Holocene circum-Mediterranean vegetation …Quaternary International 200 (2009) 4–18 Holocene...

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Quaternary International 200 (2009) 4–18

Holocene circum-Mediterranean vegetation changes: Climate forcingand human impact

Guy Jaluta,�, Jean Jacques Dedoubata, Michel Fontugneb, Thierry Ottoa

aUniversite Paul Sabatier, EcoLab, UMR 5245 CNRS-UPS-INPT, 29 rue Jeanne Marvig, F-31055 Toulouse, Cedex 4, FrancebLaboratoire des Sciences du Climat et de l’Environnment (LSCE), UMR 1572 CNRS/CEA, Avenue de la Terrasse, F-91198 Gif sur Yvette, Cedex, France

Available online 29 March 2008

Abstract

The Mediterranean climate and its variability depend on global-scale climate patterns. Close correlations appear when comparing

Holocene palaeoenvironmental data (lake levels, fluvial activity, Mediterranean surface temperature and salinity, marine sedimentation)

with the main stages of the history of the circum-Mediterranean vegetation. They indicate an evolution of the Mediterranean biome

controlled by the climate and emphasize the teleconnections between the climate of the Mediterranean area and the global climatic

system. In the circum-Mediterranean area, the Holocene can be divided into three periods: a lower humid Holocene (11 500–7000 cal BP)

interrupted by dry episodes; a transition phase (7000–5500 cal BP) during which occurred a decrease in insolation as well as the

installation of the present atmosphere circulation in the northern hemisphere; and an upper Holocene (5500 cal BP—present)

characterized by an aridification process. Throughout the Holocene, humans used and modified more or less strongly the environment

but the climatic changes were the determining factors of the evolution of the Mediterranean biome. Societies had to adapt to natural

environmental variations, their impact on the environment increasing the ecological consequences of the global changes.

r 2008 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

Due to its extension, the complexity of its relief and itsclimatic diversity, the Mediterranean area is an exceptionalregion. In its eastern part, since about 10 000 years, humansbegan to modify the ecosystems. In the Western Mediter-ranean, this impact occurred later, since about 7500–7000years, but in both cases the vegetation cover earlyunderwent the human action.

Using pollen data, numerous palynologists and phyto-geographers have considered the present vegetation coveras a result of deforestation and agriculture, the Holoceneclimate being considered as more or less stable andMediterranean (Vernet, 1973; Planchais and Duzer, 1978;Triat-Laval, 1978; Pons, 1981; Planchais, 1982; Bernardand Reille, 1987; Reille and Pons, 1992; Pons and Quezel,1998; Quezel, 1999). More recently, despite the studiesdevoted both to the relationships between the vegetationhistory and the installation of the Mediterranean climate(Huntley and Prentice, 1988; Huntley et al., 1989; Huntley,

e front matter r 2008 Elsevier Ltd and INQUA. All rights re

aint.2008.03.012

ing author. Tel.: +335 61 53 60 38; fax: +33 5 62 26 99 99.

ess: [email protected] (G. Jalut).

1990a, b; Kelly and Huntley, 1991; Terral and Arnold-Simard, 1996; Jalut et al., 1997, 2000; Terral and Mengual,1999; Sadori and Narcisi, 2001) and between climaticchanges and societies (Weiss et al., 1993; Cullen anddeMenocal, 2000; Cullen et al., 2000, 2002; deMenocal,2001; Weiss and Bradley, 2001; Catto and Catto, 2004), theHolocene climatic changes continued to be considered ashypothetical (Quezel and Medail, 2003) or of limitedimpact, the respective importance of humans and climateon the environmental changes being subject to debate(Oldfield and Dearing, 2003; de Beaulieu et al., 2005).The goal of this study is to compare in the circum-

Mediterranean area, regional palaeoenvironmental eventsto global climatic and palaeoclimatic data to estimate therespective impact of the Holocene global climatic changesand societies on the vegetation dynamics.

1.1. The Mediterranean climate

The limits of the Mediterranean area are difficult todefine. Emberger (1943) considers the limits indicated bythe plant communities as questionable and recommends

served.

dx.doi.org/10.1016/j.quaint.2008.03.012

mailto:[email protected]

ARTICLE IN PRESSG. Jalut et al. / Quaternary International 200 (2009) 4–18 5

the use of climate criteria to limit the Mediterranean area.According to Koppen (1936), a climate is Mediterraneanwhen winter precipitation is three times higher than thesummer one. Criteria vary according to the authors. It isaccepted that the Mediterranean region is characterized bydrought during the warm period (Quezel and Medail,2003), temperature and precipitation depending on theconsidered regions.

To characterize the Mediterranean stages, the mean annualtemperature, the mean of temperature maxima of the warmestmonth and of the coldest month and the mean temperatureminima of the coldest month were used (Emberger, 1930;Quezel and Barbero, 1982; Rivas-Martınez, 1982, 1987). Theamount of precipitation defined the Mediterranean bioclimates(Emberger, 1930; Quezel and Barbero, 1982; Rivas-Martınez,1982, 1987) between 100mm and more than 2000mm/y.

The climatic diversity is due to the latitudinal andlongitudinal situation of the Mediterranean and also to theorographic characteristics of the surrounding regions andto the outline complexity that influences the marine andatmospheric circulation. The Mediterranean basin is alsosituated at the transition between the middle latitudessubmitted to the North Atlantic Oscillation (NAO) and thetropical zone. The role of NAO is significant for winterprecipitation in the Western Mediterranean but weaker tothe east and the south. For winter temperatures, its role isnot important in the Western Mediterranean and is minorin the eastern part. The southern part of the Mediterraneanregion is mostly influenced by the descending branch of theHadley cell (Lionello et al., 2006; Trigo et al., 2006).

Besides NAO, other modes play an important role: theEastern Atlantic pattern (EA), the Eastern Atlantic/WesternRussia pattern (EA/WR) and the Scandinavian pattern(SCAND). In winter, EA/WR is prominent on temperatureand precipitation (Traboulsi, 2004; Trigo et al., 2006). At amore global scale, the role of ENSO on the Mediterraneanwinter precipitation is emphasized (Pozo-Vasquez et al.,2001; Alpert et al., 2006; Lionello et al., 2006) as well as therelationships between the Mediterranean climate and theSouth Asian Monsoon (Alpert et al., 2006).

