Dhole (Cuon alpinus) as a bone acumulator and new taphonomic agent. The case of the Noisetier Cave...

305

Mallye et al.

Article JTa138 All rights reserved. *E-mail: [email protected]

2012

Journal of Taphonomy

PROMETHEUS PRESS/PALAEONTOLOGICAL NETWORK FOUNDATION (TERUEL)

VOLUME 10 (ISSUE 3-4)

Available online at www.journaltaphonomy.com

Noisetier Cave (French Pyrenees) has yielded Mousterian artefacts associated with numerous faunal

remains. The faunal spectrum is dominated by chamois and ibex followed by red deer and bovids. A

previous taphonomic analysis underlined the occurrence of two distinct types of bone accumulations.

The red deer, bovid and a part of the ibex remains have been accumulated by Neanderthal. We suspected

that the bearded vultures were responsible for the chamois and some of the ibex remains. The study of

the carnivore remains illustrated the abundance of teeth and to a lesser extent bones attributed to both

Dhole (Cuon alpinus) as a Bone Accumulator and

New Taphonomic Agent?

The Case of the Noisetier Cave

(French Pyrenees)

Jean-Baptiste Mallye* Université Bordeaux 1, PACEA, UMR5199 Avenue des Facultés, 33405 Talence, France

Sandrine Costamagno

CNRS TRACES, UMR 5608, Université Toulouse 2 – Le Mirail Maison de la Recherche, 5

allées A. Machado, 31058 Toulouse cedex 9, France

Myriam Boudadi-Maligne Université Bordeaux 1, PACEA, UMR5199 Avenue des Facultés, 33405 Talence, France

Audrey Prucca

Les loups du Gévaudan, Sainte-Lucie, 48100 Saint-Léger-de-Peyre, France

Véronique Lauroulandie Université Bordeaux 1, PACEA, UMR5199 Avenue des Facultés, 33405 Talence, France

Céline Thiébaut

UMR 5608 –TRACES, Maison de la Recherche, 5 allées A. Machado,

31058, Toulouse cedex 9, France

Vincent Mourre UMR 5608 - TRACES et Inrap - Direction Interrégionale Méditerranée, 561, rue Étienne

Lenoir - KM Delta, 30900 Nîmes, France

Journal of Taphonomy 10 (3-4) (2012), 305-338.

Manuscript received 15 March 2012, revised manuscript accepted 15 November 2012.

mailto:[email protected]

306

Dhole as a bone accumulator at Noisetier Cave

published that characterise the bone

assemblages created by various carnivores.

Fewer studies have discussed the role of

large canids in the formation and

modification of bone assemblages or their

modifications (Binford, 1981; Haynes, 1983;

Klippel et al., 1987; Stiner, 1994, 2004;

Castel, 2004; Fosse et al., 2004; Campmas

& Beauval, 2008; Castel et al., 2010;

Esteban-Nadal et al., 2010; Yravedra, 2011).

The studies on bone destruction of

large and medium ungulates by large

carnivores are mainly inferred from kill

sites or scavenging sites (Haynes, 1980;

Richardson, 1980; Brain, 1981;

Blumenschine, 1988; Hill, 1989; Fosse,

1994; Nasti, 2000; Prucca, 2003; Castel,

2004; Fosse et al., 2004; Campmas &

Beauval, 2008). Most of the taphonomic

studies devoted to coproscopic analysis of

carnivore scats deal with small vertebrate

remains (Andrews & Nesbit-Evans, 1983;

Payne & Munson, 1985; Denys et al., 1992;

Denys, 2011; Andrews, 1990; Schmitt &

Juell, 1994; Mondini, 2000; Matthews,

2006; Lloveras et al., 2008a, b; Montalvo et

Introduction

Caves and rock shelters could have

alternatively been occupied by hominids and

carnivores. Consequently, stone tools and

faunal remains associated with occupants

could be mixed by post-depositional

processes (e.g. Villa & Bartram, 1996; Villa

& Soressi, 2000). This is a potential source

of important admixtures leading to

misinterpretations for instance in terms of

hominine hunting strategies or the

recognition of site function. Ongoing

researches in taphonomy aim to characterise

the accumulation produced by each bone

collectors (carnivores, raptors, humans) in

order to interpret the archaeological record.

The literature concerning damage caused by

hyenas is extremely prolific: this is due to

their ability to modify bone assemblages and

to accumulate bones inside their dens

(Brain, 1981; Haynes, 1983; Blumenschine,

1988; Hill, 1989; Cruz-Uribe, 1991;

Capaldo & Blumenschine, 1994; Pokines &

Kerbis Peterhans, 2007). In the last decade,

numerous actualistic studies have been

young and adult Cuon alpinus individuals. The identification of shed milk teeth demonstrates that this

carnivore used the cave as a nursery den. According to several authors dholes never bring back carcasses to

their dens in order to protect their offspring from other carnivores. However they tend to select an area

inside their den to defecate. We analysed modern scats of wolf in order to constitute a taphonomic

referential. Our results strongly suggest that most of the digested remains from the Noisetier Cave come

from dhole scats. This carnivore can be considered, as Binford previously suggested, as a bone accumulator

and consequently as a new taphonomic agent. Given the numerous sites where the fossil remains of this

carnivore were identified we argue that the dholes could have biased the composition of faunal spectrums

and maybe our understanding on human subsistence.

Keywords: DHOLE, CUON, COPROCOENOSIS, TAPHONOMY, NOISETIER CAVE, BONE

ACCUMULATION, DIGESTION, LATE PLEISTOCENE.

307

Mallye et al.

the frequency of digested bones is quite

comparable with that of chamois, but

anthropogenic marks are abundant. The

body part representation for the ibex shows

an intermediate pattern between the red deer

and the bovids on the one hand, and the

chamois on the other hand (Costamagno et

al., 2008). In summary, red deer and bovids

remains are mostly the result of a hunt by

the Mousterian people even if some of them

were secondly scavenged by carnivores. A

part of the ibex assemblage seems to have

been accumulated by Mousterian people

whereas the other part was introduced by a

non human accumulator, presumably the

bearded vulture.

In this paper we address the question

of the relation between the occurrence of

digested bones and the presence of different

carnivores such as the wolf (Canis lupus),

the dhole (Cuon alpinus) and the leopard

(Panthera pardus). In the light of new

actualistic studies, we test the hypothesis

previously expounded by Costamagno and

colleagues (2008) on the origin of chamois

assemblage.

Sites and Material

Noisetier Cave

Noisetier Cave (French Pyrenees) is a small

cave dwelled at a height of 825m a.s.l. and

at 145m above the Aure Valley at the

bottom of which flows the river Neste

(Figure 1). It is of 20m deep and measures 3

-4m in width and 3-6m in height. M. Allard

al., 2008; Lloveras et al., 2011) while only a

few studies concern the characterization of

ungulates bone accumulations created by

faecal deposits of large carnivores (Binford,

1981; Payne & Munson, 1985; Klippel et

al., 1987; Barja-Nunez & Corona, 2007;

Esteban-Nadal et al., 2010; Stiner et al.,

2012). The results of these studies lead them

to the same conclusions regarding the small

size of the remains, the relatively high

percentage of partially digested remains and

the possible occurrence of tooth marks.

Since its first excavation at the

beginning of the 1990’s, Noisetier cave was

interpreted as a hunting camp specialized on

chamois (Rupicapra rupicapra) and ibex

(Capra ibex) hunt (Jaubert & Bismuth,

1993). The taphonomic analysis conducted

by Costamagno and colleagues (2008) led to

modify the initial assumption regarding the

function of the site. In this study, the authors

demonstrate that a large part of the chamois

remains as well as a significant percentage

of the ibex material were not accumulated

by Neanderthal. Because of the presence of

abundant digested remains, the low

percentage of tooth marks on bones, a large

amount of short bones and cancellous bone

portions, Costamagno and colleagues (2008)

proposed that the bearded vulture (Gypaetus

barbatus) could be one of the possible

accumulators of chamois bones in the cave.

In contrast, red deer (Cervus elaphus) and

bovids are mainly represented by shaft

fragments. Short bones and cancellous bone

portions are underrepresented for these taxa

and butchery marks are extremely abundant

while carnivore marks are rare. For the ibex,

308


(Capreolus capreolus - 0.05%) and horse

(Equus caballus - 0.02%). Carnivores

represent less than 8% of number of

identified specimen for large and medium-

sized mammals. They are mostly

represented by foxes (Vulpes vulpes - 1.5%)

and bears (Ursus arctos - 3%). Weasel

(Mustela nivalis - 1%), dhole (Cuon alpinus

- 0.9%), leopard (Panthera pardus - 0.4%),

wolf (Canis lupus - 0.4%), polecat (Mustela

putorius - 0.05%) and Eurasian badger

(Meles meles – 0,02%) are represented by

fewer remains. Small games are mainly

composed of marmot (Marmotta marmotta -

3.6%) and hare (Lepus sp. - 1%).

Microvertebrates (rodents and birds)

remains are extremely abundant.

All coordinated remains and

anatomically and/or taxonomically identified

remains recovered after sieving operation

were analysed. The total number of identified

remains (NISP) at Noisetier cave is 4421

amongst which 1755 remains are attributed to

chamois. Inside the chamois assemblage, 482

remains were re-analysed in order to provide

a better characterization of the non-human

predation marks.

Les Loups du Gévaudan Park

Les Loups du Gévaudan Park is located in the

French department of Lozère (Figure 1). Since

2009, the authors have undertaken actualistic

taphonomic experimental studies there. The

aims of these studies are to characterise the

coprocoenosis generated by wolves from a

taphonomic and a micromorphologic

perspective. The wolves are fed with

first excavated the site in 1987 and 1992

(Allard, 1993, 1994). Since 2004 V.

Mourre, S. Costamagno and C. Thiébaut are

conducting new excavations and studies.

The excavation has yielded a Mousterian

industry with Discoïd and Levallois

debitage on local raw materials (Mourre et

al., 2008a,b; Thiébaut et al., 2012), bone

retouchers (Mallye et al., 2012) and human

remains (Maureille et al., 2007). Based on

AMS radiocarbon dates and

biochronological data (microfaunal and

macrofaunal remains), the archaeological

levels are attributed to a temperate phase of

the OIS 3 (see Mourre et al., 2008b for

details). The faunal spectrum is clearly

dominated by the chamois (Rupicapra

rupicapra - 64.4%), followed by the ibex

(Capra pyenaica - 12.7%), and to a lesser

extent by the red deer (Cervus elaphus -

9.3%), bovids (Bos / Bison - 1.5%), roe deer

Figure 1. Location of Noisetier cave and Les

Loups du Gévaudan Park.

309

Mallye et al.

