Pollen in hyena coprolites from Gabasa Cave (northern Spain)
←
→
Page content transcription
If your browser does not render page correctly, please read the page content below
Available online at www.sciencedirect.com
R
Review of Palaeobotany and Palynology 126 (2003) 7^15
www.elsevier.com/locate/revpalbo
Pollen in hyena coprolites from Gabasa Cave (northern Spain)
P. Gonza¤lez-Sampe¤riz a; , L. Montes b , P. Utrilla b
a
Instituto Pirenaico de Ecolog|¤a^CSIC, Campus de Aula Dei, Apdo. 202, 50.080 Zaragoza, Spain
b
Department of Antiquity Sciences (Prehistory), Pedro Cerbuna 12, 50.009 University of Zaragoza, Zaragoza, Spain
Received 6 June 2002; accepted 20 February 2003
Abstract
Pollen from hyena coprolites are useful to describe palaeovegetation and to infer palaeoclimatic conditions. In
previous studies, sedimentological and palynological analyses of the Mousterian Gabasa Cave (Pyrenean foothill
region, northeast Spain) have not been able to provide detailed palaeoenvironmental information. In this study,
coprolites of the extinct cavern hyena (Crocuta crocuta) from Gabasa Cave were used for pollen analysis with the goal
of establishing the regional environmental history between s 50 700 and s 39 900 yr BP 14 C (AMS). Eight of the
twelve coprolites analyzed contained well-preserved and rich pollen assemblages. They indicated the development of a
mosaic glacial landscape that included Pinus and Juniperus woodlands and steppes of Chenopodiaceae, Poaceae,
Artemisia, and Asteraceae. Pollen for mesophilous and thermophilous trees and shrubs are also present, suggesting the
location of nearby refugia of temperate and Mediterranean vegetation.
7 2003 Elsevier B.V. All rights reserved.
Keywords: coprolites; hyena; palaeoecology; Spain; Pleistocene; Palaeolithic; glacial refugia
1. Introduction swamps. Coprolites can be used to analyze para-
sites, the main plant taxa used for food and even
Pollen from peat bog, lacustrine and £uvial en- the season of ingestion. The results are not always
vironments, archaeological sites and even strati- as informative as in conventional palynology of
¢ed slope deposits are usually studied for palaeo- lakes and peat bog sites, but notwithstanding cop-
environmental interpretation (Gonza¤lez-Sampe¤riz rolite palynology has proven its potential in recent
et al., 2002). The analysis of coprolites may also studies (Carrio¤n et al., 2000, 2001).
provide useful information about palaeoclimate, This paper discusses the study of a collection of
palaeovegetation and even palaeoethnology (Bry- twelve coprolites from a Mousterian cave site in
ant and Holloway, 1983; Davis, 1990; Scott and northeast Spain of which eight specimens yielded
Cooremans, 1992; Carrio¤n et al., 2000, 2001), es- very well-preserved pollen. Previous studies did
pecially in arid environments without lakes or not ¢nd good correlations among the macrofau-
nal, microfaunal, palynological and sedimentolog-
ical evidence from this site (Azanza et al., 1988;
* Corresponding author.
E-mail addresses: pgonzal@posta.unizar.es (P. Gonza¤lez-
Hoyos et al., 1992). Besides, pollen percolation in
Sampe¤riz), lmontes@posta.unizar.es (L. Montes), the cave impeded more accurate interpretations
utrilla@posta.unizar.es (P. Utrilla). (Hoyos et al., 1992). Our results help to clarify
0034-6667 / 03 / $ ^ see front matter 7 2003 Elsevier B.V. All rights reserved.
doi:10.1016/S0034-6667(03)00033-2
PALBO 2527 15-8-038 P. Gonza¤lez-Sampe¤riz et al. / Review of Palaeobotany and Palynology 126 (2003) 7^15
Fig. 1. Location of the Mousterian Gabasa Cave (northern Spain).
