Revue de Paléobiologie, Genève (décembre 2004) 23 (2): 611-626
ISSN 0253-6730
Humans and Carnivores in the Early Upper Paleolithic in Portugal :
Data from Pego do Diabo Cave
Maria João VALENTE1
Abstract
During the Upper Pleistocene human hunters had to compete with large carnivores in their ecological setting. This paper analyses the
fauna from the small cave of Pego do Diabo, with an occupation classified as Aurignacian, and considers human-carnivore relationships
in the Portuguese Upper Paleolithic. Similar to other Early Upper Paleolithic cave deposits in Portugal (Salemas, Caldeirão, Casa da
Moura and Buraca Escura), Pego do Diabo faunal assemblage includes remains of several carnivore species, such as wolf, hyena and
iberian lynx.
Some evidences suggest important changes in the relationship between humans and carnivores at the end of the last glacial in Portuguese
Early Upper Paleolithic sites : first, the relative abundance of carnivore remains, associated with a high degree of carnivore-modified
bones ; second, a significant reduction in carnivore species diversity during the Solutrean and Magdalenian. A model is developed
emphasizing short and discontinuous human occupations of small cavities during the Early Upper Paleolithic, which left these sites free
for carnivore denning. These patterns changed during the Solutrean, as large carnivores started to disappear from the archaeological
record and sites containing significant numbers of carnivore-damaged fauna begin to be rare.
Key words
Upper Paleolithic, Portugal, Carnivores, Ecology.
INTRODUCTION
BACKGROUND, STRATIGRAPHY AND DATING
In the past decades, many researchers have noted that
during the Upper Pleistocene human hunters had to
compete with carnivores in their ecological setting,
frequently overlapping in the use of space and in
foraging strategies (e.g. BINFORD, 1981 ; BLASCO, 1997 ;
BLUMENSHINE, 1995 ; BLUMENSHINE & MAREAN, 1993 ;
BRANTINGHAM, 1998 ; BRUGAL & JAUBERT, 1991 ; BRUGAL
et al., 1997 ; FOSSE, 1995 ; HOCKETT, 1993 ; STINER, 1994 ;
STRAUS, 1992). This paper discusses the faunal remains
from the cave Pego do Diabo (in English meaning «deep
hollow of the Devil»). The site contains a short human
occupation classified as Aurignacian by J. ZILHÃO (1997),
and affords an opportunity to analyse human-carnivore
relationships at early Upper Paleolithic Portuguese sites.
Despite not being numerous, Pego do Diaboʼs faunal
collection shows specific taphonomic features that agree
with other Pleistocene deposits in small cave sites with
Early Upper Paleolithic assemblages (like Caldeirão,
Salemas, Cova da Moura and Buraca Escura). Data
indicate that a relatively large number of carnivore
species were in direct competition with human groups
at this time, not only for the use of space but also for the
same animal resources.
Pego do Diabo Cave (9º13ʼ6ʼʼW, 38º51ʼ52ʼʼN) is a small
cavity located in the Portuguese Estremadura about
20 km north of Lisbon (Fig. 1). The site stands in a
Turonian limestone ridge that lies on the southern slope
of the Loures River Valley, near its confluence with the
Lousa River, at 250 m a.s.l. From the cave we have a vast
view shed of the surrounding area.
The cave has two sections : a narrow passage about
13 m long and a room at the end measuring about 2 m
wide (Fig. 2). Bedrock can be seen at the entrance and is
marked by a distinct depression toward the back of the
cave. In this area there was a well-preserved sedimentary
sequence about 1.4 m deep.
Pego do Diabo was excavated in 1976 by the Grupo para
o Estudo do Paleolítico Português (GEPP, 1979) and in
the late 1980ʼs by ZILHÃO (1988 and 1997). In total, an
area of 11 m2 has been opened, representing about half of
the deposits presently accessible. In 1993, J.L. CARDOSO
published a paleontological study on the Pleistocene
Large Mammals of Portugal, in which he analized a
small sample of Pego do Diabo faunal assemblage
(only elements adequate for osteometric data) ; and in
1995, J. ZILHÃO finished is Ph.D. thesis, which included
a complete study of the lithic assemblage (published in
1997).
1
Universidade do Algarve, Campus de Gambelas, FCHS, 8000 Faro, Portugal, Email : [email protected]
Actes du XIVe Congrès UISPP « Hommes & Carnivores au Paléolithique », Liège 2001, BRUGAL, J.P. & P. FOSSE (Eds).
612
M. J. VALENTE
Fig. 1 : Approximate locations of the main archaeological sites
mentioned in the text. (1) Pego do Diabo Cave, (2)
Salemas Cave, (3) Casa da Moura Cave, (4) Lapa do
Picareiro, (5) Caldeirão Cave, (6) Buraca Escura.
The stratigraphic profile (Fig. 2) shows the 6 units
recognized by ZILHÃO (1997, 2 : 94-95). The top layer
(Level A) was only found at the back of the cave ; it
yielded historical material and has a probable funerary
occupation. Level 1 contains a mixture of modern and
Pleistocene material (including fauna) that represent
a redeposition of level 2. Level 2 comprises Upper
Paleolithic artefacts in association with bones. Level
3 also has Pleistocene fauna, and displays a distinct
coloration, probably made by manganese oxide. At the
bottom of the deposit, level 4 has rare archaeological
material and level 5 contains no material at all.
The faunal analysis and the results presented here are
from level 2. We also included in the study all the
remains from level 1 that exhibited the same surface
characteristics of level 2 (colour and fossilization).
The four radiocarbon dates obtained are shown in Table 1.
Based on laboratory information, the two younger dates
were rejected : in the ICEN-306 date of level 2, recent
carbon from the top layer probably contaminated the
sample ; in the ICEN-491 date of level 3, there was a
probable contamination incident or a deficient collagen
extraction resulting in an impure sample (ZILHÃO, op.
cit. : 98).
The accepted radiocarbon dates (ICEN-490 and ICEN732) and the lithic analysis, especially the six Dufour
bladelets, led J. ZILHÃO (1997) to classify level 2 as
Aurignacian (see Note 1). The 28.000 years BP date
was interpreted as representing the age of the human
occupation at the base of level 2, and the 23.000 years
BP date was considered the end of a slow deposition
sequence (or a contaminated sample). Consequently,
levels 3 and 4 are expected to be Middle Paleolithic, even
if the scarce lithic industry (n = 6) has no diagnostic tools
(ZILHÃO, op. cit. : 95).
