Cent. Eur. J. Geosci. • 2(2) • 2010 • 71-82DOI: 10.2478/v10085-010-0005-8

Central European Journal of Geosciences

A multidisciplinary approach to reveal the SicilyClimate and Environment over the last 20 000 years

Research Article

Alessandro Incarbona1∗, Giuseppe Zarcone1, Mauro Agate1, Sergio Bonomo2, Enrico Di Stefano1,Federico Masini1, Fabio Russo1, Luca Sineo3

1 Università di Palermo,Dipartimento di Geologia e Geodesia,Via Archirafi 22, 90123 Palermo, Italia,

2 Consiglio Nazionale delle Ricerche,Istituto per l’Ambiente Marino Costiero,Via L. Vaccara 61, 91026 Mazara del Vallo (Tp), Italia,

3 Università di Palermo,Dipartimento di Biologia animale,Via Archirafi 18, 90123 Palermo, Italia

Received 8 January 2010; accepted 9 March 2010

Abstract: We present a thorough review of the knowledge on the climate and environment in Sicily over the last 20 000years, taking into account results of several studies carried using terrestrial and marine records. We obtain acoherent framework of the most important changes succeeded in the island, even if some points need furtherinvestigation.All the reconstructions of surface temperatures of the seas and the air surrounding Sicily point out severeclimatic conditions during the last glacial period. The steppe- and semisteppe-like vegetation pattern testifies,together with additional evidence from geochemical data of lacustrine evidence, markedly arid conditions. Fi-nally, significant episodes of sea level drop connected Sicily to the Italian Peninsula and favoured the dispersionof faunal elements from southern Italy.The transition between the last glacial and the Holocene was not characterized by a gradual warming but waspunctuated by two abrupt suborbital climatic fluctuations: Bølling-Allerød (warm) and Younger Dryas (cold), asrecognized in the sediments recovered close to the northern and southern coast of Sicily. A denser arborealcover is possibly indicated by the occurrence of dormouse and Arvicola remains.Finally the sensitivity of Sicily to climate perturbations is demonstrated by the occurrence of repeated subtleclimatic anomalies during the Holocene, including the Little Ice Age, also known from historical chronicles.Forests, woods and Mediterranean maquis developed in the early-middle Holocene. Thereafter was a generaldecline of arboreal vegetation, following a general aridification trend that seems to be a common feature insouthern Europe and North Africa. Since Greek colonization (7th century before Christ), the landscape wasintensively modelled for agriculture and breeding, leading to a significant loss of vegetation cover.

Keywords: palaeoecology • palaeoclimatology • last glacial maximum • Holocene • Sicily

© Versita Warsaw

∗E-mail: alessinc@unipa.it

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A multidisciplinary approach to reveal the Sicily Climate and Environment over the last 20 000 years

1. Introduction

The island of Sicily is a segment of the NeogeneApenninic-Maghrebian fold and thrust belt. Differentstructural units are arranged in a chain south-east verg-ing over imposed on a foreland more or less deformed.This range is mainly oriented E-W and the highest peaksdo not exceed 2 000 m above sea level. Only the Etnasummit, the largest volcano in Europe, reaches 3 400 mabove sea level. During the Quaternary, Sicily has beenaffected by intense tectonic activity and local uplift [1, 2].The human history of the island was intimately tied to thewestern civilization development, through the link, amongothers, to Greek, Roman and Arabian cultures. Sicily isalso a unique place in the world for the exceptionally geo-diverse heritage, with a continuous geological record fromthe late Paleozoic to the Quaternary, with over 200 geo-sites listed by the public administration [3, 4].Several studies of international relevance have been car-ried out on Sicily or on the seas that surround it. Forinstance, data were collected on stable isotopes of la-custrine sediments, speleothems, and pollen fossil grains,while micropaleontological and geochemical data on ma-rine sediments were recovered from the Sicily Channeland the Tyrrhenian Sea (Figure 1). The interpretation ofthese data allowed gathering of a lot of information deal-ing with Sicily’s environment in the past. The aim of thepresent paper is a synthetic view of these data, in order tosketch the climatic, environmental and physiographic evo-lution undergone since the last glacial maximum (the last20 kyr). Finally, the framework of climatic and environ-mental evolution deduced for the central Mediterraneanregion is tentatively compared to other continental andmarine records, in order to recognize possible climate forc-ing mechanisms.

2. Last Glacial Maximum

Environmental conditions and the aspect of Sicily duringthe Last Glacial Maximum, about 20 kyr ago was pro-foundly different from today. Sea level was at about 110m below the present [5–8], making the island wider than17 000 km2 (from 25 883 km2 of today to 43 100 km2 ofthe last glacial maximum). Sicily was connected to theItalian peninsula and to the island of Malta, allowing thefree dispersion of continental fauna (Figure 2a).Temporary land bridges between Sicily and the ItalianPeninsula allowed the dispersion of faunas. Such a phe-nomenon made the mammalian assemblage of Sicily onlyslightly less diversified with respect to coeval southernItaly assemblages [9]. The dispersion of taxa from southern

Figure 1. Maps showing the location of sites discussed in the text.A Bathymetric map of the Mediterranean Sea. B Map ofSicily: 1 Coastal lakes of the Natural Reserve of ‘GorghiTondi’; 2 Sicily Channel Site 963 of Ocean Drilling Pro-gram; 3 Carburangeli Cave; 4 Tyrrhenian Sea Marinecore BS 79-38; 5 Coastal lake of the Natural Reserve of‘Biviere di Gela’; 6 Pergusa Lake; 7 San Teodoro Cave;8 Strait of Messina.