In order to characterise the

coprocoenosis generated by wolves, 85

droppings of Mongolian wolf were

recovered. The scats were run through a

2mm mesh sieve. All skeletal and dental

elements as well as the residual fragments

of scats were collected and provide an

assemblage of 1,314 remains. In 12 scats,

no bone could be identified. The largest

number of remains recovered in a single

scat is 125 while the minimal number of

remains is 1. On average each scat yielded

15 remains. No microvertebrate remains

were identified. The percentage of identified

remains varies from 0 to 75% with an

average of 7.2%. Amongst the identified

remains, 105 were anatomically identified

and assigned to an ungulate class size.

At the Gévaudan, legs are rarely

given to the wolves. In order to characterize

the destruction pattern for these elements,

we decided to punctually feed the wolves

with ten sheep legs. All the legs were

composed of carpals / distal row of tarsals,

metapodials and phalanges with sesamoids.

After consumption the scats were

systematically sampled throughout the week

of experimentation. They were subsequently

run through a 2mm mesh sieve to collect the

bones. Out of the ten legs given to the

wolves, only 64 bones were recovered.

These remains represent only four legs.

The overall material collected at the

Gévaudan represents nearly 7,000 bones.

Most of the remains come from the

excavation of the dunging area (81,9%)

whereas the remaining part was recovered

after sieving of the scats.

ungulates carcasses (calf, beef, pork, lamb) cut

into quarters (about 1.8kg per wolf per day).

Head and leg ends (metapodials, phalanges

and sesamoids) are rarely given. Occasionally

the whole carcasses of pig, sheep or red deer

are distributed.

Two populations of wolves retained

our attention for taphonomic studies

undertaken on the dunging areas. The first

pack is composed of 31 individuals from

Siberia whose weight varies from 30 kg

(females, N=16) to 50 kg (males, N=29). They

occupy a fenced area of 1100 square metres.

We excavated a part of a dunging area of 1

square metre and 5cm deep. The sediment was

screened using a 2mm mesh sieve. This

operation yielded a total faunal assemblage of

1,285 specimens. The second sample comes

from an enclosure of 17,000 square metres

that include 3 packs composed of 45

individuals from Mongolia. The first pack is

composed of 14 males and 9 females, the

second is composed of 6 males and 7 females

and the last one comprises 3 males and 4

females. Average weights are identical to the

previous pack. We conducted an excavation of

2 square metres and 5cm deep. Using the

same screening method we collected 4,194

remains. Of the 5,479 bone specimens

recovered from the excavation of the two

dunging areas, a sample of 1,474 remains was

studied. This sample includes all identifiable

remains and/or longer than 1cm. Of this

sample, only 134 remains were identified

anatomically and/or taxonomically. The

species identified in this sample correspond to

those given during the feeding of wolves

except for the raven (Corvus corax).

310


bones, we used the criterion of dissolution

of the cortical to determine whether the

elements were digested or not.

In order to compare the chamois

sample from Noisetier cave with our

referential framework from Gévaudan, we

only took into account the anatomically and/

or taxonomically identified remains. This

quantification method unfortunately

prevents any comparison with data from

other studies on wolf scats (Esteban-Nadal

et al., 2010; Barja Núñez & Corona, 2007).

Concerning the identified bones

exhibiting traces of acid attacks, the intensity

of the digestion was assessed by considering

the proportion of bone attacked by the gastric

acids. Thus, codes ranging from 0 to 4 were

given based on the etched surface with respect

to the total surface area. The “0” value

corresponds to an intact remain, the “1” value

is given when up to 25% of the total surface is

modified by acid attacks, “2” value when 50%

is affected, “3” when 75% is affected and “4”

value when the entire surface is attacked.

For the spongy portions, we also

quantified the intensity of the dissolution of

the cortical bone. Stage 1 was assigned to

the remains with the cortical bone merely

dissolved whereas stage 2 corresponds to

the remains with a widening of the

cancellous bone.

The occurrence of tooth marks was

recorded according to the criteria described

by Binford (1981).

The occurrence of cupules on the bone

surfaces has also been recorded. These small

cupules are ellipsoid to rounded depressions

scattered on the surface of the bones.

Method

All dental and skeletal remains were measured

for maximum length (mm) to characterize the

fragmentation of the assemblage. Various

criteria were recorded for elements identifiable

and/or greater than 1cm.

The body part representation of the

chamois assemblage from Noisetier cave have

been compared with the data given by Robert

& Vigne (2002) for bearded vulture nests

using the percentage of representation defined

by Dodson & Wexlar (1979). Since some

anatomical parts were not present on the

carcasses given to the wolves at the Gévaudan

(e.g. heads or legs), the body part

representation could not be established on

these assemblages (scats and dunging area).

To fill this gap, we compared the anatomical

profile computed of the chamois assemblage

from Noisetier Cave with the results obtained

by Esteban-Nadal and colleagues (2010) on

wild wolf and by Stiner and colleagues (2012)

on Mountain lion using the percentage of

NISP for elements grouped per body parts:

skull (cranium + mandibule), teeth, vertebrae,

ribs (ribs + costal cartilage), girdles (scapula +

pelvis), long bones (humerus + radius + ulna +

femora + tibia), metapodial (metacarpals +

metatarsial + undefined metapodial), articular

bones (carpals + tarsals + patella + sesamoids)

and phalanges.

The criteria identified by d'Errico

and Villa (1997) were used during the

identification of digested bones. Digestion

is characterized by the appearance of holes

and by a slimming of the fracture edges and

general polishing of the bones. For spongy

311

Mallye et al.

Rates Mus spretus Crocidura russula Corrected pc/c 156.12 134.05

f+h/md+mx 54.55 37.84

t+r/f+h 78.21 64.29

Results

Noisetier Cave

1. Bone collectors in Noisetier Cave

A) Cuon

At Noisetier Cave, the reconsideration of

the faunal material led to the identification

of 48 remains of dhole including twenty

deciduous teeth (Figure 2). On several teeth

We used the categories of digestion

marks first described by Robert and Vigne

(2002) and later developed by Marin Arroyo

and colleagues (2009, 2011) to establish a

first quantification of the damages observed

on the phalanges.

In addition to this quantitative study

of acid attacks, we sought to highlight the

characteristic pattern of degradation for

various skeletal elements.

Figure 2. Noisetier cave. Deciduous teeth of dhole (Cuon alpinus). 1a, b, c, d: fourth upper deciduous

teeth. 2a, b, c: third upper deciduous teeth. 3: second lower deciduous incisor. 4: third lower deciduous

incisor. 5: lower deciduous canine. 6a, b: fourth lower deciduous teeth. 7a, b: third lower deciduous

teeth. 8a, b: second lower deciduous teeth.

312


compared to wolf deciduous teeth. The

proportion between the size of the metacone

and the size of the paracone is comparable to

what is observed on adult upper carnassial.

Furthermore, the small cusp located between

the paraconule and the protocone is less

pronounced on the teeth from Noisetier Cave

than on wolf deciduous teeth.

The shape of the Ud4 from Noisetier

Cave differs from the shape of wolf

deciduous teeth. The crown portion that

surrounds the metacone is very angular on

wolf Ud4, whereas it is more rounded on

the Ud4 from Noisetier Cave. On the wolf

Ud4, accessory cusps disposed on the mesio

-lingual edge of the crown can be observed,

while these additional cusps are absent on

the Ud4 from Noisetier Cave. On the wolf

Ud4, the mesial edge is straight, whereas it

is rounded on the Ud4 from Noisetier Cave.

A “V”-shaped depression, located in the

middle of the buccal edge, separates the

metacone from the paracone on wolf Ud4,

whereas it is absent on the Ud4 from

Noisetier Cave.

All these criteria/characteristics lead

us to attribute the complete assemblage of

canid deciduous teeth from Noisetier Cave

to the dhole (Cuon alpinus).

Carnivore marks are very rare on

dhole remains. Tooth marks were identified

on only one bone and a fragment of tooth

shows digestion marks (Table 1).

B) Wolf

At Noisetier Cave, the wolf is only

represented by adult remains. Only 23 remains

could be attributed to this carnivore and it is

(NISP = 16), a root resorption can be

identified due to exfoliation process. This

process is typical of shed deciduous teeth

(Hillson, 2005). We identified incisors

(Figure 2: 3 & 4), canines (Figure 2: 5),

second (figure 2: 8), third (Figure 2: 2 and

7) or fourth deciduous (Figure 2: 1 and 6)

teeth indicating various stages of

maturation.

The canid deciduous teeth identified

at Noisetier Cave were compared (1) to

modern wolf deciduous teeth from Italy and

Portugal (NI = 5), and (2) to upper

Pleistocene teeth of adult dhole from

Malarnaud Cave (Muséum d’Histoire

naturelle de Bordeaux). According to

Hillson (2005), in canids, the morphology

of deciduous teeth is similar to the

morphology of permanent teeth.

All the canid deciduous teeth from

Noisetier Cave are smaller but look more

robust than wolf milk teeth. From a

morphological point of view, several

differences in the shape of the teeth can be

observed on the fourth lower (Ld4), the

third upper (Ud3) and the fourth upper

deciduous teeth (Ud4).

The talonid of the Ld4 presents two

cusps in wolves, while a single cusp occurs

on the canid teeth from Noisetier Cave. The

occurrence of a single cusp on the talonid of

the lower carnassials of adults is a feature

that characterises the genus Cuon (Ewer,

1998). Thus, the morphology of the Ld4

from Noisetier Cave closely matches the

morphology of dhole lower carnassials.

The Ud3 in Noisetier Cave

assemblage have a small and lower metacone

313

Mallye et al.

