mammoth-dominated sites are located in central and
eastern Europe, with concentrations in the Swabian Jura, the
Danube area and Moravian Gate, and up into Russia and Siberia.
Typically the sites are located on high river terraces or low
mountainous areas in close proximity to water. Mid-slope locations
with easy access to rivers are common. Commonly the remains are
buried in loess alternating with sandy clayey layers, but geological
details can be found in individual publications. If the abundance of
mammoth remains is related to natural disasters such as a herds
falling through the ice on a frozen river or falling into ravines while
accessing salt or mineral licks, then the bones of those mammoths
have, in general, been transported uphill after the deaths.
No mammoth megasites date to before the arrival of anatomi-
cally modern humans (AMHs) in Eurasia. The role of AMHs in the
formation of such sites is often postulated: only two mammoth
megasites may indicate activity of Neanderthals.
1.2. Other important features of the sites
Mammoth tends to dominate these faunal assemblages over-
whelmingly. The bones and individual mammoths also occur at
very high densities. For example, mammoths comprise 99% of the
fauna (Number of Individual Specimens 5860) at Krakw Spad-
zista Street (Wojtal and Sobczyk, 2005). InTrench B B1 at this site,
there was 1 individual mammoth for every 2 m
2
of excavation
(Wilczy nski et al., 2012:3638). This is comparable to the density of
mammoth individuals at Yudinovo, an Epigravettian site, in which
the pavilion area yielded 1 mammoth individual for every 1.4 m
2
(Germonpr et al., 2008: 481). Such densities contrast sharply with
those at Middle Paleolithic or Mousterian sites, in which mam-
moths are often among the most common six species in the as-
semblages, but horses, cervids, or bovids usually dominate (Patou-
Mathis, 2000). Analysis of the Stage Three Project mammalian
database, which includes 447 dated faunas from291 archaeological
and non-archaeological sites, yields a similar result. The database
contains three times as many Upper Paleolithic sites as Middle
Paleolithic ones, but an Upper Paleolithic preference for mammoths
is shown by the presence of mammoths in 4.5 times as many Upper
Paleolithic sites as Middle Paleolithic ones (Stewart, 2004). Stewart
interprets this as indicating a more systematic exploitation of
mammoths by Upper Paleolithic hominins.
Studies of the stable isotopes preserved in bones of Neander-
thals and AMHs indicate that both consumed mammoths
(Bocherens and Drucker, 2006; Bocherens, 2011). In addition to the
most common prey e horse e reindeer, rhinoceros, and bovines
were in the Neanderthal diet. Isotopic studies reveal that Nean-
derthals ate larger quantities of mammoth than zooarchaeological
studies reveal, probably because Neanderthals were reluctant to
either camp near the carcass or to transport heavy esh-bearing
mammoth bones from the kill site to camp (Bocherens, 2011: 80e
81). In contrast, Upper Paleolithic sites are also much more likely to
contain all parts of the mammoth skeleton (e.g. Wojtal and
Sobczyk, 2005; Brugre and Fontana, 2009; Bosch, 2012).
Mammoths represented a wealth of resources which were
demonstrably used in the Upper Paleolithic. Mammoths were eaten,
judging from clear cutmarks on some bones at sites such as Grub-
Kranawetberg, Krakw Spadzista Street (B), Krems-Hunddsteig,
Krems-Wachtberg, and Langmannersdorf-Lagerplatz B (summa-
rized in Bosch, 2012). Stable isotope studies conrm this interpre-
tation (Bocherens, 2011; Bocherens et al., 2013, this volume).
Mammoth bone and ivory were used to make tools and sculptures
or art objects (Conard, 2003; Antl and Fladerer, 2004; Wojtal et al.,
2012). Bone and tusks were also used as construction materials and
many Gravettian or Epigravettian mammoth megasites feature re-
mains of dwellings carefully built of mammoth bone and sometimes
supported by tusks. The frameworks were apparently covered in
mammoth hides, often with one or more hearths inside (Soffer,
1985). At least 70 mammoth bone huts are known from the
Russian Plain alone (Ward, 1998), but mammoth bone dwellings are
not conned to this region (Iakovleva and Djindjian, 2005; Svoboda
et al., 2005). Only one Mousterian site, Molodova I, level 4 contains
evidence of a similar structure (DeMay et al., 2012), but this inter-
pretation is not universally accepted (Gargett, 2011).
The representation of different skeletal elements at each site also
offers useful information. Sites where all skeletal elements are pre-
sent (such as Krems-Hundssteig, Krakw Spadzista Street, Lang-
mannersdorf, Milovice I) suggest that the mammoth(s) were killedat
or very near the site. Sites with only selected skeletal elements of
mammoths (such as Mezhirich, Yudinovo, and Berelekh) are more
suggestive of human or possibly hydraulic transport of remains.
Some suggest that the mammoth bones used for building pur-
poses were scavenged (Soffer, 1985, 2003; Haynes, 1991) from
natural deaths, citing of a lack of available wood in the mammoth
steppe environment. The frequent nding of burned mammoth
bones (e.g., Soffer et al., 1997) shows bones were also used as fuel
for re, another practice that has been related to a lack of wood.
However, recent studies of microscopic charcoal remains at several
sites suggest that birch and willow were more numerous along
river courses than previously believed and wood was used as fuel
(Beresford-Jones et al., 2010; Marquer et al., 2012).
In some of these mammoth megasites, carnivores, particularly
Arctic fox (Alopex lagopus) and wolf (Canis lupus), are unusually
well represented by many bones and high MNIs. Recent morpho-
logical and genetic studies (Germonpr et al., 2009, 2012, 2013, this
volume; Thalmann et al., 2013) show that some of the large canids
from mammoth megasites are a distinctive group which may
represent an early attempt at the domestication of the dog instead
of wolves. Some of the long bones of large canids, whether wolf or
dog, show cutmarks indicative of dismemberment, skinning, and
lleting (e.g. Wojtal and Sobczyk, 2005; Wilczy nski et al., 2012),
showing both fur and meat was used.
