Mammal Rev. 1992, Volume 22, No. 2,87-96. Printed in Great Britain

Zooarchaeology and the biogeographical history of the mammals of Corsica and Sardinia since the last ice age JEAN-DENIS VIGNE C.N.R.S. ( URA 1415), Muse'um national d'Histoire naturelle, Laboratoire d'Anatomie compare'e, 55 rue de Buffon, F-75005 Paris, France

ABSTRACT This paper presents the main results of a 10-year zooarchaeological study on the origin of the present-day terrestrial mammals of Corsica. Thousands of teeth and bones from Corsican archaeological sites comprise an almost complete succession from the end of the Pleistocene ( c . 10,000 years BP) to the present. Human activities brought about the extinction of the entire autochthonous mammalian fauna and the gradual introduction of more than 25 taxa which constitute the present wild and domestic fauna. Such a complete turnover has also been recorded on other large Mediterranean islands and on some of the smaller ones. Preliminary data indicate how these events may have occurred in Corsica. This article emphasizes the important role which zooarchaeology can play in biogeographical research.

INTRODUCTION Zoological comparisons between continental and island faunas have helped to explain the process of colonization and extinction (e.g. MacArthur & Wilson, 1967). One of the most important ways of testing these processes is to investigate the actual succession of events in the course of time, which requires data from historical, palaeontological and archaeological studies. Archaeological bones have several advantages: 1.

2.

3.

4.

The periods represented generally post-date the last ice age, thereby providing a link between palaeontological and zoological data. Well-preserved archaeological assemblages, resulting from the activity of carni- vores, owls or man are generally much larger than palaeontological ones so that an approach at the population level is possible. Dating is usually uncomplicated because of their association with cultural remains, and because various physical techniques such as radiocarbon analysis can be applied. In more-favourable situations, it is possible to trace the history of a fauna with the precision of a few hundreds (Neolithic) to a few tens (Middle Ages) of years. They yield information about people and their relationships with animals, which helps us to understand the present-day structure of a fauna. For all these reasons, zooarchaeology has already contributed to our understanding

of the process of faunal turnover on several islands, especially in the case of birds on Pacific Ocean islands (Olson & James, 1982; James et al., 1987).

The aim of this article is to provide a brief outline of my findings in Corsica undertaken during the last 10 years.

MATERIAL AND METHODS The mammal remains discussed here come from more than 80 chronostratigraphic units (Table 1) shared among 38 Corsican sites (Fig. 1). The samples span the last nine

cu P 7

Tab

le 1

6.

Mai

n ch

arac

teri

stic

s of t

he si

ze a

nd co

mpo

sitio

n of

the z

ooar

chae

olog

ical

sam

ples

stud

ied

in C

orsi

ca a

nd u

sed

in th

e pre

sent

pap

er. N

umer

ous u

npub

lishe

d dat

a on

teet

h an

d bo

nes f

rom

stra

tigra

phic

al u

nits

hav

e no

t bee

n ta

ken

into

acc

ount

in th

e co

unts

Peri

od

Num

ber o

f N

umbe

r of

N

umbe

r of

id

entif

ied

Min

imum

num

ber o

f ind

ivid

ual m

amm

als

stra

tigra

phic

ve

rteb

rate

m

amm

al

units

re

mai

ns

spec

imen

s L

arge

* Sm

all?

M

icro

- T

otal

Rec

ent (

1400

BP-

pres

ent)

Lat

e M

iddl

e A

ges

(1 10

0-14

00 B

P)

Ear

ly M

iddl

e A

ges

(160

(r11

00 B

P)

Rom

an (2

100-

1600

BP)

Ir

on A

ge (3

000-

2 10

0 B

P)

Bro

nze A

ge (4

000-

3000

BP)

Lat

e N

eolit

hic

(500

0400

0 B

P)

Mid

dle

Neo

lithi

c (6

000-

5000

BP)

E

arly

Neo

lithi

c (7

200-

6000

BP)

Pr

e-N

eolit

hic

(970

0-72

00 B

P)

Tot

al

9 13 2 5 10

12

12 5 4 3 82

718

1829

4 27

503

416

1888

40

65

9633

73

37

2985

0 72

19

1081

97

617

8123

85

19

344

852

1206

40

9 1

3692

23

346

6879

5810

1

66

284

246

131

65

97

155

114

59 5

1271

9 42

20

29

35

32

70

101

775

360

1483

1631

65

5 30

6 10

0 10

9 13

1 16

20

6 21

3003

1706

91

6 57

2 15

5 10

9 26

0 24

1 23

5 84

0 38

6

5587

*Lar

ger t

han

Dog

. ?Fr

om F

ox to

Pro

lagu

s.