Bo

dy p

arts

Lar

ge

can

id

(Dh

ole

or

Wo

lf)

Dh

ole

W

olf

L

eop

ard

Tota

l N

ISP

T

ooth

mar

ks

Dig

esti

on

mar

ks

NIS

P

Tooth

mar

ks

Dig

esti

on

mar

ks

NIS

P

Tooth

mar

ks

Dig

esti

on

mar

ks

NIS

P

Tooth

mar

ks

Dig

esti

on

mar

ks

Sku

ll

- -

- -

- -

- -

- -

- -

-

Per

man

ent

teet

h

9

- 1

19

- 1

- -

- 1

0

- -

38

Dec

idu

ou

s te

eth

- -

- 2

0

- -

- -

- 1

- -

21

Ver

teb

rae

4

1

1

2

- -

2

- -

2

1

- 1

0

Rib

s 3

- -

- -

- -

- -

- -

- 3

Ste

rneb

rae

1

- -

- -

- -

- -

- -

- 1

Hu

mer

us

- -

- 1

1

- -

- -

- -

- 1

Rad

ius

- -

- -

- -

- -

- -

- -

-

Uln

a 1

1

- -

- -

- -

- -

- -

1

Car

pal

s -

- -

- -

- -

- -

- -

- -

Met

acar

pal

s -

- -

3

- -

2

- 1

- -

- 5

Pel

vis

-

- -

1

- -

- -

- -

- -

1

Fem

ora

-

- -

- -

- 1

1

- -

- -

1

Pat

ella

e 4

1

1

- -

- -

- -

- -

- 4

Fib

ula

1

- -

- -

- 1

- -

- -

- 2

Tib

ia

- -

- -

- -

- -

- -

- -

-

Tar

sals

1

1

1

0

- -

3

- 1

1

- -

5

Met

atar

sals

-

- -

1

- -

2

- -

1

- -

4

Ph

alan

ges

6

- -

- -

- 1

1

1

1

5

2

- 2

2

Ses

amo

ids

3

- 1

1

- -

- -

- 4

- 1

8

Un

iden

tifi

ied

met

apo

dia

ls

- -

- -

- -

1

- -

- -

- 1

Tota

l 3

3

4

5

48

1

1

23

2

3

24

3

1

12

8

Ta

ble

1.

Nu

mb

er o

f id

enti

fied

rem

ain

s (N

ISP

), n

um

ber

of

too

th m

ark

s a

nd

nu

mb

er o

f d

iges

ted

ma

rks

for

each

ca

rniv

ore

fro

m N

ois

etie

r

cave

ass

emb

lag

e.

314


exceed 2cm long (Figure 4A). Among the

482 remains of chamois we re-examined,

85% present digestion marks (NISP = 410).

More than the three thirds of the digested

material are less than 2cm long and do not

exceed 4cm long (Figure 4B).

B) Body part representation

The large amount of compact bones and

phalanges was one of the main arguments used

to infer the major role played by the bearded

vulture in the chamois accumulation

(Costamagno et al., 2008). In addition chamois

long bones are only represented by epiphysis

fragments while the red deer, bovid and horse

long bones are mostly composed of shaft

fragments (Costamagno et al., 2008). When

comparing the body part profile for the

chamois from Noisetier Cave with that from

modern bearded vulture nests (Robert & Vigne,

2002) several differences can be noticed

(Figure 5). It is particularly relevant regarding

the proportion of short bones, scapulae, pelvis,

femurs and patellae (Figure 5).

difficult to discuss the body part profile (Table

1). It is however worth noting that elements of

the feet are abundant. Compared to the dhole,

neither shed milk teeth nor remains of

juveniles were identified. Tooth marks are

present on two bones and three digested

remains have been recognised namely a first

metacarpal, a talus and a first phalange.

C) Leopard

The occurrence of leopard in the

assemblage is attested by a few remains

(Table 1). They mostly consist of teeth and

foot bones. Only one deciduous tooth was

discovered indicating the presence of at

least one juvenile. The roots of the tooth are

still present which implies that this

individual died in the cave. Tooth marks

were identified on vertebrae and two

phalanges. A sesamoid is partially digested.

2. Coprolites

A large number of orange-white coloured

and parallelepiped remains with chalky and

dense texture were unearthed from the

sediment of the cave. Except for the colour,

which could be the consequence of post-

depositional processes, their aspect is closely

similar to that of coprolites (Figure 3).

3. Ungulates

A) Size of the remains

At Noisetier Cave, the overall studied

material is composed of 4421 remains

(taxonomically and/or anatomically

identified). Of this sample, 1994 remains

show digestion marks and nearly the three

quarters (74,5%) of the material do not

Figure 3. Noisetier cave: Fragments of coprolites.

315

Mallye et al.

Figure 4. Ungulate size remains according to size-classes. Digested remains in black, intact in grey. A:

complete Noisetier cave ungulate assemblage. B: Sample of chamois assemblage. C: Gévaudan sample.

Figure 5. Body part representation of chamois compared to that given by Robert & Vigne (2002) in

bearded vulture nest.

316


significant difference exists between the

two sample (Spearman’s Rho test R =

0,864, P<0,005).

C) Tooth marks

The frequency of tooth marks (Table 2

and Table 3) is significantly higher

(26.7%) in the chamois sample we

revaluated than the frequency results

obtained in the previous study (3%,

Costamagno et al., 2008). It should be

noted that this percentage is calculated by

dividing the number of chamois remains

bearing such marks (NISP=127) by the

total number of chamois remains

considering in this study and excluding

teeth (NISP=476).

When considering the anatomically

identified remains, vertebrae, phalanges

compact bones (short bones) are the three

most common elements identified in the

chamois sample while the bones of the

girdles (scapulae + pelvis) are poorly

represented. Comparing the same data with

that from mountain lion scats (Stiner et al.,

2012) some major differences could be

identified in terms of the proportion of ribs,

articular bones and phalanges. The

differences are statistically significant

(Spearman’s Rho test R= -0,28, P>0,5).

However, identical characteristics are found

in the body part profile from wild wolf scats

(data from Esteban-Nadal, 2010) body part

profile (Figure 6). No statistically

Figure 6. Body part representation of chamois compared to that given by Esteban-Nadal and colleagues

(2010) in wild wolf scats.

317

Mallye et al.

Gévaudan sample Noisetier Cave

Scats Dunging area Chamois sample

Thinning 68 80 269

Cortical dissolution**

Stage 0 12 6 28

Stage 1 35 57 172

Stage 2 32 34 207

Digestion intensity

Stage 0 24 28 69 Stage 1 20 29 54 Stage 2 11 17 92 Stage 3 16 16 70 Stage 4 34 44 191

Tooth marks** Digested 10 16 109 Not digested 4 6 18

Cupule** 18 29 176

Table 2. Characteristic of chemical and carnivore attacks in Gévaudan sample and chamois sample from

Noisetier cave. ** implies that teeth are not counted.

Body part representation Noisetier Cave: Chamois sample

NISP Digestion

marks Tooth

marks Complete

remains

Skull 8 7 2 - Teeth 6 3 - 3 Vertebrae 45 35 7 - Ribs 7 4 1 - Costal cartilage - - - - Scapula 5 5 1 - Humerus 4 4 1 - Radius 2 1 1 - Ulna 6 5 1 - Carpals 32 31 7 11 Pelvis 7 6 1 - Femora 70 58 30 - Patellae 26 24 9 6 Tibia 19 10 5 - Metapodial 10 7 2 - Tarsal 45 41 19 7 Phalanges 147 129 35 72 Sesamoids 42 39 5 32 Long bones 1 1 - -

Total 482 410 127 131

Table 3. Studied sample of chamois remains from Noisetier Cave according to body parts, number of

digested remains, number of tooth marks and number of complete remains.

318


207). Among the 410 digested chamois

remains, 65.6% show a slimming of their

fracture edges and 42.9% present cupules

on their surface (Table 2).

E) Patterns of bone destruction

- Talus

On the talus, digestion marks are always

located near the articular surfaces (Figure

7). When the degree of digestion becomes

more intense, gastric acid attacks increase

and dig the bone around the sulcus tali.

While the general shape of the bone is

preserved, compact bone disappears and

cancellous bone begins to be dug therefore

creating punctures.

D) Digestion

More than eighty percent (85%) of the

chamois sample we analysed shows

indisputable digestion marks (Table and

Table 3). The analysis of chamois digested

bones from Noisetier Cave reveals that

63.6% of them are characterised by chemical

attacks affecting more than half of their

surface (Digestion intensity, stages 3 and 4).

Almost all of digested chamois

remains from Noisetier Cave assemblage

present a loss of cortical tissue (NISP =

379). Acid attacks have dissolved only the

compact bone (stage 1, NISP= 172) but in

most cases, they have also affected spongy

bone, creating perforations (stage 2 NISP =

Figure 7. Partially-digested talus from Noisetier Cave (left) and Gévaudan sample (right). Scale bar is 1cm.

319

Mallye et al.

- Femora

Femur fragments are found in larger quantity in

Noisetier Cave assemblage. They are mostly

represented by heads which may include a shaft

fragment gradually tapering away from the

articular surface (Figure 10). This kind of

morphology (called “morphologie en clou”) has

been previously described in bearded vulture

nest accumulation (Robert & Vigne, 2002) and

observed by Costamagno and colleagues (2008).

On the distal parts of the femur, acid attacks are

located on around both sides of the condyle

surfaces. Acid attacks can also create small

perforations of the bone (Figure 11).

- Tibiae

Tibiae are often identified by the tibia tuberosity

to which is attached a portion of shaft. Acid

attacks are located around the tuberosity. On the

shaft portion a slimming of edge the fracture

edges can be observed (Figure 12).

- Calcaneus

At Noisetier Cave, two identifiable portions

of calcaneus were identified. The first is the

calcaneus head and the second, the posterior

face of the talus attached to a part of the

shaft (Figure 8). Both present acid attacks

and a slimming of the fracture edges.

- Patella

The patellae are found in large number

inside the Noisetier Cave assemblage. Thus,

the comparison between the least destroyed

patella and the more digested ones allows a

good understanding of the destruction

pattern. On the former, the digestion marks

are located on the periphery of the medial

and lateral articular surfaces. While the acid

attacks intensified, the areas located on the

periphery of the articular surfaces are

gradually removed. The patellae take then

the form of a dolly (Figure 9).

Figure 8. Comparison between partially-digested calcanei from Noisetier cave (left) and Gévaudan

sample (right). Scale bar is 1cm.

320


Figure 10. Partially digested femora

(proximal parts, “morphologie en clou”)

from Noisetier Cave (left) and Gévaudan

sample (right)

Figure 9. Partially digested patellae from Noisetier cave (left) and Gévaudan sample (right). Scale bar is

1cm.

321

Mallye et al.

Figure 11. Partially-digested femurs (distal parts) from Noisetier Cave (left) and Gévaudan sample

(right). Scale bar is 1cm.

Figure 12. Partially-digested tibias from Noisetier Cave (left) and Gévaudan sample (right). Scale bar

is 1cm.

322


Gévaudan sample

1. Coprolites

In the field (Figure 14), the scats become

quickly dry. Because of the weathering

and/or trampling effects they break up in

small fragments measuring less than 1cm

long releasing the faunal material they

contain and then get buried in the soil.

These fragments of scats were recovered

after sieving operation from both scats

and soil from the dunging area (Figure

15).

- Phalanges

In the Noisetier cave assemblage,

digestion marks observed on the first and

second phalanges are located on the shaft

and are associated with a slimming of the

fracture edges. In addition acid attacks

perforate the distal part of the first

phalanges (Costamagno et al., 2008).

Most of the third phalanges are

characterised by acid attacks located

around the articular surface. Digestion

marks are always recorded on broken

phalanges and code 3.6 (Robert & Vigne,

2002) or 36 (Marín-Arroyo et al., 2009) is

the most frequent (Figure 13).