These examples show the present teleconnections be-tween the Mediterranean basin, the middle and highlatitudes and the tropical zone. They emphasize thenecessity to analyse the Holocene climatic changes of thearea in the light of the past global climate variability.

1.2. The Mediterranean vegetation cover

Climate diversity, relief, geology and soils explain thecomplexity of the present Mediterranean vegetation sub-mitted to a strong human impact. In this study, we shallconsider only the vegetation of low and middle altitudebecause it is in this altitudinal zone that most of thepeatbogs and lakes are situated, which provided thepalaeoenvironmental data considered here.

Under the present climate conditions characterized bydrought during the hot season, sclerophyllous and ever-

green trees and shrubs dominate at low and middleelevations. In the western and central parts of the basin,Quercus ilex, Quercus suber, Quercus coccifera, Ceratonia

siliqua, Pistacia lentiscus and Olea europaea var. sylvestris

are some of the most representative species. In the easternpart, Q. suber is absent but Q. coccifera subsp. calliprinos iswell represented (Quezel and Medail, 2003).Gymnosperms are also abundant, especially Pinus

halepensis, Pinus pinaster, Pinus pinea as well as Cupressa-ceae (Juniperus and Tetraclinis). The last one is wellrepresented in northwestern Africa but rare in SouthernSpain and Cyprus.At low altitudes, deciduous broad-leaf trees develop on

soils sufficiently deep and wet to limit the effects of thesummer drought. At middle altitude and in regions whereprecipitation is high (sub-humid and humid bioclimates),deciduous Mediterranean forests with Quercus faginea, Q.

infectoria, Q. cerris, Q. ithaburensis and sub-Mediterraneanforests with Quercus pubescens, Q. fraineto and Q. trojana

are present. When exposure and soil are favourable,sclerophyllous and evergreen, deciduous broad-leaf treesand Gymnosperms can grow together. However, at lowelevation and under Mediterranean climate, when Gym-nosperms are poorly represented, sclerophyllous and ever-green trees and shrubs dominate.Beyond the Mediterranean region, at low and middle

latitudes, deciduous broad-leaf trees dominate (oaks, elms,hornbeams, lime and hazel). Contacts with the sclerophyl-lous evergreen species in the transition zones mainlydepend on exposure and soil characteristics.

1.3. Holocene vegetation changes in the Mediterranean

region

During the last decades, along the French and Catalancoast, numerous pollen analyses of continental depositsand charcoal studies from archaeological investigationswere carried out. They have shown that during the firstpart of the Holocene, deciduous broad-leaf trees werefrequently dominant, sclerophyllous and evergreen treesbeing present but more rare. During this period the climatewas considered as Mediterranean, close to that of thepresent meso-Mediterranean or supra-Mediterranean sub-humid or humid stages (Quezel and Barbero, 1982). Later,the development of the sclerophyllous and evergreenforests was interpreted as a consequence of the humanimpact, the role of the climatic changes being considered asminor or ignored (Pons and Quezel, 1998). Recent resultsallow one to reconsider these interpretations.

2. Regional examples

2.1. Villaverde (381480N–21220W, 900 m a.s.l., Spain)

Villaverde is situated in the Cubillo valley, 57 km to theSW of Albacete (Carrion et al., 2001; Carrion, 2003; Jalut,2005) (Fig. 1), in the meso-Mediterranean semi-arid stage

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(Rivas-Martınez, 1987). The pollen sequence concerns thelast ca 9700 years. Between bottom core and 7500 cal BP,during a dry period, pine dominated. Juniperus isaccompanied by some various Mediterranean and xero-phytic taxa and sparse grains of deciduous oaks, hazel andbirch. Since 7500 cal BP Pinus declines while evergreen oaks(probably Quercus cf rotundifolia) and Mediterraneandeciduous oaks (probably Quercus cf faginea) as well asmesophilous deciduous trees (Betula, Fraxinus, Corylus)expanded. Between about 6000 and 5300 cal BP, deciduoustaxa were dominant while Pinus continues to regress.Subsequently, an aridification phase began. Deciduousbroad-leaf trees regressed while sclerophyllous and ever-green species expanded, especially the Quercus cf rotundi-

folia forest. Less resistant to the increasing drought,Corylus, Alnus and Fraxinus disappeared around3600 cal BP. Among the evergreen sclerophyllous andthermophilous species, Pinus and Juniperus were welldeveloped. Until the Iron Age, the role of the climaticchanges on vegetation was prominent and the humanimpact was limited. Later, the effects of human impactadded to the effects of the aridification consequences.

2.2. Capestang (431200N–31020E, 1 m a.s.l., France)

To the NW of the Gulf of Lions, this former juxta-littoral pond is situated at the eastern extremity of aNW–SE Atlantic–Mediterranean corridor (Fig. 1), in thesub-humid meso-Mediterranean stage. The pollen sequenceconcerns the last 12 000 years (Jalut, 1995; Jalut et al.,1997, 2000). Additional pollen analysis confirmed andimproved the previous results (Jalut, 2005) (Figs. 2 and 3).

After a phase dominated by Pinus, a deciduous forestextended around 10 200 cal BP (Fig. 2). Corylus values aremore abundant than Quercus until 8500 cal BP. Thesclerophyllous and evergreen oaks (Q. ilex type includingalso Q. coccifera) as well as thermophilous and heliophi-lous taxa were present but not abundant. Ulmus and Alnus

were also present. The maximum extent of the deciduous

10 0 10

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32

N

Fig. 1. Dashed line: Limits of the Mediterranean region (Quezel and Medail, 2

4—Salinas; 5—Navarres; 6—Algendar; 7—Capestang; 8—Corchia Cave; 9—

Vico; 13—Lago Fucino; 14—Lago di Pergusa; 15—Edessa; 16—Bannock bas

Soreq Cave; 22—Hula; 23—Ghab.

oak forest occurred around 7500 cal BP. A mixed oakforest with Fraxinus, Tilia and Alnus, also including someAbies and Fagus, dominated until ca 4700 cal BP. Then,two different forests developed simultaneously: a beechforest and a Mediterranean evergreen oak forest.Nowadays, along altitudinal transects (southern slopes

of Cevennes and Massif of Alberes in Languedoc andRoussillon, Massif of Montseny in Catalunya) or infavourable topographic locations along rivers in narrowvalleys of the meso-Mediterranean area, both Fagus

sylvatica and Q. ilex, species with different ecologicalrequirements, can be observed. They are close but colonizeareas with differentiated ecological conditions, the presenceof beech being favoured or depending on atmospherichumidity.Around 3000–2800 cal BP, Fagus and the deciduous oak

forest decrease and the evergreen thermophilous orMediterranean taxa become dominant. Between the begin-ning of the Holocene and 3000 cal BP, the anthropogenicindicators (Rumex, Plantago, Cerealia, Secale) are poorlyrepresented. The regular presence of the Cerealia pollentype began around 7000 cal BP, but during the extensionphase of Fagus and Q. ilex type, the Cerealia pollen andPlantago, especially Plantago lanceolata, are less abundantand human impact does not seem responsible for thevegetation changes. It became important after 3000 cal BP.Concerning the Cerealia and Plantago lanceolata types, itmust be remembered that in the considered region,confusions are possible with the pollen of wild plants suchas Hordeum, Agropyrum and Glyceria as well as Plantago

argentea and Plantago lagopus (Beug, 2004).