History of Corsican mammals 89

Fig. 1. Distribution of archaeological sites studied in Corsica and bathymeuy to illustrate the effects of past sea-level changes. The lowest seashore regression during the last glacial (Wiirm), did not exceed 100-1 10 m, remaining inside the -200 m line of this figure.

millennia and provide relatively complete series of observations from which to deduce faunal succession. So far, more than 100,000 teeth and bones of vertebrates have been examined, from which nearly 60,000 mammalian remains have been identified. Table 1 also shows that (i) Pre-Neolithic, Iron Age and the Roman periods are less well docu- mented than other periods, (ii) the history of micromammals is poorly documented for the Neolithic period, and (iii) with the exception of Prolagus in the oldest periods, small mammals (especially small carnivores) are poorly represented through the entire sequence.

This last observation may reflect not only the natural scarcity of some of these small mammals, but also the origin (and subsequent destruction) of the samples. With the exception of the Monte-di-Tuda site (see below), all the samples come from archaeological layers where man has been the main faunal collector. The biassing influence of man upon the composition of the fauna must be taken into account in any biogeographical reconstruction.

When possible (e.g. Monte di Tuda and S. M. Lavezzi), we have been able to conclude that the micromammal remains derive from pellets of the Barn Owl Tyto alba (the only owl permanently resident on the island). Modem Tyto alba pellets provide a significant picture of the natural micromammal assemblages in Corsica (Libois, 1984).

90 J.-0. Vigne

TERMINAL PLEISTOCENE TO THE BEGINNING OF THE NEOLITHIC Corsica and Sardinia are on the same geological microplate (Fig. l), and the straits between them, whose depth does not exceed 60 my periodically disappeared during each glacial marine regression, leaving a dry land passage connecting the two islands. Studies of the fauna of Corsica should therefore also consider the fauna of Sardinia, at least for the end of the Pleistocene and for the beginning of the Holocene.

Minor late- and post-glacial tectonic movements would not have been sufficient to create a bridge between the islands and the Tuscan mainland after the middle Pleistocene (Fig. 1; Conchon, 1976).

The Terminal Pleistocene fauna The site of Maccinaggio in Corsica and several upper Pleistocene sites in Sardinia provide clear evidence of the presence, between about 20,000 and 10,000 years b.p., of at least six endemic mammalian taxa (Sondaar et al., 1986; Vigne, 1988a, 1990). These are: Episoriculus ( = ‘Nesiotites’) similis (Hensel) on Sardinia and E . corsicanus (Bate) on Corsica, two large soricid insectivores; Cynotherium sardous Studiati, a canid the size of a Fox which shows an affinity with the Jackal (Eisenmann, 1990); Prolagus sardus (Wagner), an ochotonid lagomorph the size of a big Rat; Rhagamys orthodon (Hensel), a large Field Mouse (Muridae); Microtus ( Tyrrhenicola) henseli (Forsyth Major), a large Field Vole (Arvicolidae); and Megaloceros cazioti (Deperet), a small megalocerine cervid.

It is probably necessary to add to this list a shell-eating Otter, Cyrnaonyx majori Malatesta. The possible presence of Fox Vulpes vulpes L. in the Pleistocene is unclear at the moment, because remains of this species seem to be present in the Wurm I Maccinaggio breccias, but not in any other Corsico-Sardinian Pleistocene sites. The possibility that people were present on Sardinia (and Corsica) during the upper (and even middle) Pleistocene (Sondaar et al., 1986) is now being questioned (Vigne, 1989, 1 990).

Pre-Neolithic Period We now have good evidence for the presence of Mesolithic man on Sardinia and Corsica very early in the Holocene, at about 9000 b.p. However, the classical Mesolithic cultures are not well attested so that archaeologists in Corsica call this period ‘Pre-Neolithic’ rather than Mesolithic. At this time, the Flandrian transgression re-established the Straits of Bonifacio which separate the two islands.

