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Author's personal copy
Quaternary International 264 (2012) 32e51
Contents lists available at SciVerse ScienceDirect
Quaternary International
journal homepage: www.elsevier.com/locate/quaint
Evidence of small fast game exploitation in the Middle Paleolithic of Les
Canalettes Aveyron, France
David Cochard a, Jean-Philip Brugal b, Eugène Morin c, *, Liliane Meignen d
a
Université de Bordeaux 1, UMR 5199 PACEA-PPP, Avenue des facultés, F-33405 Talence cedex, France
Maison Méditerranéenne des Sciences de l’Homme, UMR 7269, 5 rue du Château de l’horloge, Aix-en-Provence cedex 2, BP 674, 13094, France
c
Trent University, Department of Anthropology, DNA Bldg Block C, 2140 East Bank Drive, Peterborough, Ontario, Canada K9J 7B8
d
Université Nice Sophia Antipolis, Campus Saint-Jean-d’Angély SJA3 e CEPAM e UMR 6130 CNRS 24, avenue des Diables Bleus, 06357 Nice Cedex 4, France
b
a r t i c l e i n f o
a b s t r a c t
Article history:
Available online 10 February 2012
In Europe and southwest Asia, ungulates are generally very well represented in anthropogenicallyaccumulated assemblages dating to the Middle and early Late Pleistocene. In contrast, taphonomic
studies have shown that fast small-sized prey taxa, such as leporids, small carnivores and birds, were
rarely exploited during these time periods. These faunal patterns are often interpreted as indicating that
Neandertals were characterized by a narrow diet, a finding with important social and technological
implications. This paper reexamines this view using faunal data from Les Canalettes layer 4, a Middle
Paleolithic faunal assemblage in Mediterranean France dominated by rabbit remains. Multiple lines of
evidence, including the presence of cutmarks and shaft cylinders, suggest that humans accumulated
most of the rabbit specimens found in this layer. This and other comparable assemblages raise a number
of issues with respect to variations in diet breadth prior to the Upper Paleolithic.
Ó 2012 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
Ancient hominins are generally assumed to have had the
capacity for hunting small game species. This assumption is based
on the notion that small prey species are easier to catch than those
of larger body size, such as cervids and equids. However, limited
information is available on human exploitation of small game for
the periods preceding the end of the Late Pleistocene. This paper
tackles this problem by reviewing the evidence for Middle Paleolithic consumption of small fast game in western Europe and
southwest Asia using data from the Mousterian rockshelter of Les
Canalettes in southern France. The archaeozoological analysis of
rabbit remains at this site provides important new data that help to
assess change in diet breadth in Neandertal populations.
1.1. Background
Small game hunting is relatively common among non-human
primates (e.g., Teleki, 1975; Newton-Fisher et al., 2002; Watts and
Mitani, 2002) and is well documented in the ethnographic record
(e.g., Lee, 1979; Jones, 1983; Malaurie, 1989; Wadley, 2010).
* Corresponding author.
E-mail address: [email protected] (D. Cochard).
1040-6182/$ e see front matter Ó 2012 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2012.02.014
Although several studies have examined small prey exploitation
during the Upper Paleolithic (e.g., Hockett and Haws, 2002; Pérez
Ripoll, 2004; Cochard, 2005; Lloveras, 2010), little is known about
the extent of this ability during the Lower and Middle Paleolithic.
This issue is critical, given that small game exploitation has
important implications for the foraging and social organization of
past human groups (Bird and Blierge Bird, 2000; Stiner et al., 2000;
Zeanah, 2004; Lupo, 2007). The problem of the emergence of this
practice is the focus of the present article.
Small game is here defined as a species of less than 10 kg.
As pointed out by Stiner and Munro (2002), it is important to
distinguish slow or sessile small-sized species (e.g., shellfish,
tortoises)da class of game that can be gathered like plantsdfrom
fast small-sized ones (e.g., leporids, birds, small carnivores),
which require greater skills and/or more complex procurement
techniques due to the high speed they can attain during escape.
Along with variations in flight strategies and social behavior,
differences in maximum velocity require due attention, as they can
substantially affect net return rates (Bird et al., 2009; Morin, 2012).
In this discussion, consideration must also be paid to the age and
sex of the hunter, given that children and adults, as well as males
and child-rearing females, may differ significantly in their foraging
goals (Bird and Blierge Bird, 2000; Codding et al., 2010).
The present analysis focuses exclusively on fast small-sized taxa
because there are ambiguities concerning the rank of slow small-
Author's personal copy
D. Cochard et al. / Quaternary International 264 (2012) 32e51
33
Table 1
Mean and range of sex-ratios for modern wild rabbit samples captured using various methods of procurement. The values presented in this table consist of pooled data from
Smith et al. (1995:116e118, Tables 1e5). Values for “Spring traps,” “Smeuse traps,” and “Dug out/Fenn traps” were excluded due to a lack of data on location, as were values for
unspecified methods. Values for “gassing” were also excluded because this method was preceded by ferreting.
Method of procurement
Open field
Shot
Cage-trapped
Snared
Poisoned
Warren-based
Ferreted
Gin-trapped
Location
Number of case studies
Males:Females
Range of sex-ratios
Mean sex-ratio
UK, Spain, Aust., NZ
UK, Aust.
UK
Aust., NZ
23
10
1
3
14611:13922
2320:2123
219:132
232:179
0.37e0.61
0.49e0.66
e
0.54e0.62
0.51
0.52
0.62
0.56
UK, Spain
UK
10
3
3663:5284
927:1427
0.21e0.50
0.36e0.50
0.41
0.39
Abbreviations: Aust ¼ Australia; NZ ¼ New Zealand.
Table 2
Pre-Upper Paleolithic assemblages from Europe and southwest Asia with small fast game remains showing anthropic marks. The dashes in the cells indicate a lack of data.
Site, period
Lower Pleistocene
S. del Elefante TE9a
S. del Elefante TE12a
Dursunlu
Middle Pleistocene
Arago, G
Terra-Amata
Orgnac III
Gran Dolina TD10-1
Lazaret, unit 25
CII
CIII
Hayonim, 4be1
C. Bolomor XVIIc
XVIIc
XVIIc
XII
XII
XII
XI
XI
IV
IV
IV
IV
Late Pleistocene
Adaouste
Artenac, 8
Pech de l’Azé IV, 8
Les Canalettes, 4
Pié Lombard
Combe-Grenal, 24
La Crouzade, 6e8
Jonzac, Quina
Salpêtre de Pompignan, 5e9
Gabasa 1
Salzgitter-Lebenstedt
G. de l’hyène, Arcy
Cova Beneito, D4
D2
Cova Negra, IV
IIIb
IIIa
II
Pech de l’Azé I, 4
Fumane, A base
Fumane, A9
Fumane, A6eA5
Baume de Gigny
Taxon
Date (ka)
Species NISP
Cutmarks (n, %)
Burning (n, %)
References
M-sized bird
O. cuniculus
L-sized bird
<1200
1000 ?
780e990
343
75
e
1
1
1
0.2
1.3
e
0
0
e
O. cuniculus
”
”
Vulpes vulpes
O. cuniculus
C. livia
O. cuniculus
Vulpes vulpes
O. cuniculus
S-sized bird
M-sized bird
Cygnus olor
M-sized bird
O. cuniculus
O. cuniculus
Aythya sp.
Vulpes vulpes
O. cuniculus
S-sized bird
M-sized bird
400
380e320
370e300
MIS 9
170
190e150
150e130
170e70
350e300
”
”
180
”
”
<150
”
>120
”
”
”
1434
819
8878
16
942
12288
12834
18
457
9
26
1
29
135
262
202
2
789
25
184
1?
1
2
1
0
1?
2
1
23
2
4
1
3
6
28
18
1
111
1
31
0.1
0.1
0
6.2
0
0
0
5.6
5.0
22.0
15.4
100
10.3
4.4
10.7
8.9
50.0
14.1
4.0
16.8
0
23
20
0
83
0
127
3
0
0
0
0
0
0
181
106
1
481
11
106
0
2.8
0.2
0
8.8
0
1.0
16.7
0
0
0
0
0
0
69.1
52.5
50.0
61.0
44.0
57.6
D 92
G 01
G 01
B 10b
L 04
R 04
G 01
S 05
S 08, B 11
B 11
B 11
B 09, B11
B 09, B11
B11
B 11
B 10, B11
B 11
B 11
B 11
B 11
O. cuniculus
Meles meles
M-sized raptor
O. cuniculus
”
Lepus sp.
”
Vulpes vulpes
O. cuniculus
”
Cygnus sp.
Anas sp.
