RESEARCH PAPER

copy 2003 Blackwell Publishing Ltd httpwwwblackwellpublishingcomjournalsgeb

Global Ecology amp Biogeography

(2003)

12

119ndash129

Blackwell Science Ltd

Glacial refugia of temperate Mediterranean and Ibero-North African flora in south-eastern Spain new evidence from cave pollen at two Neanderthal man sites

JOSEacute S CARRIOacuteN ERRIKARTA IYLLdagger MICHAEL J WALKERDagger ALFONSO J LEGAZDagger CELIA CHAIacuteNsect and ANTONIO LOacutePEZDagger

Area de Botaacutenica Facultad de Biologiacutea Universidad de Murcia 30100 Murcia Spain

dagger

Area de Botaacutenica Facultad de Ciencias Universidad Autoacutenoma de Barcelona 01893 Bellaterra Barcelona Spain

Dagger

Area de Antropologiacutea Fiacutesica Departamento de Zoologiacutea y Antropologiacutea Fiacutesica Facultad de Biologiacutea Universidad de Murcia 30100 Murcia Spain

sect

Departamento de Informacioacuten y Documentacioacuten Facultad de Ciencias de

la Documentacioacuten Universidad de Murcia 30100 Murcia Spain

ABSTRACT

Aim

To locate glacial refugia of thermophilous plant speciesin Spain

Location

Two south-eastern Spanish Neanderthal man sitesin Murcia namely the inland Cueva Negra del Estrecho delRiacuteo Quiacutepar and the coastal Sima de las Palomas del CabezoGordo

Methods

We use pollen found in cave sediments as a sourceof palaeobotanical and palaeoecological information Thefindings are discussed with regard both to animal remainsfrom both sites and also to other refugia in south-easternSpain and elsewhere in the Iberian Peninsula

Results

Both sequences show persistence of abundant mes-othermophilous trees during the last glacial stage suggestingboth localities were reservoirs of phytodiversity and wood-land species At both sites deciduous and evergreen oaks arethe most abundant components followed by a wide variety

of deciduous trees and sclerophyllous shrubs including Ibero-North African xerothermic scrub near the coast

Conclusions

Incomplete information underlies a commonmisapprehension that Iberian glacial refugia were confinedto southernmost parts of the peninsula A rather differentpicture of Quaternary refugia emerges from considerationof pollen sequences from caves (and other inputs such asmacroscopic charcoal spatial genetic structure of present-daypopulations faunal remains and present-day distributionof thermophilous species) This picture offers a view of numer-ous viable areas for woodland species in southern Spain inaddition to others in the mountain ranges both in continentalcentral Spain and those of northern Spain these stretch fromthe Mediterranean coast of Catalonia to the westernmostextent of the Bay of Biscay

Key words

archaeobotany glacial refugia historical bio-geography Neanderthal palaeoecology Pleistocene pollenanalysis Spain

INTRODUCTION

It is of palaeoecological biogeographical and evolutionaryinterest when pollen analysis identifies localities where tem-perate and Mediterranean woodland species survived duringthe last ice age (Comes amp Kadereit 1998 Willis amp Whittaker2000) Although the Iberian Peninsula may have retainedsclerophyllous and some broad-leaved species the main

refugia for deciduous trees lay further east in the Italian andBalkan Peninsulas (Bennett

et al

1991 Magri amp Parra 1997)This picture implies Europe-wide migration of temperatewoodland species during lateglacial and early Holocene times(Huntley 1990 Brewer

et al

2002) However new palaeo-botanical findings and reinterpretation of old ones sit uneasilywith this model mdash in particular anthracological findings fromHungary (Willis

et al

2000) indicate more northerly wood-land survivals than hitherto supposed This raises a worryingquestion of whether low tree-pollen percentages owe less tolong-distance transport than to so-called lsquodeficientrsquo pollen dis-persal or production central European pollen sites might thus

Correspondence Joseacute S Carrioacuten Area de Botaacutenica Facultad deBiologiacutea Universidad de Murcia 30100 Murcia Spain E-mailcarrionumes

120

JS Carrioacuten

et al

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

be explained in terms of lateglacial regional expansion of lsquosta-tionaryrsquo populations rather than of continent-wide spreadfrom distant southern refugia

What light can southern regions of the Iberian Peninsulathrow on this question Phytodiversity there today is remark-able with respect to the rest of Europe in both arid Mediter-ranean south-eastern areas (Mota

et al

1997 Saacutenchez-Goacutemez

et al

1998) and humid Atlantic south-south-western ones(Ojeda

et al

1996) Southern Spain today moreover hasseveral thermophilous species of undoubtedly tropical or sub-tropical origin (eg

Maytenus europaeus

Withania frutescens

Periploca angustifolia

Ziziphus lotus

Chamaerops humilis

Tetraclinis articulata

Myrtus communis

Halogeton sativus

Launaea arborescens

Rhododendron ponticum

Myrica gale

Prunus lusitanica

Cosentinia vellea

Culcita macrocarpa

)implying the presence of Pleistocene refuges in the IberianPeninsula that most likely were vestigial relics of an AfricanMiocene biogeographical expansion into Europe during theso-called Messinian lsquocrisisrsquo (Arroyo 1997) How far doespalaeopalynological research corroborate these inferences

Upper Pleistocene palaeobotanical findings from Spainhave tended to be somewhat patchy up to now New pollenfindings are discussed here from two south-eastern Spanishcaves Both have Neanderthal human remains Middle Palaeo-lithic artifacts and wide faunal variety Together with findingsat other sites they show that south-eastern Spain containedsignificant centres of diversity during the last glacial stagefor both deciduous and sclerophyllous trees and shrubs includ-ing thermophilous Ibero-North African species

Study sites

Cueva Negra del Estrecho del Riacuteo Quiacutepar (lsquoBlack Cave of theRiver Quiacutepar Gorgersquo) is near the hamlet of La Encarnacioacutenwithin the township of Caravaca de la Cruz (Murcia) (Fig 1)Today the climate is subhumid to semiarid with 2600ndash2800h of sunshine annually mean January temperatures of 2

degminus

4

deg

C and mean July ones of 20

degminus

22

deg

C The north-facingrock-shelter extends some 12 m backwards from a 12-m-widemouth 4ndash5 m high

The sedimentary fill represents both erosion products of thefossiliferous sandy limestone in which the cave was formedand a significant proportion of fine angular microscopicallypitted silt-sized particles similar to wind-blown loess Thereare no sorted river gravels but thermoclastic scree is veryabundant Five lithostratigraphical units are present in thecave (Fig 2) Unit 1 is superficial disturbed powdery soil ofgrey hues containing modern artifacts and sheep droppingsUnit 2 is a compact yellowish layer comprising diffuse reddishbrown lateritic lenses eroded sand and bioclasts and 15 ofloess-size particles Its upper part is cemented by calcium car-bonate Unit 3 is a yellow to greyish sediment with 10ndash15loess-size wind-blown particles and thermoclastic scree At

the top it shows calcrete breccia layers Unit 4 includes bro-ken calcrete slabs covering sediments similar to the precedingunit Unit 5 comprises allochthnous cobbles and calcreteslabs in a loess-size mattrix (Walker

et al

1998 1999)Some 8 teeth and bones of

Homo sapiens neanderthalensis

numerous Middle Palaeolithic artifacts and abundant faunalremains have been recovered from units 2ndash5 within an area of25 square metres under methodical excavation since 1990(Walker

et al

in press)The rock-shelter lies at 780 m above sea level in a rocky

escarpment 40 m above the River Quiacutepar at 1

deg

48

prime

10

primeprime

E38

deg

2

prime

5

primeprime

N where the river leaves a gorge (lsquoEstrechorsquo) thatruns from south to north before bending sharply eastwardstoday Although formed by karstic processes in UpperMiocene biocalcarenite the rock-shelter was raised up withrespect to the river by Middle Pleistocene tectonics alongwith the rest of the east flank of the gorge such that it hascome to lie slightly higher than the glacis-terrace B sedimentsof the opposite bank to which the Cueva Negra sedimentaryfill very likely corresponds (Walker

et al

1998 1999) Glacis-terrace B is a formation of calcretes silts loess and gravelswhich occurs widely dating from the early Upper Pleistocenein the Segura and Vinalopoacute drainage basins (Walker ampCuenca 1977) Its upper units seem to be just within the rangeof radiocarbon dating around 40 000

BP

whilst lower onesformed during the period from

c

115 000ndash40 000

BP

(Cuenca

Fig 1 Study sites and location of other pollen sequences from glacialrefugia in south-eastern Spain

Glacial refugia of thermophilous flora in Spain

121

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

et al

1986) This chronostratigraphic context together withthe type of Palaeolithic industry and the abundance of ther-moclastic scree suggest that the study deposit is of last glacialage most likely early to mid Upper Pleistocene (Walker

et al

in press)

Sima de las Palomas del Cabezo Gordo (lsquoDove Hole on BigHillrsquo) is at 0

deg

53

prime

53

primeprime

W 37

deg

47

prime

54

primeprime

N in Torre Pachecotownship (Fig 1) at 80 m above sea level on the S-facing

flank of Cabezo Gordo a steep 310-metre-high hill of Permo-Triassic (Nevado-Filaacutebride) marbled limestone which risesfrom the coastal plain as an isolated block lying E-W ThelsquoSimarsquo is a natural karstic shaft that contains the remains of abreccia fill which was largely removed by iron miners Acolumn of breccia remains that runs 18 m down the shaftfrom the uppermost entrance to the floor of the MainChamber Nowadays the climate here is semiarid with 3000 h

Fig 2 Cueva Negra del Estrecho del Riacuteo Quiacutepar pollen diagram (Murcia Spain)

122

JS Carrioacuten

et al

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

of sunshine annually a winter temperature never below 10

deg

Cand a mean July temperature of 26

deg

CTwo lithostratigraphical units have been recognized in the

uppermost 18 m of the column under excavation since 1994Unit 1 is a yellowish cemented breccia and scree adhering tothe overhang Unit 2 is characterized by horizontal accumula-tion of angular scree in a clayey matrix with fine layers ofburnt soil Both units are rich in bones including remains of

Homo sapiens neanderthalensis

and Mousterian artifactsThorium-Uranium and AMS radiocarbon dating assign thesedeposits to about 60 000ndash40 000 years ago The bottom ofthe column below unit 2 has Thorium-Uranium determina-tions of about 125 000 years (Walker

et al

in press) Thestudy deposit is therefore likely to be synchronous with theCueva Negra section that is early to mid Upper PleistoceneDetails on the chronological context of both sites are pre-sented elsewhere (Walker

et al

in press)

METHODS

Sampling of vertical sections was done in close relationshipto the stratigraphy each recognizable archaeological bed

was sampled generally only one sample per bed that is at3ndash5 cm intervals Control for modern pollen rain was doneby means of two samples each made up of 5 subsamplesfrom the surface of the soil at the respective sites Laboratorytreatment was performed following the conventional HClHF and KOH method and

Lycopodium clavatum

tabletscontaining a known quantity of spores were added to eachsample prepared to enable estimation of pollen concentra-tion Pollen grains were concentrated by heavy-liquidflotation Residues were mounted in silicone oil Pollen iden-tification was performed by comparison with the referencecollection at Murcia University Identification criteria forconflicting taxa are described elsewhere (Carrioacuten 2002a)Total sums of 230ndash490 and 351ndash536 pollen grains andspores were obtained for Cueva Negra and Sima de lasPalomas respectively Pollen diagrams were constructedby using progressively the programs Tilia TiliaGraphTGView and CorelDraw 90 (Figs 2 and 3) Internal separa-tion of pollen zones has not been undertaken for the pollendiagrams because visual inspection reveals only slight varia-tion Nomenclature for plant taxa follows Saacutenchez-Goacutemez

et al

(1998)

Fig 3 Sima de las Palomas del Cabezo Gordo pollen diagram (Murcia Spain)

Glacial refugia of thermophilous flora in Spain

123

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

RESULTS

Cueva Negra pollen records

Pollen spectra are codominated by two well-differentiatedgroups of

Quercus

with percentages of around 15 and35 respectively (Fig 2) One is of deciduous species whichgiven the local limestone substrate was no doubt mainly

Quercus faginea

The other is an evergreen group whichcould indicate the presence of

Quercus ilexrotundifolia

or

Qcoccifera

Quercus

is more abundant than any other tree orshrub species although pine (

Pinus

)

juniper (

Juniperus

)and wild olive (

Olea

) consistently reach levels over 2ndash3Low frequency oscillations throughout the sections areshown for hazel (

Corylus

) birch (

Betula

) cluster pine (

Pinuspinaster

) ash (

Fraxinus

) elm (

Ulmus

) willow (

Salix

) lentisk(

Pistacia

) and

Phillyrea

Sporadically present are field maple(

Acer

) yew (

Taxus

) strawberry tree (

Arbutus

) ivy (

Hedera

)and rock-roses (

Cistus

) Although there is a significantherbaceous component of wormwood (

Artemisia

) Poaceaeand Asteraceae and to a lesser extent ChenopodiaceaeLamiaceae and Cyperaceae the abundance of those taxais nonetheless low when compared to Pleistocene pollenspectra from other Mediterranean parts of Spain (Carrioacuten

et al

2000)Pollen analyses at Cueva Negra show steppe vegetation

coexisting with woodland species Determining the geograph-ical location of plant species through pollen analysis iscomplicated due to methodological pitfalls (Birks 1993) sowe can but speculate about the composition of plant forma-tions at the time although some light may be shed on thismatter by regional topography and the ecology of those taxaidentified Thus it seems likely that upland plateaus wereopen ground where grasses and wormwood scrub (

Artemisia

)predominated sometimes with juniper bushes and occasion-ally pine trees whilst valleys and gorges (like that of theQuiacutepar) gave shelter to most of the woodland flora andmesothermophilous shrubs Thus Cueva Negra pollenfrequencies imply mixed copses of both deciduous andevergreen oaks with cluster pines for the most part alongwith representatives of other deciduous species such ashazel or beech that are no longer found there and only lingeron in sheltered valleys of the high Sierra del Segura 50 kmaway to the NW (Fig 1) Many of these trees probablybehaved as phreatophytes growing on river banks and valleyfloors near water-courses Thermophilous taxa such as

Olea

Pistacia

and

Phillyrea

show striking abundance forthe last glacial-stage mesomediterranean enclave aroundCueva Negra

Sima de las Palomas pollen records

Sima de las Palomas pollen spectra are dominated by twogroups of

Quercus

as at Cueva Negra (Fig 3) The deciduous

palynotype is more variable in shape size and exine orna-mentation perhaps reflecting a mosaic of underlying ecolo-gical factors or several species coexisting in a context wheresoils show greater diversity than at Cueva Negra (in morehumid mountainous regions than Murcia

Q faginea Qcanariensis

and

Q pyrenaica

occur today in southern regionsof the Iberian Peninsula) Pine is more prominent than atCueva Negra but which species cannot yet be determinedLikely candidates are black pine (

P nigra) Aleppo pine (Phalepensis) and umbrella pine (P pinea) anthracology hasshown P nigra occurred from c 25 000ndash13 000 year BP inPalaeolithic settlements in Alicante and Valencia (Badal ampCarrioacuten 2001)

Given that today large parts of coastal Murcia on averageget less than 200 mm of rain a year with very high levels ofevapo-transpiration (Saacutenchez-Goacutemez et al 1998) it is strik-ing that Pleistocene pollen spectra show abundant pollen ofdeciduous oaks at Sima de las Palomas alongside trees re-quiring damp-temperate conditions such as Corylus avellanaFraxinus Arbutus unedo Buxus and Betula The oaks mostlikely grew nearby because their pollen frequencies of 15ndash20 exceed those involving long-distance pollen transportto south-eastern Spanish caves (Navarro et al 2001) Waslocal climate damper than now in the upper Pleistocene Oris a combination of Holocene climate changes and anthro-pogenic intervention to blame for the recent decline in decid-uous trees These are not exclusive possibilities but the latteris supported (Carrioacuten et al in press) by a Holocene sequencefrom Gaacutedor (Almeriacutea) by archaeological findings and byhistorical evidence (Garciacutea Latorre amp Garciacutea Latorre 1996Gimeacutenez 2000)

A far broader mosaic of plant communities may beenvisaged in the local Pleistocene landscape than existstoday It would have contained both pine woods andmixed evergreen and deciduous Quercus woodland aswell as deciduous trees in shady zones gorges and besidewater-courses all contributing to a rich undergrowth ofMediterranean species with heliophilous formations onthinner soils The latter would include Periploca angustifoliaOsyris quadripartita Asphodelus Labiates Composites Cis-taceas Thymelaea hirsuta Calicotome intermedia and otherGenistas Last but not least there would have been marshlandsand coastal saltpans characterized by ChenopodiaceaeLycium and Whitania frutescens The widespread presenceof thermophytes including species such as Periploca angus-tifolia and Maytenus europaeus which readily succumb tofrost indicate that local climate can scarcely have been colderthan today Moreover those taxa along with WithaniaPistacia Phillyrea Calicotome and Osyris are all clearlyunder-represented in both external and internal pollen rain(Carrioacuten 2002a) hence their abundance in the neighbour-hood was undountedly greater than pollen spectra imply atfirst glance

124 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

DISCUSSION

Palaeoecological value of the cave pollen spectra studied

Doubts have been raised about the palaeoecological valueof palynological information obtained from archaeologicalstudies from caves and rock-shelters (eg Turner amp Hannon1988) but this issue has been discussed elsewhere (Davis1990 Carrioacuten amp Scott 1999 Carrioacuten et al 1999a) Fourpalaeopalynological aspects need stressing with regard to ourregion First it lacks lake swamp or peat deposits hencevegetational reconstruction must resort to either marinesequences (Targarona 1997) or deposits in caves or rock-shelters Secondly there are no reasons a priori to dismissresults obtained from considerations of pollen-rain repres-entativeness as regards the external vegetation At least thisis what can be concluded experimentally after analysingsurface sediments at caves of different shapes and sizesboth within our region (Navarro et al 2000 2001) andoutside it (Burney amp Burney 1993) Thirdly with regard toconventional sediments those from the very same caves havean advantage in that they are able to collect those ento-mophilous species that make up most of the local vegetationbut are conspicuous by their absence from lake-bed deposits(Carrioacuten 2002a) here indeed any chance at all of biotictransport to a site is of far more help than hindrance topalaeopalynologists

Lastly it is essential that there are palynological indicatorsboth for analytical quality and palaeoecological coherence soas to be able to detect possible skewing of pollen spectra dueto such postdepositional processes as destruction of pollenpercolation or reworking (Saacutenchez-Gontildei 1994) In thisregard it is noteworthy that in both series analysed (Figs 2and 3) have been identified very many types including somenot always readily preserved (eg Taxus Genisteae BuxusCalicotome Periploca) or whose identification demands pre-cise exine characteristics available only in nonoxidized paly-nomorphs (eg Coris Smilax Maytenus Withania) Totalconcentrations of pollen are not especially high being greaterat Sima de las Palomas (between c 9487 and 23 137 grains gminus1)than at Cueva Negra (between c 2342 and 5150 grains gminus1)but they are nevertheless comparable to those often reportedfrom surface sediments in caves (Davis 1990 Navarro et al2001) so there is no need to interpret them as being due toloss caused by destructive processes

At both sites surface pollen spectra reflect wholly modernbut never Pleistocene vegetation Cueva Negra surface samplespredominantly show Pinus (including considerable amount ofP pinaster type) Helianthemum Genisteae and Plantagowhilst Pleistocene ones show clear-cut predominance ofQuercus Poaceae Artemisia and Asteraceae (Fig 2) Simade las Palomas surface samples show greater percentage sim-ilarity but while its surface sediment lacks Quercus (absenttoday in this coastal reagion) Quercus pollen nevertheless is

very abundant in Pleistocene samples from the site Likewiseutterly absent from surface samples are other minority pollengrains identified nonetheless in the Pleistocene samples suchas Corylus Betula Fraxinus Arbutus Ulmus Salix Erica andEphedra distachya-nebrodensis (Fig 3) Last but not leastour failure to detect either bed-rock Miocene spores at CuevaNegra or Permo-Triassic ones at Sima de las Palomas is worthmentioning nor were there differences in staining or preser-vation in pollen spectra from either site such as are commonwhen reworking or sediment mixing have taken place in a cave(Carrioacuten et al 1995)

The new findings in the context of previous palynological studies

Cueva Negra pollen results are comparable with thoseslightly further west from Siles at 1320 m above sea level in anintermontane valley of the Sierra de Segura in Jaeacuten (Carrioacuten2002b) (Fig 4) In deposits dating from upper pleniglacialtimes (c 20 000ndash17 000 cal year BP) were found Pinuspinaster deciduous Quercus evergreen Quercus EricaceaeCorylus Betula and Fraxinus in pollen percentages alwaysabove 2 and frequently also Acer Taxus Arbutus BuxusSalix Ulmus Phillyrea Pistacia and Olea All of these con-tingents were better represented at Siles in lateglacial times(c 17 000ndash11 900 cal year BP) and especially in mid-Holocenetimes (c 7400ndash5300 cal year BP) although pine woodsappear to have predominated during the early Holocene(c 11 900ndash7400 cal year BP)

