how do you kill 86 mammoths

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Quaternary International xxx (2014) 1e9

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Quaternary International

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How do you kill 86 mammoths? Taphonomic investigations ofmammoth 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:MammothsDogsWolvesHuntingAge profilesTaphonomy

* Permanent address: 3140 Chatham Church Road,E-mail addresses: [email protected], pls10

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a b s t r a c t

A series of Eurasian archaeological sites formed between about 40 e 15 ka feature unusually largenumbers of mammoth remains with abundant artefacts and, often, mammoth bone dwellings. None ofthese mammoth megasites is dated prior to the appearance of modern humans in Eurasia. This unusualtype of site begs for taphonomic explanation. The large number of individual mammoths and the scarcityof carnivore toothmarks and gnawing suggest a new ability to retain kill mammoths and control ofcarcasses. Age profiles of such mammoth-dominated sites with a large minimum number of individualsdiffer statistically at the p < 0.01 level from age profiles of Loxodonta africana populations that died ofeither attritional or catastrophic causes. However, age profiles from some mammoth sites exhibit a chainof linked resemblances with each other through time and space, suggesting the transmission ofbehavioral or technological innovation. I hypothesize that this innovation may have been facilitated by anearly attempted domestication of dogs, as indicated by a group of genetically and morphologicallydistinct large canids which first appear in archaeological sites at about 32 ka B.P. (uncal). Testable pre-dictions of this hypothesis are generated based on ethnographic data.

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1. Introduction

Since the late nineteenth century, archaeological sites domi-nated by mammoth remains have been an important focus ofresearch. A key question is how the bones of large numbers ofmammoths, ranging from a minimum number of individuals (MNI)of about five to hundreds, were deposited in one place. Despitemuch investigation, the cause of death of mammoths in these siteshas remained controversial (e.g., Klima, 1954, 1963; Koz1owskiet al., 1974; Klein, 1982; Vereschagin and Baryshnikov, 1984;Soffer, 1985, 2003; Haynes, 1991; Fladerer, 2003; Wojtal andSobczyk, 2003; Svoboda et al., 2005; Wojtal, 2007; Brugère andFontana, 2009; Wojtal et al., 2012). Two dominant hypotheses arethat these sites represent natural deaths which were scavenged byhumans; or that specialized human hunting resulted in such as-semblages. Here, for convenience, I refer to such sites as mammothmegasites. An argument could be made that large sites heavilydominated by other particular species should also be termed“megasites.”

Moncure, NC 27559, [email protected].

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Many ambiguities and inconsistencies in excavation andcollection techniques used by different scholars have complicatedattempts to construct an overall consensus on the purpose, for-mation, taphonomy, and use of these megasites, which span a largegeographic and temporal range. Questions about the accuracy ofradiocarbon dates on many mammoth megasites are unresolvedbecause recent advances in decontamination, ultrafiltration, andthe use of strict criteria to eliminate samples with poor preserva-tion show that many dates in the literature calculated decades agoare 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 scholarsto attempt to understand these sites.

Fortunately, many recent researchers have compiled basic in-formation from zooarchaeological analyses of the sites, includingnumber of non-mammoth species represented, mammoth MNIs,mammoth age at death, and mammoth age profiles from individualsites. Demographic studies have gained in importance as indicatorsof 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 profiles of fossilassemblages to those of animals of known causes of death. Inparticular, profiles of the age at death of mammoths represented atvarious sites have been published and used to formulate

ths? Taphonomic investigations of mammoth megasites, Quaternary

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P. Shipman / Quaternary International xxx (2014) 1e92

interpretations about site formation processes. In an attempt tocontribute to progress in deducing the taphonomic histories ofsites, I have used these published age profiles for further analysis.

The researchers who have published the age profiles on thesesites have also commented on additional indicators of taphonomichistory, such as cutmarks, carnivore toothmarks, evidence ofburning, and skeletal representation. Virtually all sites discussedhere include lithic remains from the Gravettian or Epigravettianindustries, with the exception of Molodova I, level 4, which is aMousterian site. No lithic assemblage includes novel tool types thathave been suggested to be designed for killing mammoths. Bonesshow scarce signs of carnivore activity and human modification(usually <5% of the mammoth bones have such evidence). Thehuman modifications take the form of cutmarks, filleting marks,skinning marks, disarticulation marks, fracturing of long bones formarrow, and/or shaping of bones as tools. Some sites also includearchitectural structures such as mammoth bone huts, refuse dumpsor pits, postholes, and hearths. There is no evidence of pit traps ordrive lanes.