Figure 13. Partially-digested phalanges from Noisetier Cave (left) and Gévaudan sample (right).

Scale bar is 1cm.

323

Mallye et al.

from the dunging area (3966/5479). In other

words, remains from the scats are longer

than those from the dunging area. In the

three samples, the remains have a maximum

length of 5cm. The average percentage of

complete elements is 1.9% in the scat

sample and 1.7% in the dunging area

sample. Of the 1314 remains from the scats

sample, only 18 were complete or almost

complete. Only 26 remains were complete

among the 1474 studied remains from the

dunging area sample.

3. Digestion

The analysis of digested remains from the

dunging area sample (NISP = 852) shows

that 56,6% of the remains is affected by

chemical attacks on more than the half of

the surface. This proportion reaches 61.7%

in the scat sample (Table 2).

Almost all the digested remains from

the Gévaudan assemblage (85.8% in

dunging area and 82.7% in scats) present a

2. Size of the remains

The small size of the remains recovered at

the Gévaudan testifies of their high

fragmentation state. More than 80% of the

remains measured less than 2cm long and

never exceed 5cm long (Figure 4). At the

Gévaudan, only 57.8% of the dunging area

sample (NISP = 1474) present digestion

marks. Among the 134 identified remains

from the dunging area excavation, 79.1%

present digestion marks and more than 85%

of these remains are smaller than 3cm. Of

the 105 identified remains from the scat

sample, 77.1% show clear digestion marks

and 97.5% measured less than 3cm. In both

sample, the size of the digested remains

does not exceed 5cm long. Remains smaller

than 1cm represent nearly 50% of the scat

sample (654/1314), while they constitute

more than 70% of the material excavated

Figure 14. Gévaudan: post-depositional evolution

of wolf scats.

Figure 15. Fragments of scats (=coprolites) re-

covered after sieving operation at Gévaudan.

324


- Femora

Femur heads are poorly represented in the

Gévaudan sample. Among the 7,000 bones

recovered only one femoral head was

identified (Figure 9). A remaining fragment

of shaft is still attached above it. Digestion

marks are visible around the articular surface

of the head and fracture edges are slim.

However, distal parts of the femur are well

represented in the Gévaudan sample and all

of them present digestion marks. These

attacks are located near the articular surfaces

of the condyles on both sides. The cortical

bone has disappeared and the acid attacks

have frequently perforated the spongy bone

creating small holes (Figure 10).

- Tibia

In the Gévaudan assemblage, fragment of

tibia tuberosity have also been identified.

Once again a portion of the shaft is present

and acid attacks located around the

tuberosity cause a slimming of the shaft on

the fracture edges (Figure 11).

- Sheep’s’ legs

The overall material collected is

fragmented; only twenty-five remains out of

64 are complete or almost complete –

mainly carpals, tarsals and sesamoids. All

identified remains show digestion marks.

The slimming on the fracture edges is

observed on 34 remains. On 46 remains, the

cortical bone is dug by acid attacks (code

2). We identified tooth marks on only four

remains mainly consisting of notches. Of all

the phalanges identified after consumption

by wolves, only a third phalange is

loss of cortical tissue. Slimming of the

fracture edges of bones is high in wolf scats

(84%) and on the remains from the dunging

area (75.4%). The occurrence of cupules on

remains varies from 17.1% in the wolf scats

to 21.6% in the dunging area (Table 2).

4. Patterns of bone destruction

- Talus

In the Gévaudan sample, the tali are complete

or almost complete. The same pattern of

destruction previously noticed in the Noisetier

Cave assemblage is encountered in the

Gévaudan sample (Figure 7).

- Calcaneus

No complete calcaneus was recovered in the

studied sample. Two identifiable portions

were identified. The first is the calcaneus

head and the second is the posterior face of

the talus attached to a part of the shaft. Both

present acid attacks and a slimming of the

fracture edges. Notches are identifiable on

both sides of the tubercular process

indicating that calcanei are crushed by

wolves (Figure 8).

- Patella

The patellae are found in large number in

the Gévaudan samples. On the least

destroyed specimens, the digestion marks

are located on the periphery of the medial

and lateral articular surfaces. Subsequently,

we noticed the loss of the apex. While the

acid attacks are rising, the areas located on

the periphery of the articular surfaces are

gradually removed. The patella then takes

the form of a dolly (Figure 9).

325

Mallye et al.

(size > 40mm). These large remains are

related to long bone shaft fragments

(splinters) of the red deer, bovid, horse and

to a lesser extent to the ibex. The

examination of the surfaces on these large

remains shows many anthropogenic marks

and a lack of carnivore marks as previously

noticed (Costamagno et al., 2008). This

indicates that they were introduced in the

cave by Neanderthals rather than by a non-

human predator.

The distribution of digested remains

in the chamois sample from Noisetier Cave

is quite similar to that observed in the

Gévaudan sample: bones larger than 40mm

are absent and bones between 10 and 20mm

are the most represented.

2. Digestion

The chamois digested remains from the

Noisetier Cave are smaller in size than the

anatomically or taxonomically remains

identified in the Gévaudan sample.

On only 57.8% of the 1474 remains

from the excavation of the dunging area

digestion marks have been recognised. The

high frequency of spongy bone fragments in

the sample (28%) could explain this low

percentage. As previously noticed by

Esteban-Nadal and colleagues (2010), the

structure of spongy bones restricts the

identification of digestion marks. When

taking into account only the identified

bones, the percentage of partially-digested

remains is quite similar across the different

samples. The lowest percentages are

observed on bones from the scat sample

(77%) and from the dunging area (79%)

complete. The other thirty-three specimens

of phalanges are fragmented. Finally, in

addition to slimming of the fracture edges,

we noticed that acid attacks begin to

perforate the distal part of the first and

second phalanges (Figure 12).

Discussion

Comparison between Noisetier Cave and

wolf accumulation

1. Size of the remains

The size of remains from the Gévaudan

scats is comparable to that observed by

Esteban-Nadal and colleagues (2010) in

wild wolf scats and to that observed by

Payne and Munson (1985) in dog faeces.

However, we demonstrated that the

maximal length of bones from the scats is

significantly longer than that observed on

remains from the dunging area. This can be

explained by post depositional

fragmentation due to trampling by wolves

as previously noticed on coprolites

fragments. It is highly probable that this

phenomenon could have affected the bones

in the past. Thus, the distribution by size

class observed for the bone recovered from

the dunging area is a better reflection of

what can be found in the fossil record than

the maximum length of the remains from

the scats. The distribution of remains by

size classes in the Gévaudan sample is quite

similar to that observed in Noisetier Cave

for the total ungulates sample. However, in

the latter larger remains have been recorded

326


75.3% in scats sample and 75.4% in

dropping area). In the sample of Iberian

wolf scats (Esteban-Nadal et al., 2010) the

slimming of edges is more limited (29.5%)

but the quantification methods applied by

these authors are different. They calculate

the proportions from whole assemblage,

including the unidentified remains whereas

in our study only the identified remains

were considered.

3. Tooth marks

Tooth marks have been recognised on

26.7% of the chamois bones from

Noisetier Cave. This percentage is higher

than the percentage previously observed

by Costamagno and colleagues (2008).

Lower but similar frequencies on

identified remains have been noticed on

the Gévaudan sample, ranging from

13.3% in the scats sample to 16.4% in the

dunging area sample. These frequencies

are higher than frequencies described in

other studies (2.4% Esteban-Nadal et al.,

2010; 7.9% Barja Núñez & Corona,

2007). However, in these studies, the

occurrence of tooth marks is calculated

according to the total number of remains,

including unidentified remains.

4. Bone completeness

In the Gévaudan samples, the percentages

of complete bones are very low: 1.8% in the

dunging area and only 1.34% in the scat

sample. Both are lower than percentages

calculated by Esteban-Nadal and colleagues

(2010) in their sample (2.2%). This

difference could be related to the variability

whereas 84.7% of the chamois remains

from Noisetier Cave show digestion marks.

All these percentages are nonetheless

smaller than those obtained by Esteban-

Nadal and colleagues (2010) [92.7%] or

Barja Núñez and Corona (2007) [99.1%] in

their study of wolf scats.

Regarding the digested elements,

about the two-thirds (65.6%) of chamois

remains in Noisetier Cave are characterised

by chemical attacks that affect more than

50% of the total surface (Stage 3 and 4).

Similar proportions are found in the

Gévaudan sample (61.7% in scats sample

and 56.6% in dunging area). Concerning the

proportions of the different codes, there is

no significant difference between the two

samples of digested bones from Gévaudan

(chi-square calc = 1,676, ddl = 4, P<0,1).

However, the comparison between the

Noisetier cave assemblage and the two

samples from Gévaudan highlights some

differences. In the bone sample from

Noisetier Cave, the intensity of digestion

was further coded as 2 or 4. In the

Gévaudan sample, the highest percentages

are found for bones coded 1 and 4.

Almost all of the digested bones

from Noisetier cave and Gévaudan showed

evidence of cortical dissolution (sensu

porosity by Esteban-Nadal et al. 2010). In

both cases, this kind of attack is substantial

since not only it affects the cortical bone but

also digs into the spongy bone.

Among the 410 digested chamois

remains, 65.6% shows a slimming of edges

fracture. This proportion is slightly higher

in the samples from Gévaudan (respectively

327

Mallye et al.

& Corona’s study (2007) on Iberian wild

wolf scats.

The morphology of tibial crest is quite

similar in both Noisetier Cave and the

Gévaudan assemblages. Similarly, a

comparison between the patellas identified at

Noisetier Cave and in the Gévaudan sample

shows the same patterns of alteration.

Chemical attacks recorded on phalanges in

Noisetier cave resemble closely to chemical

attacks observed on phalanges digested by

bearded vulture described by Robert & Vigne

(2002). This is especially true if we take into

account the perforation located at the distal

end. However, this pattern have also been

recognised by Klippel and colleagues (1987)

in their study on wolf scats samples and by

Chase (1990) on coyote scat sample. To date,

our sample of phalanges consumed by wolves

is too small to consider a statistical study of

the destruction pattern as it has been

conducted for other assemblages (Robert &

Vigne, 2002; Marin Arroyo et al., 2009; Marin

Arroyo & Margalida, 2011).

Bearded vulture nest or carnivore den?

The numerous patellae identified both at

Noisetier cave and in the Gévaudan sample

could provide information about the

accumulator of chamois. In mammals, the

extremities of the main thigh muscles are

located on the patella. This bone is thus

encased in a large muscle mass and is

articulated with the femur and the tibia by

ligaments. It is therefore likely to be

swallowed on fresh carcasses. Consequently,

we suggest that the patella is more accessible

between the different referential

frameworks. Comparing to Noisetier Cave

(37%), the number of anatomically and/or

taxonomically bones that are complete in

Gévaudan sample is lower (17-19%). This

implies that chamois remains at Noisetier

cave are less fragmented than remains

identified in wolf scats.