Although not being synchronous the similarity betweenthe Cueva Negra and Siles spectra is important because bothsites lie in closely related biogeographical areas betweenwhich migration is eminently feasible (Fig 1) not for nothinghave mountains near Cueva Negra been regarded by someplant biogeographers as an eastern extension of the Sierrade Segura (Saacutenchez-Goacutemez amp Alcaraz 1993) The Siles andCueva Negra pollen records show that tree species survivedat quite high places in southern European mountains duringthe last glacial stage This agrees with a hypothesis put for-ward by Bennett et al (1991) that tree survival would havebeen especially important in mountain ranges such as thoseof the Balkans allowing rapid altitudinal displacements of treepopulations in response to climatic pulses (Willis 1994)Owing to their roughly N-S orientation the Segura moun-tains like the Balkans would have readily allowed altitudinalmovements of tree populations to take place Moreover otherindications that mountains in southern Spain containedsignificant tree reservoirs during the last ice age come fromconsideration of the genetic structure of European treepopulations today (Herraacuten et al 1999 Jimeacutenez 2000Salvador et al 2000)

Other pollen sequences in the Iberian Peninsula with com-parable incidences of mesothermophilous flora to Siles or

Glacial refugia of thermophilous flora in Spain 125

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

Cueva Negra only occur at lower altitudes as is the normelsewhere in southern Europe (Willis 1994 Leroy et al1996 Van Andel amp Tzedakis 1996 Magri amp Parra 1997)The Murcian coast offers another important pollen recordfrom the Middle-Upper Palaeolithic transition at CuevaPerneras (Carrioacuten et al 1995) Although tree pollen was lessimportant than at Sima de las Palomas Cueva Pernerasdeposits contained abundant pollen of Pinus Quercus ilex-coccifera and Oleaceae and continuous or frequent presenceof broad-leaved trees (Fraxinus Alnus Corylus JuglansUlmus Salix) and thermophytes (Myrtus Erica arborea

Pistacia Buxus Periploca Withania Lycium Ephedra fragilisCosentinia vellea Selaginella denticulata Ruta)

To the south of Murcia at San Rafael on the Almeriancoast a pollen sequence shows continuous curves for ever-green and deciduous Quercus and Olea during the last glacialmaximum and tardiglacial (Pantaleoacuten-Cano et al in press)Another pollen sequence showing lateglacial tree presencecomes from coprolites of spotted hyaena (Crocuta crocuta)at Cueva de las Ventanas in Granada where about 12 780cal year BP there were pine woods wormwood steppewith juniper grassland and mixed woodland of Quercus

Fig 4 Reference pollen diagram from the lake at Siles (Jaeacuten Spain) which includes mesothermophilous trees and shrubs (shaded zonescorrespond to the last glacial stage black dots refer to percentages below 2)

126 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

with Betula Abies Corylus Alnus Acer Taxus MyrtusBuxus Sorbus Olea Erica arborea Pistacia Ephedra fragilisViburnum Sambucus Cistus and Rhamnus

To the north of Murcia relevant pollen findings come fromsites in inland Mesomediterranean environments namely thedated Middle-Upper Palaeolithic transition at Cova Beneitoin Alicante (Carrioacuten amp Munuera 1997) and the Navarreacutespeat bog in Valencia (Carrioacuten amp van Geel 1999) Howevertheir thermophilous components show oscillations ratherthan uninterrupted presence indeed most of their deciduoustrees and Mediterranean shrubs fell away markedly duringthe upper pleniglacial stage after having advanced togetherduring oxygen-isotope stage 3 In the north-eastern Medi-terranean part of the Iberian Peninsula Abric Romaniacute nearBarcelona shows tree pollen percentages of 40ndash60 betweenabout 70 000 and 40 000 year BP with continuous presenceof Juniperus Rhamnus Quercus Olea-Phillyrea SyringaAlnus Salix Juglans Betula Fagus Betula Coriaria Pistaciaand Vitis (Burjachs amp Juliagrave 1994)

Further afield even the Cantabrian coast and adjacentmountain ranges seem to have offered refuges for trees duringthe last ice age according to pollen analyses (Dupreacute 1988Ramil-Rego et al 1998ab) and macroscopical charcoal(Uzquiano 1992) Pleniglacial pollen samples include lowfrequencies of Pinus Betula Juniperus Corylus QuercusFraxinus Alnus Ulmus Tilia Juglans Fagus and CastaneaCharcoal samples contain Pinus sylvestris P uncinataJuniperus Betula alba B pendula Corylus avellana Quercusrobur Q petraea Tilia platyphyllos T cordata Fraxinusexcelsior Sambucus nigra Viburnum tinus Cornus sanguineaQuercus ilex Fagus sylvatica Sorbus aria S aucupariaS torminalis S domestica Castanea sativa Quercus suberArbutus unedo Erica arborea Crataegus monogyna andseveral species of Prunus and Rhamnus Hunter-gatherersmay have gathered hazelnuts acorns and wild fruit (egmazzard Prunus avium) (Uzquiano 1992) The palaeobotan-ical findings concur with the genetical structure of popula-tions of Iberian brown oak (Olalde et al 2002) and somepalaeoecological inferences drawn from the abundant mega-fauna of the Biscay coast (Altuna 1972)

For continental parts of the Iberian Peniacutensula there existsboth pollen and palaeobotanical evidence of mesophiloustaxa in glacial and late glacial contexts (Dupreacute 1988 Ponsamp Reille 1988 Garciacutea-Antoacuten et al 1990 Garciacutea-Antoacuten ampSainz-Ollero 1991 Carrioacuten amp Saacutenchez-Goacutemez 1992 Peacuterez-Obiol amp Juliagrave 1994 Blanco et al 1997) When cave pollenis considered along with reference pollen sequences andcharcoal findings everything seems to point less to refugiarestricted to the far south (Brewer et al 2002) than tosurvival of stationary tree populations in many parts of thePeninsula particularly in intramontane valleys in the Baeticpre-Baetic Iberian and other coastal ranges with expansionand contraction in the central uplands

To conclude palynology at archaeological cave sites oftenraises doubts over contemporaneity between pollen and thelayer containing it and anthracology often seems closer topalaeobotanical orthodoxy than to palaeoecology Neverthe-less both approaches together can build up a solid weightof evidence Furthermore some reference pollen sequenceshave undeniable counterparts in some cave pollen sequences(Carrioacuten 2002b) (Fig 4) Our models about glacial refugiarequire far more information several sites need to be restudiedtheir chronologies need refinement and the present estimatesof continental palaeotemperatures during the upper Pleistoceneshould be viewed with caution Moreover faunal remainsespecially birds and small mammals must be taken far moreinto consideration whenever revision of glacial refugia oftemperate trees is undertaken concerning palaeoclimaticinferences

Relationships with palaeontological findings and human remains

Palynological findings at Cueva Negra del Estrecho del RiacuteoQuiacutepar and Sima de las Palomas del Cabezo Gordo sit easilywith the faunal evidence At Cueva Negra avian palaeonto-logist Anne Easthamrsquos findings (Walker et al 1998 in press)point to five different environmental biotopes coexisting nearthis upland site which today is in an open arid landscapecrossed by a small stream undeservedly called the lsquoRiverrsquoQuiacutepar These are (i) wetlands with a depth of lake-waternecessary for ruddy shelducks mallards wigeons teals gad-walls red-crested pochards common pochards ferruginousducks wild geese little stints and sandpipers (ii) riverine anddamp valley floors where soft sediments offered cover suit-able for the bee-eaters and sand martins at Cueva Negra (iii)Quercus woodland suitable for autumnal acorn-eaters suchas the caversquos Pleistocene jays and woodpigeons where itsowls nightjars woodpeckers woodlarks and several speciesof thrushes finches must have found their prey (iv) steppe andopen country preferred by larks partridges plovers choughseagles buzzards kestrels and falcons whose bones are allwell represented and (v) the craggy mountainsides andcliffs around the cave itself offering roosting places for theomnipresent choughs rock doves crag martins swallowsswifts and rock thrushes Reptiles are especially well repres-ented by tortoise remains Numerous skeletal parts of smalland large mammal species excavated support such a varietyof biotopes around the rock-shelter carnivores such ashyaena brown bear wolf a small feline omnivores such asmacaque and wild boar herbivores such as steppe rhinoceroselephantids giant deer red deer a smaller cervid aurochsbison horse and wild goat Bats hedgehogs and shrewswere also found

At Sima de las Palomas the correlation for the early tomid-upper Pleistocene between fauna and ancient habitats is

Glacial refugia of thermophilous flora in Spain 127

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

less clear-cut Cementation of excavated breccia makes forslow progress in palaeontological research Fewer bird specieshave been identified than at Cueva Negra (13 species asagainst 66) (Walker et al in press) There are of coursecraggy mountainsides around the cave Below the site thecoastal plain surrounding Cabezo Gordo nowadays dry andopen must have contained mixed woodland and grasslandwith gallery woodland and swamps beside erstwhile streamsfeeding nearby wetlands and saltmarshes behind coastal sanddunes (where the Mar Menor coastal lagoon is today) eventhough marine regressions must have repeatedly drainedthese for many millennia each time Mammals include carni-vores such as panther lynx spotted hyaena brown bearfox and badger and herbivores such as hippopotamuselephantids wild horses and asses aurochs red deer wildgoats and abundant lagomorphs Tortoise is again commonin the deposits

The two Upper Pleistocene sites considered here from apalaeoecological standpoint of flora and fauna corroboratea proposal that long-term human presence may occur prefer-entially where several different biotopes coincide

ACKNOWLEDGMENTS

Michegravele Dupreacute assisted with laboratory processing KeithBennett and an anonymous referee provided useful sugges-tions on an earlier draft Research funding was provided bythe Spanish Ministerio de Educacioacuten y Ciencia and theMurcian regional authorityrsquos Fundacioacuten Seneca (projectsBOS2000-0149 PI-1700739FS01 PB92-0971 PB98-0405)

REFERENCES

Altuna J (1972) Fauna de mamiacuteferos de los yacimientos prehistoacutericosde Guipuacutezcoa Munibe 24 1ndash464

Arroyo J (1997) Plant diversity in the region of the Strait of Gibraltara multilevel approach Lagascalia 19 393ndash404

Badal E amp Carrioacuten Y (2001) Del glaciar al interglaciar Los paisajesvegetales a partir de los restos carbonizados hallados en las cuevasde Alicante De Neandertales a Cromantildeones El inicio del pob-lamiento humano en las tierras valencianas (ed by V Villaverde)pp 21ndash40 Universidad de Valencia Valencia

Bennett KD Tzedakis PC amp Willis KJ (1991) Quaternaryrefugia of north European trees Journal of Biogeography 18 103ndash115

Birks HJB (1993) Quaternary palaeoecology and vegetationscience-current contributions and possible future developmentsReview of Palaeobotany and Palynology 79 153ndash177

Blanco E Casado MA Costa-Tenorio M Escribano R Garciacutea-Antoacuten M Geacutenova M Goacutemez A Goacutemez F Moreno JCMorla C Regato P amp Sainz H (1997) Los bosques ibeacutericosUna interpretacioacuten geobotaacutenica Planeta Barcelona

Brewer S Cheddadi R de Beaulieu JL Reille M amp Data Con-tributors (2002) The spread of deciduous Quercus throughout

Europe since the last glacial period Forest Ecology and Management156 27ndash48

Burjachs F amp Juliagrave R (1994) Abrupt climatic changes during thelast glaciation based on pollen analysis of the Abric RomaniCatalonia Spain Quaternary Research 42 308ndash315

Burney DA amp Burney LP (1993) Modern pollen deposition in cavesites experimental results from New York State New Phytolology124 523ndash535

Carrioacuten JS (2002a) A taphonomic study of modern pollenassemblages from dung and surface sediments in arid environ-ments of Spain Review of Palaeobotany and Palynology 120217ndash232

Carrioacuten JS (2002b) Patterns and processes of Late Quaternaryenvironmental change in a montane region of southwestern EuropeQuaternary Science Reviews 21 2047ndash2066

Carrioacuten JS Dupreacute M Fumanal MP amp Montes R (1995) Apalaeoenvironmental study in semi-arid southeastern Spain thepalynological and sedimentological sequence at Perneras Cave(Lorca Murcia) Journal of Archaeological Science 22 355ndash367

Carrioacuten JS amp Munuera M (1997) Upper Pleistocene palaeoenvi-ronmental change in eastern Spain new pollen analytical datafrom Cova Beneito (Alicante) Palaeogeography Palaeoclimatologyand Palaeoecology 128 287ndash299

Carrioacuten JS Munuera M Navarro C Burjachs F Dupreacute M ampWalker MJ (1999a) The palaeoecological potential of pollenrecords in caves the case of Mediterranean Spain QuaternaryScience Reviews 18 1061ndash1073

Carrioacuten JS Munuera M Navarro C amp Saacuteez F (2000) Paleo-climas e historia de la vegetacioacuten cuaternaria en Espantildea a traveacutes delanaacutelisis poliacutenico Viejas falacias y nuevos paradigmas Complutum11 1ndash28

Carrioacuten JS amp Saacutenchez-Goacutemez P (1992) Palynological data in sup-port of the survival of walnut (Juglans regia L) in the westernMediterranean area during last glacial times Journal of Biogeography19 623ndash630

Carrioacuten JS amp Scott L (1999) The challenge of pollen analysis inpalaeoenvironmental studies of hominid beds the record fromSterkfontein caves Journal of Human Evolution 36 401ndash408

Carrioacuten JS amp van Geel B (1999) Fine-resolution Upper Weichselianand Holocene palynological record from Navarreacutes (ValenciaSpain) and a discussion about factors of Mediterranean forestsuccession Review of Palaeobotany and Palynology 106 209ndash236

Comes HP amp Kadereit JW (1998) The effect of Quaternary cli-matic changes on plant distribution and evolution Trends in PlantScience 3 432ndash438

Cuenca A Pomery PJ amp Walker MJ (1986) Chronologicalaspects of the Middle Pleistocene in the coastal belt of southeasternSpain Quaternary climate in Western Mediterranean (ed byF Loacutepez-Vera) pp 353ndash363 Universidad Autoacutenoma Madrid

Davis OK (1990) Caves as sources of biotic remains in arid westernNorth America Palaeogeography Palaeoclimatology and Palae-oecology 76 331ndash348

Dupreacute M (1988) Palinologiacutea y paleoambiente Nuevos datosespantildeoles Referencias Valencia Diputacioacuten de Valencia Serviciode Investigacioacuten Prehistoacuterica Serie de Trabajos Varios no 84

Garciacutea Latorre J amp Garciacutea Latorre J (1996) Los bosques ignoradosde Almeriacutea Una interpretacioacuten histoacuterica y ecoloacutegica Historia y

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

Glacial refugia of thermophilous flora in Spain

121

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

et al

1986) This chronostratigraphic context together withthe type of Palaeolithic industry and the abundance of ther-moclastic scree suggest that the study deposit is of last glacialage most likely early to mid Upper Pleistocene (Walker

et al

in press)

Sima de las Palomas del Cabezo Gordo (lsquoDove Hole on BigHillrsquo) is at 0

deg

53

prime

53

primeprime

W 37

deg

47

prime

54

primeprime

N in Torre Pachecotownship (Fig 1) at 80 m above sea level on the S-facing

flank of Cabezo Gordo a steep 310-metre-high hill of Permo-Triassic (Nevado-Filaacutebride) marbled limestone which risesfrom the coastal plain as an isolated block lying E-W ThelsquoSimarsquo is a natural karstic shaft that contains the remains of abreccia fill which was largely removed by iron miners Acolumn of breccia remains that runs 18 m down the shaftfrom the uppermost entrance to the floor of the MainChamber Nowadays the climate here is semiarid with 3000 h

Fig 2 Cueva Negra del Estrecho del Riacuteo Quiacutepar pollen diagram (Murcia Spain)

122

JS Carrioacuten

et al

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

of sunshine annually a winter temperature never below 10

deg

Cand a mean July temperature of 26

deg

CTwo lithostratigraphical units have been recognized in the

uppermost 18 m of the column under excavation since 1994Unit 1 is a yellowish cemented breccia and scree adhering tothe overhang Unit 2 is characterized by horizontal accumula-tion of angular scree in a clayey matrix with fine layers ofburnt soil Both units are rich in bones including remains of

Homo sapiens neanderthalensis

and Mousterian artifactsThorium-Uranium and AMS radiocarbon dating assign thesedeposits to about 60 000ndash40 000 years ago The bottom ofthe column below unit 2 has Thorium-Uranium determina-tions of about 125 000 years (Walker

et al

in press) Thestudy deposit is therefore likely to be synchronous with theCueva Negra section that is early to mid Upper PleistoceneDetails on the chronological context of both sites are pre-sented elsewhere (Walker

et al

in press)

METHODS

Sampling of vertical sections was done in close relationshipto the stratigraphy each recognizable archaeological bed

was sampled generally only one sample per bed that is at3ndash5 cm intervals Control for modern pollen rain was doneby means of two samples each made up of 5 subsamplesfrom the surface of the soil at the respective sites Laboratorytreatment was performed following the conventional HClHF and KOH method and

Lycopodium clavatum

tabletscontaining a known quantity of spores were added to eachsample prepared to enable estimation of pollen concentra-tion Pollen grains were concentrated by heavy-liquidflotation Residues were mounted in silicone oil Pollen iden-tification was performed by comparison with the referencecollection at Murcia University Identification criteria forconflicting taxa are described elsewhere (Carrioacuten 2002a)Total sums of 230ndash490 and 351ndash536 pollen grains andspores were obtained for Cueva Negra and Sima de lasPalomas respectively Pollen diagrams were constructedby using progressively the programs Tilia TiliaGraphTGView and CorelDraw 90 (Figs 2 and 3) Internal separa-tion of pollen zones has not been undertaken for the pollendiagrams because visual inspection reveals only slight varia-tion Nomenclature for plant taxa follows Saacutenchez-Goacutemez

et al

(1998)

Fig 3 Sima de las Palomas del Cabezo Gordo pollen diagram (Murcia Spain)

Glacial refugia of thermophilous flora in Spain

123

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

RESULTS

Cueva Negra pollen records

Pollen spectra are codominated by two well-differentiatedgroups of

Quercus

with percentages of around 15 and35 respectively (Fig 2) One is of deciduous species whichgiven the local limestone substrate was no doubt mainly

Quercus faginea

The other is an evergreen group whichcould indicate the presence of

Quercus ilexrotundifolia

or

Qcoccifera

Quercus

is more abundant than any other tree orshrub species although pine (

Pinus

)

juniper (

Juniperus

)and wild olive (

Olea

) consistently reach levels over 2ndash3Low frequency oscillations throughout the sections areshown for hazel (

Corylus

) birch (

Betula

) cluster pine (

Pinuspinaster

) ash (

Fraxinus

) elm (

Ulmus

) willow (

Salix

) lentisk(

Pistacia

) and

Phillyrea

Sporadically present are field maple(

Acer

) yew (

Taxus

) strawberry tree (

Arbutus

) ivy (

Hedera

)and rock-roses (

Cistus

) Although there is a significantherbaceous component of wormwood (

Artemisia

) Poaceaeand Asteraceae and to a lesser extent ChenopodiaceaeLamiaceae and Cyperaceae the abundance of those taxais nonetheless low when compared to Pleistocene pollenspectra from other Mediterranean parts of Spain (Carrioacuten

et al

2000)Pollen analyses at Cueva Negra show steppe vegetation

coexisting with woodland species Determining the geograph-ical location of plant species through pollen analysis iscomplicated due to methodological pitfalls (Birks 1993) sowe can but speculate about the composition of plant forma-tions at the time although some light may be shed on thismatter by regional topography and the ecology of those taxaidentified Thus it seems likely that upland plateaus wereopen ground where grasses and wormwood scrub (

Artemisia

)predominated sometimes with juniper bushes and occasion-ally pine trees whilst valleys and gorges (like that of theQuiacutepar) gave shelter to most of the woodland flora andmesothermophilous shrubs Thus Cueva Negra pollenfrequencies imply mixed copses of both deciduous andevergreen oaks with cluster pines for the most part alongwith representatives of other deciduous species such ashazel or beech that are no longer found there and only lingeron in sheltered valleys of the high Sierra del Segura 50 kmaway to the NW (Fig 1) Many of these trees probablybehaved as phreatophytes growing on river banks and valleyfloors near water-courses Thermophilous taxa such as

Olea

Pistacia

and

Phillyrea

show striking abundance forthe last glacial-stage mesomediterranean enclave aroundCueva Negra

Sima de las Palomas pollen records

Sima de las Palomas pollen spectra are dominated by twogroups of

Quercus

as at Cueva Negra (Fig 3) The deciduous

palynotype is more variable in shape size and exine orna-mentation perhaps reflecting a mosaic of underlying ecolo-gical factors or several species coexisting in a context wheresoils show greater diversity than at Cueva Negra (in morehumid mountainous regions than Murcia

Q faginea Qcanariensis

and

Q pyrenaica

occur today in southern regionsof the Iberian Peninsula) Pine is more prominent than atCueva Negra but which species cannot yet be determinedLikely candidates are black pine (

P nigra) Aleppo pine (Phalepensis) and umbrella pine (P pinea) anthracology hasshown P nigra occurred from c 25 000ndash13 000 year BP inPalaeolithic settlements in Alicante and Valencia (Badal ampCarrioacuten 2001)

Given that today large parts of coastal Murcia on averageget less than 200 mm of rain a year with very high levels ofevapo-transpiration (Saacutenchez-Goacutemez et al 1998) it is strik-ing that Pleistocene pollen spectra show abundant pollen ofdeciduous oaks at Sima de las Palomas alongside trees re-quiring damp-temperate conditions such as Corylus avellanaFraxinus Arbutus unedo Buxus and Betula The oaks mostlikely grew nearby because their pollen frequencies of 15ndash20 exceed those involving long-distance pollen transportto south-eastern Spanish caves (Navarro et al 2001) Waslocal climate damper than now in the upper Pleistocene Oris a combination of Holocene climate changes and anthro-pogenic intervention to blame for the recent decline in decid-uous trees These are not exclusive possibilities but the latteris supported (Carrioacuten et al in press) by a Holocene sequencefrom Gaacutedor (Almeriacutea) by archaeological findings and byhistorical evidence (Garciacutea Latorre amp Garciacutea Latorre 1996Gimeacutenez 2000)