2. Material and methods
A literature search yielded information on mammoth MNIs and
age proles from fteen individual sites (Table 1). Proles of age at
P. Shipman / Quaternary International xxx (2014) 1e9 2
Please cite this article in press as: Shipman, P., How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites, Quaternary
International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048
death can be used to separate scavenged remains fromhunted ones
(Klein, 1982; Klein and Cruz-Uribe, 1984; Stiner, 1991). Such infor-
mation would clarify and possibly refute hypotheses about the
formation of these mammoth megasites. Although age proles are
often reported in the literature, they are not usually evaluated
statistically. A major aim of the research reported here was to
conduct such statistical comparisons among age proles fromfossil
sites and those from modern proboscideans with known causes of
death.
Publications on three mammoth megasites: Berelekh, Siberia;
Mezin, Ukraine; and Mezhirich, Ukraine, relied on Saunders (1977)
system of assigning age classes on the basis of mandibular tooth
eruption and wear (Fig. 1a and b). Saunders based his system on
work by Laws (1966) and Laws and Parker (1968) with Loxodonta
africana in Murchison Falls National Park, Uganda. Laws and Parker
(1968) reported the demographics of family groups, as revealed
during a culling operation. An age in African Elephant Years (AEY)
was assigned to each individual and individuals were then grouped
into four age classes each consisting of ten years, with all older
adults being grouped into the last category. The resulting age
prole serves as a model for either an intact, living elephant herd
consisting primarily of maternal or nursery groups, and the age
prole of such a herd if struck down by either catastrophic or
attritional causes that were not selective as to the age of the indi-
vidual animal. Saunders used this same procedure for ageing
mammoths and mastodons, constructing an age prole, and
interpreting the results.
In addition to representing attritional or catastrophic modes of
death, analysts often judge age proles as either U-shaped, with the
highest mortality among the very young and very old, or L-shaped,
with the highest mortality among the very young, following Klein
(1982) and Haynes (1991). These judgments are made intuitively,
not statistically as a rule.
Twelve mammoth megasites in the literature used the method
of assigning ages at death in AEY, based on eruption and wear
dental remains, as favored by Haynes (1991; Conybeare and
Haynes, 1984: Fig. 2). However, individuals were clumped into
twelve-year age classes, not ten-year age classes as Saunders had
done. This twelve-year system was developed by Haynes (1991),
based on unpublished work with L. africana by G. Craig and
descriptive studies of elephant die-offs of known cause in
Zimbabwe (Conybeare and Haynes, 1984; Haynes, 1991). Haynes
datasets yield two models representing drought deaths (Fig. 3),
which affected different age classes unequally, and culling deaths in
Zimbabwe, in which entire maternal herds were shot at one time.
Because the aging methods are well-known and well-published, I
will not repeat procedural details for age determinations here.
2.1. Statistical analyses
Haynes reference data (1991) are widely used in the literature,
so I rst conducted statistical comparisons of his two datasets.
Those derived from 291 attritional deaths in a prolonged drought
were compared with 149 deaths caused by culling of family herds.
Resemblance or difference between datasets was judged by calcu-
lating the D statistic of the KolmogoroveSmirnov test, following
Klein and Cruz-Uribe (1984). Values of D 1.64 indicate a signi-
cant difference between two samples at the p < 0.01 level. The
KolmogoroveSmirnov test is appropriate because it is sensitive to
Table 1
Mammoth megasites with published information on ages at death of individuals.
Site age (rC14 yrs) MNI mammoths
Molodova, Ukraine > 44,000 15
Vogelherd, Germany 33,000e30,000 15
Krems-Wachtberg, Austria 27,000 8
Milovice G, Czech Republic 26,000e24,000 21
Milovice I, Czech Republic 22,000 51
Krems-Hundssteig, Austria 28,000 7
Pavlov I, Czech Republic 27,000e25,000 7
Predmost, Czech Republic 26,000 105
Grub-Kranawetburg, Austria 25,000 8
Mezin, Ukraine 25,000 e 24,000 44
Krakw Spadzista Street, Poland 24,000e23,000 86
Langmannersdorf, Austria 22,000 e 20,500 27
Langmannersdorf e Lagerplatz, Austria 21,000 7
Yudinovo, Russian 16,000e12,000 35
Mezhirich, Ukraine 15,245 109
Berelekh, Siberia 13,400e10,400 166
0
5
10
15
20
25
30
35
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Age Profile
0-9 yrs
10-20 yrs
20-30 yrs
30+ yrs
0
10
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Bereleyk Mezin Mezhyrich
0-10 yrs
11-20 yrs
21-30 yrs
30+ yrs
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Mammoth Megasites
Berelekh Mezin Mezhirich
a
b
Fig. 1. a. Age prole of Loxodonta africana from Saunders (1977). This represents a
living herd of maternal groups. b. Age proles of mammoths from three sites. Age
classes were assigned according to the system developed by Saunders (1977). The
mammoth age prole from Bereleyk does not differ from the living herd prole
(compare Fig. 1a and b: KolmogoroveSmirnov D statistic 0.8, p > 0.01) or from the
age prole at Mezin (D 0.6, p > 0.01). The prole from Mezin does not differ
signicantly from the living herd prole (D 0.64, p > 0.001). The Berelekh prole
differs statistically from that at Mezhirich (D 2.52, p < 0.001). The Mezhirich prole
differs from the living herd prole (D 1.89, p < 0.01) and from the prole at Mezin
(D 1.98, p < 0.001). The Berelekh and Mezin age proles therefore suggest intact
herds, or repeated, non-selective hunting, while the Mezhirich prole may be a
palimpsest of different events.
P. Shipman / Quaternary International xxx (2014) 1e9 3
Please cite this article in press as: Shipman, P., How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites, Quaternary
International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048
rank ordering of data (i.e., changes in the number of individuals in
one age class will affect the others).