THE PLEISTOCENE FAUNA AT PEGO DO DIABO
CAVE
A very diversified Pleistocene fauna was found in the
cave, including small animals (such as birds, small
rodents, bats and amphibians), medium mammals
Table 1 : Radiocarbon dates for Pego do Diabo Cave (ZILHÃO 1997, 2: 94-95).
Provenience
Level
Sample
Lab No.
Date B.P.
M13
M11 + L11
M11 + L11
M13 + M14
2
2a
2b
3
Charcoal
Macrofauna
Macrofauna
Rabbit and macrofauna
ICEN-306
ICEN-490
ICEN-732
ICEN-491
2.410 +/- 80
23.080 +/- 490
28.120 - 780 + 860
18.630 +/- 640
Humans and Carnivores in the Early Upper Paleolithic in Portugal
613
Fig. 2 : [A] Plan view of Pego do Diabo Cave. Diagonal lines show excavated squares. [B] Stratigraphy of Pego do
Diabo Cave. (Both adapted from ZILHÃO 1997).
(mustelids and leporids), and large mammals. Excluding
microfauna and birds, fourteen taxa were identified
(Table 2). There are five species of herbivores : wild
boar (Sus scrofa), red deer (Cervus elaphus), ibex
(Capra pyrenaica), chamois (Rupicapra rupicapra) and
horse (Equus caballus). Seven species of carnivores are
present : fox (Vulpes vulpes), wolf (Canis lupus), brown
bear (Ursus arctos), hyena (cf. Crocuta crocuta spelaea),
iberian lynx (Lynx pardina), badger (Meles meles) and
another species of mustelid. Also two species of leporids
are present : rabbit (Oryctolagus cuniculus) and hare
(Lepus sp.). This faunal association does not differ from
other Portuguese collections from Initial Pleniglacial
(transition OIS 2/3, from 32 to 22 000 year BP).
Large mammals
The mammal assemblage has a total of 1472 large
mammal remains (Table 3). Of these, 450 (about 31 %)
can be identified to the species level. Whenever possible,
besides taxonomical and anatomical determination,
size and age classification were recorded. Number of
identified specimens (NISP), Minimum Number of
Individuals (MNI) and Minimum Number of Elements
(MNE) were calculated for each taxon.
Herbivore remains (NISP = 375 ; MNI = 21) are
dominated by red deer and horse, followed by caprids.
For all these species there is a significant amount of
juvenile specimens, some of them representing fetal or
neonatal individuals (see below).
An important quantity of carnivores was revealed, both
in number of individuals (NISP = 74 ; MNI = 12) and in
variety of taxa (seven different species). The carnivore
collection is dominated by iberian lynx, and followed by
wolf and fox (Table 4).
Data on animal behaviour and results on the age profiles
for each carnivore species suggest differences in the
nature of the cave occupation. Juvenile brown bear
remains are present, and it is probable that bears used the
cave to hibernate. Juvenile wolf remains are also present,
and in all probability they used the site to reproduce
and protect offspring. No juvenile hyena remains were
found, but this species is also known to den in small
614
M. J. VALENTE
Table 2 : Pego do Diabo identified mammal taxa (microfauna excluded).
Order Artiodactyla
Family Suidae
Family Cervidae
Family Bovidae
Order Perissodactyla
Order Carnivora
Family Equidae
Family Canidae
Order Lagomorpha
Family Ursidae
Family Mustelidae
Family Felidae
Family Hyaenidae
Family Leporidae
Sus scrofa, L. 1758
Cervus elaphus, L. 1758
Capra pyrenaica, SCHINZ 1838
Rupicapra rupicapra, L. 1758
Equus caballus, L. 1758
Canis lupus, L. 1758
Vulpes vulpes, L. 1758
Ursus arctos, L. 1758
Meles meles, L. 1758
Lynx pardina, TEMMINCK 1824
Cf. Crocuta crocuta spelaea, GOLDFUSS 1810
Lepus sp.
Oryctolagus cuniculus, GRAY 1874
Note: Another species of carnivore, probably a mustelid (not Meles meles), was identified.
Table 3 :
Quantifying data on the fauna present in Pego do
Diabo (level 2).
NISP
MNI
12
197
29
22
42
73
375
2
7
6
2
4
21
16
15
3
6
1
28
6
74
2
2
2
2
1
2
1
12
Sub-Total Large Mammals
Unidentified Large Mammals
TOTAL Large Mammals
450
1022
1472
33
LEPORIDS
Lepus sp.
Oryctolagus cuniculus
Sub-Total Leporids
Unidentified Leporid
TOTAL Leporid
10
539
549
158
707
2
27
39
TOTAL NISP Herbivores + Leporids
TOTAL Animal
924
2179
HERBIVORES
Sus scrofa
Cervus elaphus
Capra pyrenaica
Rupicapra rupicapra
Cf. Caprid
Equus caballus
Sub-Total Herbivores
CARNIVORES
Canis lupus
Vulpes vulpes
Ursus arctos
Meles meles
Cf. Mustelid*
Lynx pardina
Crucuta c. spelaea
Sub-Total Carnivores
Note: (*) Not Meles meles.
cavities (e.g. BARTRAM & VILLA, 1998 ; BRUGAL et al.,
1997 ; FOSSE, 1995 ; FOSSE et al., 1998). In addition, wolf
and hyena may bring herbivore bones into caves to be
consumed. Small carnivores such as the iberian lynx, fox
and badger are known to prey on smaller animals and
to take shelter in cavities either to protect themselves
or to eat. The carnivore remains in general can be due
to starvation (thus rare, this can happen to bear cubs
during hibernation ; cf. ROGERS, 1987 and 1992), to
carnivore predatory behaviour or to competition between
carnivores (interspecific) or between carnivores and
humans for the use of the cave.
Wolf, hyena and humans are the most obvious predators
of the herbivores found in the cave, and they are likely
the main contributors to the bone assemblage. But we
can also associate the iberian lynx to this predator group,
considering that under special circumstances this species
is known to prey on small herbivores (juvenile deer and
caprids ; cf. AYMERICH, 1982 ; BÉLTRAN et al., 1985 ; BRÍO,
1993 ; and DELIBES, 1980) and occasionally carrying
them into shelters (FERNANDEZ & PALOMARES, 2000). This
fact was recently documented at Buraca Escuraʼs Upper
Paleolithic levels by J.-Ph. BRUGAL (AUBRY et al., 2001).