2.3. Lago di Pergusa (371310N–141180E, 667 m a.s.l., Italy)

The lake is situated in Sicily near the town of Enna, inthe sub-humid meso-Mediterranean stage (Sadori andNarcisi, 2001). The progressive installation of a dominantdeciduous oak forest begins around 12 300 cal BP. Ever-green oaks are present but are more rare. Ulmus and

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003). Location of the cited sites: 1—Tigalmamine; 2—Siles; 3—Villaverde;

Lago dell’Accesa; 10—Lago di Mezzano; 11—Lagaccione; 12—Lago di

in; 17—Lerna; 18—Golhisar Golu; 19—Beysehir; 20—Eski Acigol; 21—

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erealia

Rum

ex

Ericaceae

Caryophyllaceae

Cistus

Phillyrea

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lmus

Quercus ilex t.

CAPESTANG (Hérault) Alt. 1.m., simplified pollen diagram1

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Calluna

Pinus

Betula

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biesFagus

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Analyst G. Jalut

Fig. 2. Simplified pollen diagram of Capestang (Herault, France). Only selected taxa are represented. The calibrated ages were calculated using the Calib

Radiocarbon Calibration Program Calib Rev 5.0.1 (Stuiver and Reimer, 1993). The mid-point (2s probability interval) was selected for these dates.

G. Jalut et al. / Quaternary International 200 (2009) 4–18 7

Corylus are also present as well as Fagus, the last oneobserved with low percentages all along the sequence.

Between about 10 000 and 4500 cal BP, a deciduous oakforest was dominant. However, around 8100–8000 cal BP,the values of Olea increased regularly while in the sametime sclerophyllous oaks (Q. suber/cerris), deciduous oaks,Corylus and Fagus decreased. Ulmus continued to develop.The changes recorded between 8000 and 7600 cal BPindicate an increasing aridity. Two successive decreases inthe pollen concentration of trees and herbs (8100 cal BPand around 7600 cal BP) are correlated with a decreasinghumidity. Their is no evidence to say whether the Olea

extension recorded around 8000 cal BP is due to a climaticchange and to a beginning of cultivation as described inEastern Spain ca 7500–6500 cal BP (Terral and Arnold-Simard, 1996).

After this arid episode, the percentages of the deciduousoaks as well as the tree pollen concentration decreasedbetween 6800 and 5700 cal BP. A last maximum of the treepollen concentration is recorded before 5500 cal BP fol-lowed by a quick decrease. The forest cover became lessdense and between 4500 and 3000 cal BP a change in theevergreen/deciduous trees and shrubs ratio is recorded.Evergreen trees became dominant from ca 3500 to2500 cal BP. Since 3600 cal BP, the increasing values of

Olea suggest a development of its cultivation favoured by adrier climate.

2.4. Lagaccione (421330N–111540E, 355 m a.s.l., Italy)

As described at lago di Pergusa, the pollen data (Magri,1999) (Fig. 1) show the development of a deciduous oakforest (Quercus robur type) with Corylus since the begin-ning of the Holocene. Between 9000 and 8000 cal BP, thespread of Fagus, poorly but regularly observed since theLateglacial, suggests, in some places, favourable humidatmospheric conditions. At the same time, evergreen oaks(Q. ilex type) expand but is less abundant than thedeciduous broad-leaf trees. Around 7900 cal BP, similarto what happened at lago di Pergusa between 8100 and7600 cal BP (Sadori and Narcisi, 2001), a bipartite decreasein trees and total pollen concentration is recorded. Later,around 4200–4100 cal BP, a second strong decrease inpollen concentration is recorded. This episode is alsoobserved in other sites of central Italy (Magri, 1999).Archaeological, palaeohydrological and sedimentologicaldata indicate that it corresponded to a clear hydrologicalchange. Lake levels were lower than the present (Giraudi,1998, 2004). The pollen data indicate a development ofhuman activity.

ARTICLE IN PRESS

Mediterranean Not MediterraneanAges BP Ages Cal BP

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2190±80

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2172

Dec. max. / Scl. min. * estimated ageDec. min. / Scl. max.

Leve

ls

Fig. 3. Pollen ratio of deciduous broad-leaf (Dec.) versus evergreen

sclerophyllous taxa (Scl.) at Capestang. The two curves represent the

minimum (min.) (solid line) and maximum (max.) (dashed line) values of

the pollen ratios. Pollen ratios are plotted logarithmically for each sample.

The vertical line represents the threshold value 0.5 obtained from the study

of the pollen content of surface samples along two transects (Bordeaux-

Narbonne-Tabernas and Peyrehorade-Narbonne (Jalut et al., 1997, 2000).

* Estimated ages from the age–depth model.

G. Jalut et al. / Quaternary International 200 (2009) 4–188

2.5. Lakes Lerna (371300N–221350E, 4 m a.s.l., Greece) and

Edessa (401480N–221030E, 300 m a.s.l., Greece)

The pollen data from lakes Lerna (Coastal plain of Gulfof Argos, Peloponnese; Jahns, 1993) and Edessa (Bottema,1974) (Fig. 1) show the dominance of the deciduous trees atthe beginning of the Holocene. Bottema (1974) considersthat Q. pubescens and higher in altitude Quercus cerris andQ. conferta were probably the most common species at thattime near Edessa. Q. pubescens and Q. frainetto could bepresent in Peloponnese (Jahns, 1993). Fraxinus, Ulmus,Tilia, Corylus and Carpinus orientalis/Ostrya were alsopresent as well as thermophilous and evergreen sclerophyl-lous trees such as Q. ilex-coccifera, Phillyrea and Olea. InPeloponnese, around 5500 cal BP, a decrease in percentagesand pollen concentrations of the deciduous oaks isrecorded. The curve of Olea becomes continuous. Slightlyafter 4000 cal BP a second decrease in the deciduous oakforest is recorded. It is contemporaneous with a strongincrease in Olea percentages probably linked to cultivation.