A. chrysaetos
O. cuniculus
”
”
”
”
”
A. chrysaetos
A. chrysaetos
A. monachus
several species
C. cygnus
120e90
e
100
MIS 5/4
80e60
MIS 4
MIS 4/3?
>49
50e35
MIS3?
MIS3?
”
MIS3?
MIS3
”
”
”
”
”
58e38
MIS3
”
40e45
33e27?
1786
149
1
1209
1292
e
85
2
2255
2658
e
e
e
955
169
368
337
94
151
3
e
e
294
1
0
1?
1
8
2
1
2
1
0
3
1
1
1
1
3
1
3
1
1
2
1
1
5
1
0
0.7
100
6.6
0.2
e
2.4
50.0
0
0.1
e
e
e
0.1
1.8
0.3
0.9
1.1
0.7
67.0
e
e
1.7
100
1
0
1
4
2
e
0
0
0
e
e
e
e
e
e
e
e
e
e
0
e
e
e
0
0.1
0
100
3.3
0.2
e
0
0
0
e
e
e
e
e
e
e
e
e
e
0
e
e
e
0
D 94
M 07
D 09
this study
G 72
C86, M 12
G 72
J 08
G 72
B 97
G 09
”
F 04
S 08
”
”
”
”
”
M 75, L 00
F 04
P 11
P 11
M 89
0
0
e
H 07
H 07
G 09
Abbreviations: Arcy, Arcy-sur-Cure; O. cuniculus, Oryctolagus cuniculus; C. livia, Columba livia; m-sized, medium-sized; C. cygnus, Cygnus cygnus; D 92, Desclaux 1992; G 09,
Güleç et al., 2009; S 08, Sanchis Serra and Fernández Peris, 2008; G 01, Guennouni, 2001; L 04, de Lumley et al., 2004; R 04, Roger, 2004, S 05, Stiner, 2005; B 09, Blasco and
Fernández Peris, 2009; B 10, Blasco et al., 2010a; B 10b, Blasco et al., 2010b; B 11, Blasco and Fernández Peris, 2012; D 94, Defleur et al., 1994; H 07, Huguet 2007; M 07, Mallye,
2007; G 73, Gerber, 1972; C 86, Chase, 1986; M 12, Morin 2012; J 08, Jaubert et al., 2008; B 97, Blasco, 1997; G 09, Gaudzinski-Windheuser and Niven, 2009; F 04, Fiore et al.,
2004; D 09, Dibble et al., 2009; M 75, Mourer-Chauviré, 1975; L 00, Laparra, 2000; P 11, Peresani et al., 2011; M 89, Mourer-Chauviré, 1989. Data presented in this study are
shown in bold.
Author's personal copy
34
D. Cochard et al. / Quaternary International 264 (2012) 32e51
sized taxa. Indeed, it has been argued that slow small-sized species
(e.g., tortoises) may be associated with higher net return rates than
several large-sized game, possibly including some ungulate taxa
(Morin, 2012). This is because slow small-sized taxa are likely to
represent high-rank prey categories for groups of people with
reduced mobility (e.g., pregnant women, children).
The consumption of small fast prey species has important socioeconomic implications, given that it may signal human-induced
resource depression (e.g., Broughton, 1999; Cannon, 2003; Munro,
2004). In Europe and southwest Asia, many specialists interpret
the substantial introduction of leporids, fish, and birds into the
human diet at the end of the Upper Paleolithic as an indication of
diet widening. Despite ongoing debates about the factors that
caused this broadening of dietary practices, it is relatively clear that
this shift had a profound impact on human societies (e.g., Stiner
et al., 2000; Jones, 2006; Richards, 2009).
The current consensus is that the western European Neandertals
were characterized by a narrow diet in which ungulates were the
main source of energy. Although this view seems broadly accurate,
some evidence suggests that this picture is incomplete, at least in
some regions (Sanchis Serra and Fernández Peris, 2008; Blasco et al.,
2010a; Blasco and Fernández Peris, 2012). In this context, the
numerous rabbit remains uncovered at Les Canalettes in France
deserve considerable attention, as they may indicate that variation
in diet breadth has been underestimated for the periods preceding
the Upper Paleolithic. However, before examining the Les Canalettes
rabbit assemblage, the issue of agency must first be raised.
1.2. Agency and procurement goals in small fast game
accumulations
In archaeological contexts, the study of small fast game remains
is complicated by several factors. Foremost among these is the issue
of the nature of the accumulation (Andrews, 1990; Hockett and
Haws, 2002). Taxa such as leporids are preyed upon by many
predators, such as the stoat (Mustela erminea), pine marten (Martes
martes), fox (Vulpes vulpes and Alopex lagopus), wildcat (Felis silvestris), wolverine (Gulo gulo), wolf (Canis lupus), golden eagle
(Aquila chrysaetos), eagle owl (Bubo bubo), buzzard (Buteo buteo),
and several other species, including humans (Valverde, 1967;
Delibes and Hiraldo, 1981; Angerbjörn and Flux, 1995). Because
many of these predators are known to den in natural
sheltersdincluding those visited by humansdremains of their
prey can be found mixed with anthropogenic refuse. This point is
important because natural deaths in rockshelters and caves can
result in significant accumulations of small species. This problem
stresses the importance of taphonomically-oriented analyses of
entire faunal assemblages recovered using modern excavation
techniques. Patterns of small fast game exploitation are best
understood when associated with a detailed analysis of the nature
and conditions of preservation of all classes of faunal remains
present in an assemblage.
Prior to the rise of taphonomic approaches, the simple association of artifacts with remains of animal species, be they large or
small, was often considered a proof of human consumption.
Research has since shown that this criterion is unsatisfactory when
used alone. Marks and bone modifications generally provide more
secure foundations for investigating anthropic exploitation of small
fast game. Cutmarks, burning, and fragmentation patterns are
particularly helpful in this regard. However, interpretation of these
forms/types of damage presents its own challenges because they
may reveal motivations other than food procurement. For instance,
parts of small prey taxa may have been used as tools and/or for
symbolic purposes. In France, the presence of cutmarks on golden
eagle (A. chrysaetos) phalanges at Pech de l’Azé I (Mourer-Chauviré,
1989; Soressi et al., 2008), Pech de l’Azé IV (GaudzinskiWindheuser and Niven, 2009), Grotte de l’Hyène at Arcy-sur-Cure
(Fiore et al., 2004), and, in Italy, at Grotta di Fumane (Fiore et al.,
2004; Peresani et al., 2011) suggest the symbolic use of raptor
claws and feathers of birds. Additional evidence for symbolic use of
Fig. 1. Location of sites mentioned in the text. 1) Cova Beneito, 2) Cova Negra, 3) Cova del Bolomor, 4) Sima del Elefante, 5) Gabasa, 6) Caune de l’Arago, 7) Jonzac, 8) Artenac, 9)
Combe-Grenal, 10) Pech de l’Azé IV, 11) Les Canalettes, 12) La Crouzade, 13) Salpêtre de Pompignan, 14) Orgnac III, 15) Adaouste, 16) Lazaret, 17) Terra-Amata, 18) Pié Lombard, 19)
Baume de Gigny, 20) Grotte de l’Hyène, Arcy-sur-Cure, 21) Salzgitter-Lebenstedt, 22) Fumane, 23) Dursunlu, 24) Hayonim.
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
raptors comes from Mousterian levels at Combe-Grenal and Les
Fieux (Morin and Laroulandie, 2012). In this debate, it is important
to note that the exploitation of pelts from leporids or small carnivores is, to the authors’ knowledge, undocumented for the periods
preceding the end of the Upper Paleolithic (Charles, 1997; Fontana,
2003; Mallye, 2007; Blasco et al., 2010a).
Burn marks can also inform the analysis of faunal assemblages.
However, these marks are often difficult to interpret because they
may result from accidental burning of faunal remains present in the
sediments (Shipman et al., 1984; Stiner, 2005). On a more positive
note, it should be pointed out that several authors have emphasized
that cooking of small prey species can sometimes be identified
based on the anatomical location of the burned patches (e.g., Vigne
and Marinval-Vigne, 1983; Speth, 2000; Laroulandie, 2001; Mallye,
2007; Lloveras et al., 2009; Royer et al., 2011). Human tooth marks,
which often take the form of narrow notches, pits or diffuse
scraping, can sometimes be found near the broken edges of bones
or on the shafts (Pérez Ripoll, 1993, 2004, 2006; Laroulandie, 2001;
Cochard, 2005; Landt, 2007; Blasco and Fernández Peris, 2012). In
leporids, human tooth marks appear to be related to marrow
extraction and meat consumption.