5. Cupules

Cupules have been identified in both the

Gévaudan sample (17-21%) and the

chamois remains from Noisetier Cave

(42,9%). It is difficult to establish clearly

their origin. Cupules can be created by the

action of gastric acids or be the result of

gnawing marks that are latter modified

during the digestion process as previously

noticed by Stiner and colleagues (2012).

6. Patterns of bone destruction

To complete our quantitative study of

digestion marks left on the ungulates

remains we compared the destruction of the

most frequently identified remains such as

carpals and tarsals but also phalanges,

patellae and tibia tuberosities in Gévaudan

sample and chamois remains from Noisetier

Cave. The remains presented in this study

belong primarily to small ungulates such as

sheep and chamois. These remains are very

small, and so can easily be ingested by

wolves.

The tali from the Noisetier Cave and

the Gévaudan sample show the same pattern

of degradation caused by carnivore teeth

and by chemical attacks. Identical patterns

of degradation are described in Barja Núñez

328


1981; Lyman, 1994; Mondini, 1995;

Mondini & Muñoz, 2008): residual

assemblage that results from a kill site and

in-situ consumption, transported assemblage

that corresponds to an accumulation of

bones from the kill site and scatological

assemblage which is the “result of the by-

products of bone ingestion and are

deposited within faeces” (Mondini &

Muñoz, 2008: 54). At Noisetier Cave, the

chamois assemblage is characterised by the

abundance of cancellous bone portions and

by the lack of long bones shaft fragments

(see Costamagno et al., 2008 for details and

proportions). In addition there is a large

amount of coprolites fragments and most of

the bones are digested. Consequently it is

highly probable that the chamois

assemblage derives from a scatological

assemblage. This assertion is reinforced by

the similarities observed between the body

part representations of the chamois sample

on the one hand and of the remains

identified in carnivores’ scats on the other

hand. In the last case the predators

unintentionally deposit faunal remains via

their scats. At Noisetier cave, the large

quantity of digested bones could be the

result of abundant scats indicating a long-

term use of the cavity by carnivores.

Which carnivore?

Eight species of carnivores have been

identified at Noisetier Cave. The smaller

taxa (weasel and polecat) cannot be

responsible of the chamois accumulation in

the cave as they prey mostly on small

to primary consumers such as terrestrial

carnivores rather than to scavengers such as

bearded vultures. In addition, the involvement

of a terrestrial carnivore in the constitution of

the chamois assemblage of Noisetier cave is

further proven by the high percentage of tooth

marks. We also demonstrate that the body part

representation of chamois assemblage of

Noisetier Cave is more closely similar to the

one provided by Esteban-Nadal and

colleagues (2010) in wolf scats than that given

by Robert and Vigne (2002) for bearded

vulture. The occurrence of coprolites has been

used to attest the occupation of the cave by the

bearded vulture (Costamagno et al., 2008).

Identical fragments were identified after the

excavation of the dunging area of wolves. In

such context the fragmentation is caused by

post-depositional processes such as

weathering or trampling by wolves for

instance. In the case of Noisetier Cave it is

likely that coprolite fragments derived from

the frequentation of the cave by inhabitants.

As previously noticed for other sites (e.g.

Stiner, 2004), the numerous fragments of

coprolites identified in Noisetier Cave indicate

long-term use of the cave as a den by

carnivores. An analysis of the geochemical

composition of these fragments compared to

modern coprolites of different species (wolf,

bearded vulture, dhole) would fully confirm

their origin.

Which kind of bone assemblage for chamois

in Noisetier cave?

Three kinds of accumulations can be related

to carnivores feeding activity (Binford,

329

Mallye et al.

marks and digestion marks. The body part

representation could hardly be discussed

due to the small size of the sample.

However, it is important to notice that some

of the undetermined canid remains belongs

to small individuals that could be related to

dholes.

Wolf is only represented by adult

remains. Only one milk tooth has been

attributed to the Leopard at Noisetier Cave.

The roots of this tooth are fully developed

without exfoliation process. If the cave was

used as a den by one of the carnivore over a

long period, remains of cubs should be

found in greater quantity as it is noticed in

cave used by bears in the context of

hibernation (e.g. David, 2002) or in hyaena

dens (e.g. Fosse, 1994). This is not the case

at Noisetier Cave.

The identification of shed milk teeth

of dhole leads us to make several

observations. To our knowledge, this is the

first time that dhole milk teeth are described

in the fossil record. According to the tooth

eruption wear pattern given by Hillson

(2005: 242) for canid, deciduous teeth are in

place at one to two months and permanent

teeth begins to erupt at five to six months to

be finally in place at six months. Few data

are available for dhole teething. However,

Sosnovskii (1967: 122) provides information

on dental eruption sequence from dholes kept

at the Moscow Zoo: “Teething begins on

Days 15 to 16, when the incisors of the upper

jaw appear. They are followed on Days 20-

21 by those of the lower jaw; canines appear

on Days 21-25; molars on Days 25-30; and

premolar on Days 30-35.” Thus, it appears

vertebrates. Large ungulate remains have

been identified in fox and badger burrows

as the result of scavenging habits of these

two carnivores (Mallye et al., 2008; Castel

et al., 2011). In these cases, such remains

are found in low quantities compared to

small vertebrates and are rarely digested. At

Noisetier Cave, cut marks have been

recorded on the only badger remain and

attest of human consumption. Traces of

digestion as well as tooth marks have been

observed on some of the fox remains and

attest of a non-human accumulation. The

large amount of bear deciduous teeth

indicate that the cave was occupied by bear

during hibernation.

Thus, three carnivores could be

responsible of chamois accumulation: the

dhole, the wolf and the leopard.

The comparison between the

chamois sample and the data provided on

wolf scats (Esteban-Nadal et al., 2010) and

mountain lion (Stiner et al., 2012) shows

greater similarities with canid scats than that

from lion. This is especially true when

looking at the proportion of ribs (higher in

lion scats) and that of short bones and

phalanges (higher in wolf scats).

On the other hand, the morphology

of digested chamois bones is quite similar to

what is observed in the Gévaudan

assemblage. Thus it is very probable that a

canid has created the chamois bone

assemblage in the cave through the

accumulation of its scats.

The analysis conducted on the

remains of three carnivores allowed us to

identify predation marks namely tooth

330


Fox, 1984). However scats are frequently

deposited in latrines and could be located in

the den (Fox, 1984; Johnsingh, 1982). Of

the 66 scats analysed, Borah and colleagues

(2009) report that bones are identified in

43.9% of the scats analysed. Then, the

accumulation of scats can produce a

coprocoenosis. Based on these ethological

data, it is very likely that the dholes used the

cave as a den and accidentally accumulated

ungulate bones by defecating.

The dhole as a new taphonomic agent

Up to now extensive research has been

conducted to characterise the bone

accumulations produced by large carnivores

such as hyaenas. Several factors led us to

underestimate the ability of dhole to create

notable bone accumulations in fossil context.

In some sites, it is possible that the remains

of dhole were not identified or were

mistaken with wolf remains. Recent studies

have identified the remains of fossil dholes

in fossil record and finally contributed to fill

this gap in our knowledge (Boudadi-

Maligne, 2010; Perez-Ripoll et al., 2010;

Pionnier-Capitan et al., 2011).

Few authors among which Simons

& Ettel (1970) for Southeast Asian sites

but also Binford (1988) in the case of

Vaufrey cave (layer VIII) have suggested

that dholes played a significant role in site

formation processes. In the latter case,

this hypothesis has been challenged

(Grayson & Delpech, 1994). However, the

work undertaken on Leporid remains

(Cochard, 2004, 2007) from the same

that tooth eruption pattern of dholes is quite

similar to that of other canids. Dhole are

known to occupy caves, rocky shelters or

burrow of their own digging to mate, give

birth and to breed the cubs (Davidar, 1974;

Johnsingh, 1982; Fox, 1984). The birthing

season occurs between November and March

with a peak in December (Davidar, 1975).

The nursing period lasts about 50 to 60 days

and thereafter adults regurgitate meat to feed

the cubs inside the den. Cubs help the adults

to kill prey at 6 to 8 months old. The

occurrence of shed milk teeth at Noisetier

Cave indicates that dhole have occupied the

cave. Based on sequences of tooth eruption,

and according to ethological data we may

conclude that this occupation may have

spanned for a long period, probably over 5

months of the year during the birthing and

weaning seasons.

Dholes are medium-size carnivores

that are widespread in South East Asia from

India through China to southern Java

(Durbin et al., 2004). Body weight for adult

males averages 20kg and 15kg for adult

females. This canid lives in pack ranging

from 5 to 11 individuals (Johnsingh, 1982)

but up to 40 individuals have been recorded

(Fox, 1984). Dholes hunt in pack and can

kill medium and large ungulates weighting

more than 200kg (Karanth & Sunquist,

1995; Venkataraman et al., 1995; Wang &

MacDonald, 2009; Thinley et al., 2011) as

well as smaller preys such as hare, birds and

rodents (Cohen, 1978; Venkataraman et al.,

1995; Borah et al. 2009).

Only occasionally dholes bring back

carcasses to their den (Johnsingh, 1982;

331

Mallye et al.

the bone accumulation process. The

constitution of a coproscopic referential

framework for this canid becomes

imperative. The continuation of such studies

will improve our understanding on human

paleoecology or human/carnivore

interactions as well as site formation

processes and site functions.

Acknowledgments

The authors would like to thank Jordi Rosell,

Enrique Baquedano, Ruth Blasco and

Edgard Camaros for organising the

International congress “Hominid Carnivore

Interaction during the Pleistocene” in Salou

where this paper was originally presented.

This research is granted by La Fondation des

Treilles project. Excavation at Le Noisetier

Cave is supported by the Ministère de la

culture et de la Communication and also the

Conseil Général des Hautes-Pyrénées. We

also would like to thank Jessica Lacarrière

who provides us supplementary data that

have stimulated our research, Walter E.

Klippel who provides us data on digested

bones. We want to thank Melvin E.

Sunquist, Leon Durbin, Lonnie I. Grassman,

Kate Jenks, Jan Kamler and Jimmy Borah

for their helpful information on Dhole scats.