1.1. Regional, temporal, and geographic distribution of mammothmegasites

Dozens of mammoth-dominated sites are located in central andeastern Europe, with concentrations in the Swabian Jura, theDanube area and Moravian Gate, and up into Russia and Siberia.Typically the sites are located on high river terraces or lowmountainous areas in close proximity to water. Mid-slope locationswith easy access to rivers are common. Commonly the remains areburied in loess alternating with sandy clayey layers, but geologicaldetails can be found in individual publications. If the abundance ofmammoth remains is related to natural disasters such as a herd’sfalling through the ice on a frozen river or falling into ravines whileaccessing salt or mineral licks, then the bones of those mammothshave, 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 theformation of such sites is often postulated: only two mammothmegasites 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 atvery high densities. For example, mammoths comprise 99% of thefauna (Number of Individual Specimens ¼ 5860) at Kraków Spad-zista Street (Wojtal and Sobczyk, 2005). In Trench Bþ B1 at this site,there was 1 individual mammoth for every 2 m2 of excavation(Wilczy�nski et al., 2012:3638). This is comparable to the density ofmammoth individuals at Yudinovo, an Epigravettian site, in whichthe pavilion area yielded 1 mammoth individual for every 1.4 m2

(Germonpré et al., 2008: 481). Such densities contrast sharply withthose 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 mammaliandatabase, which includes 447 dated faunas from 291 archaeologicaland non-archaeological sites, yields a similar result. The databasecontains three times as many Upper Paleolithic sites as MiddlePaleolithic ones, but an Upper Paleolithic preference formammothsis shown by the presence of mammoths in 4.5 times as many UpperPaleolithic sites as Middle Paleolithic ones (Stewart, 2004). Stewartinterprets this as indicating a more systematic exploitation ofmammoths by Upper Paleolithic hominins.

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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 themost common prey e horse e reindeer, rhinoceros, and bovineswere in the Neanderthal diet. Isotopic studies reveal that Nean-derthals ate larger quantities of mammoth than zooarchaeologicalstudies reveal, probably because Neanderthals were reluctant toeither camp near the carcass or to transport heavy flesh-bearingmammoth bones from the kill site to camp (Bocherens, 2011: 80e81). In contrast, Upper Paleolithic sites are also much more likely tocontain all parts of the mammoth skeleton (e.g. Wojtal andSobczyk, 2005; Brugère and Fontana, 2009; Bosch, 2012).

Mammoths represented a wealth of resources which weredemonstrably used in the Upper Paleolithic. Mammoths were eaten,judging from clear cutmarks on some bones at sites such as Grub-Kranawetberg, Kraków Spadzista Street (B), Krems-Hunddsteig,Krems-Wachtberg, and Langmannersdorf-Lagerplatz B (summa-rized in Bosch, 2012). Stable isotope studies confirm this interpre-tation (Bocherens, 2011; Bocherens et al., 2013, this volume).Mammoth bone and ivory were used to make tools and sculpturesor art objects (Conard, 2003; Antl and Fladerer, 2004; Wojtal et al.,2012). Bone and tusks were also used as construction materials andmany Gravettian or Epigravettian mammoth megasites feature re-mains of dwellings carefully built of mammoth bone and sometimessupported by tusks. The frameworks were apparently covered inmammoth hides, often with one or more hearths inside (Soffer,1985). At least 70 mammoth bone huts are known from theRussian Plain alone (Ward, 1998), but mammoth bone dwellings arenot confined to this region (Iakovleva and Djindjian, 2005; Svobodaet al., 2005). Only one Mousterian site, Molodova I, level 4 containsevidence 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 alsooffers useful information. Sites where all skeletal elements are pre-sent (such as Krems-Hundssteig, Kraków Spadzista Street, Lang-mannersdorf,Milovice I) suggest that themammoth(s)were killed ator very near the site. Sites with only selected skeletal elements ofmammoths (such as Mezhirich, Yudinovo, and Berelekh) are moresuggestive 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) fromnatural deaths, citing of a lack of available wood in the mammothsteppe environment. The frequent finding of burned mammothbones (e.g., Soffer et al., 1997) shows bones were also used as fuelfor fire, another practice that has been related to a lack of wood.However, recent studies of microscopic charcoal remains at severalsites suggest that birch and willow were more numerous alongriver 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, particularlyArctic fox (Alopex lagopus) and wolf (Canis lupus), are unusuallywell represented by many bones and high MNIs. Recent morpho-logical and genetic studies (Germonpré et al., 2009, 2012, 2013, thisvolume; Thalmann et al., 2013) show that some of the large canidsfrom mammoth megasites are a distinctive group which mayrepresent an early attempt at the domestication of the dog insteadof wolves. Some of the long bones of large canids, whether wolf ordog, show cutmarks indicative of dismemberment, skinning, andfilleting (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 andage profiles from fifteen individual sites (Table 1). Profiles of age at