The Pre-Neolithic Corsican mammalian fauna was very poor and unbalanced, with only five species: Episoriculus corsicanus, Prolagus sardus, Rhagamys orthodon, Tyrrhenicola henseli and man. Indeed Megaloceros cazioti only occurs in Sardinia at the beginning of the Pre-Neolithic, in Corbeddu Cave (Sardinia) where it constituted the main game species (Sondaar et al., 1986). This deer has not, so far, been found in any of the Corsican settlements (Vigne, 1988a). It seems probable that man rapidly hunted Megaloceros (and possibly any remaining endemic large mammals) to extinction shortly after his arrival in Corsica.

NEOLITHIC TO THE PRESENT The Neolithic culture which developed on Corsica from c. 7200 BP onwards, shows chronological and cultural similarities with the mainland Neolithic (Camps, 1988). This implies important contacts with the mainland.

History of Corsican mammals 91

... .... ... .... ........... ..... .......

Prolagus Tyrrbenicolo

Rbagomys Episoriculus Megoloceros

9000 8000 7000 6000 5000 4000 3C

? Yor les morfes Oryetologus cuniculus Rottus norvegicus Lcpus corsicanus Ursus arctos Rollus rotlus Cervus eiapbus Fdls s. I reyi Fdis s. I caius Equus co8ollus Equus asinus Muslelo nivalis Eliomys quercinus Crociduro suaveoiens Suncus elruscus Mus m domesticus Giis glis Apodemus sylvolicus Erinoceus Bos tourus europoeus

Conis familaris Vulpes vulpes Sus s scrofo Sus s domesficus Copra hircus Ovls a musimon Ovis aries Homo sapiens q. .......... ................... 1 1

.:.:.:.:.:. .:.:.:..

................. ...................

12000 1000 0 Years B.P

Fig. 2. The chronological distribution of Corsico-Sardinian terrestrial mammals since the beginning of human occupation.

From the beginning of the Neolithic to the present, two major trends characterize the history of the Corsican fauna: the progressive immigration (i.e. introduction; see below) of species, and the gradual extinction of the endemic species.

The post-glacial immigrations Chronology (Fig. 2) The first immigrants observed in the Early Neolithic at 7000 b.p. are the domestic mammals: Sheep Ovis aries L., Goats Capra hircus L., Pigs Sus scrofa domesticus Brisson, and, perhaps a little later, Cattle Bos taurus L. and Dogs Canis familiaris L. (Vigne, 1984, 1987a). The Fox Vulpes vulpes could also have been introduced at this time. These earliest immigrants undoubtedly brought about profound ecological changes, especially to the vegetation of the coastal and lowland regions.

No wild ancestor for the domestic species existed on the island, clearly indicating the efficiency of maritime transport as early as c. 7000 b.p. (Camps, 1988).

The European Hedgehog Erinaceus europaeus L., the Field Mouse Apodemus sylvaticus (L.) and the Dormouse Glisglis (L.) first appear in middle and late Neolithic layers (6000-5000 b.p.).

The Mouflon Ovis ammon musimon Pallas, whose origin from domestic Sheep is now well established on palaeontological and biochemical grounds (for a review, see Geddes, 1985), is observed for the first time in the bone samples at the end of the Neolithic. Mouflon probably originated earlier from Feral Sheep, but it is impossible to dis- tinguish between Wild and Domestic Sheep bones prior to the late Neolithic decrease in size of Domestic Sheep. The Corsican Wild Boar Sus scrofu L. is also descended from Feral Domestic Pigs, but it is impossible to know when this separation occurred due to probable interbreeding.

92 J.-0. Vigne

For the moment, remains of the House Mouse Mus musculus domesticus Rutty, the Garden Dormouse Eliomys quercinus (L.) and the two modern shrews Crocidura suaveolens (Pallas) and Suncus etruscus (Savi) are only known after the Bronze Age (since 2500 b. p.).

Horses and Donkeys appear at the end of the Iron Age (c. 2500 b.p.). The now distinctive (Arrighi & Salotti, 1988) feral population of Corsican Wild Cat Felis catus reyi Lavanden is probably descended from Domestic Cats introduced at this time or earlier, but there is no fossil record before c. AD 1300. The first remains of Red Deer Cervus elaphus L. appear during the Roman period (about 1600 b.p.; Vigne, 1988; Vigne & Marinval-Vigne, 1988; Lepetz, unpublished data), but deer might have been intro- duced at the end of the Neolithic, at least in the southern part of the island (un- published data). The earliest record of the Black Rat Rattus rattus (L.) found so far dates to c. AD 600 (Vigne & Marinval-Vigne, 1985).