A far broader mosaic of plant communities may beenvisaged in the local Pleistocene landscape than existstoday It would have contained both pine woods andmixed evergreen and deciduous Quercus woodland aswell as deciduous trees in shady zones gorges and besidewater-courses all contributing to a rich undergrowth ofMediterranean species with heliophilous formations onthinner soils The latter would include Periploca angustifoliaOsyris quadripartita Asphodelus Labiates Composites Cis-taceas Thymelaea hirsuta Calicotome intermedia and otherGenistas Last but not least there would have been marshlandsand coastal saltpans characterized by ChenopodiaceaeLycium and Whitania frutescens The widespread presenceof thermophytes including species such as Periploca angus-tifolia and Maytenus europaeus which readily succumb tofrost indicate that local climate can scarcely have been colderthan today Moreover those taxa along with WithaniaPistacia Phillyrea Calicotome and Osyris are all clearlyunder-represented in both external and internal pollen rain(Carrioacuten 2002a) hence their abundance in the neighbour-hood was undountedly greater than pollen spectra imply atfirst glance

124 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

DISCUSSION

Palaeoecological value of the cave pollen spectra studied

Doubts have been raised about the palaeoecological valueof palynological information obtained from archaeologicalstudies from caves and rock-shelters (eg Turner amp Hannon1988) but this issue has been discussed elsewhere (Davis1990 Carrioacuten amp Scott 1999 Carrioacuten et al 1999a) Fourpalaeopalynological aspects need stressing with regard to ourregion First it lacks lake swamp or peat deposits hencevegetational reconstruction must resort to either marinesequences (Targarona 1997) or deposits in caves or rock-shelters Secondly there are no reasons a priori to dismissresults obtained from considerations of pollen-rain repres-entativeness as regards the external vegetation At least thisis what can be concluded experimentally after analysingsurface sediments at caves of different shapes and sizesboth within our region (Navarro et al 2000 2001) andoutside it (Burney amp Burney 1993) Thirdly with regard toconventional sediments those from the very same caves havean advantage in that they are able to collect those ento-mophilous species that make up most of the local vegetationbut are conspicuous by their absence from lake-bed deposits(Carrioacuten 2002a) here indeed any chance at all of biotictransport to a site is of far more help than hindrance topalaeopalynologists

Lastly it is essential that there are palynological indicatorsboth for analytical quality and palaeoecological coherence soas to be able to detect possible skewing of pollen spectra dueto such postdepositional processes as destruction of pollenpercolation or reworking (Saacutenchez-Gontildei 1994) In thisregard it is noteworthy that in both series analysed (Figs 2and 3) have been identified very many types including somenot always readily preserved (eg Taxus Genisteae BuxusCalicotome Periploca) or whose identification demands pre-cise exine characteristics available only in nonoxidized paly-nomorphs (eg Coris Smilax Maytenus Withania) Totalconcentrations of pollen are not especially high being greaterat Sima de las Palomas (between c 9487 and 23 137 grains gminus1)than at Cueva Negra (between c 2342 and 5150 grains gminus1)but they are nevertheless comparable to those often reportedfrom surface sediments in caves (Davis 1990 Navarro et al2001) so there is no need to interpret them as being due toloss caused by destructive processes

At both sites surface pollen spectra reflect wholly modernbut never Pleistocene vegetation Cueva Negra surface samplespredominantly show Pinus (including considerable amount ofP pinaster type) Helianthemum Genisteae and Plantagowhilst Pleistocene ones show clear-cut predominance ofQuercus Poaceae Artemisia and Asteraceae (Fig 2) Simade las Palomas surface samples show greater percentage sim-ilarity but while its surface sediment lacks Quercus (absenttoday in this coastal reagion) Quercus pollen nevertheless is

very abundant in Pleistocene samples from the site Likewiseutterly absent from surface samples are other minority pollengrains identified nonetheless in the Pleistocene samples suchas Corylus Betula Fraxinus Arbutus Ulmus Salix Erica andEphedra distachya-nebrodensis (Fig 3) Last but not leastour failure to detect either bed-rock Miocene spores at CuevaNegra or Permo-Triassic ones at Sima de las Palomas is worthmentioning nor were there differences in staining or preser-vation in pollen spectra from either site such as are commonwhen reworking or sediment mixing have taken place in a cave(Carrioacuten et al 1995)

The new findings in the context of previous palynological studies

Cueva Negra pollen results are comparable with thoseslightly further west from Siles at 1320 m above sea level in anintermontane valley of the Sierra de Segura in Jaeacuten (Carrioacuten2002b) (Fig 4) In deposits dating from upper pleniglacialtimes (c 20 000ndash17 000 cal year BP) were found Pinuspinaster deciduous Quercus evergreen Quercus EricaceaeCorylus Betula and Fraxinus in pollen percentages alwaysabove 2 and frequently also Acer Taxus Arbutus BuxusSalix Ulmus Phillyrea Pistacia and Olea All of these con-tingents were better represented at Siles in lateglacial times(c 17 000ndash11 900 cal year BP) and especially in mid-Holocenetimes (c 7400ndash5300 cal year BP) although pine woodsappear to have predominated during the early Holocene(c 11 900ndash7400 cal year BP)

Although not being synchronous the similarity betweenthe Cueva Negra and Siles spectra is important because bothsites lie in closely related biogeographical areas betweenwhich migration is eminently feasible (Fig 1) not for nothinghave mountains near Cueva Negra been regarded by someplant biogeographers as an eastern extension of the Sierrade Segura (Saacutenchez-Goacutemez amp Alcaraz 1993) The Siles andCueva Negra pollen records show that tree species survivedat quite high places in southern European mountains duringthe last glacial stage This agrees with a hypothesis put for-ward by Bennett et al (1991) that tree survival would havebeen especially important in mountain ranges such as thoseof the Balkans allowing rapid altitudinal displacements of treepopulations in response to climatic pulses (Willis 1994)Owing to their roughly N-S orientation the Segura moun-tains like the Balkans would have readily allowed altitudinalmovements of tree populations to take place Moreover otherindications that mountains in southern Spain containedsignificant tree reservoirs during the last ice age come fromconsideration of the genetic structure of European treepopulations today (Herraacuten et al 1999 Jimeacutenez 2000Salvador et al 2000)

Other pollen sequences in the Iberian Peninsula with com-parable incidences of mesothermophilous flora to Siles or

Glacial refugia of thermophilous flora in Spain 125

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

Cueva Negra only occur at lower altitudes as is the normelsewhere in southern Europe (Willis 1994 Leroy et al1996 Van Andel amp Tzedakis 1996 Magri amp Parra 1997)The Murcian coast offers another important pollen recordfrom the Middle-Upper Palaeolithic transition at CuevaPerneras (Carrioacuten et al 1995) Although tree pollen was lessimportant than at Sima de las Palomas Cueva Pernerasdeposits contained abundant pollen of Pinus Quercus ilex-coccifera and Oleaceae and continuous or frequent presenceof broad-leaved trees (Fraxinus Alnus Corylus JuglansUlmus Salix) and thermophytes (Myrtus Erica arborea

Pistacia Buxus Periploca Withania Lycium Ephedra fragilisCosentinia vellea Selaginella denticulata Ruta)

To the south of Murcia at San Rafael on the Almeriancoast a pollen sequence shows continuous curves for ever-green and deciduous Quercus and Olea during the last glacialmaximum and tardiglacial (Pantaleoacuten-Cano et al in press)Another pollen sequence showing lateglacial tree presencecomes from coprolites of spotted hyaena (Crocuta crocuta)at Cueva de las Ventanas in Granada where about 12 780cal year BP there were pine woods wormwood steppewith juniper grassland and mixed woodland of Quercus

Fig 4 Reference pollen diagram from the lake at Siles (Jaeacuten Spain) which includes mesothermophilous trees and shrubs (shaded zonescorrespond to the last glacial stage black dots refer to percentages below 2)

126 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

with Betula Abies Corylus Alnus Acer Taxus MyrtusBuxus Sorbus Olea Erica arborea Pistacia Ephedra fragilisViburnum Sambucus Cistus and Rhamnus

To the north of Murcia relevant pollen findings come fromsites in inland Mesomediterranean environments namely thedated Middle-Upper Palaeolithic transition at Cova Beneitoin Alicante (Carrioacuten amp Munuera 1997) and the Navarreacutespeat bog in Valencia (Carrioacuten amp van Geel 1999) Howevertheir thermophilous components show oscillations ratherthan uninterrupted presence indeed most of their deciduoustrees and Mediterranean shrubs fell away markedly duringthe upper pleniglacial stage after having advanced togetherduring oxygen-isotope stage 3 In the north-eastern Medi-terranean part of the Iberian Peninsula Abric Romaniacute nearBarcelona shows tree pollen percentages of 40ndash60 betweenabout 70 000 and 40 000 year BP with continuous presenceof Juniperus Rhamnus Quercus Olea-Phillyrea SyringaAlnus Salix Juglans Betula Fagus Betula Coriaria Pistaciaand Vitis (Burjachs amp Juliagrave 1994)

Further afield even the Cantabrian coast and adjacentmountain ranges seem to have offered refuges for trees duringthe last ice age according to pollen analyses (Dupreacute 1988Ramil-Rego et al 1998ab) and macroscopical charcoal(Uzquiano 1992) Pleniglacial pollen samples include lowfrequencies of Pinus Betula Juniperus Corylus QuercusFraxinus Alnus Ulmus Tilia Juglans Fagus and CastaneaCharcoal samples contain Pinus sylvestris P uncinataJuniperus Betula alba B pendula Corylus avellana Quercusrobur Q petraea Tilia platyphyllos T cordata Fraxinusexcelsior Sambucus nigra Viburnum tinus Cornus sanguineaQuercus ilex Fagus sylvatica Sorbus aria S aucupariaS torminalis S domestica Castanea sativa Quercus suberArbutus unedo Erica arborea Crataegus monogyna andseveral species of Prunus and Rhamnus Hunter-gatherersmay have gathered hazelnuts acorns and wild fruit (egmazzard Prunus avium) (Uzquiano 1992) The palaeobotan-ical findings concur with the genetical structure of popula-tions of Iberian brown oak (Olalde et al 2002) and somepalaeoecological inferences drawn from the abundant mega-fauna of the Biscay coast (Altuna 1972)

For continental parts of the Iberian Peniacutensula there existsboth pollen and palaeobotanical evidence of mesophiloustaxa in glacial and late glacial contexts (Dupreacute 1988 Ponsamp Reille 1988 Garciacutea-Antoacuten et al 1990 Garciacutea-Antoacuten ampSainz-Ollero 1991 Carrioacuten amp Saacutenchez-Goacutemez 1992 Peacuterez-Obiol amp Juliagrave 1994 Blanco et al 1997) When cave pollenis considered along with reference pollen sequences andcharcoal findings everything seems to point less to refugiarestricted to the far south (Brewer et al 2002) than tosurvival of stationary tree populations in many parts of thePeninsula particularly in intramontane valleys in the Baeticpre-Baetic Iberian and other coastal ranges with expansionand contraction in the central uplands

To conclude palynology at archaeological cave sites oftenraises doubts over contemporaneity between pollen and thelayer containing it and anthracology often seems closer topalaeobotanical orthodoxy than to palaeoecology Neverthe-less both approaches together can build up a solid weightof evidence Furthermore some reference pollen sequenceshave undeniable counterparts in some cave pollen sequences(Carrioacuten 2002b) (Fig 4) Our models about glacial refugiarequire far more information several sites need to be restudiedtheir chronologies need refinement and the present estimatesof continental palaeotemperatures during the upper Pleistoceneshould be viewed with caution Moreover faunal remainsespecially birds and small mammals must be taken far moreinto consideration whenever revision of glacial refugia oftemperate trees is undertaken concerning palaeoclimaticinferences

Relationships with palaeontological findings and human remains

Palynological findings at Cueva Negra del Estrecho del RiacuteoQuiacutepar and Sima de las Palomas del Cabezo Gordo sit easilywith the faunal evidence At Cueva Negra avian palaeonto-logist Anne Easthamrsquos findings (Walker et al 1998 in press)point to five different environmental biotopes coexisting nearthis upland site which today is in an open arid landscapecrossed by a small stream undeservedly called the lsquoRiverrsquoQuiacutepar These are (i) wetlands with a depth of lake-waternecessary for ruddy shelducks mallards wigeons teals gad-walls red-crested pochards common pochards ferruginousducks wild geese little stints and sandpipers (ii) riverine anddamp valley floors where soft sediments offered cover suit-able for the bee-eaters and sand martins at Cueva Negra (iii)Quercus woodland suitable for autumnal acorn-eaters suchas the caversquos Pleistocene jays and woodpigeons where itsowls nightjars woodpeckers woodlarks and several speciesof thrushes finches must have found their prey (iv) steppe andopen country preferred by larks partridges plovers choughseagles buzzards kestrels and falcons whose bones are allwell represented and (v) the craggy mountainsides andcliffs around the cave itself offering roosting places for theomnipresent choughs rock doves crag martins swallowsswifts and rock thrushes Reptiles are especially well repres-ented by tortoise remains Numerous skeletal parts of smalland large mammal species excavated support such a varietyof biotopes around the rock-shelter carnivores such ashyaena brown bear wolf a small feline omnivores such asmacaque and wild boar herbivores such as steppe rhinoceroselephantids giant deer red deer a smaller cervid aurochsbison horse and wild goat Bats hedgehogs and shrewswere also found

At Sima de las Palomas the correlation for the early tomid-upper Pleistocene between fauna and ancient habitats is

Glacial refugia of thermophilous flora in Spain 127

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

less clear-cut Cementation of excavated breccia makes forslow progress in palaeontological research Fewer bird specieshave been identified than at Cueva Negra (13 species asagainst 66) (Walker et al in press) There are of coursecraggy mountainsides around the cave Below the site thecoastal plain surrounding Cabezo Gordo nowadays dry andopen must have contained mixed woodland and grasslandwith gallery woodland and swamps beside erstwhile streamsfeeding nearby wetlands and saltmarshes behind coastal sanddunes (where the Mar Menor coastal lagoon is today) eventhough marine regressions must have repeatedly drainedthese for many millennia each time Mammals include carni-vores such as panther lynx spotted hyaena brown bearfox and badger and herbivores such as hippopotamuselephantids wild horses and asses aurochs red deer wildgoats and abundant lagomorphs Tortoise is again commonin the deposits

The two Upper Pleistocene sites considered here from apalaeoecological standpoint of flora and fauna corroboratea proposal that long-term human presence may occur prefer-entially where several different biotopes coincide

ACKNOWLEDGMENTS

Michegravele Dupreacute assisted with laboratory processing KeithBennett and an anonymous referee provided useful sugges-tions on an earlier draft Research funding was provided bythe Spanish Ministerio de Educacioacuten y Ciencia and theMurcian regional authorityrsquos Fundacioacuten Seneca (projectsBOS2000-0149 PI-1700739FS01 PB92-0971 PB98-0405)

REFERENCES

Altuna J (1972) Fauna de mamiacuteferos de los yacimientos prehistoacutericosde Guipuacutezcoa Munibe 24 1ndash464

Arroyo J (1997) Plant diversity in the region of the Strait of Gibraltara multilevel approach Lagascalia 19 393ndash404

Badal E amp Carrioacuten Y (2001) Del glaciar al interglaciar Los paisajesvegetales a partir de los restos carbonizados hallados en las cuevasde Alicante De Neandertales a Cromantildeones El inicio del pob-lamiento humano en las tierras valencianas (ed by V Villaverde)pp 21ndash40 Universidad de Valencia Valencia

Bennett KD Tzedakis PC amp Willis KJ (1991) Quaternaryrefugia of north European trees Journal of Biogeography 18 103ndash115

Birks HJB (1993) Quaternary palaeoecology and vegetationscience-current contributions and possible future developmentsReview of Palaeobotany and Palynology 79 153ndash177

Blanco E Casado MA Costa-Tenorio M Escribano R Garciacutea-Antoacuten M Geacutenova M Goacutemez A Goacutemez F Moreno JCMorla C Regato P amp Sainz H (1997) Los bosques ibeacutericosUna interpretacioacuten geobotaacutenica Planeta Barcelona

Brewer S Cheddadi R de Beaulieu JL Reille M amp Data Con-tributors (2002) The spread of deciduous Quercus throughout

Europe since the last glacial period Forest Ecology and Management156 27ndash48

Burjachs F amp Juliagrave R (1994) Abrupt climatic changes during thelast glaciation based on pollen analysis of the Abric RomaniCatalonia Spain Quaternary Research 42 308ndash315

Burney DA amp Burney LP (1993) Modern pollen deposition in cavesites experimental results from New York State New Phytolology124 523ndash535

Carrioacuten JS (2002a) A taphonomic study of modern pollenassemblages from dung and surface sediments in arid environ-ments of Spain Review of Palaeobotany and Palynology 120217ndash232

Carrioacuten JS (2002b) Patterns and processes of Late Quaternaryenvironmental change in a montane region of southwestern EuropeQuaternary Science Reviews 21 2047ndash2066

Carrioacuten JS Dupreacute M Fumanal MP amp Montes R (1995) Apalaeoenvironmental study in semi-arid southeastern Spain thepalynological and sedimentological sequence at Perneras Cave(Lorca Murcia) Journal of Archaeological Science 22 355ndash367

Carrioacuten JS amp Munuera M (1997) Upper Pleistocene palaeoenvi-ronmental change in eastern Spain new pollen analytical datafrom Cova Beneito (Alicante) Palaeogeography Palaeoclimatologyand Palaeoecology 128 287ndash299

Carrioacuten JS Munuera M Navarro C Burjachs F Dupreacute M ampWalker MJ (1999a) The palaeoecological potential of pollenrecords in caves the case of Mediterranean Spain QuaternaryScience Reviews 18 1061ndash1073

Carrioacuten JS Munuera M Navarro C amp Saacuteez F (2000) Paleo-climas e historia de la vegetacioacuten cuaternaria en Espantildea a traveacutes delanaacutelisis poliacutenico Viejas falacias y nuevos paradigmas Complutum11 1ndash28

Carrioacuten JS amp Saacutenchez-Goacutemez P (1992) Palynological data in sup-port of the survival of walnut (Juglans regia L) in the westernMediterranean area during last glacial times Journal of Biogeography19 623ndash630

Carrioacuten JS amp Scott L (1999) The challenge of pollen analysis inpalaeoenvironmental studies of hominid beds the record fromSterkfontein caves Journal of Human Evolution 36 401ndash408

Carrioacuten JS amp van Geel B (1999) Fine-resolution Upper Weichselianand Holocene palynological record from Navarreacutes (ValenciaSpain) and a discussion about factors of Mediterranean forestsuccession Review of Palaeobotany and Palynology 106 209ndash236

Comes HP amp Kadereit JW (1998) The effect of Quaternary cli-matic changes on plant distribution and evolution Trends in PlantScience 3 432ndash438

Cuenca A Pomery PJ amp Walker MJ (1986) Chronologicalaspects of the Middle Pleistocene in the coastal belt of southeasternSpain Quaternary climate in Western Mediterranean (ed byF Loacutepez-Vera) pp 353ndash363 Universidad Autoacutenoma Madrid

Davis OK (1990) Caves as sources of biotic remains in arid westernNorth America Palaeogeography Palaeoclimatology and Palae-oecology 76 331ndash348

Dupreacute M (1988) Palinologiacutea y paleoambiente Nuevos datosespantildeoles Referencias Valencia Diputacioacuten de Valencia Serviciode Investigacioacuten Prehistoacuterica Serie de Trabajos Varios no 84

Garciacutea Latorre J amp Garciacutea Latorre J (1996) Los bosques ignoradosde Almeriacutea Una interpretacioacuten histoacuterica y ecoloacutegica Historia y

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

122

JS Carrioacuten

et al

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

of sunshine annually a winter temperature never below 10

deg

Cand a mean July temperature of 26

deg

CTwo lithostratigraphical units have been recognized in the

uppermost 18 m of the column under excavation since 1994Unit 1 is a yellowish cemented breccia and scree adhering tothe overhang Unit 2 is characterized by horizontal accumula-tion of angular scree in a clayey matrix with fine layers ofburnt soil Both units are rich in bones including remains of

Homo sapiens neanderthalensis

and Mousterian artifactsThorium-Uranium and AMS radiocarbon dating assign thesedeposits to about 60 000ndash40 000 years ago The bottom ofthe column below unit 2 has Thorium-Uranium determina-tions of about 125 000 years (Walker

et al

in press) Thestudy deposit is therefore likely to be synchronous with theCueva Negra section that is early to mid Upper PleistoceneDetails on the chronological context of both sites are pre-sented elsewhere (Walker

et al

in press)

METHODS

Sampling of vertical sections was done in close relationshipto the stratigraphy each recognizable archaeological bed

was sampled generally only one sample per bed that is at3ndash5 cm intervals Control for modern pollen rain was doneby means of two samples each made up of 5 subsamplesfrom the surface of the soil at the respective sites Laboratorytreatment was performed following the conventional HClHF and KOH method and