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P. Shipman / Quaternary International xxx (2014) 1e9 3

death can be used to separate scavenged remains from hunted ones(Klein, 1982; Klein and Cruz-Uribe, 1984; Stiner, 1991). Such infor-mation would clarify and possibly refute hypotheses about theformation of these mammoth megasites. Although age profiles areoften reported in the literature, they are not usually evaluatedstatistically. A major aim of the research reported here was toconduct such statistical comparisons among age profiles from fossilsites and those from modern proboscideans with known causes ofdeath.

Table 1Mammoth megasites with published information on ages at death of individuals.

Site age (rC14 yrs) MNI mammoths

Molodova, Ukraine > 44,000 15Vogelherd, Germany 33,000e30,000 15Krems-Wachtberg, Austria 27,000 8Milovice G, Czech Republic 26,000e24,000 21Milovice I, Czech Republic 22,000 51Krems-Hundssteig, Austria 28,000 7Pavlov I, Czech Republic 27,000e25,000 7P�redmostí, Czech Republic 26,000 105Grub-Kranawetburg, Austria 25,000 8Mezin, Ukraine 25,000 e 24,000 44Kraków Spadzista Street, Poland 24,000e23,000 86Langmannersdorf, Austria 22,000 e 20,500 27Langmannersdorf e Lagerplatz, Austria 21,000 7Yudinovo, Russian 16,000e12,000 35Mezhirich, Ukraine 15,245 109Berelekh, Siberia 13,400e10,400 166

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Berelekh Mezin Mezhirich

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Fig. 1. a. Age profile of Loxodonta africana from Saunders (1977). This represents aliving herd of maternal groups. b. Age profiles of mammoths from three sites. Ageclasses were assigned according to the system developed by Saunders (1977). Themammoth age profile from Bereleyk does not differ from the living herd profile(compare Fig. 1a and b: KolmogoroveSmirnov D statistic ¼ 0.8, p > 0.01) or from theage profile at Mezin (D ¼ 0.6, p > 0.01). The profile from Mezin does not differsignificantly from the living herd profile (D ¼ 0.64, p > 0.001). The Berelekh profilediffers statistically from that at Mezhirich (D ¼ 2.52, p < 0.001). The Mezhirich profilediffers from the living herd profile (D ¼ 1.89, p < 0.01) and from the profile at Mezin(D ¼ 1.98, p < 0.001). The Berelekh and Mezin age profiles therefore suggest intactherds, or repeated, non-selective hunting, while the Mezhirich profile may be apalimpsest of different events.

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 tootheruption and wear (Fig. 1a and b). Saunders based his system onwork by Laws (1966) and Laws and Parker (1968) with Loxodontaafricana in Murchison Falls National Park, Uganda. Laws and Parker(1968) reported the demographics of family groups, as revealedduring a culling operation. An age in African Elephant Years (AEY)was assigned to each individual and individuals were then groupedinto four age classes each consisting of ten years, with all olderadults being grouped into the last category. The resulting ageprofile serves as a model for either an intact, living elephant herdconsisting primarily of maternal or nursery groups, and the ageprofile of such a herd if struck down by either catastrophic orattritional causes that were not selective as to the age of the indi-vidual animal. Saunders used this same procedure for ageingmammoths and mastodons, constructing an age profile, andinterpreting the results.