My aimwas to compare the age prole of each fossil megasite to
each of Haynes datasets statistically as a means of gleaning addi-
tional data relevant to deducing the cause of death or accumulation
of the mammoths. Because large sample sizes are required by the
KolmogoroveSmirnov D statistic, Klein and Cruz-Uribe (1984:57,
59) suggest at least 25 individuals are necessary for a valid inter-
pretation of an age prole and that for maximum reliability each
sample ought to contain at least 40 individuals. I chose to discard
fossil age proles involving fewer than the minimum of 25 in-
dividuals required for reliable statistical evaluation. Only eight
fossil sites included enough aged individuals to analyze further
(Table 1, Fig. 3): Berelekh, Siberia; Krakw Spadzista Street, Poland;
Langmannersdorf, Austria; Mezhirich, Ukraine; Mezin, Ukraine;
Milovice I, Czech Republic; Predmost, Czech Republic; and Yudi-
novo, Russia. The published MNI values of mammoth specimens
that are classied into age categories ranged from 27 to 105.
The interpretation of age proles as an indicator of mortality
pattern may have special advantages in analyzing extremely large
and long-lived species. Because proboscidean teeth (and bones) are
so large, there is a much lower possibility of the destruction of teeth
of very young individuals due to their fragility when exposed to
natural (i.e., non-hominin) taphonomic forces. Even the teeth of
young proboscideans are robust and more likely to be preserved as
compared to those of smaller mammals. Fetal or very young
mammoth remains were identied from Krakw Spadzista Street,
Yudinovo, Predmost, Langmannersdorf, and Krems-Wachtberg, for
example. Further, despite the fact that the age classes of different
sites were most often assigned by different researchers, the po-
tential for errors in assigning age class is lessened because the age
classes are so broad. In contrast, in analyzing the remains of species
with annual births that reach sexual maturity at 2 years, a yearling
could possibly be mistaken for a nearly-mature individual. The
possibility of a researcher mistaking a young proboscidean for one
that is 10e12 years old is much less. Thus, potentially, analyzing age
proles of mammoths may be more reliable and informative than
analyzing age proles of some other species. The analytical proce-
dure is detailed below.
Age proles of the three mammoth megasites aged by the Laws/
Parker method were compared with age proles from Saunders
model, which represents a catastrophic death of family herds, using
the KolmogoroveSmirnov test to determine signicance. Finally,
each age prole of a mammoth megasite was then compared sta-
tistically to every other one which had been classied by the same
ageing method.
3. Results
Although many publications have compared mammoth age
proles to those published for modern elephants by Haynes (1991),
it had not been established previously that Haynes two models
differ at a statistically signicant level. This evaluation conrms
that the age proles of the attritional and catastrophic death as-
semblages presented by Haynes (1991) are statistically different,
yielding a KolmogoroveSmirnov D statistic of 2.4, well above the
p < 0.01 level. Thus, the basic premise that these two age proles
differ meaningfully and may be helpful in determining a pattern of
0
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Mammoth Megasites
0-12yrs
13-24 yrs
25-36 yrs
37-48 yrs
49-60 yrs
0
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Drought Culled
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Cause of Death
0-12 yrs
13-24 yrs
25-36 yrs
37-48 yrs
49-60 yrs
Fig. 2. a. Age proles of two populations of Loxodonta africana aged according to
Haynes (1991). One represents a population killed in a drought of several years
duration and the other, maternal herds killed in culling operations. The drought-killed
animals are disproportionately often very young, while the culled animals show a
higher proportion of middle-age or prime individuals. The drought prole differs
statistically from the culled prole (KolmogoroveSmirnov D 2.4, p > 0.001). b. Age
proles of ve different mammoth populations aged according to Haynes (1991). The
age proles from these sites all differ statistically from either the drought or the culled
model proposed by Haynes (1991) at the p > 0.01 level. The age proles of the sites are
arranged according to a pattern of resemblances of age proles. Each sites prole
differs signicantly from all others at the p > 0.01 level with the exception of the next
adjacent site, reading left to right, on the x axis. Predmost differs from all sites except
Kraw-Spadzista (D 0.128, p > 0.10). Krakw Spadzista Street differs from all sites
with the except of Predmost and Langmannersdorf (D 0.128, p > 0.10). Langman-
nersdorf differs from all sites except Krakw Spadzista Streetand two others: Milovice I
(D 0.41, p > 0.10) and Yudinovo (D 0.54, p > 0.10).
Fig. 3. The statistical comparisons of the mammoth age proles from all sites classied
according to Haynes system reveal an interesting pattern of resemblances. The top box
shows (with double-headed arrows) the age proles that resemble each other statis-
tically. The approximate dates of each site are given in the middle section of the dia-
gram. The overall geographic distribution of the sites is shown in the bottom box.
Generally each site resembles the next most recent and next closest site, moving in a
west to east direction. Langmannersdorf and Milovice I may not differ meaningfully in
age. This pattern may represent the transmission of a mammoth-killing behavior that
changes across time and space.
P. Shipman / Quaternary International xxx (2014) 1e9 4
Please cite this article in press as: Shipman, P., How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites, Quaternary
International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048
death is supported. Haynes model samples consisted of 149 (cull-
ing) and 296 (drought) individuals each.
All mammoth megasites aged using Haynes system e
Predmost, Krakw Spadzista Street, Yudinovo, Langmannersdorf,
and Milovice I e differed signicantly fromboth Haynes attritional
model and his catastrophic model at the p < 0.01 level in each case.
Two mammoth sites evaluated against Saunders model, Berelekh
and Mezin, do not differ signicantly from Saunders model and
thus represent an intact elephant herd, repeated killing of family
groups, or one entire herd killed catastrophically. Mezhirich, the
third site with a published age prole assigned according to
Saunders procedures, differs signicantly from Saunders model
with a D value of 2.52 (p < 0.01). Saunders model sample consists
of 135 individuals.
The site-by-site comparison of Paleolithic age proles to each
other yielded more promising results. Although most site-
specic age proles did not resemble each other closely, the
pattern of resemblances and differences suggested was
interesting.
The Predmost age prole differed signicantly (at the p < 0.01
level) from that at any other site, except for the age prole from
Krakw Spadzista Street. The age prole from Krakw Spadzista
Street differed from that at every other site except those from
Predmost and Langmannersdorf. The Langmannersdorf age prole,
as expected, did not differ from the prole from Krakw Spadzista
Street, nor from the proles from Yudinovo or Milovice I. The age
proles fromYudinovo and Milovice I differed signicantly fromall
others at the p < 0.01 level with the exception of the prole from
Langmannersdorf, from which each was insignicantly different.