At Edessa, the decline of the deciduous oak forest occursafter 3850 cal BP.

2.6. Eski Acigol (381330N–341320E, 1270 m a.s.l., Turkey)

Situated in Central Anatolia in the cold arid supra-Mediterranean area (Fig. 1), the site records a periodlonger than 16 000 years (Roberts et al., 2001). From theavailable radiocarbon dates, it can be assumed thatbetween 12 000 and 6000 cal BP an open formation withdeciduous oaks, Corylus, Juniperus and Pistacia wasdeveloped. Between 6800 and 6000 cal BP the lake leveldecreased and Pistacia declined as well as Corylus andUlmus some centuries later. Herbaceous and steppicchamaephytic taxa (Artemisia and Chenopodiaceae) devel-oped at the expense of Poaceae. Deciduous oaks are wellrepresented until 4500–4000 cal BP. Anthropogenic indica-tors (Rosaceae, Compositae, Plantago, Polygonum avicu-

lare and Urtica) then appear and anthropogenous steppecommunities develop.Before this last period, the synchronism between the

vegetation changes and the lake-level decreases suggeststhat the aridification induced the changes in the vegetationcover. After the initial wet part of the Holocene, a dryerphase began around 6500 cal BP. Around4500–4000 cal BP, the human impact increased the effectsof the aridification on the vegetation.

2.7. Valley of Ghab (351390N–361150E, 240 m a.s.l., Syria)

and lake Hula (331030N–351370E, 70 m a.s.l., Israel)

In NW Syria, in the Ghab valley (Fig. 1), the beginningof the Holocene is characterized by the presence of Pinus,the dominance of the evergreen oaks and Olea and thesignificant presence of deciduous oaks until ca 5600 cal BP.Then the percentages of Olea increased (Yasuda et al.,2000).To the south, at lake Hula (Baruch and Bottema, 1999)

(Fig. 1) deciduous Quercus dominate before the beginningof the Holocene. They declined at the Lateglacial–Holo-cene transition but were abundant during a great part ofthe Holocene. They regress strongly only around3900–3800 cal BP. Evergreen Quercus (Q. calliprinos type)are present before 17 000 cal BP. Their percentages increaseat ca 7300–7200 cal BP and during the historical timearound AD 650.

3. Discussion

Because of the W–E distribution of the cited examples(Fig. 1) and the complexity of the climatic teleconnec-tions (Lionello et al., 2006), the use of various proxies isnecessary to highlight, correlate and understand therelationships between Mediterranean palaeoenvironmentalevents (sometimes differing or opposite) and major globalpalaeoenvironmental events. For this reason we com-pared Mediterranean data from palynology, hydrology,

ARTICLE IN PRESS

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concentrationdecreases

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p qf g h i j k m n o

Aridification phases

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a b c d e

Events in the Mediterranean area

Dryer Wetter

r s t

Fig. 4. Correlation between Holocene climate, hydrological and pollen data concerning the Mediterranean basin. (a) Aridification phases in the Western

Mediterranean (Jalut et al., 1997, 2000). (b) Dry episode in the Eastern Mediterranean (Schilman et al., 2001); (c) Installation of the Mediterranean climate

in the Northwestern Mediterranean basin (Terral and Mengual, 1999); (d) Aridification process in the Eastern Mediterranean (Schilman et al., 2001); (e)

Phases of decreasing fluvial activity in the Western Mediterranean (Magny et al., 2002); (f) Low lake levels at lake Fucino (Giraudi, 1998, 2004); (g) Major

high lake levels at lago dell’Accesa (Magny et al., 2007; (h) Low lake levels at lago dell’Accesa (Drescher-Schneider et al., 2007); (i) Lake-level decreases in

central Italy (Magri, 1997); (j) Lake-level decreases in the Mediterranean region (Magny et al., 2002); (k) Lake-level decrease at Eski Acigol (Turkey)

(Roberts et al., 2001); (l) Sapropel S1 (Emeis et al., 2000); (m) Increasing precipitation in the Eastern Mediterranean (Soreq cave) (Bar-Matthews et al.,

2000); (n) Frequency of lacustrine and paludal Holocene deposits in Sahara (Petit-Maire and Guo, 1996); (o, p) Decrease in pollen concentration in Sicily

and Balearic Islands (o) (Sadori and Narcisi, 2001; Perez-Obiol and Sadori, 2007), in Central Italy at Lagaccione and lago di Mezzano (p) (Magri, 1999;

Sadori et al., 2004); (q) Major climatic events in Africa (Gasse, 2000); (r) Dry episodes in the domains of African and Indian monsoon (Gasse and Van

Campo, 1994; Gasse, 2000; Morrill et al., 2003); (s) Cooling phases in the sub-tropical Atlantic (deMenocal et al., 2000a); (t) Abrupt Holocene climatic

changes (Mayewski et al., 2004).

G. Jalut et al. / Quaternary International 200 (2009) 4–18 9

oceanography, speleothems and marine sedimentology tomore global phenomenons.

In the Western Mediterranean, the pollen ratio decid-uous broad-leaf/ evergreen sclerophyllous trees (Jalut et al.,1997, 2000, 2002; Jalut, 2005) indicates dry phases duringthe intervals 10 900–9700; 8400–7600; 5300–4200;4300–3400; 2850–1730 and 1300–750 cal BP (Figs. 3 and4a). Some of them are correlated with phases of decreasingfluvial activity described by Magny (1999) and Magny et al.(2002) in the Western Mediterranean around 11 500,10 500, 9000, 7000, 4000, 3000, 2000 and 800 cal BP(Fig. 4e). The succession of these phases agrees with thelake-level fluctuations at lago dell’Accesa (Tuscany, Italy)(Magny et al., 2007) where three noticeable lake-levellowerings are recorded at ca 8600–7900, 4600–4300 and3700–2800 (Drescher-Schneider et al., 2007) (Fig. 4h).These last are well correlated with the dry phases definedusing the pollen ratio (Fig. 4a). The interpretation of theevent at 11 500 cal BP is more difficult. It is considered as aphase of decreasing fluvial activity at the scale of the