In addition to representing attritional or catastrophic modes ofdeath, analysts often judge age profiles as either U-shaped, with thehighest 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 methodof assigning ages at death in AEY, based on eruption and weardental remains, as favored by Haynes (1991; Conybeare andHaynes, 1984: Fig. 2). However, individuals were clumped intotwelve-year age classes, not ten-year age classes as Saunders haddone. This twelve-year system was developed by Haynes (1991),based on unpublished work with L. africana by G. Craig anddescriptive studies of elephant die-offs of known cause inZimbabwe (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 inZimbabwe, in which entire maternal herds were shot at one time.Because the aging methods are well-known and well-published, Iwill not repeat procedural details for age determinations here.

2.1. Statistical analyses

Haynes’ reference data (1991) are widely used in the literature,so I first conducted statistical comparisons of his two datasets.Those derived from 291 attritional deaths in a prolonged droughtwere 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, followingKlein and Cruz-Uribe (1984). Values of D � 1.64 indicate a signifi-cant difference between two samples at the p < 0.01 level. TheKolmogoroveSmirnov test is appropriate because it is sensitive to


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Fig. 2. a. Age profiles of two populations of Loxodonta africana aged according toHaynes (1991). One represents a population killed in a drought of several years’duration and the other, maternal herds killed in culling operations. The drought-killedanimals are disproportionately often very young, while the culled animals show ahigher proportion of middle-age or prime individuals. The drought profile differsstatistically from the culled profile (KolmogoroveSmirnov D ¼ 2.4, p > 0.001). b. Ageprofiles of five different mammoth populations aged according to Haynes (1991). Theage profiles from these sites all differ statistically from either the drought or the culledmodel proposed by Haynes (1991) at the p > 0.01 level. The age profiles of the sites arearranged according to a pattern of resemblances of age profiles. Each site’s profilediffers significantly from all others at the p > 0.01 level with the exception of the nextadjacent site, reading left to right, on the x axis. P�redmostí differs from all sites exceptKraów-Spadzista (D ¼ 0.128, p > 0.10). Kraków Spadzista Street differs from all siteswith the except of P�redmostí and Langmannersdorf (D ¼ 0.128, p > 0.10). Langman-nersdorf differs from all sites except Kraków Spadzista Streetand two others: Milovice I(D ¼ 0.41, p > 0.10) and Yudinovo (D ¼ 0.54, p > 0.10).

Fig. 3. The statistical comparisons of the mammoth age profiles from all sites classifiedaccording to Haynes’ system reveal an interesting pattern of resemblances. The top boxshows (with double-headed arrows) the age profiles 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 awest to east direction. Langmannersdorf and Milovice I may not differ meaningfully inage. This pattern may represent the transmission of a mammoth-killing behavior thatchanges across time and space.


rank ordering of data (i.e., changes in the number of individuals inone age class will affect the others).

My aimwas to compare the age profile of each fossil megasite toeach of Haynes’ datasets statistically as a means of gleaning addi-tional data relevant to deducing the cause of death or accumulationof the mammoths. Because large sample sizes are required by theKolmogoroveSmirnov D statistic, Klein and Cruz-Uribe (1984:57,59) suggest at least 25 individuals are necessary for a valid inter-pretation of an age profile and that “for maximum reliability” eachsample ought to contain at least 40 individuals. I chose to discardfossil age profiles involving fewer than the minimum of 25 in-dividuals required for reliable statistical evaluation. Only eightfossil sites included enough aged individuals to analyze further(Table 1, Fig. 3): Berelekh, Siberia; Kraków Spadzista Street, Poland;Langmannersdorf, Austria; Mezhirich, Ukraine; Mezin, Ukraine;Milovice I, Czech Republic; P�redmostí, Czech Republic; and Yudi-novo, Russia. The published MNI values of mammoth specimensthat are classified into age categories ranged from 27 to 105.