4. Discussion and overview
As Haynes attritional and catastrophic models differ from each
other signicantly at the p < 0.01 level, the results reported here
validate the notion of using elephant age proles from populations
with known causes of death as a means of interpreting mammoth
age proles. Statistical analysis of all the mammoth megasites aged
using Haynes criteria differ signicantly from either model, sug-
gesting that many of these sites may be palimpsests accumulated
over time with a mixture of taphonomic histories, rather than
single events. This nding suggests that attempts to judge re-
semblances or differences among age proles intuitively may not
be productive.
Two mammoth megasites, Berelekh and Mezin, do not differ
from Saunders model at the p < 0.01 level. This nding indicates
that these assemblages may represent either a massive catastrophe,
which killed an entire elephant herd at once, or a repeated series of
killings of nursery herds. The third mammoth megasite aged by
Saunders criteria, Mezhirich, is signicantly different from Saun-
ders model at the p < 0.01 level, suggesting a possible palimpsest.
The chain of resemblances e or of insignicant differences e
produced by the site-by-site comparisons reproduces the approx-
imate chronological order of these sites, as judged from best
available
14
C dates (Table 1). The chain of resemblances also con-
nects sites to other sites in an east-west sequence, going from the
oldest to the youngest site (Fig. 4).
4.1. Taphonomy and possible explanations
Most mammoth megasites evaluated here do not conform to
models based on living elephants which experienced either attri-
tional or catastrophic deaths. Five possibilities might explain this
outcome.
1) Possibly mammoth dental eruption and wear does not closely
parallel that in Loxodonta, as has been assumed by both ageing
techniques. Given the relatively close phylogenetic relationship
between Mammuthus and Loxodonta, and the detailed re-
semblances in craniodental anatomy, this seems unlikely.
2) The criteria used for excavation, collection, and ageing of
these assemblages may have been inadequate to preserve
resemblances in age class proles which were actually pre-
sent at the time of site formation. Though some of the sites
were excavated and collected decades ago, and standards
have certainly changed, most or all of these assemblages have
been reanalyzed in recent times by skilled researchers. If the
collected dental remains accurately represent what was
present at the site at the time of excavation, it seems likely
that minimal taphonomic biases were introduced by the
process of collection per se. If collection standards were
biased toward more complete or more attractive specimens,
most zooarchaeological analyses will not prove helpful and
modern analysts will be unable to derive useful information
about these sites.
3) Most of these assemblages may represent a palimpsest of
different events which has weakened or destroyed the original
taphonomic signature of the age proles, to the point that sta-
tistical signicance is not achieved in these comparisons. This
nding would point to repeated reoccupation of these sites,
perhaps seasonally, with variations in subsistence strategy be-
tween occupations. This potential explanation is entirely
possible and cannot be discounted, especially given the time
span represented by these sites and their broad geographic
distribution.
4) Another potential explanation is that generalizing from modern
proboscidean behavior to ancient behaviors of different species,
across vast distances of space and time, must be viewed with
extreme caution and may be invalid. There is no obvious way to
evaluate the likelihood that this explanation is true, although
using such phylogenetic and behavioral analogues is almost
standard in the eld.
5) A nal explanation is that the chain of resemblances linking
each mammoth site to the next closest one in time and space
may reect the transmission of a specialized but changing
behavior or technology through time and space.
The fth possible explanation is the most encouraging for pa-
leoanthropologists and taphonomists hoping to understand the
past. The sites in this analysis are spread across most of the Eurasian
continent and throughout a substantial span of time. Perhaps
expecting consistency in detailed patterns of cultural activities,
such as hunting and butchering, across thousands of years and
thousands of miles is unrealistic.
The introduction or growing use of complex projectile weap-
onry by modern humans was probably important in producing the
abrupt changes in assemblages associated with hominins seen
starting about 45 ka. As Shea (2008:177) has observed, complex
projectile weaponry is a niche-broadening technology because
such weapons are light, easily carried, and they retain energy
longer in ight, allowing them to be used against larger, dangerous
prey, or other humans, with less risk of injury, enabling early
modern humans to broaden their ecological niche(s) (see also
Lombard and Phillipson, 2008). The resultant transformation from
being an ambush predator, as Neanderthals were (Finlayson, 2009),
to being long-distance hunters may have conferred considerable
benets on early modern humans, particularly in killing very large
and dangerous mammals.
I suggest here that a second advance which occurred during MIS
3 may have enhanced the advantages of improvements in lithic
P. Shipman / Quaternary International xxx (2014) 1e9 5
Please cite this article in press as: Shipman, P., How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites, Quaternary
International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048
technology. There would have been great advantages to the pres-
ence of even quasi-domesticated large canids willing to work
cooperatively with AMHs. A group of such canids that is morpho-
logically distinct from wolves and postulated to be a domesticated
dog has been identied by Germonpr et al., 2009, 2012, 2013, this
volume). Morphologically, members of this group have also been
identied at Predmost, Mezin, and Mezhirich (Germonpr et al.,
2009, 2012, 2013, this volume).
Fig. 4. Eight mammoth megasites had a sufcient number of individuals classied into age classes (N > 25) to be analyzed statistically. They vary in antiquity from about 22,000 to
10,400 years according to uncalibrated radiocarbon dates. However, recent advances in techniques for decontamination and ultraltration have not been applied to these sites so
their dates may be subject to correction when this is done. Map by Jeffery Mathison.
P. Shipman / Quaternary International xxx (2014) 1e9 6
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International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048
4.2. The domesticated canid hypothesis
In additional to the use of complex projectile technology that
facilitated distance killing, I hypothesize here that an unprece-
dented alliance between domesticated or partly domesticated ca-
nids and humans may have played a vital role in the appearance of
the large, complex, mammoth megasites in the Upper Paleolithic.