In addition to marks, fracture patterns can contribute to the
identification of the agent of accumulation. Experimental work has
shown that certain types of fractures are potentially diagnostic of
human intervention. For instance, Gourichon (1994) and
Laroulandie et al. (2008) have argued that, in birds, the overextension of the ulna during disarticulation often creates diagnostic
perforation holes (enfoncements) in the fossa olecrani region of the
humerus. Similar marks may occur in anthropogenic rabbit
assemblages. Peeling fractures, which are produced during the
snapping of bones, have also been identified in presumably humandeposited assemblages of small fast game species (Laroulandie,
2002; Sanchis Serra and Fernández Peris, 2008; Peresani et al.,
2011; Blasco and Fernández Peris, 2012). However, whether this
type of fracture is a reliable indicator of human manipulation
remains to be confirmed.
In the last few years, shaft cylindersdhere defined as long bone
shaft fragments with their full circumferencedhave received
increasing attention from archaeologists working on prehistoric
exploitation of rabbits (Hockett, 1991; Perez Ripoll, 1992; Hockett
and Bicho, 2000; Cochard, 2004; Lloveras et al., 2009). This
interest stems, in part, from Jones’s (1983) ethnoarchaeological
study of the Aché (Paraguay), which documented the frequent
occurrence of shaft cylinders in the small game (Cebus apella,
capuchin monkey, 2.5e4 kg) assemblages created by these foragers.
In Europe, rabbit assemblages assumed to have been deposited by
humans are coherent with the ethnoarchaeological evidence
presented by Jones (1983) because they tend to comprise higher
percentages (>20%) of shaft cylinders than naturally deposited
ones (typically 5% or less, e.g., Guennouni, 2001; Cochard, 2007).
The interpretation of shaft cylinders in avian assemblages is
more ambiguous because bird bones are often pneumatized
(i.e., are associated with air sacs) and show appreciable variation in
marrow distribution between species and as a function of age
(Hogg, 1984; O’Connor, 2004). As a result, considerable work is
needed to determine whether the presence of cylinders in avian
assemblages can be used as evidence for human exploitation of
marrow-bearing elements.
Unfortunately, diagnostic forms of specimen modifications are
rarely abundant in Paleolithic small game samples, even in
assemblages that were clearly deposited by humans. Obviously, this
observation complicates the analysis of human foraging strategies
(Laroulandie, 2000; Hockett and Haws, 2002; Cochard, 2004). An
additional complication is that certain ethnographic groups are
known to have frequently ingested bones from small taxa such as
35
birds (Lefevre, 1989; Malaurie, 1989). The presence of leporid and
bird bones in human coprolites dated to the Holocene in the
American southwest is consistent with ethnographic observations
(Reinhard et al., 2007). Marks of digestion made by humans are
important, as they may, to some extent, mimic marks left on faunal
remains by small predators (e.g., Jones, 1986; Crandall and Stahl,
1995; Cochard, 2005; Perez Ripoll, 2006; Landt, 2007; Lloveras
et al., 2009).
A last problem concerns the excavation methods used to collect
the faunal specimens. Several decades of research have demonstrated that the lack of sieving or the use of coarse mesh (>2 mm)
can severely distort the taxonomic representation of small prey
species in a faunal assemblage (e.g., Payne, 1972; Shaffer and
Sanchez, 1994; Cannon, 1999; Cossette, 2000; Val and Mallye,
2011). Consequently, the focus in this paper is on assemblages
that have been excavated using modern recovery techniques.
1.3. Procurement methods and their impact on net return rates
Several authors have pointed out that mass collecting can
substantially increase the net return rate of a prey taxon and move
it into the optimal diet (e.g., Madsen and Schmitt, 1998; Ugan,
2005; Jones, 2006). This argument deserves attention because it
means that the presence of small fast game in an assemblage does
not necessary entail resource depression induced by human
predation. For instance, warren-based mass harvesting of rabbits
could have been performed by groups of various sizes and
composition using nets, fences, water, fire or other approaches.
Some of these methods have allegedly low opportunity costs and
might have significantly boosted the profitability of rabbits (Jones,
2004, 2006). Consequently, identifying methods of procurement is
critical for understanding foraging strategies at Les Canalettes and
other sites.
The study of age profile and sex ratio may shed light on this
problem because distinct methods of rabbit procurement may
differentially sample classes of prey individuals. Data collected by
Smith et al. (1995) on modern samples of wild rabbits captured in
a wide range of habitats are particularly insightful in this regard, as
they show that sex-ratios are primarily influenced by whether
Table 3
Taxonomic composition of the Les Canalettes faunal assemblages. The data are from
Brugal (1993), Patou-Mathis (1993), Meignen and Brugal (2001) and include
unpublished results collected by J.-P. Brugal.
Mammuthus
Dicerorhinus hemitoechus
Equus caballus
Equus hydruntinus
Bos primigenius
Cervus elaphus
Capreolus capreolus
Capra ibex
Rupicapra rupicapra
Sus scrofa
Ursus spelaeus
Ursus arctos
Canis lupus
Vulpes vulpes
Crocuta crocuta
Panthera spelaea
Lynx spelaea
Felis silvestris
Meles meles
Oryctolagus cuniculus
Lepus europaeus
Total
Layer 2 (top)
Layer 3
1
2
114
0
33
180
2
2
13
1
2
0
0
0
0
0
0
0
0
185
0
535
0
8
498
41
112
1176
87
26
93
6
13
3
23
9
8
1
1
1
2
109
0
2217
0.2
0.4
21.3
0
6.2
33.6
0.4
0.4
2.4
0.2
0.4
0
0
0
0
0
0
0
0
34.6
0
Layer 4 (bottom)
0
0.4
22.5
1.8
5.1
53.0
3.9
1.2
4.2
0.3
0.6
0.1
1.0
0.4
0.4
0.0
0.0
0.0
0.1
4.9
0
0
1
154
6
45
324
3
5
15
4
4
1
1
1
1
0
0
0
0
1209
7
1781
0
0.1
8.6
0.3
2.5
18.2
0.2
0.3
0.8
0.2
0.2
0.1
0.1
0.1
0.1
0
0
0
0
67.9
0.4
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
Fig. 2. The area excavated in layer 4 at Les Canalettes.
procurement occurs away from the warren or not. Although
procurement method also influences sex ratios, this last factor
appears to be less critical than location (Table 1). Based on these
data, warren-based acquisition of rabbits should, assuming unbiased transport, produce an archaeological assemblage in which
females are disproportionately represented. In contrast, the aboveground hunting of solitary rabbits in the landscape should generate
assemblages with relatively even proportions of males and females
or that are dominated by adult males, as these last individuals
commonly, but not always, range farther from the warren than
females (Cowan, 1987; Dekker et al., 2006).
How can warren-based mass-harvesting of rabbits be distinguished from the warren-based hunting of solitary individuals?
According to Jones (2004, 2006), mass harvesting of rabbits at
warren sites should, all else being equal, be associated with high
proportions of females of reproductive age and at least some kittens
(<1 months). This is because these classes of individuals are, at
least during the breeding season, generally well represented in the
warren population, although there are exceptions (Biadi and Le
Gall, 1993). The prediction for the hunting of solitary individuals
is that rabbit kittens should be absent because they normally stay in
the nest chambers (Kolb, 1985).
However, it may be difficult to distinguish these predictions in
archaeological contexts due to at least three confounding issues.
First, age profiles biased against juveniles may simply track the low
availability of juveniles in winter (Hockett, 1991; West, 1997;
Hockett and Bicho, 2000). Second, the dominance of adults in rabbit
assemblages may attest to a lack of interest for the presumably lowranked juveniles. Third, the pattern may be caused by densitymediated destruction. As a result of these problems of
Table 4
Relative abundances of skeletal elements and percentages of whole specimens in the rabbit assemblage from layer 4. Classes of elements absent in the assemblage are not
shown in this table. Values for the I1 and P3 are given in parentheses for upper teeth and lower teeth, respectively.
Maxillary
Tympanic
Occipital
Pre-maxilla
Temporal
Indet. cranial fragm.