At Pego do Diabo, almost 52 % of the herbivore MNI are
represented by juveniles remains (40 % of the herbivore
NISP). It is possible that most of the medium herbivores
(caprids) and one third of the larger herbivores (the
juvenile individuals) could have been preyed on by
iberian lynx.
Brown bear was considered an unlikely candidate as
a predator since it rarely feeds on animals (even if
occasionally scavenges) and rarely transports carrion or
bones to their hibernation den. In fact, the reduction of
smells is one of its priorities to avoid attracting wolves
and humans during hibernation when a bear is often
vulnerable given its reduced metabolic state (LINNEL et
al., 2000 ; ROGERS, 1987 ; STINER, 1998 ; TIETJE & RUFF,
1980 ; also see ARGANT & PHILIPPE, 1997 on the utility
of bear fur). It is interesting to note that, contrary to the
cave bear (Ursus spelaeus), brown bears are quite rare
in strictly paleontological sites and relatively frequent in
archaeological sites, which may confirm the competition
for the use of caves between brown bears and humans
(ARGANT & PHILIPPE, 1997).
Humans and Carnivores in the Early Upper Paleolithic in Portugal
Table 4 :
Quantifying NISP and MNI for herbivores and
carnivores at Pego do Diabo.
Adult
Immat.
Total
% Herb
or Carniv
Sus scrofa
5
7
12
3,2
Cervus elaphus
114
83
197
52,5
Capra pyrenaica
14
15
29
7,7
Rupicapra r.
22
22
5,9
Cf. Caprid
28
14
42
11,2
Equus caballus
45
28
73
19,5
TOTAL
228
147
375
Sus scrofa
1
1
2
9,5
Cervus elaphus
3
4
7
33,3
Capra pyrenaica
2
4
6
28,6
Rupicapra r.
2
2
9,5
Equus caballus
2
2
4
19,1
TOTAL
10
11
21
HERBIVORES
NISP
MNI
CARNIVORES
NISP
Canis lupus
13
2
15
20,0
Vulpes vulpes
12
3
15
20,0
Ursus arctos
1
2
3
4,0
Meles meles
7
7
9,3
Cf. Mustelid
1
1
1,3
Lynx pardina
28
28
37,3
Crocuta c. s.
6
6
8,0
TOTAL
67
7
74
Canis lupus
1
1
2
16,7
Vulpes vulpes
1
1
2
16,7
Ursus arctos
1
1
2
16,7
Meles meles
2
2
16,7
Cf. Mustelid*
1
1
8,3
Lynx pardina
2
2
16,7
Crocuta c. s.
1
1
8,3
TOTAL
9
MNI
Note: (*) Not Meles meles
3
12
615
Researchers have used several criteria to identify the
main agents of herbivore bone accumulation in caves,
such as relative abundance between herbivore and
carnivore remains, presence of coprolites, presence of
artefacts, artefact abundance compared to carnivore
bones, herbivore mortality profiles, representation of
anatomical parts, and studies of bone modifications such
as gastric dissolution, gnawing damage, cut marks or
percussion marks.
The Carnivore Index is based on the ratio of herbivore
and carnivore remains (for a recent review of this
index, see BLASCO-SANCHO, 1996). In this ratio, only
the carnivores that are known to collect herbivore
bones and contribute actively to bone accumulations
were considered (for reasons explained above and by
BRUGAL et al., 1994) : wolf, hyena and lynx. This ratio
is 0,13 when using NISP [carnivore NISP (wolf, hyena
and lynx) = 49/herbivore NISP = 375]. Following STINER
(1994 :91, table 4.14) in her study of Upper Pleistocene
faunal accumulations in Italian caves, it is believable that
this result most probably corresponds to a den with fauna
collected by carnivores.
The use of the Carnivore Index, however, is somewhat
problematic, given that most carnivore remains found
in caves are probably due to in situ deaths, either by
natural causes or by carnivores preying on one another ;
therefore, under these circumstances, it is plausible to
think that the full carcass will be present in the deposit.
In contrast, herbivores are not cave users and are hunted
outside the cavity, so perhaps only a part of the carcass
will be deposited in the cave. As a result of these different
processes, we are likely to have an inflation of carnivore
remains in the faunal accumulation. Using the MNI
instead of the NISP can help to alleviate this problem
since that unit is unaffected by differential transportation
(cf. BLASCO-SANCHO, 1995). The index ratio of 0,24
[carnivore MNI (wolf, hyena and lynx) = 5/herbivore
NISP = 21] increases the probability that carnivores
were the most important agent in the accumulation of
herbivore bones at Pego do Diabo.
Another line of evidence to take into consideration is the
presence of coprolites (BARTRAM & VILLA, 1998 ; DAVID,
1994 ; FOSSE et al., 1998 ; STINER, 1994). Though few,
seven fragmented coprolites (hyena ?) were found at
Pego do Diabo Cave.
Comprehensive research of the predatory agents involved
in the deposition of bones to Pego do Diabo Cave must
include a critical evaluation of direct evidence for
human use of the cave, such as artefacts and fireplaces,
in comparison to the presence of carnivore remains.
For instance, a large quantity of lithic material coupled
with small numbers of carnivore bones normally reflects
intense human activity in the fossil assemblage (cf. DAVID,
1994 ; STINER, 1994). At Pego do Diabo, only 32 lithics
(11 tools and 21 flakes or chips) and one bone point
were recovered (ZILHÃO, 1997, 2 : 98-99). The relatively
low ratio of artefacts to carnivore NISP (33/52, or 0,63)
compares favourably to STINER ʼs (1994) ratio obtained
616
M. J. VALENTE
on Italian Pleistocene carnivore den assemblages, and
this reinforces the idea that carnivores used Pego do
Diabo cave more frequently than did humans. As to the
presence of hearths, none was found, but a few burned
bones were recovered (see below).
Herbivore mortality profiles also assist in the
identification of the agents of faunal accumulation in
caves. Most of the literature reports that wolf, hyena
and iberian lynx have a predilection for weak or young
animals, especially in large game such as red deer and
horse (on wolf see BEAUFORT, 1987 ; also see DOMINGUEZRODRIGO, 1993 for an oppositing view ; for the hyena
see DOMINGUEZ-RODRIGO, 1993 ; FOSSE, 1995 ; UADELLI,
1989 ; HILL, 1989 and KRUUK, 1972 ; for the iberian
lynx see BÉLTRAN et al., 1985 and BRÍO, 1993). On the
other hand, some modern hunter-gatherer groups favour
prime-age adults that are valuable in terms of calories
and secondary products ; however, several exceptions are
known, and thus this strategy is highly variable among
human foragers (LYMAN, 1987, 1994 ; STINER, 1990).