For convenience, I call these canids wolf-dogs here because it is
uncertain whether or not they were directly ancestral to any living
canid.
I specically identify these canids with the morphological group
rst identied by Germonpr and colleagues at Goyet Cave.
Belgium. I specically do not expect Upper Paleolithic wolf-dogs to
be the same in all respects as modern dogs. It would be unusual if
rst known domestication or domestication attempt of any species
showed all the features of later domesticates.
The Belgian morphological group of wolf-dogs is further set
apart by recent genetic work studies on three individuals, including
one clear morphological wolf-dog from Goyet Cave. These in-
dividuals possess an unusual mtDNA haplotype unknown among
modern dogs, modern wolves, or ancient wolves (Thalmann et al.,
2013). This nding shows that this maternal lineage or mtDNA
haplotype did not leave any known direct descendants; phyloge-
netically, the Belgian canids appear basal to all known other groups.
The genetic data are consistent with a scenario in which males with
this haplotype interbred with female wolves, giving rise to a pop-
ulation ancestral to either modern dogs or wolves. As Thalmann
et al. (2013) remark, the genetic data are also consistent with an
interpretation of these unusual canids (here, wolf-dogs) as an early
abortive attempt at domestication, which left few if any direct
descendants, or as a Pleistocene wolf population that also became
extinct without issue.
4.3. Predictions of the domesticated canid hypothesis
I use ethnographic analogies, particularly studies of hunters
operating with and without the assistance of dogs (Ruusila and
Pesonen, 2004; Koster, 2009; Lupo, 2011; Koster and Tankersley,
2012) to generate testable predictions of the domesticated canid
hypothesis.
The rst prediction is that additional examples of the unusual
large canids identied by morphometric and genetic techniques
will be found in mammoth megasites and will not be found in
Middle Paleolithic sites, Mousterian, or pre-AMH sites.
The second is that additional mammoth megasites will possess
specimens of such wolf-dogs. I do not predict all such sites will
yield such specimens due to differences in taphonomy and
preservation.
The third is that sites formed by hominins with wolf-dogs will
yield evidence of a larger number of kills and more meat yield than
sites formed by hominins without wolf-dogs. Ethnographically,
hunters with dogs have a markedly increased meat yield per hunt
and a decreased time of seeking prey (Ruusila and Pesonen, 2004;
Lupo, 2011; Koster and Tankersley, 2012). People have used dogs
both to locate large prey like muskoxen, elk, wolf, or polar bear and
to hold the prey in place while alerting human hunters by howling
(Arnold, 1979; Ruusila and Pesonen, 2004). Dogs have also been
used with drive lanes to trap large mammals in enclosures or drive
them off of cliffs but there is no evidence that this was done at the
mammoth megasites.
Mammoth megasites already amply fulll this prediction when
compared with Mousterian or Middle Paleolithic sites. Faster
population growth in wolf-dog using peoples is expected due to
improved nutrition and less energetic expenditure. Though esti-
mating ancient population size is inherently speculative, following
the strategy developed by Mellars and French (2011), I would
expect to see evidence of faster population growth in AMHs relative
to Neanderthals by an increasing relative number of archaeological
sites, an increasing size of archaeological sites, an increasing den-
sity of retouched stone tools at such sites, and an increasing density
of estimated meat yield relative to site size.
The fourth is that sites formed by AMHs working with wolf-dogs
should yield evidence of longer occupations than those of AMHs or
Neanderthals working without cooperative wolf-dogs. This is pre-
dicted because of the ethnographic use of dogs in guarding both
food and homesteads among the Neoeskimo (Arnold, 1979).
Retaining control of huge carcasses and diminishing raids by
scavengers would constitute a considerable advantage.
The fth is that undomesticated canids, such as wolves and
foxes, will be found in large numbers primarily in sites where wolf-
dogs are known. This prediction is based on the well-known fero-
cious territoriality of modern canids and their aggressive response
to canid intruders (Geist, 2006; Vanak and Gompper, 2009). If
domesticated or semi-domesticated canids lived with AMHs, the
wolf-dogs should react strongly to wolves or other wild canids,
alerting AMHs and giving them extra cause to kill the intruders, in
addition to the usefulness of their hides and fur.
The sixth is that these wolf-dogs will be, on average, large-
bodied and capable of transporting bones and meat of large prey
to the campsite, if it is not the killsite. Ethnographic evidence
suggests that dogs trained as pack animals are capable of carrying
loads of up to 23 kg (Turner, 2002; Fiedel, 2005; Speth et al., 2013).
If wolf-dogs assisted in transporting meat, they would effect a
signicant savings in energy expenditure by humans.
The seventh is that, as at Predmost (Bocherens et al., 2013, this
volume), future studies of stable isotopes of wolf-dogs and wolves
fromthe same sites will reveal dietary differences between the two
probably because of provisioning by humans. I do not predict
whether most of these wolf-dogs will or will not bear evidence of
dismemberment, skinning, lleting, or burning, as aged wolf-dogs,
even if domesticated, may have been eaten.
5. Conclusions
The aim of this study was to clarify the taphonomic conditions
under which mammoth megasites were formed. Analysis of Hay-
nes age proles of modern models, based on assemblages of L.
africana with known causes of death, demonstrates that attritional
and catastrophic mortality produce statistically different age pro-
les, using the KolmogoroveSmirnov D statistic as a measure of
signicance. This afrmed the appropriateness of using such
models as a method for deducing taphonomic history of such sites.
Published age proles of mammoth megasites constructed us-
ing Haynes (1991; Conybeare and Haynes, 1984) ageing techniques
were used in this study. Megasites with fewer than 25 individual
mammoths were eliminated from this study because of the de-
mands of the KolmogoroveSmirnov statistic. Comparison of age
proles of eight mammoth megasites to Haynes models revealed
that all of these megasites were signicantly different fromHaynes
models. This nding suggests the possibility of problems with the
underlying assumptions of the ageing technique, of problems with
using modern analogies in paleontology, or that the sites repre-
sented palimpsests with mixed taphonomic histories.