Thanks to Luca Sitzia for his help on figure

16, many thanks to Le Parc des Loups du

Gévaudan to welcome us, with special

mention to Sylvain Macchi. Thanks to the

two anonymous reviewers for their helpful

comments that improve a previous version

of the paper.

layer tends to support the hypothesis of

Binford (1988). Based on recent works

(Brugal & Boudadi-Maligne, 2011; Perez-

Ripoll et al., 2010) and additional

bibliographic data, we can notice that the

dhole is widely distributed in western and

central Europe on the edge of the

mountain ranges (Figure 16). In several

sites, anthropogenic marks have been

identified on dhole remains (Perez-Ripoll

et al., 2010) attesting of human origin. In

other sites, such as Amalda cave (Altuna

et al., 1990) or Boquete de Zafarraya

(Barroso Ruiz, 2003; Barroso & de

Lumley, 2005) or Gabasa (Blasco-Sancho,

1995) dhole remains are interestingly

associated with ungulates that are

identical in size (e.g. chamois and ibex) to

those identified in Noisetier cave. Hence,

bone accumulations identical to the one

described in this paper could possibly be

identified in these sites.

Conclusion

The results obtained from both Noisetier

Cave and the Gévaudan assemblage offer

new research perspectives in taphonomy

concerning the role of carnivores in the site

formation processes. We suggest that

medium-sized carnivores such as dholes

could have had a real impact on site

formation processes and that dhole could

now be considered as a potential

taphonomic agent. This carnivore has been

recognised in many Pleistocene

assemblages and could have contributed to

332


Fig

ure

16

. P

ale

og

eog

rap

hic

dis

trib

uti

on o

f dh

ole

s d

uri

ng

th

e P

leis

toce

ne

in W

este

rn E

uro

pe.

333

Mallye et al.

Arambourg, C. (1958). Les gros mammiferes des couches tayaciennes. In (Alimen, H., Arambourg,

C. & Schreuder, A., eds.) La grotte de

Fontéchevade. Paris: Archive de l'Institut de Paléontologie Humaine, pp.185-229.

Argant, A. (1991). Carnivores quaternaires de

Bourgogne. Document du laboratoire de Géologie

de Lyon, Lyon.

Barja Núñez, I. & Corona-M, E. (2007). El análysis de

excretas desde la etología y la arqueozoología. El caso del lobo ibérico. In (Corona-M., E. & Arroyo-

Carales, J., eds.) Human and Faunal Relationships

Reviewed: An Archaeozoological Approach. BAR International Series, pp.113-121.

Barroso Ruiz, C. (2003). El Pleistoceno superior de la

Cueva del Boquete de Zafarraya. Arqueología Monographías, Junta da Andalucía, Consejería de

Cultura, Sevilla.

Barroso Ruiz, C. & de Lumley, H. (2006). La Grotte du Boquete de Zafarraya (Málaga, Andalousie).

Junta de Andalucía, Sevilla.

Baryshnikov, G. F. & Tsoukala, E. (2010). New analysis of the Pleistocene carnivores from

Petralona Cave (Macedonia, Greece) based on the

collection of the Thessaloniki Aristotle University. Geobios, 43: 389-402.

Binford, L. R. (1981). Bones: Ancient Men and Modern Myths. Academic Press, New-York.

Binford, L. R. (1988). Etude taphonomique des restes

fauniques de la grotte Vaufrey, Couche VIII. In (Rigaud, J.-P., ed.) La grotte Vaufrey:

Paleoenvironnement - Chronologie - Activité

humaines. Mémoires de la Société Préhistorique Française, pp.535-563.

Blasco Sancho, F. (1995). Hombres, fieras y presas.

Estudio arqueozoológico y tafonómico del yacimiento del Paleolítico Medio de la Cueva de Gabasa 1

(Huesca). Monografías Arqueológicas, Zaragoza.

Blumenschine, R. J. (1988). An experimental model of the timing of Hominid and carnivore influence on

archaeological bone assemblages. Journal of

Archaeological Science, 15: 483-502. Bonifay, M.-F. (1971). Carnivores quaternaires du Sud

-Est de la France. Mémoire du Muséum National

d'Histoire Naturelle, Paris. Borah, J., Deka, K., Dookia, S. & Prasad Gupta, R.

(2009). Food habits of dholes (Cuon alpinus) in

Saptura Tiger Reserve, Madhya Pradesh, India. Mammalia, 73: 85-88.

Bouchud, J. (1951). Etude paléontologique de la faune

d'Isturitz. Mammalia, 15(4): 184-203.

Bouchud, P. & Bouchud, J. (1953). La faune des

grottes des Orciers et de Cottier. Bulletin de la

Société Préhistorique Française, 50: 444-457.

References

Thiollay, J.M. (1978). Les rapaces d’une zone de

contact savane-forêt en Côte d’Ivoire :

spécialisations alimentaires. Alauda, 46: 147-170.

Winkler, A. J., Denys, C. & Avery, M. (2010).

Rodentia. In: Werdelin, L., Sanders, W. J. (Eds.):

Cenozoic Mammals of Africa. University of

California Press, Berkeley, Los Angeles, London. Chap. 17: 263-304.

Adam, K. D. (1959). Mittelpleistozäne Caniden aus

dem Heppenloch bei Gutenberg (Württemberg). Stuttgarter Beiträge zur Naturkunde, 27: 1-46.

Allard, M. (1993). Fréchet-Aure, Grotte du Noisetier.

In Bilan Scientifique 1992 de la Direction Régionale de Affaires Culturelles de Midi

Pyrénées. Ministère de la Culture, pp.113-114.

Allard, M. (1994). Fréchet-Aure. Grotte de Peyrère 1 (du Noisetier). In Bilan Scientifique 1993 de la

Direction Régionale de Affaires Culturelles de

Midi Pyrénées. Ministère de la Culture, pp.156. Altuna, J. (1973). Fauna de mamíferos del yacimiento

prehistórico de Los Casares (Guadalajara). In

(Barandiarán, I., ed.) La Cueva de Los Casares (en Riba de Saelice, Gudalajara). Madrid: Excavaciones

Arqueológicas en España, pp.97–116.

Altuna, J. (1981). Restos óseos deI yacimiento prehistórico deI Rascaño (Santander). In (González

Echegaray, F. & Barandiarán, I., eds.) El

Paleolitico superior de la cueva del Rascaño (Santander). Centro de Investigacion y Museo de

Altamira, pp.223-269.

Altuna, J. (1983). Hallazgo de un cuón (Cuon alpinus Pallas) en Obarreta, Gorbea (Vizcaya). Kobie,

XIII: 141-158.

Altuna, J. (1986). The mammalian faunas from the prehistoric site of La Riera. In (Straus, L. G. &

Clark, G., eds.) La Riera Cave. Stone Age Hunter–

Gatherer Adaptations in Northern Spain. Tempe:

Anthropological Research Papers, pp.237-274.

Altuna, J., Baldeón, A. & Mariezkurrena, K. (1990). La

Cueva de Amalda (Zestoa, País Vasco): Ocupaciones Paleolíticas y Postpaleolíticas. Fundación José

Miguel de Barandiarán, San Sebastián.

Andrews, P. & Nesbit Evans, E. M. (1983). Small mammal bone accumulations produced by

mammalian carnivores. Paleobiology, 9(3): 289-307.

Andrews, P. J. (1990). Owls, caves and fossils: predation, preservation and accumulation of small

mammal bones in caves, with an analysis of Pleistocene cave faunas from Westbury-sub-

Mendip, Somerset, UK. University of Chicago

Press, Chicago.

334


Champagne, F. & Espitalié, R. (1981). Le Piage, Site préhistorique de Lot. Mémoire de la Société

Préhistorique Française, Paris.

Chase, P. G. (1990). Tool-Making Tools and Middle Paleolithic Behavior. Current Anthropology, 31(4):

443-447.

Clot, A. (1987). La grotte de la Carrière (Gerde,

Hautes-Pyrénées). Stratigraphie et paléontologie

des Carnivores. Société Ramond, Bagnère-de-

Bigorre. Cochard, D. (2004). Les léporidés dans la subsistance

paléolithique du Sud de la France. Unpublished

PhD thesis, Université Bordeaux 1. 354 p. Cochard, D. (2007). Caractérisation des apports de

Léporidés dans les sites paléolithiques et

application méthodologiques à la couche VIII de la grotte Vaufrey. In Congrès du centenaire : Un

siècle de discours scientifique en Préhistoire.

XXVIè congrès préhistorique de France - Avignon, 21-25 septembre 2004. Paris: Société Préhitorique

Française, pp.467-480.

Cohen, J. A. (1978). Cuon alpinus. Mammalian Species, 100: 1-3.

Cordy, J.-M. (1983). Découverte de Cuon alpinus

europaeus Bourguignat dans le quaternaire de Belgique. In (Poplin, F., ed.) La Faune et l'Homme

préhistoriques. Paris: Mémoire de la société préhistorique française, pp.49-54.

Costamagno, S., Robert, I., Laroulandie, V., Mourre,

V. & Thiébaut, C. (2008). Rôle du Gypaète barbu (Gypaetus barbatus) dans la constitution de

l'assemblage osseux de la grotte du Noisetier

(Fréchet-Aure, Hautes Pyrénées, France). Annales de Paléontologie, 94: 245-265.

Crégut-Bonnoure, E., Boulbes, N., Daujard, C.,

Fernandez, P. & Valensi, P. (2010). Nouvelles données sur la grande faune de l'Éémien dans le

sud-est de la France. Quaternaire, 21/3: 227-248.

Cruze-Uribe, K. (1991). Distinguish Hyena from Hominid Bone Accumulations. Journal of Field

Archaeology, 18(4): 467-486.

d'Errico, F. & Villa, P. (1997). Holes and grooves: the contribution of microscopy and taphonomy to the

problem of art origins. Journal of Human

Evolution, 33: 1-31. David, F. (2002). Les ours du Châtelperronien de la

grotte du Renne à Arcy-sur-Cure (Yonne). In

(Tillet, T. & Binford, L. R., eds.) L'Ours et l'Homme. Actes du colloque d'Auberives-en-

Royans, 1997. Liège: ERAUL, pp.185-192.

Davidar, E. R. C. (1974). Observations at the dens of

the dhole or Indian wild dog (Cuon alpinus).

Journal of the Bombay Natural History society, 71

(2): 183-187.

Boudadi-Maligne, M. (2010). Les Canis pléistocènes du Sud de la France : Approches biostratigraphique,

évolutive et biochronologique. Unpublished PhD

Thesis, Université Bordeaux 1. 446 p. Boule, M. & Villeneuve, L. d. (1927). La grotte de

l'Observatoire à Monaco. Archive de l'Institut de

paléontologie humaine, Paris.

Bourguignat, J.-R. (1875). Recherches sur les

ossements de Canidae constatés en France à l'état

fossile pendant la période quaternaire. Masson, Paris.