Brown Bears Ursus arctos L. were introduced, probably as tamed animals, at the end of the Middle Ages (c. AD 1400). Subsequently, escaped animals probably gave rise to a feral population, which was hunted to extinction by AD 1700 (Poplin, Vigne & Gauthier, 1988). Historical texts document the introduction of the Brown Hare Lepus corsicanus de Winton, from the southern half of Italy (F. Palacios, pers. comm.), shortly before the sixteenth century, and the Rabbit Oryctolagus cuniculus (L.), during the 1950s (Dubray, 1984). There are still no records to indicate when the introduction of the Weasel Mustela nivalis boccamela Bechstein, Brown Rat Rattus norvegicus (Berkenhout), or Pine Marten Martes mattes (L.) occurred. The current status of the Pine Marten in Corsica is uncertain. Cervus elaphus, while extinct since 1972 on Corsica, was re-introduced from Sardinia in 1985.

Modes of immigration : introduction Throughout the Holocene, Corsica was 80 km from the mainland. Therefore, the hypothesis that most mammals reached Corsica by active swimming or rafting (which are very rare phenomena) seems unlikely for two main reasons: 1. A colonization rate of 14 wild species within less than 10,000 years is much higher

than is to be expected from natural dispersion. 2. The present wild species do not include the more-or-less amphibious mammals such

as elephants, hippos, otters and deer which were able to swim to many of the isolated Mediterranean islands during the Pleistocene (Sondaar, 1986). The mammals of Corsica are ecological generalists (Cheylan, 1984) and are strongly

linked with human activity both ecologically and culturally. Some of them, such as Cervus elaphus, Lepus corsicanus and Oryctolagus cuniculus (and, during the Neolithic, Vulpes vulpes), are game species. Commensal small mammals such as Crocidura, Suncus, Glis, Eliomys and especially Apodemus, Mus and Rattus are better stowaways on ships than swimmers or rafters. Moreover, non-commensal or undesirable (‘vermin’, human competitors or predators, etc.) species of the nearby mainland are absent from Corsica. Taking the Insectivora as an example, the Corsican fauna only includes Crocidura and Suncus (two soricids which are closely associated with man’s surround- ings), and the Hedgehog, which probably played an important role in the Neolithic culture as shown by numerous zooarchaeological data on the continent (Vigne, 1988b); however, animals such as the Mole Talpa europea L., considered to be ‘vermin’, or animals such as Neomys or Sorex, not associated with the human environment in the Mediterranean area, are absent. The same line of reasoning applies in the case of carnivores and rodents (Vigne, 1988a).

History of Corsican mammals 93

0

10

I

E - 20 f a n 0

30

4 0 I L L L L L I 0 4 0 800 10 0 100 20 0 20 4 0 0 20 4C

Percentaae Meon no individuals ' 1-

per 21 Rodentia Insect ivora

Mammal density (number of individuals)

Fig. 3. Distribution of small mammals in the Monte di Tuda Cave (Oletta, Corsica). The lowest layer (38 cm deep) is dated to the late Iron Age (240G2100 years BP calibrated radiocarbon date). From Vigne & Marinval-Vigne (1991).

Taken together, the evidence suggests that most, if not all, modern Corsican terrestrial mammals have been introduced directly or indirectly by man (Vigne, 1987b, 1988a, 1990). The early presence of domesticated mammals clearly introduced by man, shows that transportation by boat was not only possible but effective as early as the beginning of the Neolithic.

Extinction of endemic Pleistocene stock The presence of a special pattern of burning on the distal end of bones (Vigne & Marinval-Vigne, 1983) indicates that Prolagus and even the large rodents (Rhagamys and Tyrrhenicola) were probably roasted and eaten by man. These species were no doubt subjected to severe hunting pressures from the moment that Corsica was first colonized by humans (Vigne, 1987~). The small mammals were probably also (i) pre- dated upon by some of the new immigrants such as Dog, Fox and Weasel, (ii) sub- jected to competition with domesticated hoofed-mammals and new invaders (rodents, soricids, etc.) and (iii) disadvantaged by vegetation changes brought about through pastoralism and agriculture. Nevertheless, some new findings from the 3000-year sequence at Monte di Tuda Cave (Vigne & Marinval-Vigne, 1991), show that all four autochthonous small-mammal species managed to survive human pressures for at least 8000 years. Some osteological changes (Vigne, 1987c, Vigne & Marinval-Vigne, 1990) indicate that these species may have adapted to new ecological and demographic pressures.