uncertainties regarding the plant cover in the vi- tributary of the Cinca River ^ in the Pyrenean
cinity of the Mousterian Gabasa Cave site. We foothill region of northeast Spain (Fig. 1). The
selected samples from abundant hyena coprolites basal lithology is composed of Cretaceous and
in the archaeological excavation (Blasco and Palaeocene limestones, resulting in a karstic relief
Montes, 1997) in order to perform a palynological with abundant caves, many of them containing
analysis in an attempt to provide a complete and prehistoric artifacts.
coherent picture of past regional environmental The climate of the region is of a mountain con-
and climate change. It is worth stressing that the tinental Mediterranean type. Mean annual tem-
pollen in hyena coprolites give a regional perspec- perature is 14‡C, with a large range (July 30‡C
tive of the palaeoenvironment, since these animals and January 35‡C). Mean annual precipitation
travel long distances in search of food (Scott, is about 550 mm. Maximum rainfall occurs in
1995; Carrio¤n et al., 2001). Hyena coprolites re- spring and autumn (Garc|¤a-Ruiz et al., 2000).
£ect more regional pollen sources than sediment Modern vegetation includes a Mediterranean
analysis, as is demonstrated by Scott and Klein mixed oak forest with Quercus rotundifolia and
(1981) and Scott (1987) in the South African hye- Q. faginea, especially in the shady aspects, accom-
na sites of Deelpan, Free State and Equus Cave, panied by scrubs of Juniperus communis, J. sabina,
Kalahari region. The results are especially inter- Buxus sempervirens, Pistacia lentiscus, and Ros-
esting when compared to the pollen analyses of marinus o⁄cinalis. The most frequent species of
the sedimentary matrices. the adjacent riparian communities are Populus ni-
gra, P. alba, Salix alba and Ulmus campestris.
2. Physical setting
3. Stratigraphy of the cave
Gabasa Cave (42‡00P20QN, 4‡06P20QE, 780 m
above sea level) faces south at 150 m above the Gabasa Cave has two small chambers (each
channel in the headwater of the Sosa River ^ a measuring ca. 30 m2 ) with remnants of human
PALBO 2527 15-8-03P. Gonza¤lez-Sampe¤riz et al. / Review of Palaeobotany and Palynology 126 (2003) 7^15 9
Fig. 2. Stratigraphy of Gabasa Cave. Location of the coprolites analyzed in the stratigraphic layers indicated by stars.
settlement located in the inner room. The stratig- and McIntyre (1981) and not to the beginning of
raphy (Fig. 2) shows eight archaeologically de- Isotopic Stage 3. The absolute chronology of the
¢ned beds (Hoyos et al., 1992). Layer ‘b’ is thin, cave sequence (Table 1) is somewhat imprecise
sterile, and discontinuous. The whole deposit is (minimum age), since the dates are beyond the
14
very homogeneous (except for layer ‘h’), which C time scale (Montes et al., 2000). The Gabasa
makes it di⁄cult to separate layers based on color sequence could be included in Oxygen Isotope or
and texture. According to several authors, each Marine Isotope Stage (OIS) 3 of Ruddiman and
layer corresponds to long periods of intermittent McIntyre (1981), but an older age cannot be ruled
occupation as revealed by the artifact contents out. The absence of human structures (except for
(Montes, 1988; Utrilla and Montes, 1989, 1993; a ¢replace in the ‘h’ layer), the nature of the sedi-
Blasco, 1995; Blasco et al., 1996; Montes et al., ment and the presence of Mousterian lithic tools
2000). suggest a sporadic, though persistent use of the
Chronology and climatic interpretation used in cave by one or a few human groups during the
the ¢rst sedimentological study of Gabasa Cave Mousterian period. The animal remains suggest
(Hoyos et al., 1992) were based on preliminary that the cave was used by humans during seasonal
data. After new radiocarbon dating, Montes et hunting (Blasco, 1995; Blasco et al., 1996), alter-
al. (2000) attributed the sediment of Gabasa nating with use as an animal shelter by hyenas,
Cave to the central part of the curve of Ruddiman bears and cavern lions.