Published age proles of three mammoth megasites constructed
using techniques developed by Saunders (1977) were compared to
Saunders model, which represents either an entire proboscidean
herd killed in a single episode or repeated killing of single maternal
groups over time. Both could be considered catastrophic causes of
death. Berelekh and Mezin do not differ signicantly from Saun-
ders model, suggesting a catastrophic mode of death occurred. The
P. Shipman / Quaternary International xxx (2014) 1e9 7
Please cite this article in press as: Shipman, P., How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites, Quaternary
International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048
mammoth megasite of Mezhirich differed signicantly from
Saunders model, suggesting an alternative taphonomic history
and/or a palimpsest.
The age proles of the mammoth megasites, as a group, do not
suggest a consistent taphonomic history. Comparison of mammoth
megasites to each other revealed a more interesting pattern.
Though each site differed signicantly from most other sites, a
chain of resemblances which mirrored temporal and spatial prox-
imity was found. This suggests that the sites may reect the
transmission of a technology or behavior important to the forma-
tion of such megasites through time and space. Shea and colleagues
(Shea, 2006, 2009; Shea and Sisk, 2010; Sisk and Shea, 2011) have
hypothesized that the development of complex projectile weapons
was responsible for the appearance of these megasites and for
human survival when Neanderthals went extinct.
I hypothesize that, in addition to complex projectile technology,
an alliance or collaboration between modern humans and domes-
ticated or partly-domesticated large canids best explains the
appearance of mammoth megasites, including the high numbers of
individual mammoths and canids at such sites and the close
proximity of sites to the place of mammoth death. The advantages
of projectile technology and canid collaboration may have led to the
survival of AMHs while Neanderthals went extinct. These canids
are explicitly identied with the morphologically unusual group
rst recognized by Germonpr et al. (2009) at about 32 ka B.P
(uncal) and more recently at additional sites in the Gravettian, and
Epigravettian (Germonpr et al., 2012, 2013, this volume). A
possible incipent dog has also been identied from Razboinichya
Cave, Siberia, and is dated to about 34 ka B.P. (uncal) (Ovodov et al.,
2011).
To the extent that further investigations have been conducted to
date, the group of canids or dog/wolves identied by the method-
ology of Germonpr et al. (2009) is also genetically and dietarily
distinctive from wolves (Thalmann et al., 2013; Bocherens et al.,
2013, this volume). Predictions of the domesticated canid hypoth-
esis for future research are presented.
Acknowledgments
I would like to acknowledge with gratitude Piotyr Wojtal and
Jaroslaw Wilczy nski for organizing the excellent conference The
World of Gravettian Hunters. I thank them and my other col-
leagues at the meeting for their input and helpful suggestions. I also
thank Jefferey Mathison jcmaps6@gmail.com for drawing the
beautiful map for this paper.
References
Antl, W., Fladerer, F.A., 2004. Outlook to the East: the 25 ka BP Gravettian Grub-
Kranawetberg Campsite (Lower Austria). In: Svoboda, J.A., Sedlckov, L.
(Eds.), The Gravettian Along the Danube, Proceedings of the Mikulov Confer-
ence, 20e21 November, 2002. Archeologick stav AV CR, Brno, pp. 116e130.
Arnold, C., 1979. Possible evidence of domestic dog in a Paleoeskimo context. Arctic
32 (3), 263e265.
Beresford-Jones, D., Johnson, K., Pullen, A., Pryor, A., Svoboda, J., Jones, M., 2010.
Burning wood or burning bone? A reconsideration of otation evidence from
Upper Palaeolithic (Gravettian) sites in the Moravian Gate, Czech Republic.
Journal of Archaeological Science 37, 2799e2811.
Bocherens, H., 2011. Diet and ecology of Neanderthals: implications from C and N
isotopes. In: Conard, N., Richter, J. (Eds.), Neanderthal Lifeways, Subsistence and
Technology: One Hundred Fifty Years of Neanderthal Study. Vertebrate Paleo-
biology and Paleoanthropology. Springer Science and Business Media, pp. 73e
85.
Bocherens, H., Drucker, D., 2006. Dietary competition between Neanderthals and
modern humans: insights from stable isotopes. In: Conard, N. (Ed.), When
Neanderthals and Modern Humans Met. Kerns, Verlag, pp. 129e143.
Bocherens, H., Drucker, D., Germonpr, M., Lznickov-Galetov, M., Bruzek, O.,
2013. Reconstruction of the Gravettian food-web at Predmost I using isotope
tracking of bone collagen. In: Wojtal, P. (Ed.), World of Gravettian Hunters, p. 5.
Krakow, Poland, abstract.
Bocherens, H., Drucker, D., Germonpr, M., Lznickov-Galetov, M., Bruzek, O.,
2014. Reconstruction of the Gravettian food-web at Predmost I using isotope
tracking of bone collagen. Quaternary International (this volume).
Bosch, M., 2012. Humanemammoth dynamics in the mid-Upper Palaeolithic of the
middle Danube region. Quaternary International 276e277, 170e182.
Bronk, F., Bronk Ramsey, C., Higham, T., 2007. Quality assurance of ultraltered bone
dating. Radiocarbon 49, 187e192.
Brugre, A., Fontana, L., 2009. Mammoth origin and exploitation patterns at Milo-
vice (Area G excepted). In: Oliva, M. (Ed.), Milovice: Site of the Mammoth
People below the Pavlov Hills. The Question of Mammoth Bone Structures, vol.
27(19). Moravsk Zemsk Muzeum, pp. 53e105.
Conard, N., 2003. Palaeolithic ivory sculptures from southwestern Germany and the
origins of gurative art. Nature 426, 830e832.
Conybeare, A., Haynes, G., 1984. Observations on elephant mortality and bones in
water holes. Quaternary Research 22, 189e200.
DeMay, L., Pan, S., Patou-Mathis, M., 2012. Mammoths used as food and building
resources by Neanderthals: zooarchaeological study applied to layer 4, Molo-
dova I (Ukraine). Quaternary International 276-277, 212e226.
Fiedel, S., 2005. Mans best friend and mammoths worst enemy? A speculative
essay on the role of dogs in Paleoindian colonization and megafaunal extinc-
tion. World Archaeology 37, 11e25.