Lycopodium clavatum

tabletscontaining a known quantity of spores were added to eachsample prepared to enable estimation of pollen concentra-tion Pollen grains were concentrated by heavy-liquidflotation Residues were mounted in silicone oil Pollen iden-tification was performed by comparison with the referencecollection at Murcia University Identification criteria forconflicting taxa are described elsewhere (Carrioacuten 2002a)Total sums of 230ndash490 and 351ndash536 pollen grains andspores were obtained for Cueva Negra and Sima de lasPalomas respectively Pollen diagrams were constructedby using progressively the programs Tilia TiliaGraphTGView and CorelDraw 90 (Figs 2 and 3) Internal separa-tion of pollen zones has not been undertaken for the pollendiagrams because visual inspection reveals only slight varia-tion Nomenclature for plant taxa follows Saacutenchez-Goacutemez

et al

(1998)

Fig 3 Sima de las Palomas del Cabezo Gordo pollen diagram (Murcia Spain)

Glacial refugia of thermophilous flora in Spain

123

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

RESULTS

Cueva Negra pollen records

Pollen spectra are codominated by two well-differentiatedgroups of

Quercus

with percentages of around 15 and35 respectively (Fig 2) One is of deciduous species whichgiven the local limestone substrate was no doubt mainly

Quercus faginea

The other is an evergreen group whichcould indicate the presence of

Quercus ilexrotundifolia

or

Qcoccifera

Quercus

is more abundant than any other tree orshrub species although pine (

Pinus

)

juniper (

Juniperus

)and wild olive (

Olea

) consistently reach levels over 2ndash3Low frequency oscillations throughout the sections areshown for hazel (

Corylus

) birch (

Betula

) cluster pine (

Pinuspinaster

) ash (

Fraxinus

) elm (

Ulmus

) willow (

Salix

) lentisk(

Pistacia

) and

Phillyrea

Sporadically present are field maple(

Acer

) yew (

Taxus

) strawberry tree (

Arbutus

) ivy (

Hedera

)and rock-roses (

Cistus

) Although there is a significantherbaceous component of wormwood (

Artemisia

) Poaceaeand Asteraceae and to a lesser extent ChenopodiaceaeLamiaceae and Cyperaceae the abundance of those taxais nonetheless low when compared to Pleistocene pollenspectra from other Mediterranean parts of Spain (Carrioacuten

et al

2000)Pollen analyses at Cueva Negra show steppe vegetation

coexisting with woodland species Determining the geograph-ical location of plant species through pollen analysis iscomplicated due to methodological pitfalls (Birks 1993) sowe can but speculate about the composition of plant forma-tions at the time although some light may be shed on thismatter by regional topography and the ecology of those taxaidentified Thus it seems likely that upland plateaus wereopen ground where grasses and wormwood scrub (

Artemisia

)predominated sometimes with juniper bushes and occasion-ally pine trees whilst valleys and gorges (like that of theQuiacutepar) gave shelter to most of the woodland flora andmesothermophilous shrubs Thus Cueva Negra pollenfrequencies imply mixed copses of both deciduous andevergreen oaks with cluster pines for the most part alongwith representatives of other deciduous species such ashazel or beech that are no longer found there and only lingeron in sheltered valleys of the high Sierra del Segura 50 kmaway to the NW (Fig 1) Many of these trees probablybehaved as phreatophytes growing on river banks and valleyfloors near water-courses Thermophilous taxa such as

Olea

Pistacia

and

Phillyrea

show striking abundance forthe last glacial-stage mesomediterranean enclave aroundCueva Negra

Sima de las Palomas pollen records

Sima de las Palomas pollen spectra are dominated by twogroups of

Quercus

as at Cueva Negra (Fig 3) The deciduous

palynotype is more variable in shape size and exine orna-mentation perhaps reflecting a mosaic of underlying ecolo-gical factors or several species coexisting in a context wheresoils show greater diversity than at Cueva Negra (in morehumid mountainous regions than Murcia

Q faginea Qcanariensis

and

Q pyrenaica

occur today in southern regionsof the Iberian Peninsula) Pine is more prominent than atCueva Negra but which species cannot yet be determinedLikely candidates are black pine (

P nigra) Aleppo pine (Phalepensis) and umbrella pine (P pinea) anthracology hasshown P nigra occurred from c 25 000ndash13 000 year BP inPalaeolithic settlements in Alicante and Valencia (Badal ampCarrioacuten 2001)

Given that today large parts of coastal Murcia on averageget less than 200 mm of rain a year with very high levels ofevapo-transpiration (Saacutenchez-Goacutemez et al 1998) it is strik-ing that Pleistocene pollen spectra show abundant pollen ofdeciduous oaks at Sima de las Palomas alongside trees re-quiring damp-temperate conditions such as Corylus avellanaFraxinus Arbutus unedo Buxus and Betula The oaks mostlikely grew nearby because their pollen frequencies of 15ndash20 exceed those involving long-distance pollen transportto south-eastern Spanish caves (Navarro et al 2001) Waslocal climate damper than now in the upper Pleistocene Oris a combination of Holocene climate changes and anthro-pogenic intervention to blame for the recent decline in decid-uous trees These are not exclusive possibilities but the latteris supported (Carrioacuten et al in press) by a Holocene sequencefrom Gaacutedor (Almeriacutea) by archaeological findings and byhistorical evidence (Garciacutea Latorre amp Garciacutea Latorre 1996Gimeacutenez 2000)

A far broader mosaic of plant communities may beenvisaged in the local Pleistocene landscape than existstoday It would have contained both pine woods andmixed evergreen and deciduous Quercus woodland aswell as deciduous trees in shady zones gorges and besidewater-courses all contributing to a rich undergrowth ofMediterranean species with heliophilous formations onthinner soils The latter would include Periploca angustifoliaOsyris quadripartita Asphodelus Labiates Composites Cis-taceas Thymelaea hirsuta Calicotome intermedia and otherGenistas Last but not least there would have been marshlandsand coastal saltpans characterized by ChenopodiaceaeLycium and Whitania frutescens The widespread presenceof thermophytes including species such as Periploca angus-tifolia and Maytenus europaeus which readily succumb tofrost indicate that local climate can scarcely have been colderthan today Moreover those taxa along with WithaniaPistacia Phillyrea Calicotome and Osyris are all clearlyunder-represented in both external and internal pollen rain(Carrioacuten 2002a) hence their abundance in the neighbour-hood was undountedly greater than pollen spectra imply atfirst glance

124 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

DISCUSSION

Palaeoecological value of the cave pollen spectra studied

Doubts have been raised about the palaeoecological valueof palynological information obtained from archaeologicalstudies from caves and rock-shelters (eg Turner amp Hannon1988) but this issue has been discussed elsewhere (Davis1990 Carrioacuten amp Scott 1999 Carrioacuten et al 1999a) Fourpalaeopalynological aspects need stressing with regard to ourregion First it lacks lake swamp or peat deposits hencevegetational reconstruction must resort to either marinesequences (Targarona 1997) or deposits in caves or rock-shelters Secondly there are no reasons a priori to dismissresults obtained from considerations of pollen-rain repres-entativeness as regards the external vegetation At least thisis what can be concluded experimentally after analysingsurface sediments at caves of different shapes and sizesboth within our region (Navarro et al 2000 2001) andoutside it (Burney amp Burney 1993) Thirdly with regard toconventional sediments those from the very same caves havean advantage in that they are able to collect those ento-mophilous species that make up most of the local vegetationbut are conspicuous by their absence from lake-bed deposits(Carrioacuten 2002a) here indeed any chance at all of biotictransport to a site is of far more help than hindrance topalaeopalynologists

Lastly it is essential that there are palynological indicatorsboth for analytical quality and palaeoecological coherence soas to be able to detect possible skewing of pollen spectra dueto such postdepositional processes as destruction of pollenpercolation or reworking (Saacutenchez-Gontildei 1994) In thisregard it is noteworthy that in both series analysed (Figs 2and 3) have been identified very many types including somenot always readily preserved (eg Taxus Genisteae BuxusCalicotome Periploca) or whose identification demands pre-cise exine characteristics available only in nonoxidized paly-nomorphs (eg Coris Smilax Maytenus Withania) Totalconcentrations of pollen are not especially high being greaterat Sima de las Palomas (between c 9487 and 23 137 grains gminus1)than at Cueva Negra (between c 2342 and 5150 grains gminus1)but they are nevertheless comparable to those often reportedfrom surface sediments in caves (Davis 1990 Navarro et al2001) so there is no need to interpret them as being due toloss caused by destructive processes

At both sites surface pollen spectra reflect wholly modernbut never Pleistocene vegetation Cueva Negra surface samplespredominantly show Pinus (including considerable amount ofP pinaster type) Helianthemum Genisteae and Plantagowhilst Pleistocene ones show clear-cut predominance ofQuercus Poaceae Artemisia and Asteraceae (Fig 2) Simade las Palomas surface samples show greater percentage sim-ilarity but while its surface sediment lacks Quercus (absenttoday in this coastal reagion) Quercus pollen nevertheless is

very abundant in Pleistocene samples from the site Likewiseutterly absent from surface samples are other minority pollengrains identified nonetheless in the Pleistocene samples suchas Corylus Betula Fraxinus Arbutus Ulmus Salix Erica andEphedra distachya-nebrodensis (Fig 3) Last but not leastour failure to detect either bed-rock Miocene spores at CuevaNegra or Permo-Triassic ones at Sima de las Palomas is worthmentioning nor were there differences in staining or preser-vation in pollen spectra from either site such as are commonwhen reworking or sediment mixing have taken place in a cave(Carrioacuten et al 1995)

The new findings in the context of previous palynological studies

Cueva Negra pollen results are comparable with thoseslightly further west from Siles at 1320 m above sea level in anintermontane valley of the Sierra de Segura in Jaeacuten (Carrioacuten2002b) (Fig 4) In deposits dating from upper pleniglacialtimes (c 20 000ndash17 000 cal year BP) were found Pinuspinaster deciduous Quercus evergreen Quercus EricaceaeCorylus Betula and Fraxinus in pollen percentages alwaysabove 2 and frequently also Acer Taxus Arbutus BuxusSalix Ulmus Phillyrea Pistacia and Olea All of these con-tingents were better represented at Siles in lateglacial times(c 17 000ndash11 900 cal year BP) and especially in mid-Holocenetimes (c 7400ndash5300 cal year BP) although pine woodsappear to have predominated during the early Holocene(c 11 900ndash7400 cal year BP)

Although not being synchronous the similarity betweenthe Cueva Negra and Siles spectra is important because bothsites lie in closely related biogeographical areas betweenwhich migration is eminently feasible (Fig 1) not for nothinghave mountains near Cueva Negra been regarded by someplant biogeographers as an eastern extension of the Sierrade Segura (Saacutenchez-Goacutemez amp Alcaraz 1993) The Siles andCueva Negra pollen records show that tree species survivedat quite high places in southern European mountains duringthe last glacial stage This agrees with a hypothesis put for-ward by Bennett et al (1991) that tree survival would havebeen especially important in mountain ranges such as thoseof the Balkans allowing rapid altitudinal displacements of treepopulations in response to climatic pulses (Willis 1994)Owing to their roughly N-S orientation the Segura moun-tains like the Balkans would have readily allowed altitudinalmovements of tree populations to take place Moreover otherindications that mountains in southern Spain containedsignificant tree reservoirs during the last ice age come fromconsideration of the genetic structure of European treepopulations today (Herraacuten et al 1999 Jimeacutenez 2000Salvador et al 2000)

Other pollen sequences in the Iberian Peninsula with com-parable incidences of mesothermophilous flora to Siles or

Glacial refugia of thermophilous flora in Spain 125

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

Cueva Negra only occur at lower altitudes as is the normelsewhere in southern Europe (Willis 1994 Leroy et al1996 Van Andel amp Tzedakis 1996 Magri amp Parra 1997)The Murcian coast offers another important pollen recordfrom the Middle-Upper Palaeolithic transition at CuevaPerneras (Carrioacuten et al 1995) Although tree pollen was lessimportant than at Sima de las Palomas Cueva Pernerasdeposits contained abundant pollen of Pinus Quercus ilex-coccifera and Oleaceae and continuous or frequent presenceof broad-leaved trees (Fraxinus Alnus Corylus JuglansUlmus Salix) and thermophytes (Myrtus Erica arborea

Pistacia Buxus Periploca Withania Lycium Ephedra fragilisCosentinia vellea Selaginella denticulata Ruta)

To the south of Murcia at San Rafael on the Almeriancoast a pollen sequence shows continuous curves for ever-green and deciduous Quercus and Olea during the last glacialmaximum and tardiglacial (Pantaleoacuten-Cano et al in press)Another pollen sequence showing lateglacial tree presencecomes from coprolites of spotted hyaena (Crocuta crocuta)at Cueva de las Ventanas in Granada where about 12 780cal year BP there were pine woods wormwood steppewith juniper grassland and mixed woodland of Quercus

Fig 4 Reference pollen diagram from the lake at Siles (Jaeacuten Spain) which includes mesothermophilous trees and shrubs (shaded zonescorrespond to the last glacial stage black dots refer to percentages below 2)

126 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

with Betula Abies Corylus Alnus Acer Taxus MyrtusBuxus Sorbus Olea Erica arborea Pistacia Ephedra fragilisViburnum Sambucus Cistus and Rhamnus

To the north of Murcia relevant pollen findings come fromsites in inland Mesomediterranean environments namely thedated Middle-Upper Palaeolithic transition at Cova Beneitoin Alicante (Carrioacuten amp Munuera 1997) and the Navarreacutespeat bog in Valencia (Carrioacuten amp van Geel 1999) Howevertheir thermophilous components show oscillations ratherthan uninterrupted presence indeed most of their deciduoustrees and Mediterranean shrubs fell away markedly duringthe upper pleniglacial stage after having advanced togetherduring oxygen-isotope stage 3 In the north-eastern Medi-terranean part of the Iberian Peninsula Abric Romaniacute nearBarcelona shows tree pollen percentages of 40ndash60 betweenabout 70 000 and 40 000 year BP with continuous presenceof Juniperus Rhamnus Quercus Olea-Phillyrea SyringaAlnus Salix Juglans Betula Fagus Betula Coriaria Pistaciaand Vitis (Burjachs amp Juliagrave 1994)

Further afield even the Cantabrian coast and adjacentmountain ranges seem to have offered refuges for trees duringthe last ice age according to pollen analyses (Dupreacute 1988Ramil-Rego et al 1998ab) and macroscopical charcoal(Uzquiano 1992) Pleniglacial pollen samples include lowfrequencies of Pinus Betula Juniperus Corylus QuercusFraxinus Alnus Ulmus Tilia Juglans Fagus and CastaneaCharcoal samples contain Pinus sylvestris P uncinataJuniperus Betula alba B pendula Corylus avellana Quercusrobur Q petraea Tilia platyphyllos T cordata Fraxinusexcelsior Sambucus nigra Viburnum tinus Cornus sanguineaQuercus ilex Fagus sylvatica Sorbus aria S aucupariaS torminalis S domestica Castanea sativa Quercus suberArbutus unedo Erica arborea Crataegus monogyna andseveral species of Prunus and Rhamnus Hunter-gatherersmay have gathered hazelnuts acorns and wild fruit (egmazzard Prunus avium) (Uzquiano 1992) The palaeobotan-ical findings concur with the genetical structure of popula-tions of Iberian brown oak (Olalde et al 2002) and somepalaeoecological inferences drawn from the abundant mega-fauna of the Biscay coast (Altuna 1972)

For continental parts of the Iberian Peniacutensula there existsboth pollen and palaeobotanical evidence of mesophiloustaxa in glacial and late glacial contexts (Dupreacute 1988 Ponsamp Reille 1988 Garciacutea-Antoacuten et al 1990 Garciacutea-Antoacuten ampSainz-Ollero 1991 Carrioacuten amp Saacutenchez-Goacutemez 1992 Peacuterez-Obiol amp Juliagrave 1994 Blanco et al 1997) When cave pollenis considered along with reference pollen sequences andcharcoal findings everything seems to point less to refugiarestricted to the far south (Brewer et al 2002) than tosurvival of stationary tree populations in many parts of thePeninsula particularly in intramontane valleys in the Baeticpre-Baetic Iberian and other coastal ranges with expansionand contraction in the central uplands

To conclude palynology at archaeological cave sites oftenraises doubts over contemporaneity between pollen and thelayer containing it and anthracology often seems closer topalaeobotanical orthodoxy than to palaeoecology Neverthe-less both approaches together can build up a solid weightof evidence Furthermore some reference pollen sequenceshave undeniable counterparts in some cave pollen sequences(Carrioacuten 2002b) (Fig 4) Our models about glacial refugiarequire far more information several sites need to be restudiedtheir chronologies need refinement and the present estimatesof continental palaeotemperatures during the upper Pleistoceneshould be viewed with caution Moreover faunal remainsespecially birds and small mammals must be taken far moreinto consideration whenever revision of glacial refugia oftemperate trees is undertaken concerning palaeoclimaticinferences

Relationships with palaeontological findings and human remains

Palynological findings at Cueva Negra del Estrecho del RiacuteoQuiacutepar and Sima de las Palomas del Cabezo Gordo sit easilywith the faunal evidence At Cueva Negra avian palaeonto-logist Anne Easthamrsquos findings (Walker et al 1998 in press)point to five different environmental biotopes coexisting nearthis upland site which today is in an open arid landscapecrossed by a small stream undeservedly called the lsquoRiverrsquoQuiacutepar These are (i) wetlands with a depth of lake-waternecessary for ruddy shelducks mallards wigeons teals gad-walls red-crested pochards common pochards ferruginousducks wild geese little stints and sandpipers (ii) riverine anddamp valley floors where soft sediments offered cover suit-able for the bee-eaters and sand martins at Cueva Negra (iii)Quercus woodland suitable for autumnal acorn-eaters suchas the caversquos Pleistocene jays and woodpigeons where itsowls nightjars woodpeckers woodlarks and several speciesof thrushes finches must have found their prey (iv) steppe andopen country preferred by larks partridges plovers choughseagles buzzards kestrels and falcons whose bones are allwell represented and (v) the craggy mountainsides andcliffs around the cave itself offering roosting places for theomnipresent choughs rock doves crag martins swallowsswifts and rock thrushes Reptiles are especially well repres-ented by tortoise remains Numerous skeletal parts of smalland large mammal species excavated support such a varietyof biotopes around the rock-shelter carnivores such ashyaena brown bear wolf a small feline omnivores such asmacaque and wild boar herbivores such as steppe rhinoceroselephantids giant deer red deer a smaller cervid aurochsbison horse and wild goat Bats hedgehogs and shrewswere also found

At Sima de las Palomas the correlation for the early tomid-upper Pleistocene between fauna and ancient habitats is

Glacial refugia of thermophilous flora in Spain 127

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

less clear-cut Cementation of excavated breccia makes forslow progress in palaeontological research Fewer bird specieshave been identified than at Cueva Negra (13 species asagainst 66) (Walker et al in press) There are of coursecraggy mountainsides around the cave Below the site thecoastal plain surrounding Cabezo Gordo nowadays dry andopen must have contained mixed woodland and grasslandwith gallery woodland and swamps beside erstwhile streamsfeeding nearby wetlands and saltmarshes behind coastal sanddunes (where the Mar Menor coastal lagoon is today) eventhough marine regressions must have repeatedly drainedthese for many millennia each time Mammals include carni-vores such as panther lynx spotted hyaena brown bearfox and badger and herbivores such as hippopotamuselephantids wild horses and asses aurochs red deer wildgoats and abundant lagomorphs Tortoise is again commonin the deposits

The two Upper Pleistocene sites considered here from apalaeoecological standpoint of flora and fauna corroboratea proposal that long-term human presence may occur prefer-entially where several different biotopes coincide

ACKNOWLEDGMENTS

Michegravele Dupreacute assisted with laboratory processing KeithBennett and an anonymous referee provided useful sugges-tions on an earlier draft Research funding was provided bythe Spanish Ministerio de Educacioacuten y Ciencia and theMurcian regional authorityrsquos Fundacioacuten Seneca (projectsBOS2000-0149 PI-1700739FS01 PB92-0971 PB98-0405)

REFERENCES

Altuna J (1972) Fauna de mamiacuteferos de los yacimientos prehistoacutericosde Guipuacutezcoa Munibe 24 1ndash464

Arroyo J (1997) Plant diversity in the region of the Strait of Gibraltara multilevel approach Lagascalia 19 393ndash404

Badal E amp Carrioacuten Y (2001) Del glaciar al interglaciar Los paisajesvegetales a partir de los restos carbonizados hallados en las cuevasde Alicante De Neandertales a Cromantildeones El inicio del pob-lamiento humano en las tierras valencianas (ed by V Villaverde)pp 21ndash40 Universidad de Valencia Valencia

Bennett KD Tzedakis PC amp Willis KJ (1991) Quaternaryrefugia of north European trees Journal of Biogeography 18 103ndash115

Birks HJB (1993) Quaternary palaeoecology and vegetationscience-current contributions and possible future developmentsReview of Palaeobotany and Palynology 79 153ndash177

Blanco E Casado MA Costa-Tenorio M Escribano R Garciacutea-Antoacuten M Geacutenova M Goacutemez A Goacutemez F Moreno JCMorla C Regato P amp Sainz H (1997) Los bosques ibeacutericosUna interpretacioacuten geobotaacutenica Planeta Barcelona

Brewer S Cheddadi R de Beaulieu JL Reille M amp Data Con-tributors (2002) The spread of deciduous Quercus throughout

Europe since the last glacial period Forest Ecology and Management156 27ndash48

Burjachs F amp Juliagrave R (1994) Abrupt climatic changes during thelast glaciation based on pollen analysis of the Abric RomaniCatalonia Spain Quaternary Research 42 308ndash315

Burney DA amp Burney LP (1993) Modern pollen deposition in cavesites experimental results from New York State New Phytolology124 523ndash535