The interpretation of age profiles as an indicator of mortalitypattern may have special advantages in analyzing extremely largeand long-lived species. Because proboscidean teeth (and bones) areso large, there is a much lower possibility of the destruction of teethof very young individuals due to their fragility when exposed tonatural (i.e., non-hominin) taphonomic forces. Even the teeth ofyoung proboscideans are robust and more likely to be preserved ascompared to those of smaller mammals. Fetal or very youngmammoth remains were identified from Kraków Spadzista Street,Yudinovo, P�redmostí, Langmannersdorf, and Krems-Wachtberg, forexample. Further, despite the fact that the age classes of differentsites were most often assigned by different researchers, the po-tential for errors in assigning age class is lessened because the ageclasses are so broad. In contrast, in analyzing the remains of specieswith annual births that reach sexual maturity at 2 years, a yearlingcould possibly be mistaken for a nearly-mature individual. Thepossibility of a researcher mistaking a young proboscidean for onethat is 10e12 years old is much less. Thus, potentially, analyzing ageprofiles of mammoths may be more reliable and informative thananalyzing age profiles of some other species. The analytical proce-dure is detailed below.

Age profiles of the three mammoth megasites aged by the Laws/Parker method were compared with age profiles from Saunders’model, which represents a catastrophic death of family herds, usingthe KolmogoroveSmirnov test to determine significance. Finally,each age profile of a mammoth megasite was then compared sta-tistically to every other one which had been classified by the sameageing method.

3. Results

Although many publications have compared mammoth ageprofiles to those published for modern elephants by Haynes (1991),it had not been established previously that Haynes’ two modelsdiffer at a statistically significant level. This evaluation confirmsthat the age profiles 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 thep < 0.01 level. Thus, the basic premise that these two age profilesdiffer meaningfully and may be helpful in determining a pattern of



death is supported. Haynes’ model samples consisted of 149 (cull-ing) and 296 (drought) individuals each.

All mammoth megasites aged using Haynes’ system e

P�redmostí, Kraków Spadzista Street, Yudinovo, Langmannersdorf,and Milovice I e differed significantly from both Haynes’ attritionalmodel and his catastrophic model at the p< 0.01 level in each case.Two mammoth sites evaluated against Saunders’ model, Berelekhand Mezin, do not differ significantly from Saunders’ model andthus represent an intact elephant herd, repeated killing of familygroups, or one entire herd killed catastrophically. Mezhirich, thethird site with a published age profile assigned according toSaunders’ procedures, differs significantly from Saunders’ modelwith a D value of 2.52 (p < 0.01). Saunders’ model sample consistsof 135 individuals.

The site-by-site comparison of Paleolithic age profiles to eachother yielded more promising results. Although most site-specific age profiles did not resemble each other closely, thepattern of resemblances and differences suggested wasinteresting.

The P�redmostí age profile differed significantly (at the p < 0.01level) from that at any other site, except for the age profile fromKraków Spadzista Street. The age profile from Kraków SpadzistaStreet differed from that at every other site except those fromP�redmostí and Langmannersdorf. The Langmannersdorf age profile,as expected, did not differ from the profile from Kraków SpadzistaStreet, nor from the profiles from Yudinovo or Milovice I. The ageprofiles from Yudinovo and Milovice I differed significantly from allothers at the p < 0.01 level with the exception of the profile fromLangmannersdorf, from which each was insignificantly different.

4. Discussion and overview

As Haynes’ attritional and catastrophic models differ from eachother significantly at the p < 0.01 level, the results reported herevalidate the notion of using elephant age profiles from populationswith known causes of death as a means of interpreting mammothage profiles. Statistical analysis of all the mammoth megasites agedusing Haynes’ criteria differ significantly from either model, sug-gesting that many of these sites may be palimpsests accumulatedover time with a mixture of taphonomic histories, rather thansingle events. This finding suggests that attempts to judge re-semblances or differences among age profiles intuitively may notbe productive.

Two mammoth megasites, Berelekh and Mezin, do not differfrom Saunders’ model at the p < 0.01 level. This finding indicatesthat these assemblagesmay represent either amassive catastrophe,which killed an entire elephant herd at once, or a repeated series ofkillings of nursery herds. The third mammoth megasite aged bySaunders’ criteria, Mezhirich, is significantly different from Saun-ders’ model at the p < 0.01 level, suggesting a possible palimpsest.