Table 1 4. Hyena coprolites : description and methodology
Radiocarbon dating of Gabasa Cave
Gabasa Cave Radiocarbon dating Coprolite Fifty-four coprolites or fragments were found
layers (BP) samples
in the sediment of the Gabasa Cave pit (Plate
Layer a s 39 900 H.A.1 I). The 12 largest samples were used for pollen
Layer b s 45 900 No coprolite
analysis. Most coprolites weighed less than 30 g
Layer c s 46 900 and s 47 800 No coprolite
Layer d H.D.1 (Table 2). Their characteristics and the associated
Layer e s 45 600 and s 51 900 H.E.2, H.E.4 skeleton fragments of at least 13 individuals cor-
Layer f H.F.4 respond with an age distribution that was typical
Layer g s 50 700 H.G.4, H.G.8 of hyena dens (adults and sub-adults) of Crocuta
Layer h H.H.2
crocuta spelaea, the cavern hyena from the Wu«rm
PALBO 2527 15-8-0310 P. Gonza¤lez-Sampe¤riz et al. / Review of Palaeobotany and Palynology 126 (2003) 7^15
Plate I. Spotted hyena coprolites of Gabasa Cave.
period (Blasco, 1995). This animal was larger than contamination in the ¢rst coprolites. Chemical
the modern spotted hyena (Kurte¤n, 1968) and treatment followed the conventional method (Del-
perhaps it was a hunter and not a scavenger. court et al., 1959; Moore et al., 1991; Dupre¤,
This possibility becomes crucial in the interpreta- 1992): HF, HCl, KOH, concentration with Thou-
tion of the pollen spectra. let solution with density 2, and Lycopodium cla-
The coprolite surfaces are yellowish to pale vatum tablets (Stockmarr, 1971) to calculate the
brown and they are whitish inside. The surfaces pollen concentrations.
form a dense-hard cortex that presumably pre- Hyenas digest su⁄cient organic and plant mat-
served the pollen. Thus, each coprolite was ter, including pollen, that is preserved in their
treated as a whole and was not split (Carrio¤n et faecal material (Wernet, 1955). In favourable
al., 2001). In the laboratory, the coprolites were cases, it is possible to compare the sediment and
washed with distilled water to minimize contami- coprolite pollen contents from the same locations
nation from external sources. This method ap- (southern Spain; Carrio¤n et al., 2001), and even
pears valid since we veri¢ed the absence of pollen with coprolites of di¡erent species (Canis and
Table 2
Description and pollen characteristics of the coprolites analyzed
Hyena coprolite samples Weight Taxa numbers Pollen concentration Indeterminate
(g) (grains/g) (%)
H.A.1 30 30 12 720 6
H.D.1 38 37 1 707 5
H.E.2 25 26 3 990 3
H.E.4 44 26 1 304 6
H.F.4 23 37 4 289 5
H.G.4 32 17 3 606 9
H.G.8 23 16 12 275 4
H.H.2 35 31 3 383 4
PALBO 2527 15-8-03P. Gonza¤lez-Sampe¤riz et al. / Review of Palaeobotany and Palynology 126 (2003) 7^15 11
Capra in pre-Roman settlements (Gonza¤lez-Sam- and junipers. Overall, it suggests a cold, arid cli-
pe¤riz, 2001). This approach provides useful data mate, typical of the last glacial period in continen-
to test the validity of the results from both types tal Iberia (Pons and Reille, 1986; Turner and
of pollen analysis: sediments and coprolites (Gon- Hannon, 1988; Pen‹alba, 1989; Montserrat,
za¤lez-Sampe¤riz, 2001). The coprolite pollen spec- 1992; Pe¤rez-Obiol and Julia', 1994; Carrio¤n and
tra of herbivorous animals are theoretically in£u- van Geel, 1999).