Brain, C. K. (1981). The hunters or the hunted? An

introduction to african cave taphonomy. The University of Chicago Press, Chicago & London.

Brugal, J.-P. & Boudadi-Maligne, M. (2011). Quaternary

small to large canids in europe: Taxonomic status and biochronological contribution. Quaternary

International, 243: 171-182.

Campmas, E. & Beauval, C. (2008). Consommation osseuse des carnivores : résultats de l'étude de

l'exploitation de carcasses de bieufs (Bos taurus)

par des loups captifs. Annales de Paléontologie, 94: 167-186.

Capaldo, S. D. & Blumenschine, R. J. (1994). A

quantitative diagnosis of notches made by hammerstone percussion and carnivore gnawing on

Bovid long bone. American Antiquity, 59(4): 724-748.

Cardoso, J. L. (1992). Présence de Cuon alpinus

europaeus Bourguignat, 1868 (Mammalia, Carnivora) dans le Pléistocène du Portugal.

Ciências da Terra (UNL), 11: 65-76.

Castaños, P. M. (1987). Los carnívoros prehistóricos de Vizcaya. Kobie (serie Paleoanthropología),

XVI: 7-38.

Castel, J.-C. (2004). L'influence des canidés sur la formation des ensembles archéologiques.

Caractérisation des destructions dues au Loup.

Revue de Paléobiologie, Genève, 23(2): 675-693. Castel, J.-C., Coumont, M.-P., Boudadi-Maligne, M. &

Prucca, A. (2010). Rôle des grands carnivores dans

les accumulations naturelles. Le cas des loups (Canis lupus) de l'Igue du Gral (Sauliac-sur-Célé,

Lot, France). Revue de Paléobiologie, Genève, 29

(2): 411-425. Castel, J.-C., Mallye, J.-B. & Oppliger, J. (2011). Les

petits carnivores dans leur abris temporaires: choix

des espèces et caractéristiques taphonomiques. Implications pour l'archéologie. In (Laroulandie,

V., Mallye, J.-B. & Denys, C., eds.) Taphonomie

des Petits vertébrés: Référentiels et Transferts aux fossiles. Actes de la Table Ronde du RTP

Taphonomie, Talence 20-21 octobre 2009. Oxford:

BAR International Series, pp.77-91.

335

Mallye et al.

Fosse, P. (1994). Taphonomie paléolithique: Les grands Mammifères de soleilhac (Haute-Loire) et

de Lunel-Viel (Hérault). Thèse de Doctorat de

l'université de Provence (Aix-Marseille I), Université de Provence - Aix-Marseille I. 257 p.

254 annexes.

Fosse, P., Laudet, F., Selva, N. & Warjac, A. (2004).

Premières observations néotaphonomiques sur des

assemblages osseux de Bialowieza (N. E.

Pologne): intérêts pour les gisements pléistocènes d'Europe. PALEO, 16: 91-116.

Fox, M. W. (1984). The Whistling Hunter. Field

Studies of the Asiatic Wild Dog (Cuon alpinus). State University of New York Press, Albany.

García, N. (2003). Osos y Otros Carnívoros de la

Sierra de Atapuerca. Fundacion Oso de Asturias, Ovideo.

García, N. & Arsuaga, J.-L. (1998). The carnivore

remains from the Hominid-bearing trinchera-Galería, Sierra de Atapureca, Middle Pleistocene

site (Spain). Geobios, 31: 659-674.

Grassman, L. I. J., Tewes, M. E., Silvy, N., J. & Kreetiyutanont, K. (2005). Spatial ecology and diet

of the dhole Cuon alpinus (Canidae, Carnivora) in

north central Thailand. Mammalia, 69(1): 11-20. Grayson, D. K. & Delpech, F. (1994). The evidence for

Middle Palaeolithic scavenging from Couche VIII, Grotte Vaufrey (Dordogne, France). Journal of

Archaeological Science, 21: 359-375.

Haynes, G. (1980). Prey bones and predators: Potential ecologic information from analysis of bone sites.

Ossa, 7: 75-97.

Haynes, G. (1983). A guide for differentiating mammalian carnivore taxa responsible for gnaw

damage to herbivore limb bones. Paleobiology, 9

(2): 164-172. Hill, A. (1989). Bone modification by Modern Spotted

Hyenas. In (Bonnichsen, R. & Sorg, M. H., eds.)

Bone modification. Orono, Maine: Center for the Study of the First Americans, pp.169-178.

Hillson, S. (2005). Teeth, 2nd edition. Cambridge

University Press, New York. Jánossy, D. (1986). Pleistocene Vertebrate Faunas of

Hungary. Developments in Palaeontology and

Stratigraphy. Elsevier Science, Amsterdam. Jaubert, J. & Bismuth, T. (1993). Le Paléolithique

moyen des Pyrénées centrales esquisse d'un

schéma chronologique et économique dans la perspective d'une étude comparative avec les

documents ibériques. In Actes du 118è Congrès

National des sociétés historiques et scientifiques.

Pau: CTHS, pp.9-26.

Jéquier, J.-P. (1975). Le Moustérien alpin. Révision

critique. Eburodunum Yverdon, 2: 7-130.

Davidar, E. R. C. (1975). 8. Ecology and Behavior of the Dhole or Indian Wild dog Cuon alpinus

(Pallas). In (Fox, M. W., ed.) The wild Canids.

Their Systematics, Behavioral Ecology and Evolution. Wenatchee: Dogwise Publishing,

pp.109-119.

Del Campana, D. (1923). Sopra un Cuon e una Mustela

del Quaternario di Equi (Alpi Apuane). Atti della

Reale Accademia Nazionale dei Lincei. Rendiconti

della Classe di scienze fisiche, matematiche e naturali, 32: 170-172.

Del Campana, D. (1954). Carnivori quaternari della

Tecchia e della Caverna di Equi nelle Alpi Apuane (Mustelidi, Canidi, Felidi). Palaeontographia

Italica, 44: 1-42.

Delpech, F. (1988). Les grands Mammifères, à l'exception des Ursidés. In (Rigaud, J.-P., ed.) La

Grotte Vaufrey Paléoenvironnement - Chronologie

- Activités Humaines. Mémoires de la Société Préhistorique Française, pp.213-289.

Denys, C. (2011). Des référentiels en taphonomie des

petits vertébrés: bilan et prespectives. In (Laroulandie, V., Mallye, J.-B. & Denys, C., eds.)

Taphonomie des Petits vertébrés: Référentiels et

Transferts aux fossiles. Actes de la Table Ronde du RTP Taphonomie, Talence 20-21 octobre 2009.

Oxford: BAR International Series, pp.7-22. Denys, C., Kowalski, K. & Dauphin, Y. (1992).

Mechanical and chemical alterations of skeletal

tissues in a recent Saharian accumulation of faeces from Vulpes rueppelli (Carnivora, Mammalia).

Acta Zoologica cracovensis, 35(2): 265-283.

Dodson, P. & Wexlar, D. (1979). Taphonomic investigations of owl pellets. Paleobiology, 5(3):

275-284.

Dufour, R. (1989). Les Carnivores pléistocènes de la caverne de Malarnaud (Ariège) : collection E.

Harlé, Museum d'Histoire Naturelle de

Bordeaux. Muséum d'Histoire Naturelle Bordeaux, Bordeaux.

Durbin, L. S., Venkataraman, A. B., Hedges, S. &

Duckworh, W. (2004). Dhole. Cuon alpinus (Pallas, 1811). In (Sillero-Zubiri, C., Hoffman, M.

& Macdonald, D. W., eds.) Canids: Foxes, Wolves,

Jackals and Dogs. 2004 Status Survey And Conservation Action Plan, IUCN/SSC Canid

Specialist Group. pp.210-219.

Esteban-Nadal, M., Cáceres, I. & Fosse, P. (2010). Characterization of a current coprogenic sample

originated by Canis lupus as a tool for identifying

a taphonomic agent. Journal of Archaeological Science, 37: 2959-2970.

Ewer, R. F. (1998). The carnivores. Cornell University

Press, Ithaca, New York.

336


Mallye, J.-B., Laroulandie, V. & Cochard, D. (2008). Accumulations osseuses en périphérie de terriers

de petits carnivores : les stigmates de prédation et

de fréquentation. Annales de Paléontologie, 94: 187-208.

Mallye, J.-B., Thiébaut, C., Mourre, V., Costamagno, S.,

Claud, E. & Weisbecker, P. (2012). The Mousterian

bone retouchers of Noisetier Cave: experimentation

and identification of marks. Journal of

Archaeological Science, 39(4): 1131-1142. Marín Arroyo, A. B., Fosse, P. & Vigne, J.-D. (2009).

Probable evidences of bone accumulation by

Pleistocene bearded vulture at the archaeological site of El Mirón Cave (Spain). Journal of


Marín Arroyo, A. B. & Margalida, A. (2012). Distinguish Bearded Vulture activities within

Archaeological Contexts: Identification guidelines.

International Journal of Osteoarchaeology, 22(5): 563-576.

Matthews, T. (2006). Taphonomic Characteristics of

Micrommals Predated by Small Mammalian Carnivores in South Africa: Application to Fossil

Accumulations. Journal of Taphonomy, 4(3): 143-161.

Maureille, B., Bruxelles, L., Colonge, D., Costamagno, S., Cravinho, S., Jeannet, M., Laroulandie, V.,

Thiébaut, C. & Mourre, V. (2007). Nouveaux vestiges humains moustériens de la Grotte du

Noisetier (Fréchet-Aure, Hautes-Pyrénées).

1833ème Réunion scientifique de la société d'Anthropologie de Paris (23-25 janvier 2008),

Marseille. Bulletin et Mémoires de la société

d'Anthropologie de Paris. Moigne, A.-M., Palombo, M.-R., Belda, V., Heriech-

Briki, D., Kacimi, S., Lacombat, F., de Lumley, M.

-A., Moutoussamy, J., Rivals, F., Quilès, J. & Testu, A. (2006). Les faunes de grands

mammifères de la Caune de l'Arago (Tautavel)

dans le cadre biochronologique des faunes du Pléistocène moyen italien. L'Anthropologie, 110:

788-831.

Moncel, M.-H., Brugal, J.-P., Prucca, A. & Lhomme, G. (2008). Mixed occupation during the Middle

Palaeolithic: case study of a small pit-cave-site of

Les Pêcheurs (Ardèche, south-eastern France). Journal of Anrthropological Archaeology, 27: 382-

398.

Mondini, M. (1995). Artiodactyl prey transport by foxes in Puna rock shelters. Current Anthropology,

36(3): 520-524.

Mondini, M. (2000). Tafonomia de abrigos rocosos de

la Puna. Formación de conjuntos escatológicos por

zorros y sus implicaciones arqueológicas.