The chain of resemblances e or of insignificant differences e

produced by the site-by-site comparisons reproduces the approx-imate chronological order of these sites, as judged from bestavailable 14C dates (Table 1). The chain of resemblances also con-nects sites to other sites in an east-west sequence, going from theoldest to the youngest site (Fig. 4).

4.1. Taphonomy and possible explanations

Most mammoth megasites evaluated here do not conform tomodels based on living elephants which experienced either attri-tional or catastrophic deaths. Five possibilities might explain thisoutcome.


1) Possibly mammoth dental eruption and wear does not closelyparallel that in Loxodonta, as has been assumed by both ageingtechniques. Given the relatively close phylogenetic relationshipbetween Mammuthus and Loxodonta, and the detailed re-semblances in craniodental anatomy, this seems unlikely.

2) The criteria used for excavation, collection, and ageing ofthese assemblages may have been inadequate to preserveresemblances in age class profiles which were actually pre-sent at the time of site formation. Though some of the siteswere excavated and collected decades ago, and standardshave certainly changed, most or all of these assemblages havebeen reanalyzed in recent times by skilled researchers. If thecollected dental remains accurately represent what waspresent at the site at the time of excavation, it seems likelythat minimal taphonomic biases were introduced by theprocess of collection per se. If collection standards werebiased toward more complete or more attractive specimens,most zooarchaeological analyses will not prove helpful andmodern analysts will be unable to derive useful informationabout these sites.

3) Most of these assemblages may represent a palimpsest ofdifferent events which has weakened or destroyed the originaltaphonomic signature of the age profiles, to the point that sta-tistical significance is not achieved in these comparisons. Thisfinding would point to repeated reoccupation of these sites,perhaps seasonally, with variations in subsistence strategy be-tween occupations. This potential explanation is entirelypossible and cannot be discounted, especially given the timespan represented by these sites and their broad geographicdistribution.

4) Another potential explanation is that generalizing from modernproboscidean behavior to ancient behaviors of different species,across vast distances of space and time, must be viewed withextreme caution and may be invalid. There is no obvious way toevaluate the likelihood that this explanation is true, althoughusing such phylogenetic and behavioral analogues is almoststandard in the field.

5) A final explanation is that the chain of resemblances linkingeach mammoth site to the next closest one in time and spacemay reflect the transmission of a specialized but changingbehavior or technology through time and space.

The fifth possible explanation is the most encouraging for pa-leoanthropologists and taphonomists hoping to understand thepast. The sites in this analysis are spread acrossmost of the Eurasiancontinent and throughout a substantial span of time. Perhapsexpecting consistency in detailed patterns of cultural activities,such as hunting and butchering, across thousands of years andthousands of miles is unrealistic.

The introduction or growing use of complex projectile weap-onry by modern humans was probably important in producing theabrupt changes in assemblages associated with hominins seenstarting about 45 ka. As Shea (2008:177) has observed, complexprojectile weaponry is a “niche-broadening technology” becausesuch weapons are light, easily carried, and “they retain energylonger in flight, allowing them to be used against larger, dangerousprey, or other humans, with less risk of injury”, enabling earlymodern humans to broaden their ecological niche(s) (see alsoLombard and Phillipson, 2008). The resultant transformation frombeing an ambush predator, as Neanderthals were (Finlayson, 2009),to being long-distance hunters may have conferred considerablebenefits on early modern humans, particularly in killing very largeand dangerous mammals.

I suggest here that a second advance which occurred during MIS3 may have enhanced the advantages of improvements in lithic


Fig. 4. Eight mammoth megasites had a sufficient number of individuals classified into age classes (N > 25) to be analyzed statistically. They vary in antiquity from about 22,000 to10,400 years according to uncalibrated radiocarbon dates. However, recent advances in techniques for decontamination and ultrafiltration have not been applied to these sites sotheir dates may be subject to correction when this is done. Map by Jeffery Mathison.


technology. There would have been great advantages to the pres-ence of even quasi-domesticated large canids willing to workcooperatively with AMHs. A group of such canids that is morpho-logically distinct from wolves and postulated to be a domesticated


dog has been identified by Germonpré et al., 2009, 2012, 2013, thisvolume). Morphologically, members of this group have also beenidentified at P�redmostí, Mezin, and Mezhirich (Germonpré et al.,2009, 2012, 2013, this volume).