Carrioacuten JS (2002a) A taphonomic study of modern pollenassemblages from dung and surface sediments in arid environ-ments of Spain Review of Palaeobotany and Palynology 120217ndash232

Carrioacuten JS (2002b) Patterns and processes of Late Quaternaryenvironmental change in a montane region of southwestern EuropeQuaternary Science Reviews 21 2047ndash2066

Carrioacuten JS Dupreacute M Fumanal MP amp Montes R (1995) Apalaeoenvironmental study in semi-arid southeastern Spain thepalynological and sedimentological sequence at Perneras Cave(Lorca Murcia) Journal of Archaeological Science 22 355ndash367

Carrioacuten JS amp Munuera M (1997) Upper Pleistocene palaeoenvi-ronmental change in eastern Spain new pollen analytical datafrom Cova Beneito (Alicante) Palaeogeography Palaeoclimatologyand Palaeoecology 128 287ndash299

Carrioacuten JS Munuera M Navarro C Burjachs F Dupreacute M ampWalker MJ (1999a) The palaeoecological potential of pollenrecords in caves the case of Mediterranean Spain QuaternaryScience Reviews 18 1061ndash1073

Carrioacuten JS Munuera M Navarro C amp Saacuteez F (2000) Paleo-climas e historia de la vegetacioacuten cuaternaria en Espantildea a traveacutes delanaacutelisis poliacutenico Viejas falacias y nuevos paradigmas Complutum11 1ndash28

Carrioacuten JS amp Saacutenchez-Goacutemez P (1992) Palynological data in sup-port of the survival of walnut (Juglans regia L) in the westernMediterranean area during last glacial times Journal of Biogeography19 623ndash630

Carrioacuten JS amp Scott L (1999) The challenge of pollen analysis inpalaeoenvironmental studies of hominid beds the record fromSterkfontein caves Journal of Human Evolution 36 401ndash408

Carrioacuten JS amp van Geel B (1999) Fine-resolution Upper Weichselianand Holocene palynological record from Navarreacutes (ValenciaSpain) and a discussion about factors of Mediterranean forestsuccession Review of Palaeobotany and Palynology 106 209ndash236

Comes HP amp Kadereit JW (1998) The effect of Quaternary cli-matic changes on plant distribution and evolution Trends in PlantScience 3 432ndash438

Cuenca A Pomery PJ amp Walker MJ (1986) Chronologicalaspects of the Middle Pleistocene in the coastal belt of southeasternSpain Quaternary climate in Western Mediterranean (ed byF Loacutepez-Vera) pp 353ndash363 Universidad Autoacutenoma Madrid

Davis OK (1990) Caves as sources of biotic remains in arid westernNorth America Palaeogeography Palaeoclimatology and Palae-oecology 76 331ndash348

Dupreacute M (1988) Palinologiacutea y paleoambiente Nuevos datosespantildeoles Referencias Valencia Diputacioacuten de Valencia Serviciode Investigacioacuten Prehistoacuterica Serie de Trabajos Varios no 84

Garciacutea Latorre J amp Garciacutea Latorre J (1996) Los bosques ignoradosde Almeriacutea Una interpretacioacuten histoacuterica y ecoloacutegica Historia y

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

Glacial refugia of thermophilous flora in Spain

123

copy 2003 Blackwell Publishing Ltd

Global Ecology amp Biogeography

12

119ndash129

RESULTS

Cueva Negra pollen records

Pollen spectra are codominated by two well-differentiatedgroups of

Quercus

with percentages of around 15 and35 respectively (Fig 2) One is of deciduous species whichgiven the local limestone substrate was no doubt mainly

Quercus faginea

The other is an evergreen group whichcould indicate the presence of

Quercus ilexrotundifolia

or

Qcoccifera

Quercus

is more abundant than any other tree orshrub species although pine (

Pinus

)

juniper (

Juniperus

)and wild olive (

Olea

) consistently reach levels over 2ndash3Low frequency oscillations throughout the sections areshown for hazel (

Corylus

) birch (

Betula

) cluster pine (

Pinuspinaster

) ash (

Fraxinus

) elm (

Ulmus

) willow (

Salix

) lentisk(

Pistacia

) and

Phillyrea

Sporadically present are field maple(

Acer

) yew (

Taxus

) strawberry tree (

Arbutus

) ivy (

Hedera

)and rock-roses (

Cistus

) Although there is a significantherbaceous component of wormwood (

Artemisia

) Poaceaeand Asteraceae and to a lesser extent ChenopodiaceaeLamiaceae and Cyperaceae the abundance of those taxais nonetheless low when compared to Pleistocene pollenspectra from other Mediterranean parts of Spain (Carrioacuten

et al

2000)Pollen analyses at Cueva Negra show steppe vegetation

coexisting with woodland species Determining the geograph-ical location of plant species through pollen analysis iscomplicated due to methodological pitfalls (Birks 1993) sowe can but speculate about the composition of plant forma-tions at the time although some light may be shed on thismatter by regional topography and the ecology of those taxaidentified Thus it seems likely that upland plateaus wereopen ground where grasses and wormwood scrub (

Artemisia

)predominated sometimes with juniper bushes and occasion-ally pine trees whilst valleys and gorges (like that of theQuiacutepar) gave shelter to most of the woodland flora andmesothermophilous shrubs Thus Cueva Negra pollenfrequencies imply mixed copses of both deciduous andevergreen oaks with cluster pines for the most part alongwith representatives of other deciduous species such ashazel or beech that are no longer found there and only lingeron in sheltered valleys of the high Sierra del Segura 50 kmaway to the NW (Fig 1) Many of these trees probablybehaved as phreatophytes growing on river banks and valleyfloors near water-courses Thermophilous taxa such as

Olea

Pistacia

and

Phillyrea

show striking abundance forthe last glacial-stage mesomediterranean enclave aroundCueva Negra

Sima de las Palomas pollen records

Sima de las Palomas pollen spectra are dominated by twogroups of

Quercus

as at Cueva Negra (Fig 3) The deciduous

palynotype is more variable in shape size and exine orna-mentation perhaps reflecting a mosaic of underlying ecolo-gical factors or several species coexisting in a context wheresoils show greater diversity than at Cueva Negra (in morehumid mountainous regions than Murcia

Q faginea Qcanariensis

and

Q pyrenaica

occur today in southern regionsof the Iberian Peninsula) Pine is more prominent than atCueva Negra but which species cannot yet be determinedLikely candidates are black pine (

P nigra) Aleppo pine (Phalepensis) and umbrella pine (P pinea) anthracology hasshown P nigra occurred from c 25 000ndash13 000 year BP inPalaeolithic settlements in Alicante and Valencia (Badal ampCarrioacuten 2001)

Given that today large parts of coastal Murcia on averageget less than 200 mm of rain a year with very high levels ofevapo-transpiration (Saacutenchez-Goacutemez et al 1998) it is strik-ing that Pleistocene pollen spectra show abundant pollen ofdeciduous oaks at Sima de las Palomas alongside trees re-quiring damp-temperate conditions such as Corylus avellanaFraxinus Arbutus unedo Buxus and Betula The oaks mostlikely grew nearby because their pollen frequencies of 15ndash20 exceed those involving long-distance pollen transportto south-eastern Spanish caves (Navarro et al 2001) Waslocal climate damper than now in the upper Pleistocene Oris a combination of Holocene climate changes and anthro-pogenic intervention to blame for the recent decline in decid-uous trees These are not exclusive possibilities but the latteris supported (Carrioacuten et al in press) by a Holocene sequencefrom Gaacutedor (Almeriacutea) by archaeological findings and byhistorical evidence (Garciacutea Latorre amp Garciacutea Latorre 1996Gimeacutenez 2000)

A far broader mosaic of plant communities may beenvisaged in the local Pleistocene landscape than existstoday It would have contained both pine woods andmixed evergreen and deciduous Quercus woodland aswell as deciduous trees in shady zones gorges and besidewater-courses all contributing to a rich undergrowth ofMediterranean species with heliophilous formations onthinner soils The latter would include Periploca angustifoliaOsyris quadripartita Asphodelus Labiates Composites Cis-taceas Thymelaea hirsuta Calicotome intermedia and otherGenistas Last but not least there would have been marshlandsand coastal saltpans characterized by ChenopodiaceaeLycium and Whitania frutescens The widespread presenceof thermophytes including species such as Periploca angus-tifolia and Maytenus europaeus which readily succumb tofrost indicate that local climate can scarcely have been colderthan today Moreover those taxa along with WithaniaPistacia Phillyrea Calicotome and Osyris are all clearlyunder-represented in both external and internal pollen rain(Carrioacuten 2002a) hence their abundance in the neighbour-hood was undountedly greater than pollen spectra imply atfirst glance

124 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

DISCUSSION

Palaeoecological value of the cave pollen spectra studied

Doubts have been raised about the palaeoecological valueof palynological information obtained from archaeologicalstudies from caves and rock-shelters (eg Turner amp Hannon1988) but this issue has been discussed elsewhere (Davis1990 Carrioacuten amp Scott 1999 Carrioacuten et al 1999a) Fourpalaeopalynological aspects need stressing with regard to ourregion First it lacks lake swamp or peat deposits hencevegetational reconstruction must resort to either marinesequences (Targarona 1997) or deposits in caves or rock-shelters Secondly there are no reasons a priori to dismissresults obtained from considerations of pollen-rain repres-entativeness as regards the external vegetation At least thisis what can be concluded experimentally after analysingsurface sediments at caves of different shapes and sizesboth within our region (Navarro et al 2000 2001) andoutside it (Burney amp Burney 1993) Thirdly with regard toconventional sediments those from the very same caves havean advantage in that they are able to collect those ento-mophilous species that make up most of the local vegetationbut are conspicuous by their absence from lake-bed deposits(Carrioacuten 2002a) here indeed any chance at all of biotictransport to a site is of far more help than hindrance topalaeopalynologists

Lastly it is essential that there are palynological indicatorsboth for analytical quality and palaeoecological coherence soas to be able to detect possible skewing of pollen spectra dueto such postdepositional processes as destruction of pollenpercolation or reworking (Saacutenchez-Gontildei 1994) In thisregard it is noteworthy that in both series analysed (Figs 2and 3) have been identified very many types including somenot always readily preserved (eg Taxus Genisteae BuxusCalicotome Periploca) or whose identification demands pre-cise exine characteristics available only in nonoxidized paly-nomorphs (eg Coris Smilax Maytenus Withania) Totalconcentrations of pollen are not especially high being greaterat Sima de las Palomas (between c 9487 and 23 137 grains gminus1)than at Cueva Negra (between c 2342 and 5150 grains gminus1)but they are nevertheless comparable to those often reportedfrom surface sediments in caves (Davis 1990 Navarro et al2001) so there is no need to interpret them as being due toloss caused by destructive processes

At both sites surface pollen spectra reflect wholly modernbut never Pleistocene vegetation Cueva Negra surface samplespredominantly show Pinus (including considerable amount ofP pinaster type) Helianthemum Genisteae and Plantagowhilst Pleistocene ones show clear-cut predominance ofQuercus Poaceae Artemisia and Asteraceae (Fig 2) Simade las Palomas surface samples show greater percentage sim-ilarity but while its surface sediment lacks Quercus (absenttoday in this coastal reagion) Quercus pollen nevertheless is

very abundant in Pleistocene samples from the site Likewiseutterly absent from surface samples are other minority pollengrains identified nonetheless in the Pleistocene samples suchas Corylus Betula Fraxinus Arbutus Ulmus Salix Erica andEphedra distachya-nebrodensis (Fig 3) Last but not leastour failure to detect either bed-rock Miocene spores at CuevaNegra or Permo-Triassic ones at Sima de las Palomas is worthmentioning nor were there differences in staining or preser-vation in pollen spectra from either site such as are commonwhen reworking or sediment mixing have taken place in a cave(Carrioacuten et al 1995)

The new findings in the context of previous palynological studies

Cueva Negra pollen results are comparable with thoseslightly further west from Siles at 1320 m above sea level in anintermontane valley of the Sierra de Segura in Jaeacuten (Carrioacuten2002b) (Fig 4) In deposits dating from upper pleniglacialtimes (c 20 000ndash17 000 cal year BP) were found Pinuspinaster deciduous Quercus evergreen Quercus EricaceaeCorylus Betula and Fraxinus in pollen percentages alwaysabove 2 and frequently also Acer Taxus Arbutus BuxusSalix Ulmus Phillyrea Pistacia and Olea All of these con-tingents were better represented at Siles in lateglacial times(c 17 000ndash11 900 cal year BP) and especially in mid-Holocenetimes (c 7400ndash5300 cal year BP) although pine woodsappear to have predominated during the early Holocene(c 11 900ndash7400 cal year BP)

Although not being synchronous the similarity betweenthe Cueva Negra and Siles spectra is important because bothsites lie in closely related biogeographical areas betweenwhich migration is eminently feasible (Fig 1) not for nothinghave mountains near Cueva Negra been regarded by someplant biogeographers as an eastern extension of the Sierrade Segura (Saacutenchez-Goacutemez amp Alcaraz 1993) The Siles andCueva Negra pollen records show that tree species survivedat quite high places in southern European mountains duringthe last glacial stage This agrees with a hypothesis put for-ward by Bennett et al (1991) that tree survival would havebeen especially important in mountain ranges such as thoseof the Balkans allowing rapid altitudinal displacements of treepopulations in response to climatic pulses (Willis 1994)Owing to their roughly N-S orientation the Segura moun-tains like the Balkans would have readily allowed altitudinalmovements of tree populations to take place Moreover otherindications that mountains in southern Spain containedsignificant tree reservoirs during the last ice age come fromconsideration of the genetic structure of European treepopulations today (Herraacuten et al 1999 Jimeacutenez 2000Salvador et al 2000)

Other pollen sequences in the Iberian Peninsula with com-parable incidences of mesothermophilous flora to Siles or

Glacial refugia of thermophilous flora in Spain 125

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

Cueva Negra only occur at lower altitudes as is the normelsewhere in southern Europe (Willis 1994 Leroy et al1996 Van Andel amp Tzedakis 1996 Magri amp Parra 1997)The Murcian coast offers another important pollen recordfrom the Middle-Upper Palaeolithic transition at CuevaPerneras (Carrioacuten et al 1995) Although tree pollen was lessimportant than at Sima de las Palomas Cueva Pernerasdeposits contained abundant pollen of Pinus Quercus ilex-coccifera and Oleaceae and continuous or frequent presenceof broad-leaved trees (Fraxinus Alnus Corylus JuglansUlmus Salix) and thermophytes (Myrtus Erica arborea

Pistacia Buxus Periploca Withania Lycium Ephedra fragilisCosentinia vellea Selaginella denticulata Ruta)

To the south of Murcia at San Rafael on the Almeriancoast a pollen sequence shows continuous curves for ever-green and deciduous Quercus and Olea during the last glacialmaximum and tardiglacial (Pantaleoacuten-Cano et al in press)Another pollen sequence showing lateglacial tree presencecomes from coprolites of spotted hyaena (Crocuta crocuta)at Cueva de las Ventanas in Granada where about 12 780cal year BP there were pine woods wormwood steppewith juniper grassland and mixed woodland of Quercus

Fig 4 Reference pollen diagram from the lake at Siles (Jaeacuten Spain) which includes mesothermophilous trees and shrubs (shaded zonescorrespond to the last glacial stage black dots refer to percentages below 2)

126 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

with Betula Abies Corylus Alnus Acer Taxus MyrtusBuxus Sorbus Olea Erica arborea Pistacia Ephedra fragilisViburnum Sambucus Cistus and Rhamnus

To the north of Murcia relevant pollen findings come fromsites in inland Mesomediterranean environments namely thedated Middle-Upper Palaeolithic transition at Cova Beneitoin Alicante (Carrioacuten amp Munuera 1997) and the Navarreacutespeat bog in Valencia (Carrioacuten amp van Geel 1999) Howevertheir thermophilous components show oscillations ratherthan uninterrupted presence indeed most of their deciduoustrees and Mediterranean shrubs fell away markedly duringthe upper pleniglacial stage after having advanced togetherduring oxygen-isotope stage 3 In the north-eastern Medi-terranean part of the Iberian Peninsula Abric Romaniacute nearBarcelona shows tree pollen percentages of 40ndash60 betweenabout 70 000 and 40 000 year BP with continuous presenceof Juniperus Rhamnus Quercus Olea-Phillyrea SyringaAlnus Salix Juglans Betula Fagus Betula Coriaria Pistaciaand Vitis (Burjachs amp Juliagrave 1994)

Further afield even the Cantabrian coast and adjacentmountain ranges seem to have offered refuges for trees duringthe last ice age according to pollen analyses (Dupreacute 1988Ramil-Rego et al 1998ab) and macroscopical charcoal(Uzquiano 1992) Pleniglacial pollen samples include lowfrequencies of Pinus Betula Juniperus Corylus QuercusFraxinus Alnus Ulmus Tilia Juglans Fagus and CastaneaCharcoal samples contain Pinus sylvestris P uncinataJuniperus Betula alba B pendula Corylus avellana Quercusrobur Q petraea Tilia platyphyllos T cordata Fraxinusexcelsior Sambucus nigra Viburnum tinus Cornus sanguineaQuercus ilex Fagus sylvatica Sorbus aria S aucupariaS torminalis S domestica Castanea sativa Quercus suberArbutus unedo Erica arborea Crataegus monogyna andseveral species of Prunus and Rhamnus Hunter-gatherersmay have gathered hazelnuts acorns and wild fruit (egmazzard Prunus avium) (Uzquiano 1992) The palaeobotan-ical findings concur with the genetical structure of popula-tions of Iberian brown oak (Olalde et al 2002) and somepalaeoecological inferences drawn from the abundant mega-fauna of the Biscay coast (Altuna 1972)

For continental parts of the Iberian Peniacutensula there existsboth pollen and palaeobotanical evidence of mesophiloustaxa in glacial and late glacial contexts (Dupreacute 1988 Ponsamp Reille 1988 Garciacutea-Antoacuten et al 1990 Garciacutea-Antoacuten ampSainz-Ollero 1991 Carrioacuten amp Saacutenchez-Goacutemez 1992 Peacuterez-Obiol amp Juliagrave 1994 Blanco et al 1997) When cave pollenis considered along with reference pollen sequences andcharcoal findings everything seems to point less to refugiarestricted to the far south (Brewer et al 2002) than tosurvival of stationary tree populations in many parts of thePeninsula particularly in intramontane valleys in the Baeticpre-Baetic Iberian and other coastal ranges with expansionand contraction in the central uplands

To conclude palynology at archaeological cave sites oftenraises doubts over contemporaneity between pollen and thelayer containing it and anthracology often seems closer topalaeobotanical orthodoxy than to palaeoecology Neverthe-less both approaches together can build up a solid weightof evidence Furthermore some reference pollen sequenceshave undeniable counterparts in some cave pollen sequences(Carrioacuten 2002b) (Fig 4) Our models about glacial refugiarequire far more information several sites need to be restudiedtheir chronologies need refinement and the present estimatesof continental palaeotemperatures during the upper Pleistoceneshould be viewed with caution Moreover faunal remainsespecially birds and small mammals must be taken far moreinto consideration whenever revision of glacial refugia oftemperate trees is undertaken concerning palaeoclimaticinferences

Relationships with palaeontological findings and human remains

Palynological findings at Cueva Negra del Estrecho del RiacuteoQuiacutepar and Sima de las Palomas del Cabezo Gordo sit easilywith the faunal evidence At Cueva Negra avian palaeonto-logist Anne Easthamrsquos findings (Walker et al 1998 in press)point to five different environmental biotopes coexisting nearthis upland site which today is in an open arid landscapecrossed by a small stream undeservedly called the lsquoRiverrsquoQuiacutepar These are (i) wetlands with a depth of lake-waternecessary for ruddy shelducks mallards wigeons teals gad-walls red-crested pochards common pochards ferruginousducks wild geese little stints and sandpipers (ii) riverine anddamp valley floors where soft sediments offered cover suit-able for the bee-eaters and sand martins at Cueva Negra (iii)Quercus woodland suitable for autumnal acorn-eaters suchas the caversquos Pleistocene jays and woodpigeons where itsowls nightjars woodpeckers woodlarks and several speciesof thrushes finches must have found their prey (iv) steppe andopen country preferred by larks partridges plovers choughseagles buzzards kestrels and falcons whose bones are allwell represented and (v) the craggy mountainsides andcliffs around the cave itself offering roosting places for theomnipresent choughs rock doves crag martins swallowsswifts and rock thrushes Reptiles are especially well repres-ented by tortoise remains Numerous skeletal parts of smalland large mammal species excavated support such a varietyof biotopes around the rock-shelter carnivores such ashyaena brown bear wolf a small feline omnivores such asmacaque and wild boar herbivores such as steppe rhinoceroselephantids giant deer red deer a smaller cervid aurochsbison horse and wild goat Bats hedgehogs and shrewswere also found

At Sima de las Palomas the correlation for the early tomid-upper Pleistocene between fauna and ancient habitats is

Glacial refugia of thermophilous flora in Spain 127

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

less clear-cut Cementation of excavated breccia makes forslow progress in palaeontological research Fewer bird specieshave been identified than at Cueva Negra (13 species asagainst 66) (Walker et al in press) There are of coursecraggy mountainsides around the cave Below the site thecoastal plain surrounding Cabezo Gordo nowadays dry andopen must have contained mixed woodland and grasslandwith gallery woodland and swamps beside erstwhile streamsfeeding nearby wetlands and saltmarshes behind coastal sanddunes (where the Mar Menor coastal lagoon is today) eventhough marine regressions must have repeatedly drainedthese for many millennia each time Mammals include carni-vores such as panther lynx spotted hyaena brown bearfox and badger and herbivores such as hippopotamuselephantids wild horses and asses aurochs red deer wildgoats and abundant lagomorphs Tortoise is again commonin the deposits