The extinction of those four small mammals has not yet been radiocarbon dated, but probably occurred sometime between the arrival of Rattus rattus (probably during the Roman period) and the present time. In northern Corsica at least, extinction may be correlated with drastic deforestation as evidenced by the reversal in proportions between Mus musculus and Apodemus sylvaricus (Fig. 3). It seems that extinction of those four species was brought about by a major increase in agriculture, the effects of

94 J.-0. Vigne T

20

Ln Y

a QJ

Ln ”.. 0

0 D L

5 10 2

Fig. 4. Changes in the frequency of terrestrial mammal species on Corsica since the end of the Pleistocene. Dates are not calibrated. The confidence intervals reflect uncertainty about the stratigraphical distribution of some species. MA= Middle Ages. See text for description of the numbers.

which were probably aggravated by the appearance of Rattus rattus, an aggressive competitor (Vigne & Marinval-Vigne, 1991).

BIOGEOGRAPHICAL PROCESSES ON CORSICA AND THE OTHER MEDITERRANEAN ISLANDS The introduction by man of all modern terrestrial mammals on Corsica finally resulted in the complete demise of the Pleistocene fauna. The relatively recent origin of the present fauna explains the absence on the island of endemism at the species level. This huge extinction-immigration process has resulted in a three- to fivefold increase in mammal diversity (two- to fourfold if only wild mammals are considered). As Cheylan (1984) has shown, the resulting modern fauna is much richer than would be predicted by the MacArthur & Wilson theory of island biogeography, and, according to this theory, the Corsican fauna may be considered to be supersaturated (Vigne, 1987b).

The development of the Corsican mammalian fauna may be summarized in five steps (Fig. 4; Vigne, 1990): 1.

2.

3.

4.

5 .

a stablebut disharmonic late Pleistocene endemic fauna which consisted of no more than seven to ten species; human colonization, about 10,000 years ago, which resulted in the rapid extinction of all the large species; between 9000 and 8000 b.p. the fauna became especially poor, with no more than five species: man and four small mammals; during the Neolithic, the number of species rapidly increased as a result of human introductions to reach a plateau in the Bronze Age; from the Iron Age and Roman periods onwards faunal diversity again increased as a result of exchange between mainland and islands through human maritime activities. Extinctions of the endemic species Prolagus, Rhagamys, Tyrrhenicola and Episoriculus and, more recently, of large mammals such as Brown Bear and Red

History of Corsican mammals 95

Deer, as well as the decline in some taxa such as Mouflon and Brown Hare and the difficulties in introducing new game species such as Rabbit, all suggest that the present fauna is supersaturated. Is the history of the Corsican fauna analogous to that of other Mediterranean islands?

Although each island must be considered as a special case, the answer is definitely yes, at least as far as the main patterns are concerned. A complete replacement, during the Holocene, of an impoverished, endemic and unbalanced mammalian fauna by one which is considered to be super-saturated and poorly endemic probably occurred on all the larger Western (Vigne & Alcover, 1985) and Eastern Mediterranean islands: on the Balearic Islands (Alcover, 1980; Alcover, Moya-Sola & Pons-Moya, 1981), Malta (Boessneck & Kuver, 1970; Storch, 1970), Crete (compare Pleistocene (Dermitzakis & de Vos, 1987) with Neolithic (Jarman & Jarman, 1968) and present faunas), and on Cyprus (Davis, 1984; Simmons, 1988; Held, 1989). The same general scenario probably occurred on many of the smaller islands, as has been demonstrated on Pantelleria (Felten & Storch, 1970), Provenqal islands (Cheylan, 1984), on a Corsican islet (Lavezzi; Vigne & Cheylan, 1989) and on the Tunisian islet of Zembra (Vigne, 1988~).

CONCLUSION This study has emphasized the important role man has played in the development of the fauna of islands, since as long ago as the beginning of the Holocene, 10,000 years ago.

The study of the mechanisms of faunal turnover on both large and small islands is still in progress and should improve our understanding of both human behaviour patterns and micro-evolution following the arrival of a new invader species (Vigne & Marinval-Vigne, 199 1). Zooarchaeology thus serves both anthropology and bio- geography and, it is hoped, the management and preservation of living species.