4.2. The domesticated canid hypothesis

In additional to the use of complex projectile technology thatfacilitated 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 ofthe large, complex, mammoth megasites in the Upper Paleolithic.For convenience, I call these canids wolf-dogs here because it isuncertain whether or not they were directly ancestral to any livingcanid.

I specifically identify these canids with the morphological groupfirst identified by Germonpré and colleagues at Goyet Cave.Belgium. I specifically do not expect Upper Paleolithic wolf-dogs tobe the same in all respects as modern dogs. It would be unusual iffirst known domestication or domestication attempt of any speciesshowed all the features of later domesticates.

The Belgian morphological group of wolf-dogs is further setapart by recent genetic work studies on three individuals, includingone clear morphological wolf-dog from Goyet Cave. These in-dividuals possess an unusual mtDNA haplotype unknown amongmodern dogs, modern wolves, or ancient wolves (Thalmann et al.,2013). This finding shows that this maternal lineage or mtDNAhaplotype 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 inwhichmales withthis haplotype interbred with female wolves, giving rise to a pop-ulation ancestral to either modern dogs or wolves. As Thalmannet al. (2013) remark, the genetic data are also consistent with aninterpretation of these unusual canids (here, wolf-dogs) as an earlyabortive attempt at domestication, which left few if any directdescendants, or as a Pleistocene wolf population that also becameextinct without issue.

4.3. Predictions of the domesticated canid hypothesis

I use ethnographic analogies, particularly studies of huntersoperating with and without the assistance of dogs (Ruusila andPesonen, 2004; Koster, 2009; Lupo, 2011; Koster and Tankersley,2012) to generate testable predictions of the domesticated canidhypothesis.

The first prediction is that additional examples of the unusuallarge canids identified by morphometric and genetic techniqueswill be found in mammoth megasites and will not be found inMiddle Paleolithic sites, Mousterian, or pre-AMH sites.

The second is that additional mammoth megasites will possessspecimens of such wolf-dogs. I do not predict all such sites willyield such specimens due to differences in taphonomy andpreservation.

The third is that sites formed by hominins with wolf-dogs willyield evidence of a larger number of kills and more meat yield thansites formed by hominins without wolf-dogs. Ethnographically,hunters with dogs have a markedly increased meat yield per huntand a decreased time of seeking prey (Ruusila and Pesonen, 2004;Lupo, 2011; Koster and Tankersley, 2012). People have used dogsboth to locate large prey like muskoxen, elk, wolf, or polar bear andto hold the prey in place while alerting human hunters by howling(Arnold, 1979; Ruusila and Pesonen, 2004). Dogs have also beenused with drive lanes to trap large mammals in enclosures or drivethem off of cliffs but there is no evidence that this was done at themammoth megasites.

Mammoth megasites already amply fulfill this prediction whencompared with Mousterian or Middle Paleolithic sites. Fasterpopulation growth in wolf-dog using peoples is expected due toimproved nutrition and less energetic expenditure. Though esti-mating ancient population size is inherently speculative, following


the strategy developed by Mellars and French (2011), I wouldexpect to see evidence of faster population growth in AMHs relativeto Neanderthals by an increasing relative number of archaeologicalsites, an increasing size of archaeological sites, an increasing den-sity of retouched stone tools at such sites, and an increasing densityof estimated meat yield relative to site size.

The fourth is that sites formed by AMHsworking with wolf-dogsshould yield evidence of longer occupations than those of AMHs orNeanderthals working without cooperative wolf-dogs. This is pre-dicted because of the ethnographic use of dogs in guarding bothfood and homesteads among the Neoeskimo (Arnold, 1979).Retaining control of huge carcasses and diminishing raids byscavengers would constitute a considerable advantage.