Mortality profiles are generally based on the analysis of
tooth wear, and we have distinguished four age classes
for the herbivores : infant, juvenile, adult and old (see
BLASCO-SANCHO, 1995). Although Pego do Diaboʼs
collection is relatively small (see Tables 4, 5), several
interesting trends can be noted. These results, displayed
in Figure 3, show herbivore mortality curves with a
bimodal distribution dominated by very young (infant)
and prime-age adults for deer, horse and ibex. These data
also inform on seasonality considerations. The presence
of infants with only deciduous dentition (a few with dPʼs
still erupting and others with dPʼs already erupted but
without wear) suggest fetal or neonatal animals. Red
deer remains include two left dP2 completely unworn and
one right dP2 with intermediate stage wear, representing
three infant individuals (fetal to a few weeks old) and a
season of death between late spring and summer (LEGGE
& ROWLEY-CONWY, 1988 ; MARIEZKURRENA, 1983). Horse
deciduous remains include two dP2 with intermediate
use wear (infant), also matching late spring or summer
Table 5: Teeth NISP for each herbivore and carnivore taxon at Pego do Diabo.
Deciduous teeth
Incisors/Canines
Pmʼs
Mʼs
Pmʼs / Mʼs
TOTAL
Deciduous teeth
Incisors/Canines
Pmʼs
Mʼs
TOTAL
Sus scrofa
5
1
3
9
Canis lupus
1
1
1
1
4
Cf. Caprid
4
2?
Equus caballus
19
2
6
12
2
32
55
Cervus elaphus
51
3
11
33
21
119
Capra pyrenaica
15
2
2
7
3
29
Rupicapra r.
Vulpes vulpes
4
1
2
Ursus arctos
2
Lynx pardina
Crocuta
3
5
1
1
7
2
8
2
2
5
5
1
13
Fig. 3 : Mortality profiles for wild boar, red deer, ibex, chamois and horse. Age classes: infant, juvenile, adult and old.
Humans and Carnivores in the Early Upper Paleolithic in Portugal
killing. Ibex remains include two left unworn dP2 and
two left intermediately worn dP3, suggesting late spring
to summer kills (PEREZ RIPPOL, 1988). Finally, wild boar
displays one infant individual with unworn dPʼs, but since
this species has a reproduction period extending between
February and June, its season of death can correspond to a
period that goes from late winter/early spring to summer
(BULL & PAYNE, 1982). In sum, the abundance of isolated
teeth does not allow us a detailed study of the herbivore
population age, but the use wear features on deciduous
teeth (unworn or intermediately worn) suggests that 40 %
of the deer, 50 % of the horse and 66 % of the ibex were
killed in late spring or summer. This interpretation does
not apply to adult prey, which could have been hunted at
any other season, but implies preference for temperate
seasons at least for the predation of immature animals.
Defining presence and absence of some anatomical parts
may help identify the agents of bone deposition. Most
predator lairs are known to contain a large number of
complete or nearly complete herbivore long bones,
whereas human sites present a different pattern with
cranial elements predominating and higher long bone
breakage [for hyenas see DOMINGUEZ-RODRIGO, 1994 ;
FOSSE, 1995 ; HILL, 1989 ; for wolves see DOMINGUEZ-
617
RODRIGO, 1994 ; HAYNES, 1980 ; STINER, 1994 ; for the
iberian lynx there are no adequate data, but for other
felids see NASTI, 1996 (puma) and RUITER & BERGER,
2000 (leopard)].
At Pego do Diabo, the distal elements of the limbs
are particularly prevalent, in contrast to the axial
and cranial bones (see Table 6). Red deer exhibits an
overrepresentation of the appendicular extremities, such
as metapodials and phalanges. Horse is represented
primarily by limb extremities and other forelimb
elements (radii, ulnae and carpals). Wild boar displays
few remains, but again long bone ends predominate.
Finally, caprids are represented mainly by humeri,
astragali, metapodials and phalanges.
Aware of the possibility that these results could have
been caused by post depositional factors that affect
fragmentation, particular attention was devoted to
unidentifiable bone fragments ; again, considerable
numbers of long bones were present.
The evidence shows an overrepresentation of some
anatomical parts, especially appendicular extremities,
together with a few forelimb bones. These skeletal
segments do not represent the highest nutritional value
of the carcass, but they may correspond to carnivores
preying on leftovers discarded in the cave.
Table 6 : Bone NISP for each herbivore and carnivore taxon at Pego do Diabo.
Sus
Antler/horn
Maxilla
Mandible
Vertebra
Innominate
Scapula
Humerus
Radius
Ulna
Carpal
Metacarpal
Femur
Tibia
Calcaneus
Astragalus
Tarsal
Metatarsal
Sesamoid
Metapode
1 Phalanx
2 Phalanx
3 Phalanx
Sub-total
TOTAL
Cervus
Rupicap
1
(2)
2+(1)
3+(1)
1
1
(1)
(3)
3
78
(1)
Note: (x) immature
2+(2)
4+(3)
2
1
(1)
3+(1)
1
5
1
6+(2)
5+(11)
8+(6)
1+(3)
45+(33)
Equus
2
1
1+(1)
(1)
(1)
Cf. Caprid
(2)
3
(1)
1
(1)
1
2
2
9
1+(3)
2+(3)
4
2
18+(12)
9
30
3
Canis
Vulpes
Ursus
Meles Cf. Mustelid
(1)
1
2
1
(4)
2+(3)
1
2
1
(1)
(1)
1
1
1
1
1
1
3
2
1
1
2
1
1
8
1
7
8
1
7
11
Crocuta
1
1
2
4
1
(1)
9+(10) 10+(1)
19
4
1
1
Lynx
1
1
2
4
6
1
1
1
1
1
20
4
1
20
4
618
M. J. VALENTE
Bone modifications are the best diagnostic tool to identify
the accumulator or modifier of the faunal assemblage
(HAYNES, 1983). Table 7 summarizes bone NISP
displaying gastric dissolution, gnawing damage, cut
marks and burning. There is direct evidence for carnivore
activity, such as remains with gastric dissolution (mostly
in red deer but also in chamois ; see Fig. 4), scoring,
punctures and a shortage of long bone ends. In contrast,
there are a few clear cut-marks, normally on long bone
shafts (Fig. 5 and 6). The only identified species that have
cut-marks are the chamois, all on shafts, and red deer, on
a distal antler branch.