Finlayson, C., 2009. The Humans Who Went Extinct. Oxford University Press,
Oxford.
Fladerer, F.A., 2003. A calf-dominated mammoth age prole from the 27 ky BP
stadial Krems-Wachtberg site in the middle Danube valley. In: Reumer, J.W.F.,
Vos, J., Mol, D. (Eds.), Advances in Mammoth Research (Proceedings of the
Second International Mammoth Conference, Rotterdam, May 16e20, 1999),
pp. 135e158.
Gargett, R., 2011. One Mammoth Steppe Too Far. The Subversive Archaeologist
(Blog). http://www.thesubversivearchaeologist.com/2011/12/one-mammoth-
steppe-too-far.html. December 21, 2011.
Geist, V., 2006. When do Wolves Become Dangerous to Humans?. http://www.
vargfakta.se/wp-content/uploads/2012/05/Geist-when-do-wolves-become-
dangerous-to-humans-pt-1.pdf.
Germonpr, M., Rikknen, J., Lznickov-Galetov, M., Sablin, M., 2014. Mandibles
from Palaeolithic dogs and Pleistocene wolves at Predmost, the Czech Republic.
Quaternary International (this volume).
Germonpr, M., Rikknen, J., Lznickov-Galetov, M., Sablin, M., 2013. Mandibles
from Palaeolithic dogs and Pleistocene wolves at Predmost, the Czech Republic.
In: Wojtal, P. (Ed.), The World of Gravettian Hunters, Abstract.
Germonpr, M., Lznickov-Galetov, M., Sablin, M., 2012. Palaeolithic dog skulls at
the Gravettian Predmost site, the Czech Republic. Journal of Archaeological
Science 39, 184e202.
Germonpr, M., Sablin, M.V., Stevens, R.E., Hedges, R.E.M., Hofreiter, M., Stiller, M.,
Despres, V.R., 2009. Fossil dogs and wolves from Palaeolithic sites in Belgium,
the Ukraine, and Russia: osteometry, ancient DNA and stable isotopes. Journal
of Archaeological Science 36, 473e490.
Germonpr, M., Sablin, M., Khlopachev, G., Grigorieva, G., 2008. Possible evidence of
mammoth hunting during the Epigravettian at Yudinovo, Russian Plain. Journal
of Anthropological Archaeology 27, 475e492.
Haynes, G., 1991. Mammoths, Mastodons and Elephants; Biology, Behavior, and the
Fossil Record. Cambridge University Press, Cambridge.
Higham, T., Basell, L., Jacobi, R., Wood, R., Bronk, F., Bronk Ramsey, C., Conard, N.,
2012. Testing models for the beginnings of the Aurignacian and the advent of
gurative art and music: the radiocarbon chronology of Geienklsterle. Jour-
nal of Human Evolution 62, 664e676.
Iakovleva, L., Djindjian, F., 2005. New data on Mammoth bone settlements of
Eastern Europe in the light of the new excavations of the Gontsy site (Ukraine).
Quaternary International 126e128, 195e207.
Klein, R., 1982. Age (mortality) proles as a means of distinguishing hunted species
from scavenged ones in Stone Age archaeological sites. Paleobiology 8, 151e
158.
Klein, R., Cruz-Uribe, K., 1984. The Analysis of Animal Bones from Archeological
Sites. University of Chicago Press, Chicago.
Klima, B., 1954. Palaeolithic huts at Doln Vestonice, Czechoslovakia. Antiquity 109,
4e14.
Klima, B., 1963. Doln Vestonice: Vyzkum Taboriste Lovcu Mamutu v Letech 1947e
1952. Nakladatelsv Cekeslovensk Akademie Ved, Prague.
Koster, J., 2009. Hunting dogs in the Lowland Neotropics. Journal of Anthropological
Research 65, 575e610.
Koster, J., Tankersley, K., 2012. Heterogeneity of hunting ability and nutritional
status among domestic dogs in lowland Nicaragua. Proceedings of the National
Academy of Sciences 109 (8), 463e470.
Koz1owski, J., Kubibak, H., Sachse-Koz1owska, E., van Vliet, B., Zakrzewska, G., 1974.
Upper Palaeolithic site with dwellings of mammoth bones, Krakw Spadzista
Street B. Folia Quaternaria 36, 1.
Laws, R., 1966. Age criteria for the African elephant, Loxodonta a. africana. East
African Wildlife Journal IV, 5e13.
Laws, R., Parker, I., 1968. Recent studies on elephant populations in East Africa.
Symposium of the Zoological Society of London 20, 1e59.
Lombard, M., Phillipson, L., 2008. Indications of bow and stone-tipped arrow use
64,000 years ago in KwaZulu-Natal, South Africa. Antiquity 84 (325), 635e
648.
Lupo, K., 2011. A dog is for hunting. In: Albarella, U., Trentacoste, A. (Eds.), Ethno-
zooarchaeology. Oxbow Press, Oxford, pp. 4e12.
P. Shipman / Quaternary International xxx (2014) 1e9 8
Please cite this article in press as: Shipman, P., How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites, Quaternary
International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048
Marquer, L., Lebretona, V., Otto, T., Valladas, H., Haesaerts, P., Messager, E.,
Nuzhnyif, D., Pan, S., 2012. Charcoal scarcity in Epigravettian settlements with
mammoth bone dwellings: the taphonomic evidence from Mezhyrich
(Ukraine). Journal of Archaeological Science 39, 109e120.
Mellars, P., French, J., 2011. Tenfold population increase in Western Europe at the
NeandertaletoeModern Human transition. Science 333, 623e627.
Ovodov, N., Crockford, S., Kuzmin, Y., Higham, T., Hodgins, G., 2011. A 33,000-year-
old incipient dog from the Altai Mountains of Siberia: evidence of the earliest
domestication disrupted by the Last Glacial Maximum. PLoS ONE 6 (7), 1e7.
Patou-Mathis, M., 2000. Neanderthal subsistence behaviours in Europe. Interna-
tional Journal of Osteoarchaeology 10, 379e395.