Upper teeth
Mandible
Lower teeth
Atlas
Cervic. vert. IIIeVII
Thoracic vertebrae
Lumbar vertebrae
Sacrum
Ribs
Scapula
Humerus
Radius
Ulna
Metacarpal II
Metacarpal III
Metacarpal IV
Pelvis
Femur
Tibio-fibula
Calcaneum
Talus
Metatarsal II
Metatarsal III
Metatarsal IV
Metatarsal V
Phalanx I
Phalanx II
Phalanx III
Total
NISP
MNE
Abundance in a skeleton
%MAU
n whole
% whole (NISP)
% whole (MNE)
17
1
2
8
4
4
112
110
419
1
2
8
25
4
13
76
37
84
57
12
9
7
95
42
166
32
2
14
18
21
13
47
4
1
1627
15
1
2
8
4
e
112 (60)
90
419 (107)
1
2
8
24
4
7
70
29
56
42
12
9
7
59
18
87
31
2
14
18
21
13
44
4
1
1394
2
2
2
2
2
e
16 (2)
2
12 (2)
1
5
22
7
1
24
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
18
18
18
156
14
0.9
1.9
7.5
3.7
e
13.1 (16.8)
84.1
65.3 (100)
1.9
0.7
0.7
6.4
7.5
0.5
65.4
27.1
52.3
39.3
11.2
8.4
6.5
55.1
16.8
81.3
29
1.9
13.1
16.8
19.6
12.1
4.6
0.4
0.1
e
6
e
e
e
e
e
e
0
e
1
1
2
15
1
0
0
1
0
0
9
5
6
0
1
0
25
2
6
5
2
6
38
3
1
136
35.3
e
e
e
e
e
e
0.0
e
100.0
50.0
25.0
60.0
25.0
0.0
0.0
2.7
0.0
0.0
75.0
55.6
85.7
0.0
2.4
0.0
78.1
100.0
42.9
27.8
9.5
46.2
80.9
75.0
100.0
8.4
40.0
e
e
e
e
e
e
0.0
e
100.0
50.0
25.0
62.5
25.0
0.0
0.0
3.4
0.0
0.0
75.0
55.6
85.7
0.0
5.6
0.0
80.6
100.0
42.9
27.8
9.5
46.2
86.4
75.0
100.0
9.8
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
equifinality, age profiles must be interpreted with great caution. In
the following section, the evidence for anthropic use of small fast
game is reviewed for the period preceding the Upper Paleolithic.
1.4. Small fast game exploitation during the Lower and Middle
Paleolithic
In Europe and southwest Asia, there is scant evidence for human
use of small fast game prior to the Upper Paleolithic (Stiner et al.,
2000; Guennouni, 2001; Cochard, 2004; Costamagno and
Laroulandie, 2004; Laroulandie, 2004; Sanchis Serra and
Fernández Peris, 2008; Blasco and Fernández Peris, 2012; Morin,
2012). This is also the case for fish, which are poorly represented
in Lower and Middle Paleolithic sites with few exceptions (Le Gall,
1990, 2000; Roselló Izquierdo and Morales Muñiz, 2005). The
following discussion will principally be concerned with birds and
leporids, given that signs of human exploitation are anecdotal or
lacking for other classes of fast small-bodied taxa (e.g., small
carnivores, fish).
Lower and Middle Pleistocene assemblages comprising remains
of small fast game species with anthropic marks are listed in
Table 2. At the Lazaret cave (France, Fig. 1), Bouchud (1969) argued
that humans deposited the large bird assemblagesdwhich are
dominated by the alpine chough (Pyrrhocorax graculus) and the
rock dove (Columba livia)duncovered in this sequence. Subsequent
work by Mourer-Chauviré (1975), Vilette (1993), and Roger (2004)
on larger samples confirmed the abundance of birds at this site.
However, despite the very large samples, the taphonomic analysis
of these remains only revealed the presence of ambiguous
37
cutmarks on a single rock dove humerus (Roger, 2004:206). These
findings suggest that non-human predators were mostly, if not
entirely, responsible for the deposition of avian remains at the
Lazaret cave. Similar conclusions have been reached concerning the
bird assemblages from l’Arago (Desclaux, 1992) and Grotte Vaufrey
level VIII (Laroulandie, 2010).
A lack of taphonomic studies prevents the interpretation of the
moderately to very large bird assemblages from l’Hortus (alpine
chough: MNI ¼ 49, rock dove: NMI ¼ 23, partridge: MNI ¼ 18,
Mourer-Chauviré, 1972), Lunel-Viel (partridge: MNI ¼ 35), l’Abîme
de la Fage (partridge: MNI ¼ 1100), and Orgnac III (partridge:
MNI ¼ 91, Mourer-Chauviré, 1975). A comparable lack of data limits
the analysis of the avifaunas from Cova Negra in Spain and Gorham’s Cave in Gibraltar (Eastham, 1989, 1997). In contrast,
preliminary results indicate the presence of possible cutmarks on
an indeterminate large bird bone in the Lower Paleolithic site of
Dursunlu in Turkey (Güleç et al., 2009).
The picture appears to be similar for leporids. In France, the
Middle Pleistocene sites of Terra-Amata, Orgnac III, and Lazaret
cave comprise relatively large samples of rabbits (Guennouni,
2001). However, only a very small number of cutmarks, several of
which are dubious, were identified in the assemblages from these
sites (Table 2). Burned rabbit remains are generally slightly more
abundant at these sites. Additional work will be needed to fully
document the nature of the leporid assemblages at these sites.
Located in the Valencia region in eastern Spain, the Middle
Pleistocene sequence of Cova del Bolomor stands in sharp contrast
with the previous sites, as it provides compelling evidence for
human consumption of both leporids and birds. Anthropic marks,
Fig. 3. Patterns of skeletal representation in the rabbit assemblage from layer 4. NISP values are given above the bars.
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38
D. Cochard et al. / Quaternary International 264 (2012) 32e51
including cutmarks and shaft cylinders, are attested in the rabbit
assemblage from level XVIIc dated to ca. 350 kya (Sanchis Serra and
Fernández Peris, 2008). The same level also shows indication of
anthropogenic use of Passeriformes, Phasianidae and Anas sp.
(Blasco and Fernández Peris, 2012). At the same site, level XII is
consistent with this trend, given that cutmarks were identified on 6
remains of Oryctolagus cuniculus, 1 remain of Galliformes, 2 of Anas
sp. and 1 remain of Cygnus olor (Blasco et al., 2010a; Blasco and
Fernández Peris, 2012). In the overlying level XI, 8.9% of the avian
remains (NISP ¼ 202, all attributed to diving ducks Aythya) and
10.7% of the rabbit remains (NISP ¼ 262) are cut-marked (Blasco
and Fernández Peris, 2009). Lastly, in level IV, cutmarks and
anthropogenic forms of bone damage, such as peeling, shaft
cylinders, burning and tooth marks were observed on O. cuniculus
and bird (Passeriformes, Corvidae, Pyrrhocorax sp., Galliformes,
Phasianidae, Columba sp., Anas sp., Aythya sp.) specimens (Blasco
and Fernández Peris, 2012). Overall, because the cutmarks are
frequently found on meat-bearing elements, Cova Bolomor
constitutes a strong and relatively unique case for diet widening
during the Middle Pleistocene.
The general pattern for the early and middle Late Pleistocene is
similar, although there are some minor changes. As shown in
Table 2, small carnivore remains in these periods rarely bear cutmarks, including those from Cova Bolomor. The evidence from
southwest Asia is coherent with this general picture (Stiner, 2005).
Likewise, there is very little evidence for a dietary use of birds
during the early and middle Late Pleistocene, with the exception of
Cova Bolomor (see above). As emphasized above, in the majority of
cases, the few cutmarks observed on bird bones dated to these time
periods have been attributed to symbolic behavior rather than food
procurement (e.g., Peresani et al., 2011; Morin and Laroulandie
2012).
Concerning leporids, specimens with cutmarks occur in Spain at
Cova Bolomor, Cova Negra, Gabasa 1, and Cova Beneito, and in
France, at Pié Lombard, Combe-Grenal, Salpêtre de Pompignan, and
Les Canalettes layer 4 (this study). However, archaeozoological
studies of the Spanish sites have shown that carnivore marks are
common in the rabbit samples, which suggests that humans played
a minor role in the accumulations (Villaverde et al., 1996; Lloveras
et al., 2011). This generalization does not apply to Cova Bolomor, as
damage generated by non-human predators is rare in the rabbit
assemblages from this site (percentages of carnivore modifications
between 0 and 5.2%, Blasco and Fernández Peris, 2012). A different
interpretation may hold for some of the French sites, as exemplified
by the layer 4 assemblage from Les Canalettes.
2. Material and methods
Les Canalettes is a mid-altitude (700 m) rockshelter located at
the southern border of the Massif Central in the Aveyron region of
France. The site faces south-southwest and lies underneath a 4-m
high overhang that provides 60e70 m2 of protected area. The
first excavations at Les Canalettes (conducted by G. Costantini and
M. Delclaud) were initiated shortly after the discovery of the site in
Fig. 4. Skeletal representation by portion in the rabbit assemblage from layer 4.
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
Table 5
% MAU values in the rabbit assemblage from layer 4 compared to bone density values
(g/cm3) as calculated by Pavao and Stahl (1999:56e57).