enced by diet regime and plant availability. In Mesophilous vegetation occurred in the pollen
archaeological settlements on the island of Mal- catchments. It includes Betula, Corylus, Ulmus,
lorca, coprolite pollen analysis from the endemic Fagus, and the Mediterranean Quercus ilex^cocci-
goat, Myotragus balearicus, demonstrated that fera type and Quercus faginea^pubescens type. The
this extinct ruminant had a monospeci¢c diet pollen content of the hyena coprolites is probably
based on Buxus. Thus, the extinction of M. ba- derived from a complex landscape, including a
learicus has been related with the extinction of number of biotopes, and trapped by hyenas while
Buxus in the island (Yll Aguirre et al., 2001). traveling up to 50 km away from their dens and
Since hyenas are not herbivorous, the ingestion eating a variety of fauna (Mills, 1989). In the
of pollen and spores might be accidental or indi- same archaeological layer, the pollen spectra
rect through respiration (air) in addition to the show a large variability. Other studies of pollen
oral system (water, ingestion of vegetable matter in spotted hyena coprolites also provide informa-
via the stomach contents of prey) (Carrio¤n et al., tion about a regional, heterogeneous plant com-
2001). In this study, the information obtained position, as in Las Ventanas Cave, southern Spain
from coprolites is not used to describe the diet (Carrio¤n et al., 2001), dated to the end of the
but to help understand the palaeoenvironment. Pleistocene. This site was characterized by patchy
The pollen spectra of hyena coprolites in two grass vegetation, heliophytic herbaceous plants
Palaeolithic settlements in Auvergne ^ Saint-Hip- like Artemisia and Chenopodiaceae, pine forests
polyte (Puy-de-Do“me) and Cha“telperron (Allier) ^ and glacial plant refugia. All these features were
in France also proved to be representative of the present in several hyena coprolites from the same
pollen rain as they compared well with the pollen layer.
spectra of the sediment samples (Vivent, 1989). The suggested plant composition varies in all
This author observed the absence of pollen con- layers at Gabasa, but the climatic conditions in-
tamination during the digestive process in the hy- terpreted were similar. In the top layer (‘a’ layer),
ena. Frequently, there is no over- or underrepre- pollen analysis suggests more temperate and hu-
sentation of speci¢c taxa in the spore^pollen mid conditions (Fig. 3) instead of arid and cold/
contents of coprolites, but there may be a slight cool conditions. The ‘a’ layer had a higher pro-
enrichment in entomophilous pollen and shrubs portion of arboreal pollen, especially Pinus (60%
(Girard, 1987; Burjachs, 1988). of the total pollen content) and Quercus ilex^coc-
cifera type (10%). In Mediterranean Europe, ex-
cept in the case of glacial refugia situations, the
5. Palynological results increase of Quercus ilex^coccifera type is usually
interpreted as belonging to interglacial or rela-
In general, the pollen spectra in the studied tively warm periods (Wijmstra, 1969; Woillard,
coprolites (Fig. 3) indicate a patchy landscape 1978; Pe¤rez-Obiol, 1987; Dupre¤, 1988; Burjachs,
composed of steppe and forest areas distributed 1989; Carrio¤n et al., 1993).
according to topography (especially altitude and It is interesting that the bottom and middle of
aspect) similar to the current foothill landscape of the sequence had an unusual proportion of Che-
the Pyrenees. The region is transitional between nopodiaceae in the ¢rst case, and Plantago in
the Ebro Valley and the mountains. The steppes layer ‘e’. These plants are appreciated by rumi-
were composed of chenopods, composites, nants, particularly Plantago, because their leaves
grasses, and Artemisia, with patches of pines are very nutritious. The high percentages of these
PALBO 2527 15-8-0312 P. Gonza¤lez-Sampe¤riz et al. / Review of Palaeobotany and Palynology 126 (2003) 7^15
Fig. 3. Pollen diagram of hyena coprolites from Gabasa Cave (northern Spain).