Archaeofauna, 9: 151-164.

Johnsingh, A. J. T. (1982). Reproductive and social behaviour of the Dhole, Cuon alpinus (Canidae).

Journal of Zoology, London, 198: 443-463.

Kahlke, H. D. (1961). Revision der Säugetierfaunen der klassischen deutschen Pleistozän-Fundestellen

von Süßenborn, Mosbach und Taubach. Geologie,

10: 493-532.

Karanth, U. K. & Sunquist, L. E. (1995). Prey

Selection by Tiger, Leopard and Dhole in Tropical

Forests. The Journal of Animal Ecology, 64(4): 439-450.

Kawanishi, K. & Sunquist, M. E. (2008). Food habits

and activity patterns of the Asiatic golden cat (Catopuma temminckii) and the dhole (Cuon

alpinus) in a primary rainforest of Peninsula

Malaysia. Mammal Study, 33: 173-177. Klippel, W. E., Snyder, L. M. & Parmalee, P. W.

(1987). Taphonomy and Archaeologically

recovered Mammal Bone from Southeast Missouri. Journal of Ethnobiology, 7(2): 155-169.

Kumaraguru, A., Saravanamuthu, R., Brinda, K. &

Asokan, S. (2011). Prey preference of large carnivores in Anamalai Tiger Reserve, India.

European Journal of Wildlife Research, 57: 627-

637. Lloveras, L., Moreno Garcia, M. & Nadal, J. (2008b).

Taphonomic study of leporid remains accumulated by the Spanish Imperial Eagle (Aquila adalberti).

Geobios, 41: 91-100.

Lloveras, L., Moreno Garcia, M. & Nadal, J. (2011). Feeding the Foxes: An experimental study to

assess their taphonomic signature on Leporid

R em a in s . In t e rn a t io n a l Jo u rn a l o f osteoarchaeology, 22(5): 577-590.

Lloveras, L., Moreno-García, M. & Nadal, J. (2008a).

Taphonomic analysis of leporid remains obtained from modern Iberian Lynx (Lynx pardinus) scats.

Journal of Archaeological Science, 35: 1-13.

Lyman, R. L. (1994). Vertebrate Taphonomy. Cambridge University Press, Cambridge.

Malez, M. (1974). Noviji Rezultati Istrazivanja

Paleolitika u Velikoj pecini, Veternici i Sandalji. - Neue Ergebnisse der Paläolithikum-Forschungen

in Velika Pecina, Veternica und Sandalja

(Kroatien). Jugoslavenske Akademija Znanosti i Umjetnosti: 7-44.

Malez, M. (1975). Die Höhle Vindija - Eine neue

Fundstelle fossiler Hominiden in Kroatien. Bulletin Scientifique, Section A, 20: 139-140.

Malez, M. & Malez, V. (1989). The Upper Pleistocene

Fauna from the Neanderthal Man Site in Krapina

(Croatia, Yugoslavia). Geoloski vjesnik instituta za

geoloska istrazivanja u zagrebu i hrvatskog

geoloskob drustva, 42: 49-57.

337

Mallye et al.

Pokines, J. T. & Kerbis Peterhans, J. C. (2007). Spotted hyena (Crocuta crocuta) den use and taphonomy

in the Masai Mara National Reserve, Kenya.

Journal of Archaeological Science, 34: 1914-1931. Prucca, A. (2003). Caractérisation de l'impact des

Loups sur les ossements d'Herbivores (Cerfs de

Virginie, Orignaux, Bisons) : Étude des

modifications infligées par des Loups captifs et

sauvages Nord-Américains. Unpublished Master

thesis, Université de Provence. 135 p. Richardson, P. R. K. (1980). Carnivore Damage to

Antelope Bones and its Archaeological Implications.

Palaeontologica Africana, 23: 109-125. Robert, I. & Vigne, J.-D. (2002). The Bearded Vulture

(Gypaetus barbatus) as an Accumulator of

archaeological Bones. Late Glacial Assembalges and Present-day Reference Data in Corsica

(Western Mediterranean). Journal of

Archaeological Science, 29: 763-777. Schmitt, D. N. & Juell, K. E. (1994). Toward the

Identification of coyote Scatological Faunal

accumulations in Archaeological Contexts. Journal of Archaeological Science, 21(2): 249-262.

Simons, E. L. & Ettel, P. C. (1970). Gigantopithecus.

Scientific American: 77-85. Sosnovskii, I. P. (1967). Bredding the Red dog or

dhole Cuon alpinus at Moscow Zoo. International Zoo Yearbook, 7: 120-122.

Stiner, M. C. (1994). Honor among Thieves. A

zooarchaeological study of Neandertal ecology. Princeton University Press, Princeton.

Stiner, M. C. (2004). Comparative ecology and

taphonomy of spotted hyenas, humans, and wolves in Pleistocene Italy. Revue de Paléobiologie,

Genève, 23(2): 771-785.

Stiner, M. C., Munro, N. D. & Sanz, M. (2012). Carcass damage and digested bone from mountain lions

(Felis concolor): implications for carcass persistence

on landscapes as a function of prey age. Journal of Archaeological Science, 39: 896-907.

Thenius, E. (1954). Die Caniden (Mammalia) aus dem

Altquartär von Hundsheim (Niederösterreich) nebst Bemerkungen zur Stammesgeschichte der

Gattung Cuon. Neues Jahrbuch für Geologie und

Paläontolologie, 99: 230-286. Thiébaut, C., Mourre, V., Chalard, P., Colonge, D.,

Coudenneau, A., Deschamps, M. & Sacco-

Sonador, A. (2012). Lithic technology of the final Mousterian on both sides of the Pyrenees.

Quaternary International, 247: 182-198.

Thinley, P., Kamler, J. F., Wang, S. W., Lham, K.,

Stenkewitz, U. & Macdonald, D. W. (2011). Seasonal

diet of dholes (Cuon alpinus) in northwestern Bhutan.

Mammalian Biology, 76: 518-520.

Montalvo, C. I., Pessino, M. E. M. & Bagatto, F. C. (2008). Taphonomy of the bones of rodents

consumed by Andean hog-nosed skunks

(Conepatus chinga, Carnivora, Mephitidae) in central Argentina. Journal of Archaeological

Science, 35: 1481-1488.

Mourre, V., Costamagno, S., Bruxelles, L., Colonge, D.,

Cravinho, S., Laroulandie, V., Maureille, B., Thiébaut,

C. & Viguier, J. (2008a). Exploitation du milieu

montagnard dans le Moustérien final: La grotte du Noisetier à Fréchet-Aure (Pyrénées centrales

françaises). In (Grimaldi, S., Perrin, T. & Guilaine, J.,

eds.) Mountains environments in Prehistoric Europe. BAR International Series, pp.1-10.

Mourre, V., Costamagno, S., Thiébaut, C., Allard, M.,

Bruxelles, L., Colonge, D., Cravinho, S., Jeannet, M., Juillard, F., Laroulandie, V. & Maureille, B.

(2008b). Le site moustérien de la Grotte du

Noisetier à Fréchet-Aure (Hautes-Pyrénées) - premiers résultats des nouvelles fouilles. In

(Jaubert, J., Bordes, J.-G. & Ortega, I., eds.) Les

sociétés du Paléolithique dans un Grand Sud-Ouest de la France : nouveaux gisements,

nouveaux résultats, nouvelles méthodes. Mémoire

de la Société préhistorique française, pp.189-202. Nasti, A. (2000). Modification of Vicuña carcasses in

high-altitude deserts. Current Anthropology, 41(2): 279-283.

Nehring, A. (1891). Diluviale Reste von Cuon, Ovis,

Saiga, Ibex und Rupicapra aus Mähren. Neues Jahrbuch fiir Mineralogie, Geologie und

Paleontologie, 2: 107-155.

Pailhaugue, N. (1995). La faune de la salle Monique, Grotte de la Vache (Alliat, Ariège). Bulletin de la

société Préhistorique de l'Ariège, L: 225-289.

Payne, S. & Munson, P. J. (1985). Ruby an dhow many Squirrels? The destruction of Bones by Dogs. In

(Fieller, N. R. J., Gilbertson, D. E. & Ralph, N. G.

A., eds.) Palaeobiological Investigations. Research Design Method and Data Analysis. BAR

international Series, pp.31-39.

Pérez Ripoll, M., Morales Pérez, J. V., Sanchis Serra, A., Aura Tortosa, J. E. & Sarrión Montañana, I.

(2010). Presence of the genus Cuon in upper

Pleistocene and initial Holocene sites of the Iberia Penninsula: new remains identified in

archaeological contexts of the Mediterranean

region. Journal of Archaeological Science, 37(3): 437-450.

Pionnier-Capitan, M., Bémilli, C., Bodu, P., Célérier,

G., Ferrié, J.-G., Fosse, P., Garcia, M. & Vigne, J.-

D. (2011). New evidence for Upper Palaeolithic

dogs in South-Western Europe. Journal of


338


Trinkaus, E., Marks, A. E., Brugal, J.-P., Bailey, S. E., Rink, W. J. & Richter, D. (2003). Later Middle

Pleistocene human remains from the Almonda

Karstic system, Torres Novas, Portugal. Journal of Human evolution, 45: 219-226.

Venkataraman, A. B., Arumugam, R. & Sukumar, R.

(1995). The foraging ecology of dhole (Cuon

alpinus) in Mudumalai Sanctuary, southern India.

Journal of Zoology, London, 237: 543-561.

Villa, P. & Bartram, L. E. (1996). Flaked bone from hyena den. PALEO, 8: 143-159.

Villa, P. & Soressi, M. (2000). Stone tool in carnivore

sites: the case of Bois Roche. Journal of Archaeological Research, 56: 187-211.

Wang, S. W. & Macdonald, D. W. (2009). Feeding habits and niche partitioning in a predator guild

composed of tigers, leopards and dholes in a

temperate ecosystem in central Bhutan. Journal of Zoology, 277: 275-283.

Wetzel, R., Bosinski, G. & Filzer, P. (1969). Die

Bocksteinschmiede im Lonetal (Markung

Rammingen, Kreis Ulm). Veröffentlichungen des

Staatlichen Amtes für Denkmalpflege Stuttgart / A,

Vor- und Frühgeschichte, Stuttgart. Yravedra, J., Lagos, L. & Bárcena, F. (2011). A

taphonomic study of Wild Wolf (Canis lupus)

modification of horse bones in Northwestern Spain. Journal of Taphonomy, 9: 37-65.

Dhole (Cuon alpinus) as a bone acumulator and new taphonomic agent. The case of the Noisetier Cave...

Documents

Transcript of Dhole (Cuon alpinus) as a bone acumulator and new taphonomic agent. The case of the Noisetier Cave...