Mandible
Incisor portion (DN1)
Molar portion (DN2)
Coronoid process (DN5)
Atlas (AT1)
Axis (AX1)
Vertebrae (LU1)
Rib (RI1)
Scapula
Proximal end (SP1)
Fossa (SP3)
Humerus
Proximal epiphysis (HU1)
Proximal end (HU2)
Distal end (HU4)
Distal epiphysis (HU5)
Radius
Proximal epiphysis (RA1)
Proximal end (RA2)
Distal end (RA4)
Distal epiphysis (RA5)
Ulna
Proximal end (UL1)
Distal end (UL4)
Metacarpal (MC1)
Sacrum (SC1)
Innominate
Distal ilium (IL1)
Proximal ilium (IL2)
Proximal ischium (IS2)
Distal ischium (IS1)
Femur
Proximal epiphysis (FE1)
Proximal end (FE2)
Distal end (FE5)
Distal epiphysis (FE6)
Tibia
Proximal epiphysis (TI1)
Proximal end (TI2)
Shaft (TI3)
Distal end (TI5)
Distal epiphysis (TI6)
Calcaneum (CA2)
Talus (AS1)
Metatarsal (MT1)
Phalanx I (PH1)
Density values (g/cm3)
%MAU
0.43
0.74
0.14
0.33
0.46
0.35
0.04
0.64
0.84
0.03
0.02
0.00
0.06
0.00
0.33
0.09
0.65
0.07
0.43
0.25
0.40
0.40
0.01
0.05
0.27
0.21
0.14
0.15
0.12
0.11
0.51
0.52
0.27
0.21
0.20
0.14
0.12
0.42
0.34
0.03
0.11
0.07
0.38
0.45
0.37
0.17
0.13
0.55
0.38
0.13
0.27
0.28
0.26
0.63
0.07
0.11
0.08
0.03
0.54
0.33
0.30
0.26
0.44
0.32
0.28
0.11
0.10
0.12
0.32
0.81
0.55
0.12
0.29
0.02
0.20
0.04
1964. More systematic excavations followed between 1980 and
1997 (Meignen, 1993). These excavations, which were concentrated
in the central zone of the rockshelter, enlarged the excavated area
to z30 m2. Three stratigraphic layersd4, 3, and 2, from bottom to
topdwere then defined. The material presented here derives from
Meignen’s excavations.
The Les Canalettes layers contain rich and well-preserved faunal
assemblages associated with the Typical Mousterian (Meignen,
1996; Meignen and Brugal, 2001). Lithic analyses indicate that
raw materials were mostly obtained from geological deposits
located near the site. Climatic data and a TL date of
73,500 6000 ka (Valladas and Joron, 1993) on burnt flint from
layer 2 suggest that the site was occupied at the end of MIS 5a
(71e85 ka). The composition of the faunal and anthracological
assemblages (Vernet, 1993; Théry-Parisot, 1998) indicates that
environmental conditions during the human occupations were
relatively temperate, although slightly cooler and drier than at
present (Meignen and Brugal, 2001).
Approximately 3000 ungulate specimens were identified at
least to the genus level at Les Canalettes. These remains represent
approximately 10% of the total faunal sample (Brugal, 1993;
39
Meignen and Brugal, 2001). Data based on tooth eruption point to
the procurement of ungulates between spring and the beginning of
autumn (Patou-Mathis, 1993; Meignen and Brugal, 2001). Carnivores are poorly represented in the faunal assemblages and few
specimens show signs of their activities. The ungulate assemblages
appear to be mostly anthropogenic, a phenomenon that is characteristic of several French Mousterian sites (e.g., David and Poulain,
1990; Morin, 2004; Costamagno et al., 2005).
Remains of small vertebrates, including a large number of rabbit
specimens, were found mixed with those of ungulates in all of the
Les Canalettes layers. Patou-Mathis (1993) conducted a preliminary
analysis of the rabbit assemblages from layers 3 (NISP ¼ 109) and 2
(NISP ¼ 185) and concluded that the majority of the rabbits died
naturally in the rockshelter, possibly in their burrow, although she
pointed out that some of the specimens from layer 3 may reflect
human procurement (Patou-Mathis, 1993:202). This analysis
focuses on the previously unstudied large rabbit assemblage from
layer 4. This layer reaches a maximum thickness of 80 cm and was
exposed over a surface of 30 m2; it consists of heterometric gravels
in a brown sandy matrix.
Experimental studies (Shaffer and Sanchez, 1994; Cannon, 1999;
Val and Mallye, 2011) and observations on fossil collections have
shown that the recovery of small rabbit elements such as
phalanges, metacarpals, carpals, and patellae are greatly affected by
recovery methods. At Les Canalettes, the use of a 2-mm mesh
screen likely significantly reduced the loss of these small rabbit
elements, as this mesh size is relatively effective at capturing small
rabbit specimens. Although not conclusive, the large number of
isolated teeth (NISP ¼ 237) identified in layer 4 is consistent with
this interpretation. Nevertheless, some small rabbit remains were
likely missed during the excavations.
The age profile of rabbits in layer 4 was examined using the
degree of bone fusion. In rabbits, sexual maturity and adult body
size are reached at approximately 5 months of age (Biadi and Le
Gall, 1993); the proximal radius and distal humerus are already
fused, while the ossification of the proximal femur, distal tibia, and
proximal ulna isdor is in the process of beingdcompleted. By
about 9 months old, the long bone epiphyses are all fused with few
exceptions (Bujalska et al., 1965; Broekhuizen and Maaskamp,
Fig. 5. Correlation between the relative abundances of rabbit skeletal parts in layer 4
and bone density (g/cm3) values. The data are from Table 4.
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40
D. Cochard et al. / Quaternary International 264 (2012) 32e51
1979; Driver, 1985; Callou, 2003; Gardeisen and Valenzuela Lamas,
2004). As a result of these osteological developments, data on bone
fusion can be used to distinguish various age classes, such as
“immatures” (<3 months), “sub-adults” (5e9 months) and “adults”
(>9 months). Although additional information on age profile can be
obtained by assessing the proportion of vertebral epiphyses that are
ossified to the centrum, the low numbers of vertebrae in the sample
prevented the application of this last method.
Because humans often fracture bones to extract marrow, variations in the degree of element completeness may yield information on human behavior. This issue was assessed in layer 4 by
dividing the number of whole specimens for a given element by the
NISP count for that element. Values were then multiplied by 100 to
obtain percentages of whole specimens (hereafter referred to as %
whole specimens). To ensure that the results are robust, %whole
specimens were also calculated relative to MNE counts. The head
parts were excluded from the analysis, given that it comprises
fragile (e.g., vomer, ascending ramus) and dense (e.g., petrosal)
parts.
Breakage patterns in layer 4 were also examined using Villa’s
and Mahieu’s (1991) classification, which focuses on three
features of long bone shaft fragments: the overall morphology and
the angle of the fracture, and the shape of the fracture’s edges.
Previous studies indicate that this classification may help to
distinguish leporid assemblages resulting from natural death (e.g.,
Coudoulous II, Les Rameaux) from carnivore- (e.g., Les Rochers de
Villeneuve, Grotte Vaufrey) and human-accumulated (e.g., BoisRagot, La Faurelie II) leporid assemblages (Cochard, 2004).
3. Results
In the Les Canalettes layer 4, leporids account for 67.9% of the
total NISP (Table 3). These remains are less abundant in the upper
level 3 (4.9% of total NISP) and moderately frequent in the uppermost level 2 (34.6% of total NISP). The overwhelming majority of
the leporid remains from layer 4 were attributed to the rabbit
(Oryctolagus cunniculus, NISP ¼ 1209, MNI ¼ 56). Hare remains
(Lepus europaeus, NISP ¼ 7) are rare and were not taken into
account in this analysis. The faunal assemblage from layer 4
comprises 557 ungulate remains identified at least to the genus
level. The ungulate sample is dominated by red deer (Cervus elaphus, 58.2% of ungulate NISP) followed by horse (Equus caballus,
27.6% of ungulate NISP). Several species of carnivores are present at
Les Canalettes. Remains of these species are rare in layers 4
(NISP ¼ 8/1781 or 0.4%) and 2 (NISP ¼ 2/535 or 0.4%) and slightly
more common in the intermediate level 3 (NISP ¼ 61/2217 or 2.8%).
In layer 4, 36.7% (NISP ¼ 444) of the rabbit remains were found
in three adjacent squares (A6, A7, B7) located in the northeastern
section of the rockshelter (Fig. 2). This zone of high rabbit density
(130 remains/m2) was possibly larger in the past, given that it was
partially truncated by a looting episode (area: 4 m2). The other
remains are evenly distributed, although densities are slightly
Fig. 6. Degree of element completeness in the rabbit assemblage from layer 4. Numbers of complete elements are given above the bars.