PALBO 2527 15-8-03P. Gonza¤lez-Sampe¤riz et al. / Review of Palaeobotany and Palynology 126 (2003) 7^15 13
two taxa in hyena coprolites are interpreted as the more regional pollen sources than the sediment
result of consumption of some ruminant stomach matrix, but very little is known about pollen in-
contents. The Mousterian cavern hyena was larg- corporation and post-depositional processes in
er than the modern hyena and it could have chos- hyena coprolites. The pollen analyses presented
en its meat food before other mammals (except in this paper are a small new contribution to
felines). These hyenas would have been able to this debate.
eat more stomachs and intestines and, conse-
quently, ingest more pollen grains from these se-
lected taxa (Kruuk, 1972). On the other hand, the Acknowledgements
prey of the hyena mainly consumed grasses, and
grass swards are excellent traps for regional pollen We would like to thank the people and institu-
(Gutie¤rrez et al., 1998). tions that collaborated with the following proj-
ects: DIGICYT (PB93/0307) ‘Transpyrenees com-
munication during the Lateglacial and the
6. Conclusions beginning of the Holocene’ and DGES (PB97/
1030) ‘The Ebro valley: a communication road
Pollen analysis can be performed on samples to southern Spain during the passage of Lategla-
from many types of sites and materials, including cial to the Holocene’. We also acknowledge the
faecal material or coprolites. In this paper, the Diputacio¤n General de Arago¤n and the Pyrenean
pollen content of hyena coprolites from the Institute of Ecology (CSIC) for ¢nancial support.
Mousterian Gabasa Cave (northern Spain) pro- The pollen were analyzed at the History and
vided information about the palaeoenvironment Geography Sciences Laboratory of the University
and palaeoclimate, and reinforced previous ar- of Zaragoza. We are grateful to Jose¤ S. Carrio¤n
chaeological, palynological, sedimentological and (University of Murcia), Miche'le Dupre¤ (Univer-
chronological studies. It also showed the existence sity of Valencia) and Jose¤ M. Garc|¤a-Ruiz and
of pollen percolation in the stratigraphy of the Blas Valero-Garce¤s (Pyrenean Institute of Ecol-
cave. ogy^CSIC), for thoughtful suggestions and opin-
Our results are in agreement with the revised ions.
chronology (Montes et al., 2000) for the cave
and various regional environmental and climatic
interpretations (Pe¤rez-Obiol, 1987; Dupre¤, 1988; References
Burjachs, 1989; Carrio¤n et al., 1993, 2001).
The pollen analysis of hyena coprolites in Ga- Azanza, B., Baldellou, V., Cuchi, J.A., Lo¤pez, P., Montes, L.,
basa Cave suggests an arid and cold/cool climate Utrilla, P., 1988. Cronoestratigraf|¤a de la cueva musteriense
de los Moros (Gabasa, Huesca). Cuaternario y Geomorfo-
except in the upper layer (‘a’), which suggests a
log|¤a, vol. 2 (1^4). AEQUA, pp. 1^12.
probably warmer climate due to the relatively Blasco, M.F., 1995. Hombres, ¢eras y presas. Estudio arqueo-
high proportion of Quercus ilex^coccifera type. zoolo¤gico y tafono¤mico del yacimiento del Paleol|¤tico Medio
The rest of the layers show a complex landscape de la cueva de Gabasa I (Huesca). Departimento de Ciencias
formed by xerophyte-steppe herbaceous plants de la Antigu«edad (Area de Prehistoria), Universidad de Zar-
agoza, Monograf|¤as Arqueolo¤gicas 38, 205 pp.
and forester patches (essentially coniferous). The
Blasco, M.F., Montes, L., Utrilla, P., 1996. Deux mode'les de
presence of a proportion of mesophilous vegeta- strategie occupationelle dans le Mouste¤rien Tardif de la val-
tion suggests the occurrence of refugia of temper- le¤e de l’Ebre: les grottes de Pen‹a Miel et Gabasa. In: Car-
ate trees which were probably related with favor- bonell, E., Vaquero, M. (Eds.), The Last Neandertals, The
able topographic conditions in the Ebro valley. First Anatomically Modern Humans, Barcelona, pp. 289^
The varied composition of the regional vegetation 313.