The two Upper Pleistocene sites considered here from apalaeoecological standpoint of flora and fauna corroboratea proposal that long-term human presence may occur prefer-entially where several different biotopes coincide

ACKNOWLEDGMENTS

Michegravele Dupreacute assisted with laboratory processing KeithBennett and an anonymous referee provided useful sugges-tions on an earlier draft Research funding was provided bythe Spanish Ministerio de Educacioacuten y Ciencia and theMurcian regional authorityrsquos Fundacioacuten Seneca (projectsBOS2000-0149 PI-1700739FS01 PB92-0971 PB98-0405)

REFERENCES

Altuna J (1972) Fauna de mamiacuteferos de los yacimientos prehistoacutericosde Guipuacutezcoa Munibe 24 1ndash464

Arroyo J (1997) Plant diversity in the region of the Strait of Gibraltara multilevel approach Lagascalia 19 393ndash404

Badal E amp Carrioacuten Y (2001) Del glaciar al interglaciar Los paisajesvegetales a partir de los restos carbonizados hallados en las cuevasde Alicante De Neandertales a Cromantildeones El inicio del pob-lamiento humano en las tierras valencianas (ed by V Villaverde)pp 21ndash40 Universidad de Valencia Valencia

Bennett KD Tzedakis PC amp Willis KJ (1991) Quaternaryrefugia of north European trees Journal of Biogeography 18 103ndash115

Birks HJB (1993) Quaternary palaeoecology and vegetationscience-current contributions and possible future developmentsReview of Palaeobotany and Palynology 79 153ndash177

Blanco E Casado MA Costa-Tenorio M Escribano R Garciacutea-Antoacuten M Geacutenova M Goacutemez A Goacutemez F Moreno JCMorla C Regato P amp Sainz H (1997) Los bosques ibeacutericosUna interpretacioacuten geobotaacutenica Planeta Barcelona

Brewer S Cheddadi R de Beaulieu JL Reille M amp Data Con-tributors (2002) The spread of deciduous Quercus throughout

Europe since the last glacial period Forest Ecology and Management156 27ndash48

Burjachs F amp Juliagrave R (1994) Abrupt climatic changes during thelast glaciation based on pollen analysis of the Abric RomaniCatalonia Spain Quaternary Research 42 308ndash315

Burney DA amp Burney LP (1993) Modern pollen deposition in cavesites experimental results from New York State New Phytolology124 523ndash535

Carrioacuten JS (2002a) A taphonomic study of modern pollenassemblages from dung and surface sediments in arid environ-ments of Spain Review of Palaeobotany and Palynology 120217ndash232

Carrioacuten JS (2002b) Patterns and processes of Late Quaternaryenvironmental change in a montane region of southwestern EuropeQuaternary Science Reviews 21 2047ndash2066

Carrioacuten JS Dupreacute M Fumanal MP amp Montes R (1995) Apalaeoenvironmental study in semi-arid southeastern Spain thepalynological and sedimentological sequence at Perneras Cave(Lorca Murcia) Journal of Archaeological Science 22 355ndash367

Carrioacuten JS amp Munuera M (1997) Upper Pleistocene palaeoenvi-ronmental change in eastern Spain new pollen analytical datafrom Cova Beneito (Alicante) Palaeogeography Palaeoclimatologyand Palaeoecology 128 287ndash299

Carrioacuten JS Munuera M Navarro C Burjachs F Dupreacute M ampWalker MJ (1999a) The palaeoecological potential of pollenrecords in caves the case of Mediterranean Spain QuaternaryScience Reviews 18 1061ndash1073

Carrioacuten JS Munuera M Navarro C amp Saacuteez F (2000) Paleo-climas e historia de la vegetacioacuten cuaternaria en Espantildea a traveacutes delanaacutelisis poliacutenico Viejas falacias y nuevos paradigmas Complutum11 1ndash28

Carrioacuten JS amp Saacutenchez-Goacutemez P (1992) Palynological data in sup-port of the survival of walnut (Juglans regia L) in the westernMediterranean area during last glacial times Journal of Biogeography19 623ndash630

Carrioacuten JS amp Scott L (1999) The challenge of pollen analysis inpalaeoenvironmental studies of hominid beds the record fromSterkfontein caves Journal of Human Evolution 36 401ndash408

Carrioacuten JS amp van Geel B (1999) Fine-resolution Upper Weichselianand Holocene palynological record from Navarreacutes (ValenciaSpain) and a discussion about factors of Mediterranean forestsuccession Review of Palaeobotany and Palynology 106 209ndash236

Comes HP amp Kadereit JW (1998) The effect of Quaternary cli-matic changes on plant distribution and evolution Trends in PlantScience 3 432ndash438

Cuenca A Pomery PJ amp Walker MJ (1986) Chronologicalaspects of the Middle Pleistocene in the coastal belt of southeasternSpain Quaternary climate in Western Mediterranean (ed byF Loacutepez-Vera) pp 353ndash363 Universidad Autoacutenoma Madrid

Davis OK (1990) Caves as sources of biotic remains in arid westernNorth America Palaeogeography Palaeoclimatology and Palae-oecology 76 331ndash348

Dupreacute M (1988) Palinologiacutea y paleoambiente Nuevos datosespantildeoles Referencias Valencia Diputacioacuten de Valencia Serviciode Investigacioacuten Prehistoacuterica Serie de Trabajos Varios no 84

Garciacutea Latorre J amp Garciacutea Latorre J (1996) Los bosques ignoradosde Almeriacutea Una interpretacioacuten histoacuterica y ecoloacutegica Historia y

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

124 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

DISCUSSION

Palaeoecological value of the cave pollen spectra studied

Doubts have been raised about the palaeoecological valueof palynological information obtained from archaeologicalstudies from caves and rock-shelters (eg Turner amp Hannon1988) but this issue has been discussed elsewhere (Davis1990 Carrioacuten amp Scott 1999 Carrioacuten et al 1999a) Fourpalaeopalynological aspects need stressing with regard to ourregion First it lacks lake swamp or peat deposits hencevegetational reconstruction must resort to either marinesequences (Targarona 1997) or deposits in caves or rock-shelters Secondly there are no reasons a priori to dismissresults obtained from considerations of pollen-rain repres-entativeness as regards the external vegetation At least thisis what can be concluded experimentally after analysingsurface sediments at caves of different shapes and sizesboth within our region (Navarro et al 2000 2001) andoutside it (Burney amp Burney 1993) Thirdly with regard toconventional sediments those from the very same caves havean advantage in that they are able to collect those ento-mophilous species that make up most of the local vegetationbut are conspicuous by their absence from lake-bed deposits(Carrioacuten 2002a) here indeed any chance at all of biotictransport to a site is of far more help than hindrance topalaeopalynologists

Lastly it is essential that there are palynological indicatorsboth for analytical quality and palaeoecological coherence soas to be able to detect possible skewing of pollen spectra dueto such postdepositional processes as destruction of pollenpercolation or reworking (Saacutenchez-Gontildei 1994) In thisregard it is noteworthy that in both series analysed (Figs 2and 3) have been identified very many types including somenot always readily preserved (eg Taxus Genisteae BuxusCalicotome Periploca) or whose identification demands pre-cise exine characteristics available only in nonoxidized paly-nomorphs (eg Coris Smilax Maytenus Withania) Totalconcentrations of pollen are not especially high being greaterat Sima de las Palomas (between c 9487 and 23 137 grains gminus1)than at Cueva Negra (between c 2342 and 5150 grains gminus1)but they are nevertheless comparable to those often reportedfrom surface sediments in caves (Davis 1990 Navarro et al2001) so there is no need to interpret them as being due toloss caused by destructive processes

At both sites surface pollen spectra reflect wholly modernbut never Pleistocene vegetation Cueva Negra surface samplespredominantly show Pinus (including considerable amount ofP pinaster type) Helianthemum Genisteae and Plantagowhilst Pleistocene ones show clear-cut predominance ofQuercus Poaceae Artemisia and Asteraceae (Fig 2) Simade las Palomas surface samples show greater percentage sim-ilarity but while its surface sediment lacks Quercus (absenttoday in this coastal reagion) Quercus pollen nevertheless is

very abundant in Pleistocene samples from the site Likewiseutterly absent from surface samples are other minority pollengrains identified nonetheless in the Pleistocene samples suchas Corylus Betula Fraxinus Arbutus Ulmus Salix Erica andEphedra distachya-nebrodensis (Fig 3) Last but not leastour failure to detect either bed-rock Miocene spores at CuevaNegra or Permo-Triassic ones at Sima de las Palomas is worthmentioning nor were there differences in staining or preser-vation in pollen spectra from either site such as are commonwhen reworking or sediment mixing have taken place in a cave(Carrioacuten et al 1995)

The new findings in the context of previous palynological studies

Cueva Negra pollen results are comparable with thoseslightly further west from Siles at 1320 m above sea level in anintermontane valley of the Sierra de Segura in Jaeacuten (Carrioacuten2002b) (Fig 4) In deposits dating from upper pleniglacialtimes (c 20 000ndash17 000 cal year BP) were found Pinuspinaster deciduous Quercus evergreen Quercus EricaceaeCorylus Betula and Fraxinus in pollen percentages alwaysabove 2 and frequently also Acer Taxus Arbutus BuxusSalix Ulmus Phillyrea Pistacia and Olea All of these con-tingents were better represented at Siles in lateglacial times(c 17 000ndash11 900 cal year BP) and especially in mid-Holocenetimes (c 7400ndash5300 cal year BP) although pine woodsappear to have predominated during the early Holocene(c 11 900ndash7400 cal year BP)

Although not being synchronous the similarity betweenthe Cueva Negra and Siles spectra is important because bothsites lie in closely related biogeographical areas betweenwhich migration is eminently feasible (Fig 1) not for nothinghave mountains near Cueva Negra been regarded by someplant biogeographers as an eastern extension of the Sierrade Segura (Saacutenchez-Goacutemez amp Alcaraz 1993) The Siles andCueva Negra pollen records show that tree species survivedat quite high places in southern European mountains duringthe last glacial stage This agrees with a hypothesis put for-ward by Bennett et al (1991) that tree survival would havebeen especially important in mountain ranges such as thoseof the Balkans allowing rapid altitudinal displacements of treepopulations in response to climatic pulses (Willis 1994)Owing to their roughly N-S orientation the Segura moun-tains like the Balkans would have readily allowed altitudinalmovements of tree populations to take place Moreover otherindications that mountains in southern Spain containedsignificant tree reservoirs during the last ice age come fromconsideration of the genetic structure of European treepopulations today (Herraacuten et al 1999 Jimeacutenez 2000Salvador et al 2000)

Other pollen sequences in the Iberian Peninsula with com-parable incidences of mesothermophilous flora to Siles or

Glacial refugia of thermophilous flora in Spain 125

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

Cueva Negra only occur at lower altitudes as is the normelsewhere in southern Europe (Willis 1994 Leroy et al1996 Van Andel amp Tzedakis 1996 Magri amp Parra 1997)The Murcian coast offers another important pollen recordfrom the Middle-Upper Palaeolithic transition at CuevaPerneras (Carrioacuten et al 1995) Although tree pollen was lessimportant than at Sima de las Palomas Cueva Pernerasdeposits contained abundant pollen of Pinus Quercus ilex-coccifera and Oleaceae and continuous or frequent presenceof broad-leaved trees (Fraxinus Alnus Corylus JuglansUlmus Salix) and thermophytes (Myrtus Erica arborea

Pistacia Buxus Periploca Withania Lycium Ephedra fragilisCosentinia vellea Selaginella denticulata Ruta)

To the south of Murcia at San Rafael on the Almeriancoast a pollen sequence shows continuous curves for ever-green and deciduous Quercus and Olea during the last glacialmaximum and tardiglacial (Pantaleoacuten-Cano et al in press)Another pollen sequence showing lateglacial tree presencecomes from coprolites of spotted hyaena (Crocuta crocuta)at Cueva de las Ventanas in Granada where about 12 780cal year BP there were pine woods wormwood steppewith juniper grassland and mixed woodland of Quercus

Fig 4 Reference pollen diagram from the lake at Siles (Jaeacuten Spain) which includes mesothermophilous trees and shrubs (shaded zonescorrespond to the last glacial stage black dots refer to percentages below 2)

126 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

with Betula Abies Corylus Alnus Acer Taxus MyrtusBuxus Sorbus Olea Erica arborea Pistacia Ephedra fragilisViburnum Sambucus Cistus and Rhamnus

To the north of Murcia relevant pollen findings come fromsites in inland Mesomediterranean environments namely thedated Middle-Upper Palaeolithic transition at Cova Beneitoin Alicante (Carrioacuten amp Munuera 1997) and the Navarreacutespeat bog in Valencia (Carrioacuten amp van Geel 1999) Howevertheir thermophilous components show oscillations ratherthan uninterrupted presence indeed most of their deciduoustrees and Mediterranean shrubs fell away markedly duringthe upper pleniglacial stage after having advanced togetherduring oxygen-isotope stage 3 In the north-eastern Medi-terranean part of the Iberian Peninsula Abric Romaniacute nearBarcelona shows tree pollen percentages of 40ndash60 betweenabout 70 000 and 40 000 year BP with continuous presenceof Juniperus Rhamnus Quercus Olea-Phillyrea SyringaAlnus Salix Juglans Betula Fagus Betula Coriaria Pistaciaand Vitis (Burjachs amp Juliagrave 1994)

Further afield even the Cantabrian coast and adjacentmountain ranges seem to have offered refuges for trees duringthe last ice age according to pollen analyses (Dupreacute 1988Ramil-Rego et al 1998ab) and macroscopical charcoal(Uzquiano 1992) Pleniglacial pollen samples include lowfrequencies of Pinus Betula Juniperus Corylus QuercusFraxinus Alnus Ulmus Tilia Juglans Fagus and CastaneaCharcoal samples contain Pinus sylvestris P uncinataJuniperus Betula alba B pendula Corylus avellana Quercusrobur Q petraea Tilia platyphyllos T cordata Fraxinusexcelsior Sambucus nigra Viburnum tinus Cornus sanguineaQuercus ilex Fagus sylvatica Sorbus aria S aucupariaS torminalis S domestica Castanea sativa Quercus suberArbutus unedo Erica arborea Crataegus monogyna andseveral species of Prunus and Rhamnus Hunter-gatherersmay have gathered hazelnuts acorns and wild fruit (egmazzard Prunus avium) (Uzquiano 1992) The palaeobotan-ical findings concur with the genetical structure of popula-tions of Iberian brown oak (Olalde et al 2002) and somepalaeoecological inferences drawn from the abundant mega-fauna of the Biscay coast (Altuna 1972)

For continental parts of the Iberian Peniacutensula there existsboth pollen and palaeobotanical evidence of mesophiloustaxa in glacial and late glacial contexts (Dupreacute 1988 Ponsamp Reille 1988 Garciacutea-Antoacuten et al 1990 Garciacutea-Antoacuten ampSainz-Ollero 1991 Carrioacuten amp Saacutenchez-Goacutemez 1992 Peacuterez-Obiol amp Juliagrave 1994 Blanco et al 1997) When cave pollenis considered along with reference pollen sequences andcharcoal findings everything seems to point less to refugiarestricted to the far south (Brewer et al 2002) than tosurvival of stationary tree populations in many parts of thePeninsula particularly in intramontane valleys in the Baeticpre-Baetic Iberian and other coastal ranges with expansionand contraction in the central uplands

To conclude palynology at archaeological cave sites oftenraises doubts over contemporaneity between pollen and thelayer containing it and anthracology often seems closer topalaeobotanical orthodoxy than to palaeoecology Neverthe-less both approaches together can build up a solid weightof evidence Furthermore some reference pollen sequenceshave undeniable counterparts in some cave pollen sequences(Carrioacuten 2002b) (Fig 4) Our models about glacial refugiarequire far more information several sites need to be restudiedtheir chronologies need refinement and the present estimatesof continental palaeotemperatures during the upper Pleistoceneshould be viewed with caution Moreover faunal remainsespecially birds and small mammals must be taken far moreinto consideration whenever revision of glacial refugia oftemperate trees is undertaken concerning palaeoclimaticinferences

Relationships with palaeontological findings and human remains

Palynological findings at Cueva Negra del Estrecho del RiacuteoQuiacutepar and Sima de las Palomas del Cabezo Gordo sit easilywith the faunal evidence At Cueva Negra avian palaeonto-logist Anne Easthamrsquos findings (Walker et al 1998 in press)point to five different environmental biotopes coexisting nearthis upland site which today is in an open arid landscapecrossed by a small stream undeservedly called the lsquoRiverrsquoQuiacutepar These are (i) wetlands with a depth of lake-waternecessary for ruddy shelducks mallards wigeons teals gad-walls red-crested pochards common pochards ferruginousducks wild geese little stints and sandpipers (ii) riverine anddamp valley floors where soft sediments offered cover suit-able for the bee-eaters and sand martins at Cueva Negra (iii)Quercus woodland suitable for autumnal acorn-eaters suchas the caversquos Pleistocene jays and woodpigeons where itsowls nightjars woodpeckers woodlarks and several speciesof thrushes finches must have found their prey (iv) steppe andopen country preferred by larks partridges plovers choughseagles buzzards kestrels and falcons whose bones are allwell represented and (v) the craggy mountainsides andcliffs around the cave itself offering roosting places for theomnipresent choughs rock doves crag martins swallowsswifts and rock thrushes Reptiles are especially well repres-ented by tortoise remains Numerous skeletal parts of smalland large mammal species excavated support such a varietyof biotopes around the rock-shelter carnivores such ashyaena brown bear wolf a small feline omnivores such asmacaque and wild boar herbivores such as steppe rhinoceroselephantids giant deer red deer a smaller cervid aurochsbison horse and wild goat Bats hedgehogs and shrewswere also found

At Sima de las Palomas the correlation for the early tomid-upper Pleistocene between fauna and ancient habitats is

Glacial refugia of thermophilous flora in Spain 127

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

less clear-cut Cementation of excavated breccia makes forslow progress in palaeontological research Fewer bird specieshave been identified than at Cueva Negra (13 species asagainst 66) (Walker et al in press) There are of coursecraggy mountainsides around the cave Below the site thecoastal plain surrounding Cabezo Gordo nowadays dry andopen must have contained mixed woodland and grasslandwith gallery woodland and swamps beside erstwhile streamsfeeding nearby wetlands and saltmarshes behind coastal sanddunes (where the Mar Menor coastal lagoon is today) eventhough marine regressions must have repeatedly drainedthese for many millennia each time Mammals include carni-vores such as panther lynx spotted hyaena brown bearfox and badger and herbivores such as hippopotamuselephantids wild horses and asses aurochs red deer wildgoats and abundant lagomorphs Tortoise is again commonin the deposits

The two Upper Pleistocene sites considered here from apalaeoecological standpoint of flora and fauna corroboratea proposal that long-term human presence may occur prefer-entially where several different biotopes coincide

ACKNOWLEDGMENTS

Michegravele Dupreacute assisted with laboratory processing KeithBennett and an anonymous referee provided useful sugges-tions on an earlier draft Research funding was provided bythe Spanish Ministerio de Educacioacuten y Ciencia and theMurcian regional authorityrsquos Fundacioacuten Seneca (projectsBOS2000-0149 PI-1700739FS01 PB92-0971 PB98-0405)

REFERENCES

Altuna J (1972) Fauna de mamiacuteferos de los yacimientos prehistoacutericosde Guipuacutezcoa Munibe 24 1ndash464

Arroyo J (1997) Plant diversity in the region of the Strait of Gibraltara multilevel approach Lagascalia 19 393ndash404

Badal E amp Carrioacuten Y (2001) Del glaciar al interglaciar Los paisajesvegetales a partir de los restos carbonizados hallados en las cuevasde Alicante De Neandertales a Cromantildeones El inicio del pob-lamiento humano en las tierras valencianas (ed by V Villaverde)pp 21ndash40 Universidad de Valencia Valencia

Bennett KD Tzedakis PC amp Willis KJ (1991) Quaternaryrefugia of north European trees Journal of Biogeography 18 103ndash115

Birks HJB (1993) Quaternary palaeoecology and vegetationscience-current contributions and possible future developmentsReview of Palaeobotany and Palynology 79 153ndash177

Blanco E Casado MA Costa-Tenorio M Escribano R Garciacutea-Antoacuten M Geacutenova M Goacutemez A Goacutemez F Moreno JCMorla C Regato P amp Sainz H (1997) Los bosques ibeacutericosUna interpretacioacuten geobotaacutenica Planeta Barcelona

Brewer S Cheddadi R de Beaulieu JL Reille M amp Data Con-tributors (2002) The spread of deciduous Quercus throughout

Europe since the last glacial period Forest Ecology and Management156 27ndash48

Burjachs F amp Juliagrave R (1994) Abrupt climatic changes during thelast glaciation based on pollen analysis of the Abric RomaniCatalonia Spain Quaternary Research 42 308ndash315

Burney DA amp Burney LP (1993) Modern pollen deposition in cavesites experimental results from New York State New Phytolology124 523ndash535

Carrioacuten JS (2002a) A taphonomic study of modern pollenassemblages from dung and surface sediments in arid environ-ments of Spain Review of Palaeobotany and Palynology 120217ndash232

Carrioacuten JS (2002b) Patterns and processes of Late Quaternaryenvironmental change in a montane region of southwestern EuropeQuaternary Science Reviews 21 2047ndash2066