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
41
higher (60 remains/m2) in squares D5, E4, G5 and G6 than in the
remaining squares (40 remains/m2). In contrast to level 3, where
some specimens were uncovered in anatomical connection (PatouMathis, 1993), no articulated elements were observed during the
excavation of layer 4.
3.1. Anatomical representation
Skeletal representation in the rabbit assemblage from Les
Canalettes was assessed using NISP and MNE (including its derived
form, %MAU). Concerning this last measure, it should be noted that
the values correspond to the highest tally obtained for an element,
irrespective of bone portion or landmark (Table 4). The data, which
are shown in Fig. 3, indicate a clear dominance of teeth and long
bone elements. In contrast, vertebrae, ribs, and foot bones are
poorly represented. The presence of a calcrete crust on a small
number of specimens in layer 4 (see below) possibly decreased the
degree of identification for certain elements. Bone fragmentation,
particularly vertebrae and metapodials, might have had a similar
effect. However, these problems are limited in layer 4 and their
impact on the anatomical profile is probably minor.
It may also be instructive to examine variations in skeletal
representation at a finer scale. Fig. 4 shows %MAU values by portion
for ten classes of elements. In general, long bones are marked by the
under-representation of at least one, and sometimes both, of their
epiphyses, which creates an over-representation of the shaft
portion. This pattern is manifest for the tibia, femur, and humerus.
The low representation of extremities is also apparent for the
scapula and innominates. Concerning the mandible, the ascending
ramus is very weakly represented in layer 4 relative to other
portions of the same element.
These anatomical patterns may be attributed to several factors,
including density-mediated attrition, a phenomenon that causes
the destruction of the more porous parts of the skeleton. To explore
this issue, the %MAU values for the layer 4 assemblage were
compared with mineral bone density, as measured using photon
densitometry (Table 5). The data show no correlation between the
two variables (rs ¼ 0.14, p < 0.41, Fig. 5). Assuming that complete
rabbits were introduced into the siteda reasonable assumption
with small-sized prey taxadthe statistical analysis of the density
data suggests that density-mediated destruction had a limited
impact on the rabbit assemblage.
An analysis of 11 categories of skeletal elements (femur, tibia,
metatarsal, humerus, radius, ulna, scapula, innominate, mandibular
bone, upper teeth, lower teeth, the latter two regardless of whether
they were still in their bony sockets or not) in seven squares (A6, A7,
B7, D5, E4, G5, G6) did not reveal any significant pattern with
respect to the spatial distribution of the specimens (results not
shown). In most squares, humeri and femurs outnumber mandibles, lower teeth, and tibia. However, innominates are the most
common element in square G6 (NISP ¼ 13), while mandibles are
poorly represented in E4 (NISP ¼ 3).
3.2. Patterns of long bone fragmentation
As mentioned earlier, the pattern of long bone fragmentation is
an important criterion that may help identify the agent that
deposited the rabbit specimens from layer 4. To shed light on this
issue, three aspects of the assemblage are successively examined
here: the percentage of whole elements, the abundance of dryversus green-bone fractures, and the percentage of shaft cylinders.
As shown in Fig. 6, long bones, scapulae, and innominates are
rarely complete in the assemblage (NISP: 2/557 or 0.4%, MNE: 2/361
or 0.5%, Table 4). In contrast, vertebrae, ribs, and foot bones were
frequently found whole (NISP: 128/233 or 54.9%; MNE: 128/222 or
Fig. 7. Ternary plots providing information on patterns of bone fragmentation in the
rabbit assemblage from layer 4: A) fracture outline, B) fracture angle, C) fracture’s
edges.
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42
D. Cochard et al. / Quaternary International 264 (2012) 32e51
57.7%). Interestingly, the data suggest a negative relationship
between the percentage of whole specimens and the size of the
samples (NISP: rs ¼ 0.69, p 0.001; MNE: rs ¼ 0.60, p 0.01).
These results indicate that elements that are common in the rabbit
assemblage also tend to be more fragmented.
Similar patterns of fragmentation have been observed in other
rabbit assemblages of Late Pleistocene age (e.g., Jullien and Pillard,
1969; Séronie-Vivien, 1971; Gerber, 1972). These patterns have, in
certain contexts, been attributed to marrow processing (Hockett,
1991). However, additional lines of evidence are needed to
support this argument, given that fragmentation is often severe
(with %whole values <10% for long bone elements) in carnivore
scats (Andrews and Nesbit Evans, 1983; Payne and Munson, 1985;
Schmitt and Juell, 1994; Cochard, 2007; Lloveras et al., 2007).
Moreover, various pre- (trampling, weathering) and post-burial
(e.g., scree, sediment compaction) processes can also fragment
specimens (Cochard, 2004; Barisic, 2006).
These problems of equifinality can partly be solved by assessing
whether the fractures occurred on dry or green bone. This information is useful because human and non-human predators are
unlikely to have deliberately fractured dry bones, as these are
largely devoid of soft tissues and juices. The fracture patterns
observed in layer 4 are reminiscent of those documented in
carnivore and anthropic assemblages. Indeed, in a sample of 334
ancient fractures, 51.8% present a curved or V-shaped fracture
outline (Fig. 7A), morphologies that are frequently associated with
green-bone fractures. The sample is also dominated by right (43.4%)
and oblique (40.4%) angles (Fig. 7B). The majority of the fracture
edges show a smooth texture (53.6%), although jagged edges
(29.9%) are also common (Fig. 7C). Because these patterns are
globally concordant with green-bone fractures, it seems reasonable
to conclude that, in layer 4, post-depositional processes have had
limited effects on specimen fragmentation.
As discussed earlier, shaft cylinders (or tubes) often occur in
relatively high frequencies in rabbit assemblages presumably
deposited by humans. The layer 4 assemblage is in agreement with
this pattern, given that 39.4% (151/383) of the long bone specimens
are cylinders (Table 6). Although all classes of long bones are
represented, most of them (104/151 or 68.9%) are tibia cylinders.
The prevalence of tibia specimens in the cylinder sample may be
explained by the high marrow utility of this element, especially
when compared to the radius and ulna (Hockett and Bicho, 2000).
The destruction of long bone extremities, either through biting or
the use of stone hammers, possibly accounts for the low representation of epiphyses relative to shaft portions in the layer 4
assemblage.
Fig. 8 shows that the tibia cylinders are relatively long in layer 4
with an average length of 41.8 mm (n ¼ 104, standard deviation of
12.7) for the whole tibia sample. For comparison, whole diaphyses
measure z60 mm in the reference collection consulted. The high
values recorded in layer 4 may confirm the anthropogenic origin of
the specimens because rabbit shaft cylinders are typically short in
natural deposits (Hockett, 1993; Brugal, 2006). Unfortunately, no
quantified data are available at the moment to support this
inference.
Other patterns are consistent with anthropic breakage. In layer
4, the olecranon region of the ulna is generally broken (22/32 or
68.8%). This observation, combined with instances of breakage near
the radial fossa region of the distal humerus (4/28 or 14.3%), may
attest to the overextension of the elbow during disarticulation,
which occurs when the joint is bent in the direction opposite to that
of natural flexion (Gourichon, 1994; Laroulandie et al., 2008;
Peresani et al., 2011). Damage on the tuberosity of the tibia (6/10 or
60.0%) possibly indicates that severe pressure was applied against
the cranial side of the distal femur during disarticulation of the
knee. Likewise, the frequent absence of trochanters (7/12 or 58.3%)
on the proximal femur may evidence disarticulation of the hip joint.
However, the possibility that these forms of damage were inflicted
during marrow cracking or by post-depositional processes cannot
be dismissed.
3.3. Bone surface modifications
As a complement to the study of anatomical representation and
fragmentation patterns, the taphonomic analysis of layer 4
included the observation of bone surface modifications produced
by abiotic processes and non-human and human agents.
In the sample (NISP ¼ 936, teeth excluded), 211 bones show
signs of modification by abiotic processes (Table 7). The majority of
these modifications correspond to grooves (Fig. 9A) and shallow
striations (Fig. 9B) presumably caused by contact with sedimentary
particles or stone fragments. Shaft portions frequently display
parallel striations perpendicular to the longitudinal axis of the
element. These marks are similar to those visible in the naturally
accumulated hare (Lepus timidus) assemblage from Coudoulous II
(Cochard, 2004). Because trampling normally produces scratches
with random orientations (Barisic, 2006), the parallel striations
identified in layer 4 are here tentatively attributed to “geological”
movements of the specimens within the matrix (especially
compaction). The same process may account for the superficial loss
of bone material observed on the prominences of several elements
(e.g., the condyles and epicondyles of the femur, the trochlea of the
humerus, the edges of the acetabulum and ischium). Other types of
abiotic damage, such as manganese stains (5.4%), weathering
(1.4%), and calcite coating (1.3%), occur at lower frequencies in the
sample.