Blasco, M.F., Montes, L., 1997. Los hie¤nidos del yacimiento
is probably re£ected in the pollen composition of Musteriense de Gabasa I (Huesca, Espan‹a). Instituto de
the hyena coprolites due to the mobility of these Estudios Altoaragoneses, Huesca, Bolskan 14, pp. 9^27.
animals. It is clear that hyena coprolites re£ect Bryant, V.M., Jr., Holloway, R.G., 1983. The role of palynol-
PALBO 2527 15-8-0314 P. Gonza¤lez-Sampe¤riz et al. / Review of Palaeobotany and Palynology 126 (2003) 7^15
ogy in archaeology. Adv. Archeol. Method Theor. 6, 191^ Gutie¤rrez, A., D|¤ez, M.J., Carrio¤n, J.S., 1998. Primeros datos
224. sobre la deposicio¤n de polen actual en el Parque Natural de
Burjachs, F., 1988. Ana¤lisis pol|¤nico de los niveles cera¤micos los Alcornocales, Ca¤diz, a trave¤s de la evaluacio¤n de cepel-
de la cova 120 (Alta Garrotxa, Catalunya). VI Simposio de lones de gram|¤neas. In: Fombella, M.A. (Ed.), Actas XII
Palinolog|¤a de APLE, Ed. Universidad de Salamanca, Acta Simposio APLE, Leo¤n, p. 118.
Salmanticencia 65, pp. 285^290. Hoyos, M., Utrilla, P., Montes, L., Cuchi, J.A., 1992. Estrati-
Burjachs, F., 1989. Palinologia dels dolmens de l’Alt Emporda¤ graf|¤a, sedimentolog|¤a y paleoclimatolog|¤a de los depo¤sitos
i dels diposits quaternaris de la cova de l’Arbreda i del Pla musterienses de la Cueva de los Moros de Gabasa. Cuater-
de l’Estany. Evolucio¤ del paitsage vegetal i del clima des de nario y Geomorfolog|¤a, vol. 6 (1^4), AEQUA, pp. 143^156.
fa me¤s de 140.000 anys al N.E. de la Pen|¤nsula Ibe¤rica. Kruuk, H., 1972. The Spotted Hyena: A Study of Predation
Ph.D. Thesis, Universidad Auto¤noma de Barcelona, Bella- and Social Behaviour. University of Chicago Press, Chicago,
terra, 324 pp. IL.
Carrio¤n, J., van Geel, B., 1999. Fine-resolution Upper Weich- Kurte¤n, B., 1968. In: Weidenfeld, Nicholson (Eds.), Pleisto-
selian and Holocene palynological record fron Navarre¤s cene Mammals of Europe. London.
(Valencia, Spain) and a discussion about factors of Medi- Mills, M.G.L., 1989. The comparative behavioural ecology of
terranean forest sucession. Rev. Palaeobot. Palynol. 106, hyenas: the importance of diet and food dispersion. In:
209^236. Gittleman, J.L. (Ed.), Carnivore, Behaviour, Ecology and
Carrio¤n, J., Fumanal, M.P., Iturbe, G., 1993. La secuencia Evolution. Chapman and Hall, London, pp. 125^142.
pol|¤nica de Cova Beneito en su marco litoestratigra¤¢co, ar- Montes, L., 1988. El Musteriense en la cuenca del Ebro. De-
queolo¤gico y geocronolo¤gico. Universidad de Valencia, Es- partimento de Ciencias de la Antigu«edad, Universidad de
tudios sobre Cuaternario, pp. 139^148. Zaragoza, Monograf|¤as Arqueolo¤gicas 28, 326 pp.