The fifth is that undomesticated canids, such as wolves andfoxes, 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 responseto canid intruders (Geist, 2006; Vanak and Gompper, 2009). Ifdomesticated or semi-domesticated canids lived with AMHs, thewolf-dogs should react strongly to wolves or other wild canids,alerting AMHs and giving them extra cause to kill the intruders, inaddition 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 preyto the campsite, if it is not the killsite. Ethnographic evidencesuggests that dogs trained as pack animals are capable of carryingloads of up to 23 kg (Turner, 2002; Fiedel, 2005; Speth et al., 2013).If wolf-dogs assisted in transporting meat, they would effect asignificant savings in energy expenditure by humans.

The seventh is that, as at P�redmostí (Bocherens et al., 2013, thisvolume), future studies of stable isotopes of wolf-dogs and wolvesfrom the same sites will reveal dietary differences between the twoprobably because of provisioning by humans. I do not predictwhether most of these wolf-dogs will or will not bear evidence ofdismemberment, skinning, filleting, 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 conditionsunder which mammoth megasites were formed. Analysis of Hay-nes’ age profiles of modern models, based on assemblages of L.africana with known causes of death, demonstrates that attritionaland catastrophic mortality produce statistically different age pro-files, using the KolmogoroveSmirnov D statistic as a measure ofsignificance. This affirmed the appropriateness of using suchmodels as a method for deducing taphonomic history of such sites.

Published age profiles of mammoth megasites constructed us-ing Haynes (1991; Conybeare and Haynes, 1984) ageing techniqueswere used in this study. Megasites with fewer than 25 individualmammoths were eliminated from this study because of the de-mands of the KolmogoroveSmirnov statistic. Comparison of ageprofiles of eight mammoth megasites to Haynes’ models revealedthat all of these megasites were significantly different from Haynes’models. This finding suggests the possibility of problems with theunderlying assumptions of the ageing technique, of problems withusing modern analogies in paleontology, or that the sites repre-sented palimpsests with mixed taphonomic histories.

Published age profiles of threemammothmegasites constructedusing techniques developed by Saunders (1977) were compared toSaunders’ model, which represents either an entire proboscideanherd killed in a single episode or repeated killing of single maternalgroups over time. Both could be considered catastrophic causes ofdeath. Berelekh and Mezin do not differ significantly from Saun-ders’model, suggesting a catastrophic mode of death occurred. The



mammoth megasite of Mezhirich differed significantly fromSaunders’ model, suggesting an alternative taphonomic historyand/or a palimpsest.

The age profiles of the mammoth megasites, as a group, do notsuggest a consistent taphonomic history. Comparison of mammothmegasites to each other revealed a more interesting pattern.Though each site differed significantly from most other sites, achain of resemblances which mirrored temporal and spatial prox-imity was found. This suggests that the sites may reflect thetransmission of a technology or behavior important to the forma-tion of suchmegasites through time and space. Shea and colleagues(Shea, 2006, 2009; Shea and Sisk, 2010; Sisk and Shea, 2011) havehypothesized that the development of complex projectile weaponswas responsible for the appearance of these megasites and forhuman 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 theappearance of mammoth megasites, including the high numbers ofindividual mammoths and canids at such sites and the closeproximity of sites to the place of mammoth death. The advantagesof projectile technology and canid collaborationmay have led to thesurvival of AMHs while Neanderthals went extinct. These canidsare explicitly identified with the morphologically unusual groupfirst recognized by Germonpré et al. (2009) at about 32 ka B.P(uncal) and more recently at additional sites in the Gravettian, andEpigravettian (Germonpré et al., 2012, 2013, this volume). Apossible “incipent dog” has also been identified from RazboinichyaCave, Siberia, and is dated to about 34 ka B.P. (uncal) (Ovodov et al.,2011).

To the extent that further investigations have been conducted todate, the group of canids or dog/wolves identified by the method-ology of Germonpré et al. (2009) is also genetically and dietarilydistinctive 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 andJaroslaw Wilczy�nski for organizing the excellent conference “TheWorld of Gravettian Hunters”. I thank them and my other col-leagues at themeeting for their input and helpful suggestions. I alsothank Jefferey Mathison [email protected] for drawing thebeautiful map for this paper.

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