western Mediterranean (Magny et al., 2002) and as a phaseof major high level when considering the lake-levelfluctuations at lago dell’Accesa (Magny et al., 2007)(Fig. 4g).The upper limit of the dry interval 8400–7600 cal BP, the

last of the humid Holocene phase, also corresponds to amajor oceanographic circulation change recorded in thewestern Mediterranean at 7700 cal BP (Jimenez-Espejoet al., 2007). Around 7700–7600 cal BP, a decrease in treepollen percentages and concentration is recorded in centralItaly, at Lagaccione (Magri, 1999) (Fig. 4p) as well as apollen concentration minimum at lago di Vico (Magri andParra, 2002). The dry phase 4300–3400 cal BP is contem-poraneous both with a decrease in pollen concentrationrecorded in central Italy at Lagaccione (Magri, 1999)(Fig. 4p) and with the hydrological changes at lago Fucinoand in other lakes in central Italy at 4200–4100 cal BP(Fig. 4f) (Giraudi, 1998, 2004). Three phases identifiedby Magny et al. (2002) at 11 500, 9000 and 7000 cal BP(Fig. 4e) were not found using the pollen ratio in the


palynological records. This difference as well as theopposite record of the phase at 11 500 cal BP confirm theobservations by Magny et al. (2003) and Jimenez-Espejoet al. (2007) who emphasize that different, opposite orlacking responses to a same climate forcing mechanism canbe observed in the Mediterranean area. Differences andsimilarities might help to characterize the limits of the pastclimatic domains.

In addition to characterization of the Holocene dryepisodes, the studies based on the pollen ratio also showthat the Mediterranean climate became established follow-ing a south–north gradient (Jalut et al., 1997, 2000; Jalut,2005).

In the region of the Gulf of Lions, pollen andhydrological data show a bipartition of the Holocene: alower part characterized by a wet climate (Jalut et al.,2005), and an upper part that was drier. These two periodswere not homogeneous. In the Western Mediterranean, thepollen data show, during the humid part of the Holocene,the existence of two dry episodes (10 900–9700 and8400–7600 cal BP) (Fig. 4a), which can be correlated withthe arid phases 1 and 2 of Tigalmamine (Morocco, Lambet al., 1995), respectively. The first corresponded to a phaseof decreasing fluvial activity in the Western Mediterranean(Magny et al., 2002) (Fig. 4e). The second, recorded atSalinas (Jalut et al., 1997, 2000), is well correlated with thedry episode described around 8500–8100 cal BP at Siles(Sierra de Segura, S-E Spain) by Carrion (2002, 2003) andwith the cooling period recorded at lago di Mezzano and inmany other regional archives between 8200 and 7800 cal BP(Ramrath et al., 2000). The dry event recorded near8100 cal BP in central Italy, at lago di Vico (Magri andParra, 2002), might also be a local consequence of thisregional arid episode not clearly recorded in the spe-leothems of the Corchia cave (Zanchetta et al., 2007).Between these two dry episodes, the palynological dataindicate wetter conditions prevailing in the westernMediterranean. To the east, in Crete, they show oppositeresults, the period 9460–8395 cal BP corresponding to thedriest conditions of the Holocene (Bottema and Sarpaki,2003). The study of the fluvial activity in the westernMediterranean also shows a decrease around 9000 cal BP(Magny et al., 2002) (Fig. 4e), and at lago dell’Accesa(Tuscany, Italy), a maximal lake-level lowstand is alsorecorded at ca 9200–7700 cal BP (Magny et al., 2007). Thislast, contrary to the maximum lake-level highstandobserved at ca 9000–8200 cal BP at lago di Perguza (Sicily)(Sadori and Narcisi, 2001), illustrates again the possibilityof opposite responses to a same climate forcing mechanism.

At Capestang, during the second part of the Holocene,the extension of the evergreen sclerophyllous trees around5000–4500 cal BP (Fig. 3) can be correlated with the phaseof decreasing lake levels in the Western Mediterranean andNorthern Africa between 5000–4000 and 5500–4500,respectively (Magny et al., 2002). The beginning ofthe dry phase 5300–4200 cal BP (Jalut et al., 1997,2000) corresponds to the end of the humid episode

5600–5300 cal BP described in the Alps (Magny and Haas,2004; Magny et al., 2006). The following dry phase(4300–3400 cal BP) was marked by climatic fluctuations(Fig. 3). These two dry episodes (5300–4200 and4300–3400 cal BP) can be correlated with lake loweringssuggested at Siles, respectively, between 5400–4800 and4300–3800 cal BP, with peaks at ca 5200 and 4100 cal BP,by Pseudoschizaea (Carrion, 2002).Around 3000–2900 cal BP, the strengthening of the

Mediterranean climate conditions (Fig. 3) agrees with theresults by Terral and Mengual (1999) concerning olivecharcoals, who suggest the installation of the Mediterra-nean climate in the Western Mediterranean, between 39and 441N, around 3000 cal BP (Fig. 4c).With respect to the NW of the Mediterranean basin, SE

Spain was characterized by a Mediterranean climate sincethe beginning of the Holocene (Pons and Reille, 1988;Esteban Amat, 1995; Jalut et al., 1997, 2000, 2005;Pantaleon-Cano, 1998, 2003; Jalut, 2005). As emphasizedby Magny et al. (2002), due to the general atmosphericcirculation, the pollen ratio at Salinas (Jalut et al., 1997,2000) (Fig. 1) agrees with the evolution of the lake levels inNorthern Africa while to the north, the data fromCapestang show a better correlation with the WesternMediterranean.The hypothesis of a climatic link between southern Spain

and Northern Africa is strengthened by data concerningthe transport of pollen of Cedrus from North Africa to thenorthern bank of the Mediterranean (Magri and Parra,2002). This would explain the existence of a Mediterraneanvegetation cover since the beginning of the Holocene insouthern Spain (Jalut et al., 1997, 2000).The data from Sicily, Greece and NW Syria also show a

bipartite Holocene. At Eski Acigol, according to thechronology, a lake-level decrease between 6800 and6000 cal BP (Fig. 4k) indicates the beginning of thearidification (Fontugne et al., 1999; Kuzucuoglu et al.,1999). It can be correlated with the decrease in tree pollenconcentration found at lago di Pergusa (Sicily) (Fig. 4o). InBulgaria, in the mountain (Bozilova and Tonkov, 2000)and in the coastal area of the Black Sea (Bozilova andBeug, 1992), the period 6700–6600 cal BP also correspondsto significant vegetational changes.A strong decrease in tree pollen concentration is

recorded near 5500 cal BP at lago di Pergusa as well as aretreat of the deciduous oaks around 5500 and 5600 cal BPin Peloponnese and 5600 cal BP in NW Turkey. Acorrelation of these events with the aridification phase5300–4200 cal BP of the Western Mediterranean (Jalut etal., 1997, 2000; Jalut, 2005) (Fig. 4a) can be proposed.Later, the extension of Olea in Sicily around 3600 cal BP,then of the evergreen sclerophyllous taxa from 3000 to2500 cal BP onwards, agree with the period of installationof the Mediterranean climate around 3000 cal BP (Terraland Mengual, 1999).What was the impact of humans during these different

periods?