Carrio¤n, J.S., Scott, L., Hu¡man, T., Dreyer, C., 2000. Pollen Montes, L., Utrilla, P., Hedges, R., 2000. Le passage Pale¤oli-
analysis of Iron Age cow dung in Southern Africa. Veg. thique Moyen-Pale¤olithique Supe¤rieur dans la valle¤e de
Hist. Archaeobot. 9, 239^249. l’Ebre (Espagne). Datations radiome¤triques des grottes de
Carrio¤n, J.S., Riquelme, J.A., Navarro, C., Munuera, M., Pen‹a Miel et Gabasa. In: Zilhao, J., Aubry, T., Carvalho,
2001. Pollen in hyena coprolites re£ects late glacial land- A. (Eds.), Les premiers hommes modernes de la Pe¤ninsule
scape in southern Spain. Palaeogeogr. Palaeoclimatol. Pa- Ibe¤rique. Actes du Colloque de la Commission VIII de
laeoecol. (in press). l’UISPP, Trabalhos d’Arqueologia 17, Lisboa, pp. 87^102.
Davis, O.K., 1990. Caves as sources of biotic remains in arid Montserrat, J., 1992. Evolucio¤n glaciar y postglaciar del clima
western North America. Palaeogeogr. Palaeoclimatol. Pa- y la vegetacio¤n en la vertiente sur del Pirineo. Monograf|¤as
laeoecol. 76, 331^348. del I.P.E.^CSIC, Zaragoza, 147 pp.
Delcourt, A., Mullenders, W., Pie¤rard, P., 1959. La pre¤para- Moore, P., Webb, J.A., Collinson, A., 1991. An Illustrated
tion des spores et des grains de pollen actuels et fossiles. Guide to Pollen Analysis. Hodder and Stroughton, London,
Nat. belg. 40, 91^120. 216 pp.
Dupre¤, M., 1988. Palinolog|¤a y paleoambiente. Nuevos datos Pen‹alba, C., 1989. Dynamique de la ve¤ge¤tation tardiglaciare et
espan‹oles. Referencias. Serie de trabajos varios, S.I.P. Va- holoce'ne du centre^nord de l’Espagne d’apre's l’analyse pol-
lencia, 160 pp. linique. Ph.D. Thesis, Universite¤ d’Aix-Marseille.
Dupre¤, M., 1992. Palinolog|¤a. Sociedad Espan‹ola de Geomor- Pe¤rez-Obiol, R., 1987. Evolucio¤ del paitsage vegetal quaternari
folog|¤a, 30 pp. en zones d’Olot i Sils. Ph.D. Thesis, Universidad Auto¤noma
Garc|¤a-Ruiz, J.M., Arna¤ez, J., White, S., Lorente, A., Be- de Barcelona.
guer|¤a, S., 2000. Uncertainty assessment in the prediction Pe¤rez-Obiol, R., Julia', R., 1994. Climatic changes on the Ibe-
of extreme rainfall events: an example from the Central rian Peninsula recorded in a 30.000-yr pollen record from
Spanish Pyrenees. Hydrol. Proc. 14, 887^898. Lake Banyoles. Quat. Res. 41, 91^98.
Girard, M., 1987. Contenu vegetal: pollen, spores. In: Re- Pons, A., Reille, M., 1986. Nouvelles recherches pollenanaly-
nault-Miskovsky, J. (Ed.), Ge¤ologie de la Pre¤histoire. Asso- tiques a' Padul (Granada): la ¢n du dernier glacier et l’Ho-
ciation pour l’e¤tude de l’environnement ge¤ologique de la loce'ne. In: Lo¤pez Vera, F. (Ed.), Quaternary Climate in the
Pre¤histoire, Paris, pp. 587^617. Western Mediterranean. Universidad Auto¤noma de Madrid,
Gonza¤lez-Sampe¤riz, P., 2001. Ana¤lisis palinolo¤gico aplicado a pp. 405^420.
la reconstruccio¤n paleoclima¤tica en medios mediterra¤neos y Ruddiman, W.F., McIntyre, A., 1981. The North Atlantic
eurosiberianos. Unpubl. Ph.D. Thesis, Universidad de Zar- Ocean during the last deglaciation. Palaeogeogr. Palaeocli-
agoza, 354 pp. matol. Palaeoecol. 35, 14^214.