Damage produced by plants and animals has also been identified at Les Canalettes (Table 7). Marks ascribed to plant growth are
fairly common in layer 4, recorded on 27.8% of the rabbit specimens.
These marks vary randomly in the sample, which is confirmed by
a lack of correlation between the percentage of root-marked
specimens per element and the NISP counts for these elements
(rs ¼ 0.21, p < 0.40). In layer 4, root marks often take the form of
sinuous lines, isolated pits, and small patches (Fig. 10). Importantly,
these marks rarely (49/972 or 5.0%) cover more than 1/3 of the
specimen surfaces.
Table 6
Types of long bone fragments in layer 4 at Les Canalettes. Shaft cylinders are shown in bold.
Humerus
Whole bone
Proximal end
Distal end
Shaft cylinder
Shaft fragment
Total NISP
Radius
Ulna
Femur
Tibia
Total
n
%
n
%
n
%
n
%
n
%
n
%
1
2
27
5
2
37
2.7
5.4
73.0
13.5
5.4
100.0
0
55
11
17
0
83
e
66.3
13.3
20.5
e
100.0
0
39
2
15
0
56
e
69.6
3.6
26.8
e
100.0
1
11
5
10
15
42
2.4
26.2
11.9
23.8
35.7
100.0
0
16
15
104
30
165
e
9.7
9.1
63.0
18.2
100.0
2
123
60
151
47
383
0.5
32.1
15.7
39.4
12.3
100.0
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
Fig. 8. A sample of rabbit tibia shaft cylinders from layer 4.
43
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44
D. Cochard et al. / Quaternary International 264 (2012) 32e51
Table 7
Bone surface modifications in the rabbit assemblage from layer 4. The sample
comprises 936 specimens (teeth are excluded).
Agent
Type of damage
NISP
% sample
Sedimentary abrasion
Manganese stains
Weathering (stage 1)
Calcrete coating
135
51
13
12
14.4
5.4
1.4
1.3
Root etching
Digested
Carnivore tooth marks
Insectivore tooth marks
Cutmarks
Burning
Bite marks
260
7
1
1
9
4
1
27.8
0.7
0.1
0.1
1.0
0.4
0.1
Abiotic
Biotic
non-human
human
The very low incidence (0.7%, Table 7) of marks of gnawing and
digestion suggests that non-human predators played, at most,
a minor role in the deposition of the rabbit assemblage. The ravaged
specimens (three lower incisors, two metacarpals, two
innominates, two calcanei, and one radius) include bones and teeth
that have been hollowed out (Fig. 11AeB), remains with polished
and rounded edges (Fig. 11C), and bones with puncture marks
(Fig. 11D). The marks shown in Fig. 11, particularly the tooth
punctures, are attributed to the activities of small carnivores.
Carnivore-modified remains show no clear spatial patterning in
layer 4, B6 (n ¼ 2) being the only square with more than one of
these specimens. Lastly, the small gnaw marks and notches
(<1 mm) recorded on a scapula fragment adjacent to the supraglenoid tuberosity (Fig. 12) were ascribed, based on Andrews’s
(1990:6) actualistic observations, to the activity of a shrew (Sorex
sp.).
The rabbit assemblage from layer 4 comprises several bones
with unambiguous anthropic modifications (Table 7). Of particular
interest within this sample are cutmarks, which were identified on
six hindlimb elements (two innominates, two femurs, and two
tibiae), two humeri, and one metatarsal V (Fig. 13). All of these
specimens belong to adults, except for a proximal femur assigned to
a sub-adult. The slicing marks on the caudal side of the femurs
(Fig. 13AeB) and on the lateral sides of the pelvic girdle (Fig. 13CeD)
probably reflect the removal of the gluteal muscles, while the ones
placed on the caudal and lateral sides of the tibial shafts
(Fig. 13EeF) are arguably associated with the removal of the flexor
muscles. The cutmarks recorded on the midshaft of the humeri
(Fig. 13GeH) suggest the defleshing of forearm muscles. Lastly, the
deep and transverse cutmarks on the palmar side of the metatarsal
V (Fig. 13I) possibly attest to the disarticulation of the feet or
removal of the skin.
Despite the possible existence of a hearth and the recovery of
a moderately large quantity of charcoal (Théry-Parisot, 1998), rabbit
specimens are rarely burned in layer 4. Only four burned bones,
from as many elements (scapula, ulna, tibia, and indeterminate
metatarsal), are documented in the rabbit assemblage. The brown/
black color of the burned patches points to low-fire temperature,
save for a white, presumably calcinated, tibia fragment. Burning on
the distal metatarsal is limited to areas adjacent to the fracture on
the shaft. The olecranon portion of the proximal ulna is brown,
while the articulatory end and the proximal shaft are black. Both
specimens suggest that some of their portions were protected from
the fire by soft tissues. Lastly, a shaft fragment, probably of rabbit
(Fig. 14), shows what appears to be human bite marks (Aura Tortosa
et al., 2002; Cochard, 2005; Perez Ripoll, 2006). Assuming that the
interpretation of this specimen is correct, these marks may indicate
that the Les Canalettes foragers sometimes used their teeth to
fracture rabbit long bones.
3.4. Age profile
Fig. 9. Natural striations observed on rabbit bones from layer 4: A) grooves, B) shallow
striations.
When weaned rabbits (<1 month of age) leave the warren, the
permanent teeth are all in place (Dice and Dice, 1941; Callou, 2003).
Therefore, deciduous teeth may be important indicators as they are
likely to occur in appreciable numbers at nursing sites. In layer 4,
these teeth are absent. Moreover, very low percentages of unfused
proximal radii (3.5%) and distal humeri (4.2%) point to a low incidence of immature rabbits (<3 months of age) in the assemblage
(Table 8). These data do not match the expected age profile for
a warren.
The relative abundances of unfused proximal ulnas (25.0%) and
proximal femurs (42.9%) attest to a moderate representation of subadults (0e5 months) in layer 4. Although the percentage is
considerably lower for the distal tibia (13.3%), comparisons using
a test statistic (denoted ts, Sokal and Rohlf, 1969:607e610) based on
the arcsine transformation that examines the equality of percentages indicate that, within this age class, none of the differences
with the distal tibia are significant (proximal ulna versus distal
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
45
Fig. 10. Bone surfaces of rabbit remains modified by plant growth: A and D) sinuous lines, B) isolated pits, C) small patches.
tibia: ts ¼ 0.78, p < 0.44; proximal femur versus distal tibia:
ts ¼ 1.49, p < 0.14). The use of a broader age class (0e9 months)
does not result in a significant change in the proportion of juveniles, which suggests that few 5e9 month-old rabbits are present in
the rabbit assemblage (23.5% in the 0e5 month age class versus
23.3% in the 0e9 month age class; ts ¼ 0.02, p < 0.99).
Overall, the proportion of rabbit juveniles and young adults is
low in layer 4, given that 87.6% (127/145) of the long bone epiphyses are fused in this level. This trend is compatible with the
generally adult-dominated rabbit assemblages accumulated by
humans (Hockett, 1991). However, because some large non-human
predators (e.g., wolf, eagle) also primarily target adults (Cochard,
2004, 2008), in layer 4, the age profile for rabbits is consistent
with anthropic accumulation.
Given the dominance of adults, it may be argued that the rabbits
were mostly deposited during the warm season (spring-summer)
in layer 4. Although plausible, this argument must be viewed with
great caution because rabbit can produce several litters per year,
including, in favorable years, the early or late winter (Flux, 1965;
Boyd and Myhill, 1987; Gibb, 1990). Density-mediated
destructionda process that seems to have had a minor impact on
the rabbit assemblage at Les Canalettesdcan also depress the
relative abundance of rabbit kittens in a sample.
3.5. Sex-ratio
Recently, Jones (2006) presented a new method for distinguishing males from females in European modern rabbits using
measurements made on fully fused distal humeri. Her method,
which focuses on the interpretation of scatterplots, compares the
distal breadth and the trochlear breadth of distal humeri (Fig. 15). In
layer 4, these measurements were made on 21 specimens. The
results cannot be directly compared with those of Jones (2006)
partly due to body size differences between the two samples.