At Capestang, during the extension of the evergreen oakforest since 4500 cal BP, the anthropogenic indicatorsregress (Jalut, 1995) (Fig. 2). In Spain at Navarres(Fig. 1) (Carrion and Dupre, 1996), the sclerophyllousspecies develop before the anthropogenic indicators. Theirlow representation since 5700 cal BP indicates a reducedhuman activity, which poorly affected the environment. Inthe Balearic Islands (Minorca, Algendar: Yll et al., 1997)(Fig. 1), the spread of Olea and other sclerophyllous taxaoccurrs around 5700 cal BP, before the increase in anthro-pogenic indicators and the extension of the disturbed areasdated around 4500 cal BP. At Majorca, the fall in Buxus

and Corylus percentages between 6800 and 4500 cal BP ismore related to a decrease in precipitation than to a humanimpact, this last having accelerated the vegetation changes(Perez-Obiol et al., 1996; Perez-Obiol and Sadori, 2007); inItaly, at lago di Mezzano (Fig. 1) (Sadori et al., 2004), theabrupt fall of the pollen concentrations around 3800 cal BP(Fig. 4p) is prior to the extension of the human impactdated in two periods, around 3600 and 3200 cal BP.

Around 5900 cal BP, in southeast Bulgaria, along theBlack Sea coast (Bozilova and Beug, 1992) anthropogenicindicators are rare and the archaeological data indicatesparse human settlements only until ca 4700 cal BP. To theeast, in Anatolia, at Eski Acigol, the changes in vegetationcover observed between 6800 and 6000 cal BP are also olderthan the first strong anthropogenic disturbances recordedca 4500–4000 cal BP. Similar data are observed to thesouth-west at Golhisar Golu (Eastwood et al., 1999) whereevidence for human impact is not convincing during thefirst half of the Holocene, despite the regional abundanceof archaeological and archaeobotanical data since theNeolithic. Pollen evidences of the agricultural practicesappear only from ca 3200 cal BP onwards, during theBeysehir Occupation phase (Eastwood et al., 1998, 1999),despite the early agriculture developed in NW Syria around10 000 cal BP (Yasuda et al., 2000) and more generally inthe Fertile Crescent. These examples show that Holoceneclimatic changes induced vegetation modifications in thecircum-Mediterranean area before the first significantanthropogenic impacts.

Additional data on speleothems, marine sedimentation,salinity and sea surface temperature of the Mediterraneanemphasize the prominent part of the climate on theenvironmental changes and the links with the globalchanges.

In western Africa, from the Sahelian zone to the north ofSahara, a humid period is recorded between 11 500 and5500 cal BP (Petit-Maire and Guo, 1996) (Fig. 4n). It wasinterrupted by dry episodes of various durations (Gasse,2000). During this phase, between ca 9500 and 7000 cal BP(Fontugne et al., 1994) sapropels (S1) were deposited in theEastern Mediterranean (Fig. 4l). To the W, in the anoxicbasin of Bannock (south eastern Ionian Sea) (Fig. 1), coreBAN 8409 GC (Cheddadi et al., 1991) shows two parts(S1a and S1b), which indicate a climatic change around9000–8900 cal BP. The pollen analysis of the deposits shows

a climatic optimum between 10 000 and 8000 cal BP,characterized by the spread of the oak forests favouredby abundant moisture. The maximum of Quercus isrecorded around 8000 cal BP and the greatest summermoisture ca 7000 cal BP.These results are confirmed by recent studies concerning

the Corchia cave (Central Italy) (Zanchetta et al., 2007)(Fig. 1). They indicate, in the Western Mediterranean,increasing precipitation during deposition of Sapropel S1and a maximum rainfall between 8900 and 7300 cal BP.From comparisons with continental data (Greece, Botte-ma, 1974; Dalmatia, Beug, 1967, 1975), Cheddadi et al.(1991) consider that during the oak maximum, deciduousoaks were dominant. Evergreen oaks prevailed after7000 cal BP during a drying phase. These interpretationsagree with recent pollen data (Jalut et al., 2005). Theincrease in humidity indicated by the spread of oaks iscorrelated, in core BAN 8409 GC, with sapropels (S1),and the maximum summer rainfall suggests a continentalclimate.In comparison with these data, the study of speleothems

of Soreq cave (Fig. 1) (Ayalon et al., 1999; Bar-Matthewset al., 2000) shows an increase in precipitation during theperiod 8500–7000 cal BP (Fig. 4j). Bar-Matthews et al.(2000) define the phase of sapropels S1 as a humid periodwith frostless winters and wet summers without a dryperiod. This conclusion does not agree with the existence ofa Mediterranean climate in the considered region duringthis period, but supports the pollen and palaeoclimaticdata from Bannock basin. According to Neumann et al.(2007), full Mediterranean climatic conditions prevailed inthe near east during the last 6500 years, which implies aregional climatic change around 7000–6000 cal BP. Duringthe previous humid period, a massive input of fresh waterdue to the high precipitation explains why the salinity ofthe surface water of the Mediterranean Sea and the RedSea was lower than the present (Cheddadi and Rossignol-Strick, 1995; Kallel et al., 1997; Bar-Matthews et al., 2000;Emeis et al., 2000; Paul et al., 2001; Arz et al., 2003;Principato et al., 2003).During this first part of the Holocene (10 000–