ACKNOWLEDGMENTS I am grateful to the Mammal Society for inviting me to present this paper to the Symposium ‘Mammals and Archaeology’ (London, November 1990), and to S. Davis, who improved an early draft of the manuscript.

REFERENCES Alcover, J.-A. (1980) Note on the origin of the present mammalian fauna from the Balearic and Pityusic

Alcover, J.-A., Moya-Sola, S. & Pons-Moya, J. (1981) Les Quimeres delpassat. Editorial Mool, Mallorca. Arrighi, J. & Salotti, M. (1988) Le chat sauvage (Felis silvesrris) en Corse: une certitude. Mammalia, 52 ,

Boessneck, J. & Kiiver, M. (1970) Alluviale Tierknochenfunde aus der Char Dalam-Hohle, Malta.

Camps, G. (1988) Prehistoire d’une ile. Les origines de la Corse. Errance, Paris. Cheylan, G. (1984) Les Mammiferes des iles de Provence et de Mediterranee occidentale: un exemple de

Conchon, 0. (1976) The human settlement of Corsica: palaeogeographic and tectonic considerations.Journal

Islands. Miscel-Lania Zoologia, 6,141-149.

123-125.

Senkenbergiana Biologica, 51,147-158.

peuplernent insulaire non equilibre? Revue d’Ecologie (Terre V i e ) , 39,37-54.

of Human Evolution, 5,241-248.

Khirokitia f Chybrei 1977-1 981, pp. 147-179. Recherches sur les Civilisations, Paris. Davis, S. J. M. (1984) Khirokitia and its mammal remains: a Neolithic Noah’s Ark. In: Fouilles recentes ci

Dermitzakis, M. D.8r de Vos, J. (19878jj Faunal successions and the evolution ofmammals in Crete during the

Dubrsy, D. (1984) Repartition des Lagomorphes (Lievre commun et Lapin de garenne) en Corse. Bulletin

Eisenmann, V. (1990) Caracteres juveniles et affinites systematiques du crPne de Cynocherium sardous, canide

Felten, H. & Storch, G. (1970) Kleinsauger von den italienischen Mittelmeer-Inseln Pantelleria und

Geddes, D. (1985) Mesolithic Domestic Sheep in West Mediterranean Europe. Journal of Archaeological

Pleistocene. Neue Jahrbuch Geologie und Palaontologic Abhandlungen, 173,337-408.

Mensuel de I‘Ofice national de la Chasse, 85,3638.

endemique pleistocene de Sardaigne. Compres Rendus de I’Acadimie des Sciences, serie II, 310,433439.

Lampedusa. Senckenbergiana Biologica, 51, 159-173.

Science, 12,2543.

96 J.-D. Vigne Held, S. 0. (1989) Colonization cycles on Cyprus. 1 : The biogeographic and paleontological foundation of early

prehistoric settlement. Report of the Department of Antiquities Cyprus. James, H. F., Stafford, T. W., Steadman, D. W., Olson, S. L., Martin, P. S., Jull, A. J. T. & McCoy, P. C.

(1987) Radiocarbon dates on bones of extinct birds from Hawaii. Proceedings of the National Academy of Sciences U . S . A . , 84,2350-2354.

Jarman, M. R. & Jarman, H. N. (1968) The fauna and economy of Early Neolithic Knossos. The Annual of the British School at Athens, 63,241-264.

Libois, R. M. (1984) Le Regime alimentaire de la chouette effraie. Cahiers d’Ethologie appliquie, 4 (Z), 1-202. MacArthur, R. H. & Wilson, E. 0. (1967) The Theory of Island Biogeography. Princeton University Press,

Princeton, NJ. Olson, S. L. &James, H. F. (1982) Fossil birds from the Hawaiian Islands: evidence for wholesale extinction

by man before Western contact. Science, 217,633-635. Poplin, F., Vigne, J.-D. & Gauthier, A. (1988) Donnees nouvelles sur I’ours (Ursus arctos, Carnivora,

Mammalia) en Corse. Bulletin d’Ecologie, 19, 189-194. Simmons, A. H. (1988) Extinct Pygmy Hippopotamus and Early Man in Cyprus. Nature, 333,554-557. Sondaar, P.-Y. (1986) The Island Sweepstake. Natural History, 95 (9), 52-57. Sondaar, P.-Y., Sanges, M., Kotsakis, T. & de Boer, P. L. (1986) The Pleistocene Deer Hunter of Sardinia.