Although no hearths were found in the cave, some
carbonised remains were found. A total of 14 bones was
charred, but only two were identified taxonomically and
to element, and both were carnivores : a wolf maxilla
fragment and a proximal radius of lynx.
Fig. 6: Anthropical modifications: indeterminate long bone
fragment with cut-marks.
Table 7 : Bone NISP damage in Pego do Diabo Cave.
HERBIVORES
Sus scrofa
Cervus el.
Rupicapra r.
Cf. Caprid
Equus c.
Dissol.
Gnawed
9
1
1
1?
7
2
6
2
CARNIVORES
Canis lupus
Vulpes v.
Ursus arctos
Meles m.
Lynx pardina
Crocuta
Cutmarks
Fig. 5: Anthropical modifications : indeterminate long bone
fragment with cut-marks.
Burned
1
2
1?
1?
1
1
1?
Unidentified
71
10
4+1?
12
TOTAL
82
27+3?
7+2?
14
Fig. 4 : Carnivore damage: digested red deer bones (hyena
scats?).
Leporids : rabbits and hares
Approximately seven hundred leporid bones have been
recovered from the Early Upper Paleolithic level. These
bones represent more than four hundred elements, or
27 rabbits and two hares (Tables 3 and 8).
Both small carnivores such as lynx, wildcat, fox or
badger, or larger ones like wolves, are known to hunt
rabbits and accumulate their bones in shelters (e.g.
DELIBES & HIRALDO, 1981 ; GIL-SANCHEZ et al., 1999 ;
HENRY et al., 1988 ; JAKSIC & SORIGUER, 1981 ; MARTIN et
al., 1995 ; ALOMARES et al., 2001 ; REVILLA & PALOMARES,
2002). As HOCKETT (1991, 1993, 1995 ; also HOCKETT &
HAWS, 2002) pointed out, assemblages created by smaller
carnivores may be characterized by the accumulation of
entire rabbit carcasses, with large numbers of complete
or nearly complete limb elements usually with one of
the epiphysis attached to long bones. On the other hand,
Humans and Carnivores in the Early Upper Paleolithic in Portugal
619
Table 8 : Leporid bone NISP and NME, and NISP damage at Pego do Diabo.
Oryctolagus cuniculus
NISP
Adult
Juve Infant
Head
Maxilla
Mandible
Scapula
Humerus
Radius
Ulna
Metacarpal
Vertebra
Rib
Sacrum
Innominate
Femur
Tibia
Calcaneus
Astragalus
Metatarsal
Metapode
Phalanx
Unidentified
1
6
21
9
15
21
22
23
21
5
2
43
19
28
36
9
124
TOTAL
500
1
1
1
1
7
3
4
1
1
1
3
1
14
95
MNE
Adult
1
3
11
5
12
13
17
22
17
3
2
17
9
16
36
9
83
Juve
1
1
1
1
6
3
Infant
2
1
1
Lepus sp.
NISP
Adult Juve
1
1
1
1
MNE
Adult
Juve
1
1
1
1
Damage
Punct Burned
2
1
1
1
1
1
1
14
1
2
1
1
1
1
1
1
1
1
1
2
1
2
1
10
2
9
2
2
1
1
96
28
11
372
27
wolves tend to bite and chew rabbit bones leaving less
complete elements, and humans usually create limb
shafts, breaking the proximal and distal ends of long
bones (e.g. humeri, femora and tibiae) to extract bone
marrow. Sometimes humans also leave cut-marks on
rabbit bones, predominantly on mandibles and limbs,
as has been shown for the Magdalenian levels at Lapa
do Suão (about 15 % of the collection shows cut-marks,
many of them seen only under a microscope ; HAWS &
VALENTE, 2001 ; VALENTE, 2000 ; also see PÉREZ-RIPOLL,
1992 for higher percentages).
In the Pego do Diabo assemblage, all skeletal bones are
equally present, showing no differential transportation of
any part of the rabbit carcasses. There are no cut-marks
and most of the limb bones have at least one of their
ends (Fig. 7), reinforcing the absence of anthropic and
large carnivore activity. There are also two limb bones
that show opposite tooth punctures on the distal end (in
particular the femora ; see Fig. 8), which is typical of
small carnivore activity, such as lynx and fox (HOCKETT,
1993, 1999). The potential human influence is limited to
four burned bones. Five remains display the characteristic
singular puncture of raptor activity.
As mentioned before, there is a small number of hare
bones present. None of these specimens represents an
entire element, and all long bones have both or one of its
7
8
4
epiphyses. One proximal tibia exhibits a single puncture,
typical of raptor damage.
CONCLUSIONS ON PEGO DO DIABOʼS
ASSEMBLAGE
Gathering all the evidence, Pego do Diaboʼs Early Upper
Paleolithic faunal assemblage can be characterized
as a palimpsest of different depositional agents, most
possibly affected by bioturbation and sedimentary
or hydrological transportation. This interpretation is
confirmed by the radiocarbon dates and reinforced by the
noted differences in mortality age profiles (bimodal), and
in particular by the probable multiple numbers of agents
that accumulated and modified bones, such as carnivores,
raptors and humans.
In 1997, ZILHÃO said that Pego do Diabo Cave was
sporadically used by small human groups, who organised
and maintained their hunting equipment (1997, 2 : 99).
According to him, the large carnivores found in the cave
accumulated most of the faunal remains. This study
does not contradict his interpretations, but the processes
that produced the Pego do Diabo faunal assemblage
seem to be more complex ; besides wolf and hyena, we
should associate humans, iberian lynx and other small
620
M. J. VALENTE
Fig. 7 : Carnivore damage : rabbit femura (upper row) and
tibiae (lower row).
Fig. 8 : Carnivore damage : rabbit femur with opposite
punctures by small carnivore.
all the herbivore species represented contain remains
displaying carnivore gnawing. Hyenas are surely one of
the carnivore agents that modified the herbivore remains,
as gastric action is one of their signatures on chamois and
red deer remains. Wolves and lynxes are a possibility as
well ; the latter may have preyed on immature red deer
and horse, and possibly caprids. In addition, raptors,
foxes, badgers, and above all lynxes, are potential
accumulators of the leporid bone assemblage.