8000 cal BP), which corresponded to the Climatic Opti-mum, the global mean temperature was estimated to be2 1C higher than the present and the precipitation washigher as well (COHMAP Members, 1988; Petit-Maireet al., 2000). In SE Spain and Maghreb, pollen data showthat a summer drought could have existed, in the otherperi-Mediterranean regions, they indicate either a no-Mediterranean climate or a sub-Mediterranean/supra-Mediterranean bioclimate favourable for deciduousbroad-leaf trees and shrubs (Jalut et al., 2005).In the northern hemisphere, around 6600 cal BP, the

present atmospheric circulation mode was established(Schulz and Paul, 2002). Between 7000 and 6500 cal BP,the increase in sea surface temperature in the Ionian seaand in the eastern Mediterranean, to the SE of Cyprus(Emeis et al., 2000), indicates a rise of the mean annual


atmospheric temperature. This is synchronous with thehydrological changes of Eski Acigol (Fontugne et al., 1999;Kuzucuoglu et al., 1999) and with the decrease in the pollenconcentration in Sicily and in the Balearic Islands around6800–6000 cal BP (Yll et al., 1997; Perez-Obiol and Sadori,2007) (Fig. 4o). This climate change was followed by ageneral decrease in lake level in the Mediterranean region(Fig. 4j) and a strong modification of the humansettlements in Sahara (Vernet and Faure, 2000; Magnyet al., 2002). The drying of the lakes of the Western andEastern Great Ergs increases ca 4500 cal BP (Petit-Maireet al., 1991; Callot and Fontugne, 1992; Petit-Maire andGuo, 1996). It is during this period that populationspoured into the valleys. Sumerian, Egyptian and Induscivilizations appeared, their main preoccupation being tomanage irrigation and water reserves. The occurrences, theimportance and the synchronism of these events suggest aglobal climatic control of these changes.

Studies in the sub-tropical Atlantic (deMenocal et al.,2000a) show moderate cooling phases correlated withperiods of climatic instability in Greenland and sub-polarNorth Atlantic occurring with a cyclicity of around 15007500 years (Fig. 4s).

A comparable relationship is also demonstrated betweenthe climatic changes in the Mediterranean region, thethermohaline circulation and the atmospheric circulationin the Northern Atlantic (Bond et al., 1997; Cullen anddeMenocal, 2000; Rohling et al., 2002; Kim et al., 2004;Lamy et al., 2006: Ziv et al., 2006). The rhythm of theHolocene climatic changes (Bond et al., 1997; Magny,2004; Mayewski et al., 2004; Schulz et al., 2004) (Fig. 4t)also emphasizes the role of the changes in insolation (vanGeel et al., 1996; Kallel et al., 2000; Bond et al., 2001;Haigh, 2001; Magny, 2004; Lamy et al., 2006). Its decreasearound 5500 cal BP corresponded to the acceleration of thearidification phase observed since 8000–7000 cal BP inNorthern Africa (Haynes et al., 1989; Lezine, 1989; Lezineand Casanova, 1989; Petit-Maire and Guo, 1996) and alsorecorded in the Mediterranean marine cores ca 8500 cal BP.However, the solar forcing was not the unique cause of therapid and wide changes more probably linked to theinteractions between the modifications in the insolation,the vegetation cover and the oceanic circulation (deMeno-cal et al., 2000b; Kohfeld and Harrison, 2000).

When referring to the present climatic teleconnectionsbetween the Mediterranean basin, the middle and highlatitudes and the tropical zone (Alpert et al., 2006; Lionelloet al., 2006), the period 7000–5500 cal BP can be consideredas a transition phase during which some of theseteleconnections were established or strengthened. Thecorrelations between the Mediterranean basin, the Tropicsand the Monsoons domains occurred early. The dryepisodes recorded in the domain of the Indian and Africanmonsoon during the intervals 11 000–9500 and8000–7000 cal BP (Caratini et al., 1981, 1994; Gasse andVan Campo, 1994) (fig. or 8400–8000 (Tropical Africa,Gasse, 2000) (Fig. 4r) are synchronous with the dry phases

10 900–9700 and 8400–7600 cal BP described in the Wes-tern Mediterranean (Jalut et al., 1997, 2000) (Fig. 4a).Later, a good correlation is also observed between the dryphases recorded in the Western Mediterranean around5300–4200, 4300–3400 and 1300–750 cal BP (Jalut et al.,1997, 2000), and the events recorded ca. 5000–4500 and1300 cal BP for the Asiatic monsoon (Morrill et al., 2003)and ca 4200–4000 cal BP for the African monsoon, north-ern domain (Gasse, 2000) (Fig. 4q and r).In the Western Mediterranean, the dry phase recorded at

Capestang around 4500–4000 cal BP (Fig. 2) correspondsto a clear evolution towards Mediterranean climaticconditions (Jalut et al., 2000) (Fig. 4a). It is contempora-neous with the cooling of the surface water of the NorthernAtlantic around 4200 cal BP (Bond et al., 1997), recordedboth by pollen, hydrological, archaeological and spe-leothem data over the world (Caratini et al., 1994; Magri,1999; Magri and Sadori, 1999; Gasse, 2000; Magny et al.,2002; Sadori et al., 2004; Drysdale et al., 2006) (Fig. 4).In Mesopotamia, during about 400 years, this event

corresponded to an abrupt aridification of the floodingplains and the neighbouring regions (Weiss et al., 1993)recorded by aeolian and archaeological deposits accumu-lated in marine sediments of the Gulf of Oman (Cullenet al., 2000; deMenocal, 2001). Because of the correlationbetween the arid event and the extinction of the Akkadianempire, this abrupt aridification was sometimes consideredas the cause of this collapse (Cullen et al., 2000;deMenocal, 2001). However, the abrupt fall of this empirewas more probably due to political reasons such as anarchyand wars against Elamites and the king of Mari (Roux,1995). The climatic stress probably worsened the effects ofthese conflicts.In China, this period corresponded to an accentuated

drought in the north and to floodings in the south. Thismight have contributed to the extinction of the NeolithicCultures around the Chinese Central Plain (Wu and Liu,2004).On the western coast of India around 4000 cal BP, a

noticeable decrease in precipitation is recorded (Caratini etal., 1991, 1994; Bentaleb et al., 1993, 1997). This climaticdegradation is also observed to the north in Rajasthan(Singh et al., 1974, 1990; Bryson and Swain, 1981;Krishnamurty et al., 1981; Courty, 1990). It correspondedto the end of the Harappan Culture. The increasingdrought culminating around 4000 cal BP is observed tothe north of Arabia as well as in numerous sites of thetropical zone (Caratini et al., 1994; Gasse and Van Campo,1994).In the Western Mediterranean, from ca 4500 cal BP, the

progressive evolution towards Mediterranean climateconditions determined a higher frequency of dry summersunfavourable for cattle breeding at low altitude. In theMediterranean Pyrenees, around 2200m a.s.l., this climaticevolution was one of the possible causes of the develop-ment of the seasonal grazing in altitude (Esteban Amat,1995; Rendu et al., 1996; Galop, 1997; Jalut et al., 1997,