Geobios, 19, 17-25. Storch, G. (1970) Holozane Kleinsaugerfunde aus der Ghar Dalam-Hohle, Malta (Mammalia: Insectivora,

Chiroptera, Rodentia). Senckenbergiana Biologica, 51,135-145. Vigne, J.-D. (1984) Premieres donnees sur les debuts de l’elevage du Mouton, de la Chevre et du Porc dans le

sud de la Corse (France). British Archaeological Reports, International Series, 202 (3), 47-65. Vigne, J.-D. (1987a) L’exploitation des ressources alimentaires carnees en Corse, du VIIe au IVe millenaire.

In: Premiires communautis paysannes en Miditerranie occidentale (Acres coll. Int. Montpellier, 1983) (ed. by J. Guilaine, J. Courtin & J.-L. Vernet), pp. 193-199. CNRS, Paris.

Vigne, J.-D. (1987b) L’origine du peuplement mammalien de la Corse: quelques reflexions biogeographi- ques. Bulletin de la Sociiti Zoologique de France, 111,165-178.

Vigne, J.-D. (1987~) L’extinction holocene du fonds de peuplement mammalien indigene des iles de Mediterranee occidentale. Memoires de la Sociiri Geologique de France, N.S. , 150,167-177.

Vigne, J.-D. (1988a) Les Mammijires post-glaciaires de Corse. Etude archiozoologique (26e suppl. Gallia Prihistoire) . CNRS, Paris.

Vigne, J.-D. (1988b) Apports de la biogeographie insulaire a la connaissance de la place des Mammiferes sauvages dans les societes neolithiques mediterraneennes. Anthropozoologica, 8,31-52.

Vigne, J.-D. (1988~) Donnees preliminaires sur I’histoire du peuplement mammalien de I’ilot de Zembra (Tunisie). Mammalia, 52,567-574.

Vigne, J.-D. (1989) Le peuplement paleolithique des iles: le debat s’ouvre en Sardaigne. Les Nouvelles de -I‘Archiologie, 35,3-2. ~

Viune, 1.-D. (1990) BioaeoaraDhical Historv of the Mammals on Corsica (and Sardinia) since the Final - - - Pleistocene. In:.Biogiog&phicaI Aspects if Insularity (International Symposium, Roma, 1987). Att i dei Convegni Lincei, 85,370-392.

Vigne, J.-D. & Alcover, J.-A. (1985) Incidence des relations historiques entre 1’Homme et 1’Animal dans la composition actuelle du peuplement amphibien, reptilien et mammalien des iles de Mediterranee occidentale. Acres du 1 1 Oime Congris national des Sociitis Savantes (Montpellier, 1985), Section Sciences, 2,79-91.

Vigne, J.-D. & Cheylan, G. (1989) Ile Lavezzu. Compte rendu des travaux archeozoologiques dans lachapelle Santa Maria. Travaux Scientifiques du Parc nature1 Rigional et des Reserves naturelles de Corse, 24,3541.

Vigne, J.-D. & Marinval-Vigne, M.-C. (1983) Methode pour la mise en evidence de la consommation du petit gibier. British Archaeological Reports, International Series, 163 (l) , 239-242.

Vigne, J.-D. & Marinval-Vigne, M.-C. (1985) Le Rat en Corse au VI‘ siecle apres J.-C.? Mammalia, 49, 138-139.

Vigne, J.-D. & Marinval-Vigne, M.-C. (1988) Contribution a la connaissance du cerf de Corse (Cervus elaphus, Artiodactyla, Mammalia) et de son histoire. Bulletin d’Ecologie, 19, 177-187.

Vigne, J.-D. & Marinval-Vigne, M.-C. (1990) Nouvelles donnees sur l’histoire des musaraignes en Corse (Insectivora, Mammalia). Vie Q Milieu, 40,207-212.

Vigne, J.-D. & Marinval-Vigne, M.-C. (1991) Reflexions ecologiques sur le renouvellement holocene des micromammiferes en Corse: les donnkes preliminaires des fossiles du Monte di Tuda. In: Le Rongeur et I‘Espace (ed. by M. Le Berre & L. Le Guelte), pp. 183-193. Raymond Chabaud, Paris.