Carrioacuten JS Dupreacute M Fumanal MP amp Montes R (1995) Apalaeoenvironmental study in semi-arid southeastern Spain thepalynological and sedimentological sequence at Perneras Cave(Lorca Murcia) Journal of Archaeological Science 22 355ndash367

Carrioacuten JS amp Munuera M (1997) Upper Pleistocene palaeoenvi-ronmental change in eastern Spain new pollen analytical datafrom Cova Beneito (Alicante) Palaeogeography Palaeoclimatologyand Palaeoecology 128 287ndash299

Carrioacuten JS Munuera M Navarro C Burjachs F Dupreacute M ampWalker MJ (1999a) The palaeoecological potential of pollenrecords in caves the case of Mediterranean Spain QuaternaryScience Reviews 18 1061ndash1073

Carrioacuten JS Munuera M Navarro C amp Saacuteez F (2000) Paleo-climas e historia de la vegetacioacuten cuaternaria en Espantildea a traveacutes delanaacutelisis poliacutenico Viejas falacias y nuevos paradigmas Complutum11 1ndash28

Carrioacuten JS amp Saacutenchez-Goacutemez P (1992) Palynological data in sup-port of the survival of walnut (Juglans regia L) in the westernMediterranean area during last glacial times Journal of Biogeography19 623ndash630

Carrioacuten JS amp Scott L (1999) The challenge of pollen analysis inpalaeoenvironmental studies of hominid beds the record fromSterkfontein caves Journal of Human Evolution 36 401ndash408

Carrioacuten JS amp van Geel B (1999) Fine-resolution Upper Weichselianand Holocene palynological record from Navarreacutes (ValenciaSpain) and a discussion about factors of Mediterranean forestsuccession Review of Palaeobotany and Palynology 106 209ndash236

Comes HP amp Kadereit JW (1998) The effect of Quaternary cli-matic changes on plant distribution and evolution Trends in PlantScience 3 432ndash438

Cuenca A Pomery PJ amp Walker MJ (1986) Chronologicalaspects of the Middle Pleistocene in the coastal belt of southeasternSpain Quaternary climate in Western Mediterranean (ed byF Loacutepez-Vera) pp 353ndash363 Universidad Autoacutenoma Madrid

Davis OK (1990) Caves as sources of biotic remains in arid westernNorth America Palaeogeography Palaeoclimatology and Palae-oecology 76 331ndash348

Dupreacute M (1988) Palinologiacutea y paleoambiente Nuevos datosespantildeoles Referencias Valencia Diputacioacuten de Valencia Serviciode Investigacioacuten Prehistoacuterica Serie de Trabajos Varios no 84

Garciacutea Latorre J amp Garciacutea Latorre J (1996) Los bosques ignoradosde Almeriacutea Una interpretacioacuten histoacuterica y ecoloacutegica Historia y

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

Glacial refugia of thermophilous flora in Spain 125

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

Cueva Negra only occur at lower altitudes as is the normelsewhere in southern Europe (Willis 1994 Leroy et al1996 Van Andel amp Tzedakis 1996 Magri amp Parra 1997)The Murcian coast offers another important pollen recordfrom the Middle-Upper Palaeolithic transition at CuevaPerneras (Carrioacuten et al 1995) Although tree pollen was lessimportant than at Sima de las Palomas Cueva Pernerasdeposits contained abundant pollen of Pinus Quercus ilex-coccifera and Oleaceae and continuous or frequent presenceof broad-leaved trees (Fraxinus Alnus Corylus JuglansUlmus Salix) and thermophytes (Myrtus Erica arborea

Pistacia Buxus Periploca Withania Lycium Ephedra fragilisCosentinia vellea Selaginella denticulata Ruta)

To the south of Murcia at San Rafael on the Almeriancoast a pollen sequence shows continuous curves for ever-green and deciduous Quercus and Olea during the last glacialmaximum and tardiglacial (Pantaleoacuten-Cano et al in press)Another pollen sequence showing lateglacial tree presencecomes from coprolites of spotted hyaena (Crocuta crocuta)at Cueva de las Ventanas in Granada where about 12 780cal year BP there were pine woods wormwood steppewith juniper grassland and mixed woodland of Quercus

Fig 4 Reference pollen diagram from the lake at Siles (Jaeacuten Spain) which includes mesothermophilous trees and shrubs (shaded zonescorrespond to the last glacial stage black dots refer to percentages below 2)

126 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

with Betula Abies Corylus Alnus Acer Taxus MyrtusBuxus Sorbus Olea Erica arborea Pistacia Ephedra fragilisViburnum Sambucus Cistus and Rhamnus

To the north of Murcia relevant pollen findings come fromsites in inland Mesomediterranean environments namely thedated Middle-Upper Palaeolithic transition at Cova Beneitoin Alicante (Carrioacuten amp Munuera 1997) and the Navarreacutespeat bog in Valencia (Carrioacuten amp van Geel 1999) Howevertheir thermophilous components show oscillations ratherthan uninterrupted presence indeed most of their deciduoustrees and Mediterranean shrubs fell away markedly duringthe upper pleniglacial stage after having advanced togetherduring oxygen-isotope stage 3 In the north-eastern Medi-terranean part of the Iberian Peninsula Abric Romaniacute nearBarcelona shows tree pollen percentages of 40ndash60 betweenabout 70 000 and 40 000 year BP with continuous presenceof Juniperus Rhamnus Quercus Olea-Phillyrea SyringaAlnus Salix Juglans Betula Fagus Betula Coriaria Pistaciaand Vitis (Burjachs amp Juliagrave 1994)

Further afield even the Cantabrian coast and adjacentmountain ranges seem to have offered refuges for trees duringthe last ice age according to pollen analyses (Dupreacute 1988Ramil-Rego et al 1998ab) and macroscopical charcoal(Uzquiano 1992) Pleniglacial pollen samples include lowfrequencies of Pinus Betula Juniperus Corylus QuercusFraxinus Alnus Ulmus Tilia Juglans Fagus and CastaneaCharcoal samples contain Pinus sylvestris P uncinataJuniperus Betula alba B pendula Corylus avellana Quercusrobur Q petraea Tilia platyphyllos T cordata Fraxinusexcelsior Sambucus nigra Viburnum tinus Cornus sanguineaQuercus ilex Fagus sylvatica Sorbus aria S aucupariaS torminalis S domestica Castanea sativa Quercus suberArbutus unedo Erica arborea Crataegus monogyna andseveral species of Prunus and Rhamnus Hunter-gatherersmay have gathered hazelnuts acorns and wild fruit (egmazzard Prunus avium) (Uzquiano 1992) The palaeobotan-ical findings concur with the genetical structure of popula-tions of Iberian brown oak (Olalde et al 2002) and somepalaeoecological inferences drawn from the abundant mega-fauna of the Biscay coast (Altuna 1972)

For continental parts of the Iberian Peniacutensula there existsboth pollen and palaeobotanical evidence of mesophiloustaxa in glacial and late glacial contexts (Dupreacute 1988 Ponsamp Reille 1988 Garciacutea-Antoacuten et al 1990 Garciacutea-Antoacuten ampSainz-Ollero 1991 Carrioacuten amp Saacutenchez-Goacutemez 1992 Peacuterez-Obiol amp Juliagrave 1994 Blanco et al 1997) When cave pollenis considered along with reference pollen sequences andcharcoal findings everything seems to point less to refugiarestricted to the far south (Brewer et al 2002) than tosurvival of stationary tree populations in many parts of thePeninsula particularly in intramontane valleys in the Baeticpre-Baetic Iberian and other coastal ranges with expansionand contraction in the central uplands

To conclude palynology at archaeological cave sites oftenraises doubts over contemporaneity between pollen and thelayer containing it and anthracology often seems closer topalaeobotanical orthodoxy than to palaeoecology Neverthe-less both approaches together can build up a solid weightof evidence Furthermore some reference pollen sequenceshave undeniable counterparts in some cave pollen sequences(Carrioacuten 2002b) (Fig 4) Our models about glacial refugiarequire far more information several sites need to be restudiedtheir chronologies need refinement and the present estimatesof continental palaeotemperatures during the upper Pleistoceneshould be viewed with caution Moreover faunal remainsespecially birds and small mammals must be taken far moreinto consideration whenever revision of glacial refugia oftemperate trees is undertaken concerning palaeoclimaticinferences

Relationships with palaeontological findings and human remains

Palynological findings at Cueva Negra del Estrecho del RiacuteoQuiacutepar and Sima de las Palomas del Cabezo Gordo sit easilywith the faunal evidence At Cueva Negra avian palaeonto-logist Anne Easthamrsquos findings (Walker et al 1998 in press)point to five different environmental biotopes coexisting nearthis upland site which today is in an open arid landscapecrossed by a small stream undeservedly called the lsquoRiverrsquoQuiacutepar These are (i) wetlands with a depth of lake-waternecessary for ruddy shelducks mallards wigeons teals gad-walls red-crested pochards common pochards ferruginousducks wild geese little stints and sandpipers (ii) riverine anddamp valley floors where soft sediments offered cover suit-able for the bee-eaters and sand martins at Cueva Negra (iii)Quercus woodland suitable for autumnal acorn-eaters suchas the caversquos Pleistocene jays and woodpigeons where itsowls nightjars woodpeckers woodlarks and several speciesof thrushes finches must have found their prey (iv) steppe andopen country preferred by larks partridges plovers choughseagles buzzards kestrels and falcons whose bones are allwell represented and (v) the craggy mountainsides andcliffs around the cave itself offering roosting places for theomnipresent choughs rock doves crag martins swallowsswifts and rock thrushes Reptiles are especially well repres-ented by tortoise remains Numerous skeletal parts of smalland large mammal species excavated support such a varietyof biotopes around the rock-shelter carnivores such ashyaena brown bear wolf a small feline omnivores such asmacaque and wild boar herbivores such as steppe rhinoceroselephantids giant deer red deer a smaller cervid aurochsbison horse and wild goat Bats hedgehogs and shrewswere also found

At Sima de las Palomas the correlation for the early tomid-upper Pleistocene between fauna and ancient habitats is

Glacial refugia of thermophilous flora in Spain 127

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

less clear-cut Cementation of excavated breccia makes forslow progress in palaeontological research Fewer bird specieshave been identified than at Cueva Negra (13 species asagainst 66) (Walker et al in press) There are of coursecraggy mountainsides around the cave Below the site thecoastal plain surrounding Cabezo Gordo nowadays dry andopen must have contained mixed woodland and grasslandwith gallery woodland and swamps beside erstwhile streamsfeeding nearby wetlands and saltmarshes behind coastal sanddunes (where the Mar Menor coastal lagoon is today) eventhough marine regressions must have repeatedly drainedthese for many millennia each time Mammals include carni-vores such as panther lynx spotted hyaena brown bearfox and badger and herbivores such as hippopotamuselephantids wild horses and asses aurochs red deer wildgoats and abundant lagomorphs Tortoise is again commonin the deposits

The two Upper Pleistocene sites considered here from apalaeoecological standpoint of flora and fauna corroboratea proposal that long-term human presence may occur prefer-entially where several different biotopes coincide

ACKNOWLEDGMENTS

Michegravele Dupreacute assisted with laboratory processing KeithBennett and an anonymous referee provided useful sugges-tions on an earlier draft Research funding was provided bythe Spanish Ministerio de Educacioacuten y Ciencia and theMurcian regional authorityrsquos Fundacioacuten Seneca (projectsBOS2000-0149 PI-1700739FS01 PB92-0971 PB98-0405)

REFERENCES

Altuna J (1972) Fauna de mamiacuteferos de los yacimientos prehistoacutericosde Guipuacutezcoa Munibe 24 1ndash464

Arroyo J (1997) Plant diversity in the region of the Strait of Gibraltara multilevel approach Lagascalia 19 393ndash404

Badal E amp Carrioacuten Y (2001) Del glaciar al interglaciar Los paisajesvegetales a partir de los restos carbonizados hallados en las cuevasde Alicante De Neandertales a Cromantildeones El inicio del pob-lamiento humano en las tierras valencianas (ed by V Villaverde)pp 21ndash40 Universidad de Valencia Valencia

Bennett KD Tzedakis PC amp Willis KJ (1991) Quaternaryrefugia of north European trees Journal of Biogeography 18 103ndash115

Birks HJB (1993) Quaternary palaeoecology and vegetationscience-current contributions and possible future developmentsReview of Palaeobotany and Palynology 79 153ndash177

Blanco E Casado MA Costa-Tenorio M Escribano R Garciacutea-Antoacuten M Geacutenova M Goacutemez A Goacutemez F Moreno JCMorla C Regato P amp Sainz H (1997) Los bosques ibeacutericosUna interpretacioacuten geobotaacutenica Planeta Barcelona

Brewer S Cheddadi R de Beaulieu JL Reille M amp Data Con-tributors (2002) The spread of deciduous Quercus throughout

Europe since the last glacial period Forest Ecology and Management156 27ndash48

Burjachs F amp Juliagrave R (1994) Abrupt climatic changes during thelast glaciation based on pollen analysis of the Abric RomaniCatalonia Spain Quaternary Research 42 308ndash315

Burney DA amp Burney LP (1993) Modern pollen deposition in cavesites experimental results from New York State New Phytolology124 523ndash535

Carrioacuten JS (2002a) A taphonomic study of modern pollenassemblages from dung and surface sediments in arid environ-ments of Spain Review of Palaeobotany and Palynology 120217ndash232

Carrioacuten JS (2002b) Patterns and processes of Late Quaternaryenvironmental change in a montane region of southwestern EuropeQuaternary Science Reviews 21 2047ndash2066

Carrioacuten JS Dupreacute M Fumanal MP amp Montes R (1995) Apalaeoenvironmental study in semi-arid southeastern Spain thepalynological and sedimentological sequence at Perneras Cave(Lorca Murcia) Journal of Archaeological Science 22 355ndash367

Carrioacuten JS amp Munuera M (1997) Upper Pleistocene palaeoenvi-ronmental change in eastern Spain new pollen analytical datafrom Cova Beneito (Alicante) Palaeogeography Palaeoclimatologyand Palaeoecology 128 287ndash299

Carrioacuten JS Munuera M Navarro C Burjachs F Dupreacute M ampWalker MJ (1999a) The palaeoecological potential of pollenrecords in caves the case of Mediterranean Spain QuaternaryScience Reviews 18 1061ndash1073

Carrioacuten JS Munuera M Navarro C amp Saacuteez F (2000) Paleo-climas e historia de la vegetacioacuten cuaternaria en Espantildea a traveacutes delanaacutelisis poliacutenico Viejas falacias y nuevos paradigmas Complutum11 1ndash28

Carrioacuten JS amp Saacutenchez-Goacutemez P (1992) Palynological data in sup-port of the survival of walnut (Juglans regia L) in the westernMediterranean area during last glacial times Journal of Biogeography19 623ndash630

Carrioacuten JS amp Scott L (1999) The challenge of pollen analysis inpalaeoenvironmental studies of hominid beds the record fromSterkfontein caves Journal of Human Evolution 36 401ndash408

Carrioacuten JS amp van Geel B (1999) Fine-resolution Upper Weichselianand Holocene palynological record from Navarreacutes (ValenciaSpain) and a discussion about factors of Mediterranean forestsuccession Review of Palaeobotany and Palynology 106 209ndash236

Comes HP amp Kadereit JW (1998) The effect of Quaternary cli-matic changes on plant distribution and evolution Trends in PlantScience 3 432ndash438

Cuenca A Pomery PJ amp Walker MJ (1986) Chronologicalaspects of the Middle Pleistocene in the coastal belt of southeasternSpain Quaternary climate in Western Mediterranean (ed byF Loacutepez-Vera) pp 353ndash363 Universidad Autoacutenoma Madrid

Davis OK (1990) Caves as sources of biotic remains in arid westernNorth America Palaeogeography Palaeoclimatology and Palae-oecology 76 331ndash348

Dupreacute M (1988) Palinologiacutea y paleoambiente Nuevos datosespantildeoles Referencias Valencia Diputacioacuten de Valencia Serviciode Investigacioacuten Prehistoacuterica Serie de Trabajos Varios no 84

Garciacutea Latorre J amp Garciacutea Latorre J (1996) Los bosques ignoradosde Almeriacutea Una interpretacioacuten histoacuterica y ecoloacutegica Historia y

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

126 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

with Betula Abies Corylus Alnus Acer Taxus MyrtusBuxus Sorbus Olea Erica arborea Pistacia Ephedra fragilisViburnum Sambucus Cistus and Rhamnus

To the north of Murcia relevant pollen findings come fromsites in inland Mesomediterranean environments namely thedated Middle-Upper Palaeolithic transition at Cova Beneitoin Alicante (Carrioacuten amp Munuera 1997) and the Navarreacutespeat bog in Valencia (Carrioacuten amp van Geel 1999) Howevertheir thermophilous components show oscillations ratherthan uninterrupted presence indeed most of their deciduoustrees and Mediterranean shrubs fell away markedly duringthe upper pleniglacial stage after having advanced togetherduring oxygen-isotope stage 3 In the north-eastern Medi-terranean part of the Iberian Peninsula Abric Romaniacute nearBarcelona shows tree pollen percentages of 40ndash60 betweenabout 70 000 and 40 000 year BP with continuous presenceof Juniperus Rhamnus Quercus Olea-Phillyrea SyringaAlnus Salix Juglans Betula Fagus Betula Coriaria Pistaciaand Vitis (Burjachs amp Juliagrave 1994)

Further afield even the Cantabrian coast and adjacentmountain ranges seem to have offered refuges for trees duringthe last ice age according to pollen analyses (Dupreacute 1988Ramil-Rego et al 1998ab) and macroscopical charcoal(Uzquiano 1992) Pleniglacial pollen samples include lowfrequencies of Pinus Betula Juniperus Corylus QuercusFraxinus Alnus Ulmus Tilia Juglans Fagus and CastaneaCharcoal samples contain Pinus sylvestris P uncinataJuniperus Betula alba B pendula Corylus avellana Quercusrobur Q petraea Tilia platyphyllos T cordata Fraxinusexcelsior Sambucus nigra Viburnum tinus Cornus sanguineaQuercus ilex Fagus sylvatica Sorbus aria S aucupariaS torminalis S domestica Castanea sativa Quercus suberArbutus unedo Erica arborea Crataegus monogyna andseveral species of Prunus and Rhamnus Hunter-gatherersmay have gathered hazelnuts acorns and wild fruit (egmazzard Prunus avium) (Uzquiano 1992) The palaeobotan-ical findings concur with the genetical structure of popula-tions of Iberian brown oak (Olalde et al 2002) and somepalaeoecological inferences drawn from the abundant mega-fauna of the Biscay coast (Altuna 1972)

For continental parts of the Iberian Peniacutensula there existsboth pollen and palaeobotanical evidence of mesophiloustaxa in glacial and late glacial contexts (Dupreacute 1988 Ponsamp Reille 1988 Garciacutea-Antoacuten et al 1990 Garciacutea-Antoacuten ampSainz-Ollero 1991 Carrioacuten amp Saacutenchez-Goacutemez 1992 Peacuterez-Obiol amp Juliagrave 1994 Blanco et al 1997) When cave pollenis considered along with reference pollen sequences andcharcoal findings everything seems to point less to refugiarestricted to the far south (Brewer et al 2002) than tosurvival of stationary tree populations in many parts of thePeninsula particularly in intramontane valleys in the Baeticpre-Baetic Iberian and other coastal ranges with expansionand contraction in the central uplands

To conclude palynology at archaeological cave sites oftenraises doubts over contemporaneity between pollen and thelayer containing it and anthracology often seems closer topalaeobotanical orthodoxy than to palaeoecology Neverthe-less both approaches together can build up a solid weightof evidence Furthermore some reference pollen sequenceshave undeniable counterparts in some cave pollen sequences(Carrioacuten 2002b) (Fig 4) Our models about glacial refugiarequire far more information several sites need to be restudiedtheir chronologies need refinement and the present estimatesof continental palaeotemperatures during the upper Pleistoceneshould be viewed with caution Moreover faunal remainsespecially birds and small mammals must be taken far moreinto consideration whenever revision of glacial refugia oftemperate trees is undertaken concerning palaeoclimaticinferences

Relationships with palaeontological findings and human remains

Palynological findings at Cueva Negra del Estrecho del RiacuteoQuiacutepar and Sima de las Palomas del Cabezo Gordo sit easilywith the faunal evidence At Cueva Negra avian palaeonto-logist Anne Easthamrsquos findings (Walker et al 1998 in press)point to five different environmental biotopes coexisting nearthis upland site which today is in an open arid landscapecrossed by a small stream undeservedly called the lsquoRiverrsquoQuiacutepar These are (i) wetlands with a depth of lake-waternecessary for ruddy shelducks mallards wigeons teals gad-walls red-crested pochards common pochards ferruginousducks wild geese little stints and sandpipers (ii) riverine anddamp valley floors where soft sediments offered cover suit-able for the bee-eaters and sand martins at Cueva Negra (iii)Quercus woodland suitable for autumnal acorn-eaters suchas the caversquos Pleistocene jays and woodpigeons where itsowls nightjars woodpeckers woodlarks and several speciesof thrushes finches must have found their prey (iv) steppe andopen country preferred by larks partridges plovers choughseagles buzzards kestrels and falcons whose bones are allwell represented and (v) the craggy mountainsides andcliffs around the cave itself offering roosting places for theomnipresent choughs rock doves crag martins swallowsswifts and rock thrushes Reptiles are especially well repres-ented by tortoise remains Numerous skeletal parts of smalland large mammal species excavated support such a varietyof biotopes around the rock-shelter carnivores such ashyaena brown bear wolf a small feline omnivores such asmacaque and wild boar herbivores such as steppe rhinoceroselephantids giant deer red deer a smaller cervid aurochsbison horse and wild goat Bats hedgehogs and shrewswere also found