2000). Around 5000–4000 cal BP, data from the Alboransea indicate, before a continuous decreasing tendency, ahigh sea surface temperature (Martrat et al., 2004). It canbe assumed that the contemporaneous spread of Fagus onthe nearby mountains and probably along flood plains wasfavoured by a strong evaporation responsible for anincrease in atmospheric humidity and precipitation on thereliefs from southern Spain to Gulf of Lions. In theBalearic Islands (Minorca and Majorca) (Yll et al., 1994;Perez-Obiol and Sadori, 2007), pollen data reinforce thishypothesis. They show the rare presence of Fagus in thetwo islands around 7800–7100 cal BP then a maximumrecorded at Majorca around 5000 cal BP. At low eleva-tions, the increase in temperature and summer droughtwould have favoured the synchronous extension of theevergreen sclerophyllous trees.

If at low elevation the increasing drought coulddetermine changes in the agricultural practices andseasonal migrations in the mountains, it cannot beconsidered as the unique factor responsible for thecolonization of the European mountains by humans. Asdescribed in Argolide (Greece) (Angel, 1972; Jahns, 1993),demographic growth cannot be excluded.

After 4500–4000 cal BP, two main aridification phasesare recorded by pollen data in the Western Mediterranean,between 2850–1730 and 1300–750 cal BP (Fig. 4a). The firstis correlated with the dry episode recorded from marinecores in the Eastern Mediterranean between 3000 and1700 cal BP (Schilman et al., 2001) (Fig. 4b), and thesecond with the short Medieval climatic optimum at theend of the 9th century (Jalut et al., 2002) (Fig. 4a). Theybelong and are the last expression of the aridificationprocess that began around 8000–7000 cal BP (Fontugne etal., 1994; Schilman et al., 2001) (Fig. 4d).

4. Conclusion

The Holocene was a period of climatic instability inwhich three phases can be distinguished (Fig. 4).

The first (11 500–7000 cal BP) was mostly humid andfavourable for the development of a vegetation coverdominated by deciduous broad-leaf trees. Various regionalclimates prevailed such as an attenuated oceanic type in theWestern and Central Mediterranean, characterized byshort dry summer periods and abundant precipitation inautumn, spring and winter (supra-Mediterranean type). Tothe SW of the basin, in SE Spain, the pollen data suggestMediterranean conditions since the beginning of theHolocene. In other zones, irregular summer drought withmore or less high precipitation during spring, autumn andwinter (sub-Mediterranean type, Ozenda, 1994) could haveexisted as well as continental tendencies with summerprecipitation.

The second phase (7000–5500 cal BP) was marked by theprogressive decrease in insolation at the high latitudes ofthe northern hemisphere. This event is considered as apossible cause of the rapid climatic changes recorded

around 6000–5000 and 3500–2500 cal BP (Mayewski et al.,2004) (Fig. 4t). It corresponded to the end of the humidperiod in Northern Africa, especially in Sahara (Hayneset al., 1989; Lezine, 1989; Lezine and Casanova, 1989;Petit-Maire and Guo, 1996). The phase was also marked byminor humid episodes in western Asia and in northernAfrica (Gasse and Van Campo, 1994).The third phase (5500 cal BP-Present) was marked by an

increasing aridification demonstrated by well-correlatedpollen, hydrological and marine data (Fontugne et al.,1994; Schilman et al., 2001) (Fig. 4d). The climatic origin ofthis change is well established. The consequences for thevegetation cover were a decline of the deciduous broad-leaftrees and a spread of the evergreen sclerophyllous taxa.Examples demonstrate that such changes could haveoccurred before the first human impacts.The period 4500–4000 cal BP corresponded in the Wes-

tern Mediterranean to the beginning of the evolutiontowards full Mediterranean climatic conditions. In the NWof the Mediterranean basin they were installed around3000 cal BP, during the arid episode 3500–2500 cal BP. Thelast dry phase recorded in this region (1300–750 cal BP) issynchronous with the climatic change recorded at highlatitude around 1200–1000 cal BP (Mayewski et al., 2004)(Fig. 4t).Nowadays, at a monthly or annual scale, the variations

of the Mediterranean climate are linked to global tele-connections. The examples previously discussed (Fig. 4)show that such links have also existed during the pre-industrial period of the Holocene (Schmidt et al., 2004).With respect to the chronology of the main global

events, there is a diversity of responses and minorchronological gaps, which could be due to the variousregional impacts of the global events, to the time ofresponse of the ecosystems and finally, but not the last inimportance, to the relative uncertainty of the dating.In the Mediterranean basin and elsewhere, these global

events influenced the evolution of the societies, which, insome cases, had to adapt to not disappear (Weiss andBradley, 2001). In the Northern and Southern Alps,between 4300 cal BP and AD 800, good concordancesbetween GRIP data (Johnsen et al., 2001) and agriculturaldata show the role of climate as a possible determiningfactor of the agricultural activities (Tinner et al., 2003).It is obvious that because of their land use, the new

agricultural techniques and materials, the demographicgrowth, humans have accentuated the effects of theclimatic constraints on the vegetation cover, especiallyduring the late Holocene. Since the Roman period, thehuman impact was so strong that the associated changes inalbedo are considered as a possible cause of the aridifica-tion during the last two millennia (Reale and Dirmeyer,2000). Therefore it was obviously an additional cause ofthe vegetation changes. But, as emphasized by Sadori(2007), when considering the Holocene, the major syn-chronous changes in vegetation throughout the Mediterra-nean basin could not have been caused by humans alone.


The development of the cultural landscapes induced theformation of new plant communities: the anthropogeniccommunities. Different from the previous and in equili-brium with the new climatic conditions, they nowcharacterize the Mediterranean area ‘‘a biologic unitmodelled on the climate of which it is the living expression’’(Emberger, 1943).

Acknowledgements

The authors express their sincere thanks to Dr. L. Sadorifor critically reviewing the manuscript and offering helpfulremarks for its improvement.

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