Gonza¤lez-Sampe¤riz, P., Valero-Garce¤s, B., Garc|¤a-Ruiz, J.M., Scott, L., 1987. Pollen analysis of hyena coprolites and sedi-
Mart|¤, C., 2002. Lateglacial and Holocene vegetational ments from Equus cave, Taung, southern Kalahari (South
change in the Pyrenees and the Central Ebro valley (NE Africa). Quat. Res. 28, 144^156.
Spain). In: Quaternary Climatic Changes and Environmen- Scott, L., 1995. Palaeoenvironmental conditions in South Afri-
tal Crises in the Mediterranean Region, Alcala¤ de Henares, ca at the Pleistocene^Holocene transition. Quat. Sci. Rev.
Madrid; 15^18 July 2002 (CD-rom) (in press). 14, 937^947.
PALBO 2527 15-8-03P. Gonza¤lez-Sampe¤riz et al. / Review of Palaeobotany and Palynology 126 (2003) 7^15 15
Scott, L., Klein, R.G., 1981. A hyena-accumulated bone as- Vivent, D., 1989. Analyses polliniques de coprolites d’hye'nes
semblage from Late Holocene deposits at Deelpan, Orange de deux sites pale¤lolithiques d’Auvergne (Saint Hippolyte,
Free State. Ann. S. Afr. Mus. 86, 217^227. Puy-de-Do“me et Cha“telperron, Allier). Revue d’Arche¤ologie
Scott, L., Cooremans, B., 1992. Pollen in recent Procavia (hy- du centre de la France 28 (2), 229^235.
rax), Petromus (dassie rat) and bird dung in South Africa. Wernet, P., 1955. Relief d’hye'nes quaternaires des loess
J. Biogeogr. 19, 205^215. d’Achenheim: matie're premie're de l’industrie osseuse hu-
Stockmarr, J., 1971. Tablets with spores used in absolute pol- maine. Bull. de l’Association Philomatique de l’Alsace et
len analysis. Pollen Spores 13, 614^621. de Lorraine 9 (3), 150^156.
Turner, C., Hannon, G.E., 1988. Vegetational evidence for late Wijmstra, T.A., 1969. Palynology of the ¢rst 30 m of a 120 m
Quaternary climatic changes in southwest Europe in relation deep section in NE Greece. Acta Bot. Neerl. 18, 511^527.
to the in£uence of the North Atlantic Ocean. Philos. Trans. Woillard, G., 1978. Grande Pile peat bog: a continuous pollen
R. Soc. Lond. B 318, 451^485. record for the last 140.000 years. Quat. Res. 9, 1^21.
Utrilla, P., Montes, L., 1989. La grotte mouste¤rienne de Ga- Yll Aguirre, E., Pantaleo¤n-Cano, J., Roure, J., 2001. Ana¤lisis
basa (Huesca, Espagne). In: Otte, M. (Ed.), L’homme de pol|¤nico de coprolitos de Myotragus balearicus: La extincio¤n
Ne¤anderthal, vol. 6 (La subsistance). Lie'ge, pp. 145^153. de especies animales y vegetales durante el Holoceno en
Utrilla, P., Montes, L., 1993. El ¢nal del Musteriense en el Mallorca (Islas Baleares). In: Fombella Blanco, M.A., Fer-
valle del Ebro: Datos y re£exiones. In: Cabrera, V. (Ed.), na¤ndez Gonza¤lez, D., Valencia Barrera, R. (Eds.), Palinolo-
El origen del hombre moderno en el suroeste de Europa. g|¤a: Diversidad y Aplicaciones. Actas XII Simposio de
UNED, Madrid, pp. 219^246. APLE, Leo¤n, pp. 205^212.
PALBO 2527 15-8-03You can also read