It is also plausible to think that encounters among
carnivores occurred ; perhaps a few of them had mortal
consequences, thereby producing at least part of the
carnivore bone assemblage found inside the cave (two
carnivore bones show possible gnawing marks, one
belonging to wolf, the other to hyena).
The same kind of competition for the use of the cavity
was established between humans and carnivores, since
we found some artefacts and a few herbivore remains
with unmistakable anthropic modifications. Of all the
herbivores, chamois seems to have the highest number of
evidence for human action : prime age prey (the literature
suggests that hyenas, wolves and lynxes prey primarily
on immature herbivores), cut-marks, and a shortage
of typical carnivore modifications. Plus, even if most
caprid bones were not identified to the species level,
some of them may be chamois, including some elements
with high economic utility (e.g. pelvis and humerus ; cf.
BINBFORD, 1978 ; METCALFE & JONES, 1988). The evidence
also suggests that humans modified red deer remains.
It is difficult to detail human intervention in Pego do
Diaboʼs assemblage and numerous hypothesis could be
offered. Humans could have transported animal bones
into the cave as a result of hunting activities or secondary
exploitation of carcasses found in the vicinity ; or,
alternatively, humans could have recovered some parts
of carcasses brought to the cave by carnivores, either to
use them as food (BRUGAL & JAUBERT, 1991) or for other
purposes, such as manufacturing bone tools.
THE NATURE OF HUMAN-CARNIVORE
RELATIONSHIP DURING THE EARLY UPPER
PALEOLITHIC IN PORTUGAL
carnivores, as well as raptors as probable contributors of
bones to the assemblage.
Inter-species competition for the use of the cave was
pronounced. It is clear that during the Upper Pleistocene
several carnivore species (at least hyenas, wolves, bears,
lynxes, foxes and mustelids) used Pego do Diabo as
shelter and/or den, often transporting herbivore prey
remains inside the cave. With the exception of wild boar,
The consecutive use of Pego do Diabo Cave both by
human communities and carnivores is something we
also can perceive in other settlements dated to the Early
Upper Paleolithic in Portugal. Here, we do not have
many archaeological contexts dated from this period,
and those with preserved and studied fauna are even
fewer (see Note 2). Besides Pego do Diabo, we presently
have five cave sites that are early Upper Paleolithic in
age, all in Central Portugal (see Fig. 1 for locations) :
Salemas, Caldeirão, Casa da Moura, Buraca Escura and
Lapa do Picareiro. In the absence of quantitative data, a
Humans and Carnivores in the Early Upper Paleolithic in Portugal
taxonomical list for these sites (and others) is presented
in Appendix 1. Radiometric dates are documented in
Appendix 2.
Salemas Cave is a thin gallery (1 m in width and 30 m
in length) located on a limestone cornice situated not
far from Pego do Diabo. Level III was classified as
Perigordian by ZBYSZEWSKI and ROCHE in the 1960ʼs and
1970ʼs (ZBYSZEWSKI, 1963), but recently ZILHÃO (1997,
2 : 91) re-evaluated the lithic industry, classifying it as
Aurignacian and Early Gravettian. The faunal collection
was never completely studied, but CARDOSO (1993)
analysed the specimens that allowed paleontological and
osteometrical observations (n = 142). Species present
and their relative frequencies are : Felis sylvestris 14,8 %, Lynx pardina - 21,8 %, Canis lupus - 14,3 %,
Vulpes vulpes - 2,1 %, Ursus arctos - 1,4 %, Sus scrofa
- 4,9 %, Cervus elaphus - 31 %, Rupicapra rupicapra
- 7,7 %, and Bos primigenius - 1,4 % (p. 544). No exact
information on the age of the animals is available, but
the existing data show an assemblage similar to Pego do
Diaboʼs level 2.
Caldeirão Cave is a narrow meandering gallery (3 m
in maximum width and 20 m in length) located on
the slope of a small valley at the bottom of which a
stream flows into the Nabão River. The site has Early
Upper Paleolithic occupations : levels Jb and Ja were
respectively classified as Early and Final Gravettian by
ZILHÃO (1997, 2 : 118, 121). He also concluded that the
human role in the bone accumulation was rather modest.
Recent studies by S. DAVIS (2002) showed that large
carnivores, especially hyena, but also cave lion, were
common in these levels. Other carnivores are bear, lynx,
and fox. Some indicators suggest that the cave, during the
early periods of occupation (Middle Paleolithic and Early
Upper Paleolithic), was used as a hyena den : presence
of Crocuta remains, hyena coprolites and gastric etched
bones associated to a relatively high number of juvenile
remains of red deer and equids, scarce burned remains
and a small density of lithic materials (opposed to a high
number of faunal remains). Based on the increasing ratio
of rabbit remains to herbivore remains from the older
Mousterian and Early Upper Paleolithic levels to the
Magdalenian ones, it is also suggested that there was a
gradual increase of hunting pressure on the environment,
maybe due to human population increasing.
At Cesareda plateau, on its north edge, is Casa da Moura
Cave, with a 50 m sinuous gallery. Its outer room (the
one with cultural deposits, measuring about 10 x 20 m)
has a disturbed layer (1b) with materials that according
to ZILHÃO (1997, 2) correspond to three occupations,
Early and Final Gravettian (corresponding to the base
of layer 1b) and Solutrean (top of layer 1b). Excavated
by Delgado in the 19th century, level 1b delivered scant
evidence for human occupation and a faunal assemblage
dominated by rabbit and a wide variety of carnivores
(iberian lynx, leopard, wild cat, hyena, european polecat,
fox, wolf, bear) along with horse, red deer, ibex, chamois
and wild boar (ROCHE, 1951). More recent studies by
621
CARDOSO (1993) and Altuna (on a small sample from
a test excavation made in 1987 by STRAUS ; STRAUS et
al. 1988) again emphasized the high frequency of wolf
remains. The available data strongly suggest that during
the Pleistocene Casa da Moura was essentially a wolf den
intermingled with some human occupation.