Indeed, the rabbits from Les Canalettes appear to have been
significantly larger than those in the modern sample. These
differences were not unexpected, given that the body size of rabbits
tends, in agreement with Bergmann’s rule, to increase with
increasing latitude and with decreasing mean annual temperature
(Sharples et al., 1996; Davis and Moreno Garcia, 2007). From this
perspective, the cooler conditions of the early Late Pleistocene
relative to the present might explain why the rabbits from Les
Canalettes had a large body size. As a consequence of this limitation, the results must be considered tentative.
If the proposed limits for the distributions of males and females
are correct, the layer 4 assemblage would be strongly dominated by
females (17/21 or 81.0%) during the occupation of layer 4. The
implications of these patterns are examined in the following
section.
4. Discussion
In Europe and southwest Asia, humans seldom consumed fast
small-sized prey species prior to the Upper Paleolithic (e.g.,
Cochard and Brugal, 2004; Morin, 2012). When these taxa are
present in Middle Paleolithic archeofaunas, taphonomic analyses
generally indicate that they were deposited by carnivores or
raptors. Cova Bolomor in eastern Spain departs from this trend as it
shows evidence for regular exploitation of fast small-sized species
(leporids and birds) during the Middle Pleistocene (Sanchis Serra
and Fernández Peris, 2008; Blasco and Fernández Peris, 2009;
Blasco and Fernández Peris, 2012).
As at Cova Bolomor, the early Late Pleistocene assemblage from
Les Canalettes layer 4 contrasts with most pre-Upper Paleolithic
assemblages of Europe and southwest Asia. The fauna from layer 4
is atypical in being dominated by rabbits (67.9% of total NISP),
Author's personal copy
Fig. 11. Ravaged rabbit specimens: AeB) hollowed-out specimens, C) specimen with polished and rounded edges, D) puncture marks.
Fig. 12. Scapula fragment with tooth marks presumably made by a shrew (Sorex sp.).
especially adults, the majority of which appears to have been
accumulated by humans. As discussed above, this argument finds
support in a low representation of carnivores (<1% of total NISP),
a high percentage of shaft cylinders (39.4%), and a rarity of ravaged
specimens (0.7%). Although not abundant, the presence of
unambiguous butchery marks (1.0%) supports an anthropogenic
origin for the rabbit assemblage. As pointed out earlier, the low
frequency of cutmarks in layer 4 does not contradict this view, as
these marks are rarely abundant in Upper Paleolithic and Holocene rabbit assemblages attributed to human predation (Hockett
and Bicho, 2000; Sanchis Serra and Fernández Peris, 2008;
Blasco and Fernández Peris, 2012). Yet, although the anthropogenic origin of the assemblage seems well supported, more data
will be needed to clarify how rabbits were processed in layer 4 at
Les Canalettes.
In the Les Canalettes sequence, ungulate species are abundant
with only minor variations in taxonomic composition (when leporids are excluded). Relative to layer 4, rabbits are less abundant in
layers 3 and 2 and are mostly represented by complete long bones.
Moreover, juveniles are better represented in the rabbit samples
from layers 3 and 2 than in layer 4, which may indicate that they
were naturally accumulated in the former layers (Patou-Mathis,
1993).
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
47
Fig. 13. Cutmarks recorded on rabbit bones from layer 4: AeB) femur, CeD) innominates, EeF) tibia, GeH) humerus, I) metatarsal.
From a temporal perspective, the pattern of rabbit exploitation documented at Les Canalettes is atypical, as no comparable
data exist outside eastern Spain and southern France (Cochard,
2004; Morin, 2012). In this respect, the layer 4 assemblage
from Les Canalettes presents greater affinities with the Middle
Pleistocene assemblages of Cova Bolomor and, to a lesser extent,
the early Upper Paleolithic assemblages from Klissoura layer V
in Greece (Starkovich, 2009) and l’Arbreda level H in Spain
(Maroto et al., 1996; Lloveras, 2010) than with coeval Mousterian sites.
Despite the limitations of the data, the rabbit assemblage from
Les Canalettes layer 4 shows a strong over-representation of
females and an absence of rabbit kittens. The prevalence of adult
females in the sample may reflect the procurement of solitary or
small groups of rabbits in warren patches. However, because the
presumably low-ranked kittens might have been discarded at the
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48
D. Cochard et al. / Quaternary International 264 (2012) 32e51
Fig. 15. Sex-ratio in the rabbit assemblage from layer 4 based on a plot of the trochlear
breadth versus the distal breadth of distal humeri. The two ellipses and dashed lines
show the distribution of data points (excluding one outlier) and the best fit obtained by
Jones (2006) for females (upper ellipse) and males (lower ellipse) using a modern
reference collection of domestic rabbits. Note that rabbit females tend to be larger than
males.
5. Conclusion
Fig. 14. Possible human bite marks on an unidentified long bone shaft fragment
attributed to a leporid species (layer 4).
point of capture and because this class of individuals is preferentially affected by density-mediated attritiondincluding carnivore
ravagingdwarren-based mass harvesting of rabbits cannot be
entirely ruled out in the present case, particularly if procurement
occurred outside of the birthing season. For these reasons, mass
harvesting remains a plausible hypothesis for explaining the origin
of the layer 4 assemblage.
Table 8
Percentages of unfused long bone epiphyses in layer 4 at Les Canalettes. Percentages
based on very small samples (10) are shown in italic.
Age class
Element
Immatures
proximal radius
(0e3 months) distal humerus
sub-total
Sub-adults
proximal femur
(0e5 months) distal tibia
proximal ulna
sub-total
Very young to
distal radius
young adults proximal humerus
(0e9 months) distal femur
proximal tibia
sub-total
Grand Total
Age of
full fusion
# of # unfused % unfused
ends
>3 months
>3 months
57
24
81
7
15
12
34
10
3
6
11
30
145
>5 months
>5 months
>5 months
>9
>9
>9
>9
months
months
months
months
2
1
3
3
2
3
8
1
2
3
1
7
18
3.5
4.2
3.7
42.9
13.3
25.0
23.5
10.0
66.7
50.0
9.1
23.3
12.4
In Europe and southwest Asia, assemblages pre-dating the
Upper Paleolithic generally indicate that animal proteins and fat
were mostly derived from medium- to large-sized ungulates. The
occupation from layer 4 at Les Canalettes contrasts with this
picture, as is the case for several of the earlier occupations from
Cova Bolomor. At Les Canalettes, a taphonomic study of the rabbit
assemblage from layer 4 suggests habitual consumption of this
species by Neandertals. This inference is based not only on the high
taxonomic representation of rabbit remains in this layer, but also on
specific patterns of damage observed on bone elements (e.g., the
presence of cut-marked bones and shaft cylinders). Mortality and
sex-ratio data were also used to examine methods of rabbit
procurement. In layer 4, the data appear consistent with the
procurement of solitary or small groups of rabbits in warren
patches, although the possibility of warren-based mass hunting of
rabbits cannot be excluded.
The densities of animal species likely affected how human groups
exploited their environment in the past. Given the general focus on
medium- and large-sized ungulates during the Middle Paleolithic,
the recurrent use of a fast small-sized species as a source of food at
Les Canalettes may appear anomalous, particularly given that leporid skeletal elements were not exploited for symbolic purposes
prior to the Upper Paleolithic. The presence of leporids at Les
Canalettes might have been influenced by the function of the site,
the seasonality and duration of the occupations, as well as the sex
and age composition of the human groups, among others (e.g., Bietti,
2000; Brugal, 2000; Cochard and Brugal, 2004). As pointed out by
Binford (2001), the social context is a critical variable influencing
subsistence. This aspect should receive considerable attention in
future studies of small fast game assemblages, especially those
focusing on dietary intensification. The faunal patterns uncovered at
Les Canalettes emphasize the existence of complex foraging patterns
among Neanderthal groups, perhaps not unlike those documented
in coeval and later modern humans.
Both Les Canalettes and Cova Bolomor show signs of
consumption of fast small-sized taxa that anticipate similar
assemblages by tens of thousands of years. Although the former
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D. Cochard et al. / Quaternary International 264 (2012) 32e51
sites may be viewed as failed or anecdotal attempts toward dietary
intensification, they indicate that diet breadth changed in nonlinear ways over time and that foraging strategies were more
varied during the Middle and early Late Pleistocene than generally
appreciated (Morin, 2012). These sites also demonstrate that the
dietary use of small fast game species was not limited to anatomically modern humans and was potentially a habitual practice
among some archaic humans. This leads to the conclusion that the
exploitation of fast small-bodied taxa was largely constrained by
social and environmental factors rather than by biological differences between human populations.
Acknowledgments
We would like to thank Ruth Blasco and an anonymous reviewer
for their valuable comments. These comments considerably
improved the interpretations presented in this paper.
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