At Sima de las Palomas the correlation for the early tomid-upper Pleistocene between fauna and ancient habitats is

Glacial refugia of thermophilous flora in Spain 127

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

less clear-cut Cementation of excavated breccia makes forslow progress in palaeontological research Fewer bird specieshave been identified than at Cueva Negra (13 species asagainst 66) (Walker et al in press) There are of coursecraggy mountainsides around the cave Below the site thecoastal plain surrounding Cabezo Gordo nowadays dry andopen must have contained mixed woodland and grasslandwith gallery woodland and swamps beside erstwhile streamsfeeding nearby wetlands and saltmarshes behind coastal sanddunes (where the Mar Menor coastal lagoon is today) eventhough marine regressions must have repeatedly drainedthese for many millennia each time Mammals include carni-vores such as panther lynx spotted hyaena brown bearfox and badger and herbivores such as hippopotamuselephantids wild horses and asses aurochs red deer wildgoats and abundant lagomorphs Tortoise is again commonin the deposits

The two Upper Pleistocene sites considered here from apalaeoecological standpoint of flora and fauna corroboratea proposal that long-term human presence may occur prefer-entially where several different biotopes coincide

ACKNOWLEDGMENTS

Michegravele Dupreacute assisted with laboratory processing KeithBennett and an anonymous referee provided useful sugges-tions on an earlier draft Research funding was provided bythe Spanish Ministerio de Educacioacuten y Ciencia and theMurcian regional authorityrsquos Fundacioacuten Seneca (projectsBOS2000-0149 PI-1700739FS01 PB92-0971 PB98-0405)

REFERENCES

Altuna J (1972) Fauna de mamiacuteferos de los yacimientos prehistoacutericosde Guipuacutezcoa Munibe 24 1ndash464

Arroyo J (1997) Plant diversity in the region of the Strait of Gibraltara multilevel approach Lagascalia 19 393ndash404

Badal E amp Carrioacuten Y (2001) Del glaciar al interglaciar Los paisajesvegetales a partir de los restos carbonizados hallados en las cuevasde Alicante De Neandertales a Cromantildeones El inicio del pob-lamiento humano en las tierras valencianas (ed by V Villaverde)pp 21ndash40 Universidad de Valencia Valencia

Bennett KD Tzedakis PC amp Willis KJ (1991) Quaternaryrefugia of north European trees Journal of Biogeography 18 103ndash115

Birks HJB (1993) Quaternary palaeoecology and vegetationscience-current contributions and possible future developmentsReview of Palaeobotany and Palynology 79 153ndash177

Blanco E Casado MA Costa-Tenorio M Escribano R Garciacutea-Antoacuten M Geacutenova M Goacutemez A Goacutemez F Moreno JCMorla C Regato P amp Sainz H (1997) Los bosques ibeacutericosUna interpretacioacuten geobotaacutenica Planeta Barcelona

Brewer S Cheddadi R de Beaulieu JL Reille M amp Data Con-tributors (2002) The spread of deciduous Quercus throughout

Europe since the last glacial period Forest Ecology and Management156 27ndash48

Burjachs F amp Juliagrave R (1994) Abrupt climatic changes during thelast glaciation based on pollen analysis of the Abric RomaniCatalonia Spain Quaternary Research 42 308ndash315

Burney DA amp Burney LP (1993) Modern pollen deposition in cavesites experimental results from New York State New Phytolology124 523ndash535

Carrioacuten JS (2002a) A taphonomic study of modern pollenassemblages from dung and surface sediments in arid environ-ments of Spain Review of Palaeobotany and Palynology 120217ndash232

Carrioacuten JS (2002b) Patterns and processes of Late Quaternaryenvironmental change in a montane region of southwestern EuropeQuaternary Science Reviews 21 2047ndash2066

Carrioacuten JS Dupreacute M Fumanal MP amp Montes R (1995) Apalaeoenvironmental study in semi-arid southeastern Spain thepalynological and sedimentological sequence at Perneras Cave(Lorca Murcia) Journal of Archaeological Science 22 355ndash367

Carrioacuten JS amp Munuera M (1997) Upper Pleistocene palaeoenvi-ronmental change in eastern Spain new pollen analytical datafrom Cova Beneito (Alicante) Palaeogeography Palaeoclimatologyand Palaeoecology 128 287ndash299

Carrioacuten JS Munuera M Navarro C Burjachs F Dupreacute M ampWalker MJ (1999a) The palaeoecological potential of pollenrecords in caves the case of Mediterranean Spain QuaternaryScience Reviews 18 1061ndash1073

Carrioacuten JS Munuera M Navarro C amp Saacuteez F (2000) Paleo-climas e historia de la vegetacioacuten cuaternaria en Espantildea a traveacutes delanaacutelisis poliacutenico Viejas falacias y nuevos paradigmas Complutum11 1ndash28

Carrioacuten JS amp Saacutenchez-Goacutemez P (1992) Palynological data in sup-port of the survival of walnut (Juglans regia L) in the westernMediterranean area during last glacial times Journal of Biogeography19 623ndash630

Carrioacuten JS amp Scott L (1999) The challenge of pollen analysis inpalaeoenvironmental studies of hominid beds the record fromSterkfontein caves Journal of Human Evolution 36 401ndash408

Carrioacuten JS amp van Geel B (1999) Fine-resolution Upper Weichselianand Holocene palynological record from Navarreacutes (ValenciaSpain) and a discussion about factors of Mediterranean forestsuccession Review of Palaeobotany and Palynology 106 209ndash236

Comes HP amp Kadereit JW (1998) The effect of Quaternary cli-matic changes on plant distribution and evolution Trends in PlantScience 3 432ndash438

Cuenca A Pomery PJ amp Walker MJ (1986) Chronologicalaspects of the Middle Pleistocene in the coastal belt of southeasternSpain Quaternary climate in Western Mediterranean (ed byF Loacutepez-Vera) pp 353ndash363 Universidad Autoacutenoma Madrid

Davis OK (1990) Caves as sources of biotic remains in arid westernNorth America Palaeogeography Palaeoclimatology and Palae-oecology 76 331ndash348

Dupreacute M (1988) Palinologiacutea y paleoambiente Nuevos datosespantildeoles Referencias Valencia Diputacioacuten de Valencia Serviciode Investigacioacuten Prehistoacuterica Serie de Trabajos Varios no 84

Garciacutea Latorre J amp Garciacutea Latorre J (1996) Los bosques ignoradosde Almeriacutea Una interpretacioacuten histoacuterica y ecoloacutegica Historia y

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

Glacial refugia of thermophilous flora in Spain 127

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

less clear-cut Cementation of excavated breccia makes forslow progress in palaeontological research Fewer bird specieshave been identified than at Cueva Negra (13 species asagainst 66) (Walker et al in press) There are of coursecraggy mountainsides around the cave Below the site thecoastal plain surrounding Cabezo Gordo nowadays dry andopen must have contained mixed woodland and grasslandwith gallery woodland and swamps beside erstwhile streamsfeeding nearby wetlands and saltmarshes behind coastal sanddunes (where the Mar Menor coastal lagoon is today) eventhough marine regressions must have repeatedly drainedthese for many millennia each time Mammals include carni-vores such as panther lynx spotted hyaena brown bearfox and badger and herbivores such as hippopotamuselephantids wild horses and asses aurochs red deer wildgoats and abundant lagomorphs Tortoise is again commonin the deposits

The two Upper Pleistocene sites considered here from apalaeoecological standpoint of flora and fauna corroboratea proposal that long-term human presence may occur prefer-entially where several different biotopes coincide

ACKNOWLEDGMENTS

Michegravele Dupreacute assisted with laboratory processing KeithBennett and an anonymous referee provided useful sugges-tions on an earlier draft Research funding was provided bythe Spanish Ministerio de Educacioacuten y Ciencia and theMurcian regional authorityrsquos Fundacioacuten Seneca (projectsBOS2000-0149 PI-1700739FS01 PB92-0971 PB98-0405)

REFERENCES

Altuna J (1972) Fauna de mamiacuteferos de los yacimientos prehistoacutericosde Guipuacutezcoa Munibe 24 1ndash464

Arroyo J (1997) Plant diversity in the region of the Strait of Gibraltara multilevel approach Lagascalia 19 393ndash404

Badal E amp Carrioacuten Y (2001) Del glaciar al interglaciar Los paisajesvegetales a partir de los restos carbonizados hallados en las cuevasde Alicante De Neandertales a Cromantildeones El inicio del pob-lamiento humano en las tierras valencianas (ed by V Villaverde)pp 21ndash40 Universidad de Valencia Valencia

Bennett KD Tzedakis PC amp Willis KJ (1991) Quaternaryrefugia of north European trees Journal of Biogeography 18 103ndash115

Birks HJB (1993) Quaternary palaeoecology and vegetationscience-current contributions and possible future developmentsReview of Palaeobotany and Palynology 79 153ndash177

Blanco E Casado MA Costa-Tenorio M Escribano R Garciacutea-Antoacuten M Geacutenova M Goacutemez A Goacutemez F Moreno JCMorla C Regato P amp Sainz H (1997) Los bosques ibeacutericosUna interpretacioacuten geobotaacutenica Planeta Barcelona

Brewer S Cheddadi R de Beaulieu JL Reille M amp Data Con-tributors (2002) The spread of deciduous Quercus throughout

Europe since the last glacial period Forest Ecology and Management156 27ndash48

Burjachs F amp Juliagrave R (1994) Abrupt climatic changes during thelast glaciation based on pollen analysis of the Abric RomaniCatalonia Spain Quaternary Research 42 308ndash315

Burney DA amp Burney LP (1993) Modern pollen deposition in cavesites experimental results from New York State New Phytolology124 523ndash535

Carrioacuten JS (2002a) A taphonomic study of modern pollenassemblages from dung and surface sediments in arid environ-ments of Spain Review of Palaeobotany and Palynology 120217ndash232

Carrioacuten JS (2002b) Patterns and processes of Late Quaternaryenvironmental change in a montane region of southwestern EuropeQuaternary Science Reviews 21 2047ndash2066

Carrioacuten JS Dupreacute M Fumanal MP amp Montes R (1995) Apalaeoenvironmental study in semi-arid southeastern Spain thepalynological and sedimentological sequence at Perneras Cave(Lorca Murcia) Journal of Archaeological Science 22 355ndash367

Carrioacuten JS amp Munuera M (1997) Upper Pleistocene palaeoenvi-ronmental change in eastern Spain new pollen analytical datafrom Cova Beneito (Alicante) Palaeogeography Palaeoclimatologyand Palaeoecology 128 287ndash299

Carrioacuten JS Munuera M Navarro C Burjachs F Dupreacute M ampWalker MJ (1999a) The palaeoecological potential of pollenrecords in caves the case of Mediterranean Spain QuaternaryScience Reviews 18 1061ndash1073

Carrioacuten JS Munuera M Navarro C amp Saacuteez F (2000) Paleo-climas e historia de la vegetacioacuten cuaternaria en Espantildea a traveacutes delanaacutelisis poliacutenico Viejas falacias y nuevos paradigmas Complutum11 1ndash28

Carrioacuten JS amp Saacutenchez-Goacutemez P (1992) Palynological data in sup-port of the survival of walnut (Juglans regia L) in the westernMediterranean area during last glacial times Journal of Biogeography19 623ndash630

Carrioacuten JS amp Scott L (1999) The challenge of pollen analysis inpalaeoenvironmental studies of hominid beds the record fromSterkfontein caves Journal of Human Evolution 36 401ndash408

Carrioacuten JS amp van Geel B (1999) Fine-resolution Upper Weichselianand Holocene palynological record from Navarreacutes (ValenciaSpain) and a discussion about factors of Mediterranean forestsuccession Review of Palaeobotany and Palynology 106 209ndash236

Comes HP amp Kadereit JW (1998) The effect of Quaternary cli-matic changes on plant distribution and evolution Trends in PlantScience 3 432ndash438

Cuenca A Pomery PJ amp Walker MJ (1986) Chronologicalaspects of the Middle Pleistocene in the coastal belt of southeasternSpain Quaternary climate in Western Mediterranean (ed byF Loacutepez-Vera) pp 353ndash363 Universidad Autoacutenoma Madrid

Davis OK (1990) Caves as sources of biotic remains in arid westernNorth America Palaeogeography Palaeoclimatology and Palae-oecology 76 331ndash348

Dupreacute M (1988) Palinologiacutea y paleoambiente Nuevos datosespantildeoles Referencias Valencia Diputacioacuten de Valencia Serviciode Investigacioacuten Prehistoacuterica Serie de Trabajos Varios no 84

Garciacutea Latorre J amp Garciacutea Latorre J (1996) Los bosques ignoradosde Almeriacutea Una interpretacioacuten histoacuterica y ecoloacutegica Historia y

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

128 JS Carrioacuten et al

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

medio ambiente en el territorio almeriense (ed by A Saacutenchez-Picoacuten)pp 99ndash126 Universidad de Almeriacutea Servicio de PublicacionesAlmeriacutea

Garciacutea-Antoacuten M Morla C amp Sainz-Ollero H (1990) Considera-ciones sobre la presencia de algunos taacutexones relictos terciariosdurante el Cuaternario en la Peniacutensula Ibeacuterica Boletiacuten de la RealSociedad Espantildeola de Historia Natural (Seccioacuten de Biologiacutea) 8695ndash105

Garciacutea-Antoacuten M amp Sainz-Ollero H (1991) Pollen records from theMiddle Pleistocene Atapuerca site (Burgos Spain) Palaeogeogra-phy Palaeoclimatology and Palaeoecology 85 199ndash206

Gimeacutenez E (2000) Bases botaacutenico-ecoloacutegicas para la restauracioacutende la cubierta vegetal de la Sierra de Gaacutedor (Almeriacutea) PhD ThesisUniversidad de Almeriacutea

Herraacuten A Espinel S amp Goicoechea PG (1999) Utilizacioacuten delpolimorfismo del ADN de cloroplastos para definir regiones deprocedencia materna en los robles blancos de la PeniacutensulaIbeacuterica Investigacioacuten en Agronomiacutea y Recursos Forestales 8139ndash150

Huntley B (1990) Dissimilarity mapping between fossil and con-temporary pollen spectra in Europe for the past 13 000 yearsQuaternary Research 33 360ndash376

Jimeacutenez MP (2000) Genetic variability of Quercus suber (cork oak)studied by isozymes and chloroplast DNA Design of conservationstrategies PhD Thesis Universidad Politeacutecnica de Madrid

Leroy SAG Giralt S Francus P amp Seret G (1996) The high-sensitivity of the palynological record in the Vico Maar lacustrinesequence (Latium Italy) highlights the climatic gradient throughEurope for the last 90 ka Quaternary Science Reviews 15 189ndash201

Magri D amp Parra I (1997) Rifugi mediterranei di vegetazionearborea nel Tardo-Quaternario Atti del 4deg Colloquio su Approccimetodologici per la definizione dellrsquoambiente fisico e biologicomediterraneo pp 1ndash17 Castro Marina

Mota J Cabello J Cueto M Goacutemez F Gimeacutenez E amp Pentildeas J(1997) Datos sobre la vegetacioacuten del sureste de Almeriacutea (Desiertosde Tabernas Karst en Yesos de Sorbas y Cabo de Gata) Universidadde Almeriacutea Servicio Publicaciones Almeriacutea

Navarro C Carrioacuten JS Munuera M amp Prieto AR (2001) Cavesurface pollen and the palynological potential of karstic cavesediments in palaeoecology Review of Palaeobotany and Palynology117 245ndash265

Navarro C Carrioacuten JS Navarro J Munuera M amp Prieto AR(2000) An experimental approach to the palynology of cave depositsJournal of Quaternary Science 15 603ndash619

Ojeda F Marantildeoacuten T amp Arroyo J (1996) Patterns of ecologicalchorological and taxonomic diversity at both sides of the Strait ofGibraltar Journal of Vegetation Science 7 63ndash72

Olalde M Herraacuten A Espinel S amp Goicoechea P (2002) Whiteoaks phylogeography in the Iberian Peninsula Forest Ecology andManagement 156 89ndash102

Pantaleoacuten-Cano J Yll EI Peacuterez-Obiol R amp Roure JM (inpress) Palynological evidence for vegetational history in semi-aridareas of the western Mediterranean (Almeriacutea Spain) The Holocenein press

Peacuterez-Obiol R amp Juliagrave R (1994) Climatic change on the IberianPeninsula recorded in a 30 000-yr pollen record from lake BanyolesQuaternary Research 41 91ndash98

Pons A amp Reille M (1988) The Holocene and Upper Pleistocenepollen record from Padul (Granada Spain) a new study Palaeo-geography Palaeoclimatology and Palaeoecology 66 243ndash263

Ramil-Rego P Muntildeoz-Sobrino C Rodriacuteguez-Guitiaacuten M ampGoacutemez-Orellana L (1998a) Differences in the vegetation of theNorth Iberian Peninsula during the last 16 000 years Plant Ecology138 41ndash62

Ramil-Rego P Rodriacuteguez-Guitiaacuten amp MMuntildeoz-Sobrino C(1998b) Sclerophyllous vegetation dynamics in the north of theIberian Peninsula during the last 16000 years Global EcologyBiogeographical Letters 7 335ndash351

Salvador L Aliacutea R Aguacutendez D amp Gil L (2000) Genetic variationand migration pathways of maritime pine (Pinus pinaster Ait) inthe Iberian Peninsula Theoretical and Applied Genetics 100 89ndash95

Saacutenchez-Goacutemez P amp Alcaraz F (1993) Flora vegetacioacuten y paisajevegetal de las Sierras de Segura Orientales Instituto de EstudiosAlbacetenses Albacete

Saacutenchez-Goacutemez P Guerra J Coy E Hernaacutendez A Fernaacutendez Samp Carrillo AF (1998) Flora de Murcia Diego Mariacuten Murcia

Saacutenchez-Gontildei MF (1994) The identification of European upperpalaeolithic interstadials from cave sequences Aspects of archaeo-logical palynology methodology and applications (ed by OKDavis) AASP Contribution Series 29 161ndash182

Targarona J (1997) Climatic and oceanographic evolution ofthe Mediterranean Region over the last Glacial-Interglacialtransition A palynological approach LPP Contribution Series 7Utrecht

Turner C amp Hannon GE (1988) Vegetational evidence for lateQuaternary climatic changes in southwest Europe in relation to theinfluence of the North Atlantic Ocean Philosophical Transactionsof the Royal Society of London Series B 318 451ndash485

Uzquiano P (1992) The Late Glacial Postglacial transition in theCantabrian Cordillera (Asturias and Cantabria Spain) based oncharcoal analysis Palaios 7 540ndash547

Van Andel TH amp Tzedakis PC (1996) Palaeolithic landscapes ofEurope and environs 150 000ndash25 000 years ago an overviewQuaternary Science Reviews 15 481ndash500

Walker MJ amp Cuenca A (1977) Nuevas fechas C-14 para el sectorde Murcia y Alicante Trabajos Sobre Neogeno-Cuaternario 6309ndash317

Walker MJ Gibert J Rodriacuteguez Estrella T Eastham ACarrioacuten JS Yll E Legaz A Loacutepez A Loacutepez M amp Romero G(in press) Neanderthals and their landscapes aspects of researchat Sima de las Palomas del Cabezo Gordo and Cueva Negra delEstrecho del Riacuteo Quiacutepar in the context of middle palaeolithic andNeanderthal sites in the Segura drainage basin and adjacent areasof southeastern Spain Dynamics in the Middle Paleolithic andMiddle Stone Age Vol 2 (ed by N Conard) Tuumlbingen KernsVerlag lsquoTuumlbingen Studies in Prehistoryrsquo

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IEastham A Ribot F Arribas A Cuenca A Saacutenchez-CabezaJ-A Garciacutea-Orellana J Gibert L Albaladejo S amp Andreu JA(1998) Two SE Spanish middle palaeolithic remains Sima de lasPalomas del Cabezo Gordo and Cueva Negra del Estrecho del RiacuteoQuiacutepar (Murcia province) Internet archaeology issue 5 lthttpintarchacukjournalissue5walker_indexhtmlgt

Glacial refugia of thermophilous flora in Spain 129

copy 2003 Blackwell Publishing Ltd Global Ecology amp Biogeography 12 119ndash129

Walker MJ Gibert J Saacutenchez F Lombardi AV Serrano IGoacutemez A Eastham A Ribot F Arribas A Cuenca AAlbadalejo S amp Andreu JA (1999) Excavations at new sites ofearly man in Murcia Sima de las Palomas del Cabezo Gordo andCueva Negra del Estrecho del Riacuteo Quiacutepar de la EncarnacioacutenHuman Evolution 14 99ndash123

Willis KJ (1994) The vegetational history of the Balkans QuaternaryScience Reviews 13 769ndash788

Willis KJ Rudner E amp Suumlmegi P (2000) The full-glacial forests ofcentral and southeastern Europe Quaternary Research 53 203ndash213

Willis KJ amp Whittaker RJ (2000) The refugial debate Science287 1406ndash1407

BIOSKETCH

Joseacute S Carrioacuten teaches plant evolution in the University of Murcia His research has focused on Quaternary palaeoecology and climate in arid regions of Spain and South Africa studying the pollen cryptogam spores and other microfossils preserved in the sediments of lakes marshes and caves including coprolites and middens Current topics include the study of thresholds of long-term vegetation change and the location of glacial refugia of thermophilous flora in the Mediterranean