Buraca Escura is a small karstic cavity situated on the
southern slope of the Poio Novo valley. It has four
levels with Upper Pleistocene occupations, dated from
the Gravettian, Final Gravettian and Proto-Solutrean,
all with rare artefacts amongst an abundant faunal
assemblage (AUBRY et al., 2001). The faunal remains
were fully analysed by J.-Ph. BRUGAL and recently
published (op. cit.), displaying a relatively high
percentage of carnivores. In the top layer (level 2a)
horse dominates (37,2 % of the NISP), followed by
ibex (31,4 %) and red deer (14 %). Carnivore bones are
present in very small quantities, most of them belonging
to wolf (2,5 %, 3 specimens). All of the herbivore species
are represented both by adults and immature (sometimes
fetal or neonatal) individuals. Nearly half of the postcranial horse remains exhibit cut-marks, and a significant
quantity of caprid bones display evidence of carnivore
action. In layer 2b more than half the herbivore remains
are from ibex (56,1 %). Auroch (14,3 %), red deer (9,7 %)
and horse (8,1 %) are also present and lynx dominates
the carnivore assemblage (3,4 %). There is a large
percentage of immature herbivore individuals, mostly
fetal or neonatal ibex. With the exception of auroch, all
species display typical carnivore modifications and a few
ibex and auroch remains show cut-marks. Two species
are well represented in level 2e : ibex (72,9 %) and lynx
(11,2 %). Small quantities of horse (3,8 %), red deer (2 %)
and wild cat (0,8 %) were also found. Most of the ibex
bones belong to infant (foetal or neonatal) or juvenile
individuals, and carnivores modified an important number
of ibex remains. Finally, layer 2f had a small amount of
ibex remains, totalling two individuals, one adult and
the other a juvenile. Three specimens show carnivore
modifications. Based on these data (zooarchaeological,
few artefacts and burned bones present), AUBRY et al.
(2001 : 41) concluded that in Buraca Escura Cave, Upper
Paleolithic human occupations were short and intermixed
with carnivore lairs.
The Lapa do Picareiro assemblage displays different
features. J. HAWS (2000) summarized the evidence for
carnivore activity in all the Pleistocene cave deposits
as two small marten-sized teeth, a few bones that show
limited punctures and a possible auroch bone that appears
to be acid-etched. None of these features, however, were
found on bones from the supposed Gravettian level (J)
(see Note 2). This layer contained only two identifiable
remains belonging to red deer and caprid. In contrast,
leporid remains dominate all the assemblage, including
the Gravettian occupation. HOCKETT & BICHO (2000)
and BICHO et al. (2000 ; in preparation) point out that
the leporid collection shows no clear evidence for the
natural accumulation of rabbit bones (no puncture marks,
622
M. J. VALENTE
corrosion from gastric fluids, thinning or polishing) ; in
addition, limb elements and cranial bones are abundantly
represented which is typical of human accumulations.
In sum, the researchers concluded that intense rabbit
hunting occurred and rabbit constituted a principal
subsistence item near Picareiro Cave during much of the
Upper Palaeolithic.
important presence of carnivore remains (in number,
but especially in diversity of species) and carnivore
bone modifications. The overall pattern provides us
with a scenario consistent with short and discontinuous
human occupations intermixed with carnivore dens.
Furthermore, the evidence suggests that hyena, wolf and
other large carnivores, lynx and other small carnivores,
and humans, were competitors for the use of the caves as
shelters during this time.
The pattern in human-carnivore relationships seems to
change during the Final Gravettian/Proto-Solutrean,
when sites with significant evidence for carnivore
occupation are much less common and the carnivores do
not seem to play a big role in the bone accumulations. In
Central Portugal, Lapa do Picareiro (level J and I, if we
accept their Gravettian attribution ; BICHO et al., 2000
and in preparation) and Anecrial Cave [level 2 ; ZILHÃO,
CONCLUDING REMARKS
During the Initial Pleniglacial (before Last Glacial
Maximum), Central Portugal had a large number of cave
sites with archaeological deposits dating from the Early
Upper Paleolithic. These occupations are characterized
by a small amount of direct human occupation (notably
artefacts and bone modifications), and by a relatively
9c
FG?
I/H/Fc/Fb/Fa
PS/ES/
/MS/US
(Solutrean)*
HERBIVORES
Sus scrofa
Cervus elaphus
Capreolus capreolus
Bos primigenius
Capra pyrenaica
Rupicapra rupicapra
Equus caballus
CARNIVORES
Canis lupus
Vulpes vulpes
Ursus arctos
Meles meles
Panthera spelaea
Panthera pardus
Lynx pardina
Felis sylvestris
Crocuta c.spelaea
Buraca Grande
2e
FG
Lapa do Picareiro
J
MG?
Anecrial
2f
G
Buraca Escura
Buraca Grande
1b
EG
Caldeirão
Buraca Escura
Caldeirão
Jb/Ja
EG/FG
(EUP)*
Lapa do Picareiro
III
A
+
EG
Buraca Escura
2
A
Casa da Moura
Level
Techno-complex
Salemas
Pego do Diabo
Appendix 1 : Portuguese Upper Pleistocene Fauna present in Early and Middle Upper Palaeolithic sites, listed by ungulates, carnivores
and small mammals (microfauna excluded).
2b/2a
PS
2
PS
I
PS
9b/9a/8
FG/US?
?
?
?
?
?
?
?
SMALL MAMMALS
Lepus sp.
Oryctolagus cuniculus
Note: A – Aurignacian, EG – Early Gravettian, FG – Final Gravettian, EUP – Early Upper Paleolithic, G – Gravettian, MG – Middle
Gravettian, PS – Proto –Solutrean, ES – Early Solutrean, MS – Middle Solutrean, US – Upper Solutrean, ? – Unsure presence of the
species, (---)* – Caldeirão Cave cultural units according to DAVIS 2002.
Humans and Carnivores in the Early Upper Paleolithic in Portugal
623
Appendix 2 : Radiometric Dates (14C) for the mentioned sites (Initial and Middle Upper Paleolithic sites with fauna).
Site
Level
Tecno-complex
Material
Lab. No.
Age B.P
Buraca Escura
Buraca Escura
Buraca Escura
Buraca Grande
Buraca Grande
Caldeirão
Caldeirão
Caldeirão
Caldeirão
Caldeirão
Caldeirão
Caldeirão
Casa da Moura
Pego do Diabo
Pego do Diabo
Pego do Diabo
Salemas
Salemas
Salemas
2a
2e
2f
9a
9b
Fa topo
Fa topo
Fc
H
H
I
Jb
1b
2a
2b
2b
VS (II)
VS (II)
VS (II)
Proto-Solutream
Final Gravettian
Gravettian
Upper Solutrean?