Ruusila, V., Pesonen, M., 2004. Interspecic cooperation in human (Homo sapiens)
hunting: the benet of a barking dog (Canis familiaris). Annales Zoologici Fen-
nici 41, 545e549.
Saunders, J., 1977. A model for man-mammoth relationships in Late Pleistocene
North America. Canadian Journal of Anthropology 1 (1), 87e98.
Shea, J., 2009. The impact of projectile weaponry on late Pleistocene hominin
evolution. In: Hublin, J., Richards, M. (Eds.), The Evolution of Hominin Diets.
Springer, New York, pp. 189e199.
Shea, J., Sisk, M., 2010. Projectile technology and Homo sapiens dispersal from Africa
to Western Eurasia. PaleoAnthropology 2010, 100e122.
Sisk, M., Shea, J., 2011. The African origin of complex projectile technology: an
analysis using tip cross-sectional area and perimeter. International Journal of
Evolutionary Biology, 968012.
Soffer, O., 1985. The Upper Paleolithic of the Central RussianPlain. Academic Press, N.Y.
Soffer, O., 2003. Mammoth bone accumulations: death sites? kill sites? dwellings?
In: Vasilev, S., Soffer, O., Koz1owski, J. (Eds.), Perceived Landscapes and Built
Environments: The Cultural Geography of Late Paleolithic Eurasia. Archaeo-
press, Oxford, pp. 39e46.
Soffer, O., Adovasio, J., Kornietz, N., Velichko, A., Gribchenko, Y., Lenz, B., Suntsov, V.,
1997. Cultural stratigraphy at Mezhirich, an Upper Palaeolithic site in Ukraine
with multiple occupations. Antiquity 71, 48e62.
Speth, J., Newlander, K., White, A., Lemke, A., Anderson, L., 2013. Early Paleoindian
big-game hunting in North America: provisioning or Politics? Quaternary In-
ternational 285, 111e139.
Stewart, J., 2004. Neanderthal-Modern Human competition? A comparison be-
tween the mammals associated with Middle and Upper Palaeolithic Industries
in Europe during OIS 3. International Journal of Osteoarchaeology 14, 178e189.
Stiner, M., 1991. Human Predators and Prey Mortality. Westview Press, Boulder,
Colorado.
Svoboda, J., Pan, S., Wojtal, P., 2005. Mammoth bone deposits and subsistence
practices during Mid-Upper Palaeolithic in Central Europe: three cases from
Moravia and Poland. Quaternary International 126e128, 209e221.
Thalmann, O., Shapiro, B., Cui, P., Schuenemann, V., Sawyer, S., Greeneld, D.,
Germonpr, M., Sablin, M., Lpez-Girldez, F., Domingo-Roura, X., Napierala, H.,
Uerpmann, H.-P., Loponte, D., Acosta, A., Giemsch, L., Schmitz, R.,
Worthington, B., Buikstra, J., Druzhkova, A., Graphodatsky, A., Ovodov, N.,
Wahlberg, N., Freedman, A., Schweizer, R., Koepi, K.-P., Leonard, J., Meyer, M.,
Krause, J., Pbo, S., Green, R., Wayne, R., 2013. Complete mitochondrial ge-
nomes of ancient canids suggest a European origin of domestic dogs. Science
342, 871e874.
Turner, C., 2002. Teeth, needles, dogs and Siberia: bioarchaeological evidence for
the colonization of the New World. In: Jablonski, N. (Ed.), The First Americans:
The Pleistocene Colonization of the New World. California Academy of Sciences,
San Francisco, pp. 123e158.
Vanak, A.T., Gompper, M., 2009. Dogs as carnivores: their role and function in
intraguild competition. Mammal Review 39, 265e283.
Vereschagin, N., Baryshnikov, G., 1984. Quaternary mammalian extinctions
in northern Eurasia. In: Martin, P., Klein, R. (Eds.), Quaternary Extinc-
tions: A Prehistoric Revolution. University of Arizona Press, Tucson,
pp. 483e516.
Voorhies, M.R., 1969. Taphonomy and Population Dynamics of an Early Pliocene
Vertebrate Fauna, Knox County, Nebraska. University of Wyoming. Contribu-
tions to Geology, Special Paper no. 1, 69 pp.
Ward, P., 1998. The Call of Distant Mammoths: With the Ice Age Mammals Dis-
appeared. Springer-Verlag, New York.
Wilczy nski, J., Wojtal, P., Sobczyk, K., 2012. Spatial organization of the Gravettian
mammoth hunters site at Krakw Spadzista (southern Poland). Journal of
Archaeological Science 39, 3627e3642.
Wojtal, P., Wilczy nski, J., Boche nski, Z., Svoboda, J., 2012. The scene of spectacular
feasts: animal remains from Pavlov I south-east, the Czech Republic. Quaternary
International 252, 122e141.
Wojtal, P., 2007. Zooarchaeological Studies of the Late Pleistocene Sites in Poland.
Institute of Systematics and Evolution of Animals. Polish Academy of Sciences,
Poland.
Wojtal, P., Sobczyk, K., 2003. Taphonomy of the Gravettian site - Krakw Spadzista
Street (B). In: Reumer, J., de Vos, J., Mol, D. (Eds.), Advances in Mammoth
Research (Proceedings of the Second International Mammoth Conference),
Rotterdam, The Netherlands, pp. 557e561.
Wojtal, P., Sobczyk, K., 2005. Man and woolly mammoth at the Krakw Spadzista
Street (B); taphonomy of the site. Journal of Archaeological Science 32 (2), 193e
206.
Wood, R., Barroso-Ruz, C., Caparrs, M., Jord Pardo, J., Galvn Santos, B.,
Higham, T., 2013. Radiocarbon dating casts doubt on the late chronology of the
Middle to Upper Palaeolithic transition in southern Iberia. Proceedings of the
National Academy of Science USA 110 (8), 2781e2786.
P. Shipman / Quaternary International xxx (2014) 1e9 9
Please cite this article in press as: Shipman, P., How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites, Quaternary
International (2014), http://dx.doi.org/10.1016/j.quaint.2014.04.048