Final Gravettian
Upper Solutrean
Upper Solutrean
Upper Solutrean
Middle Solutrean
Middle Solutrean
Proto-Solutrean
Early Gravettian
Early and Final Gravettian
Aurignacian
Aurignacian
Aurignacian
Proto-Solutrean
Proto-Solutrean
Proto-Solutrean
Bone
Bone
Bone
Charcoal
Charcoal
Bone
Charcoal
Bone
Bone
Bone
Bone
Bone
Bone
Bones
Charcoal
Bones
Bones
Bones
Bones
OxA-5524
OxA-5523
GifA-97258
Gif-9502
GifA-93048
OxA-1938
ICEN-295
OxA-2510
OxA-2511
OxA-1939
OxA-1940
OxA-5542
TO-1102
ICEN-490
ICEN-306
ICEN-732
ICEN-376
ICEN-385
ICEN-367
21,820 ± 200
22,700 ± 240
26,560 ± 450
17,850 ± 200
23,920 ± 300
20,400 ± 270
21,200 + 1,800 – 2,300
18,840 ± 200
20,530 ± 270
19,900 ± 260
22,900 ± 380
26,020 ± 320
25,090 ± 220
23,080 ± 490
2,400 ± 80
28,120 + 860 - 780
20,250 ± 320
19,220 ± 300
17,770 ± 420
Evaluation
A
A
A
A
A
A
A
A
A
A
A
A
?
?
R
A
?
?
?
Note: A – Accepted result; R – Rejected result; ? – Unsure result.
1997 and BRUGAL (pers. comm.)] seem to be the oldest
contexts representing this “new reality”, soon to be a
standard during the Solutrean (see Appendix 1).
This situation during the Early Upper Paleolithic
(Aurignacian and Early Gravettian) can be attributed
either to low density of human population at that time
(STRAUS et al., 2000 ; BICHO, 2000), to geomorphological
factors affecting site preservation and/or to changes in
human use of the landscape (ZILHÃO, 1997, 2001 ; see
also VILLAVERDE et al., 1996 for Middle-Upper Paleolithic
transition in central Spanish Mediterranean zone).
Finally, we cannot overlook that by the Solutrean, large
carnivores such as leopard (Panthera pardus), cave
lion (Panthera spelaea) and hyena (Crocuta crocuta),
decrease or disappear from Portuguese archaeological
contexts.
In Europe, leopard was a frequent presence after the
last interglacial (IS 5e) and appears to become extinct
during the beginning of the Upper Paleolithic (CASTAÑOS,
1990 ; GUÉRIN & PATOU-MATHIS, 1996). In contrast, in
Iberia, this species was common in the northern area
(most especially during the Aurignacian ; cf. ALTUNA &
MARIEZKURRENA, 1988), where it seems to survive until
Magdalenian times (CASTAÑOS, 1990). CARDOSO (1993)
identified most of the known Portuguese specimens
[Lorga de Dine and Furninha dated from the Early
Glacial ; Casa da Moura, Fontainhas (probably Solutrean
levels ; see CARDOSO, op. cit. : 446), Pedreira das Salemas,
Figueira Brava and Escoural dated from the Late Würm]
and, recently, DAVIS (2002) also documented the presence
of leopard specimens in Cardeirão solutrean levels.
Cave lion was widespread in European assemblages
during the last glaciation. In Portugal it was found in five
cave sites [Lorga de Dine (Early Würm), and Pedreira
das Salemas, Figueira Brava, and Escoural (Late Würm),
according to CARDOSO 1993 ; and in Caldeirão Early
Upper Paleolithic levels, see DAVIS, 2002] and seems to
disappear after the Initial Pleniglacial.
Crocuta crocuta is one of the most regular taxa during
the European Middle and Upper Pleistocene. In the
meridional regions of Iberia, the presence of this species
does not seem to be as regular as in the northern areas ;
still, in Portugal, the hyena was found in some cave
sites (most data in CARDOSO, 1993) : Crocuta crocuta
intermedia in two Early Würm sites (Lorga de Dine and
Furninha) ; and Crocuta crocuta spelaea in nine sites
dated from the Late Würm (Caldeirão, Columbeira,
Fontainhas, Lapa da Rainha, Porto Covo, Algar de
Cascais, Figueira Brava, Escoural and Pego do Diabo).
The lack of detailed information does not allow
final conclusions on the causes for the reduction or
disappearance of large carnivores from Portuguese
archaeological contexts. Nonetheless, factors like human
competition and/or changes in ecological conditions
such as a loss of herbivore biomass and the reduction
of herbivore prey availability may help to explain the
observed record (VALENTE & BRUGAL, 2002 ; BRUGAL &
VALENTE, in press).
624
M. J. VALENTE
Notes :
1. Some researchers maintain doubts on the existence
of an Aurignacian techno-complex in Portugal (see
BICHO, 1999, 2000 ; RAPOSO, 2000).
2. Only sites with occupations classified as Aurignacian
and Early or Middle Gravettian, thus dating until circa
22.000 BP (i.e. until the end of Initial Pleniglacial and
before the Last Glacial Maximum). Level J of Lapa
do Picareiro was also included. This layer has no
radiometric dates, but according to N. BICHO (pers.
comm.) its material displays some technological
differences from level I, attributed to Final Gravettian.
Therefore, Level J can be preliminarily classified as
Middle Gravettian.
ACKNOWLEDGEMENTS
This paper is a revised summary of my Masters thesis
presented to the Faculdade de Letras da Universidade
de Lisboa, funded by Praxis XXI - Junta Nacional de
Investigação Científica e Tecnológica. The study also
beneficiated from two scholarships : one from Centre
National de la Recherche Scientifique and Instituto
de Cooperação Científica e Tecnológica Internacional
(Proc. 423/França ; dir. of J. ZILHÃO and J.-Ph. BRUGAL) ;
the other from Ambassade de France au Portugal
and Instituto de Cooperação Científica e Tecnológica
Internacional (Proc. 001 A0, Dossier nº 97/001 ; dir. of
L. RAPOSO and J.-Ph. BRUGAL).
I am indebted to Museu Nacional de Arqueologia and
to João ZILHÃO for making the Pego do Diabo collection
available for study. I would like to thank Jean-Philip
BRUGAL for inviting me to participate in the UISPP 2001
symposium at Liège. I am particularly grateful to JeanPhilip BRUGAL, Nuno BICHO and Bryan HOCKETT for their
comments on the manuscript.
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