Italy led to a large turnover, as can be seen in the com-pletely renewed Castello Faunal Complex (20-9 kyr BP).Endemic large mammals such as elephants, fallow deerand large predators disappeared, as well as most of theendemic small mammals, replaced instead by equids, reddeer and aurochs. The arrival of horses and the abundanceof the vole Microtus (Terricola) ex gr. savii, supports theoccurrence of an open landscape in Sicily [9].All the available information confirms that severe cold cli-matic conditions characterized Sicily 20 kyr ago, evenif only in a few cases absolute temperature estimateswere proposed. Based on the equilibrium line altitudeof glaciers, the air temperature reconstruction was lowerthan 7-8°C, with respect to today [10].A larger number of studies deals with sea surface tem-perature (SST) estimates of the surrounding seas. [11, 12]report SSTs between about 10°C - 15°C, on the basisof planktonic foraminifera assemblages. Their reconstruc-tions clearly diverge on the spatial cooling trend respec-tively westward and northward. The geographical differ-ence is again reported for summer glacial SSTs, whichindeed are extremely dissimilar from a numerical point ofview, at about 16°C - 20°C for [11] and from about 23°C -25°C for [12].More recently, planktonic foraminifera assemblages esti-

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Alessandro Incarbona, Giuseppe Zarcone, Mauro Agate, Sergio Bonomo,Enrico Di Stefano, Federico Masini, Fabio Russo, Luca Sineo

Figure 2. Physiographic evolution of Sicily, based on eustaticchanges: A Last Glacial Maximum, sea level at -118 m; B Transition between the Last Glacial Period andHolocene, sea level at -90 m; C Early-middle Holocene,sea level at -13 m.

mated annual SSTs colder than 2-6°C, with possibly majortemperature anomalies during the summer (Figure 3) [13–15]. Much more pronounced is instead the temperaturedrop of 12-13°C hypothesized by alkenone data in theTyrrhenian Sea for the uppermost part of the last glacialperiod [16, 17].Another interesting point comes from coccolithophoredata, and in particular from the absence of species spe-cialized to inhabit the lower and upper photic zone. Thisfact indicates that during the last glacial period the sum-mer thermocline never reached the lower photic zone andimplies that there was a lower temperature difference be-tween atmosphere and sea surface (colder air temperaturein the summer) and a strengthened atmospheric circulation(strong winds and storm episodes) [18–20].Today’s atmospheric circulation in the MediterraneanBasin, in particular in the western-central sector, is sea-sonally controlled. This area is under the influence ofrainy westerlies in the winter. In the summer, because of

the establishing of the Azorean High, westerlies blow ata higher latitude, provoking a high pressure regime anddrought. Information on northern hemisphere atmosphericcirculation is recorded in Greenland ice cores [21–23]. Inparticular, high values of Na+ in ice cores have been ob-served during deep Iceland low periods, when enhancedNorth Atlantic winter atmospheric circulation allows thetransport of sea salt to central Greenland. High values ofnon-sea salt K+ are linked to periods of a strengthenedSiberian High. Both of them are proxies for the strengthof atmospheric circulation in the northern hemisphere [24]and have been associated with periods of expanded andstrengthened polar vortex [22, 25]. During the glacialperiod, both Na+ and K+ values in Greenland ice coreGISP2 are much higher (Figure 3), supporting the sig-nificant strengthening of the atmospheric circulation, withstrong action of westerlies, the most common phenomenonof perturbation in the central Mediterranean.The strengthened atmospheric circulation is further con-firmed by oceanographic circulation studies on theMediterranean Sea. In fact, the circulation system ofthe Mediterranean Sea develops in three layers and isaffected by deep water production, which occurs in thenorthern basin sectors, mainly in the Gulf of Lions andthe Adriatic Sea. Deep water forms during these shortepisodes of blowing northerlies [26, 27]. All data indicatethat during the last glacial period, Mediterranean watercirculation was strengthened, implying a higher volumeproduction of deep waters, triggered by a stronger andmore prolonged wind action in the northern part of thebasin [28–32].The study of sediments and fossil pollen grains from Per-gusa Lake offers a snapshot of the vegetation pattern andof the precipitation regime that was established 20 kyrago. Vegetation was characterized by a high abundanceof Artemisia and Chenopodiaceae, indicating a steppeor semi-steppe environment [33]. Steppe environment,even with minor local variations, seemed to be a com-mon feature in southern Europe, since it was recognizedamong others in the Iberian Peninsula, southern Italy andGreece [34–37]. However, some mesophilous and ther-mophilous species of Angiosperme trees survived in Sicilyand could have behaved as a refugia area, favouring therapid central Mediterranean re-colonization once climateamelioration happened [33]. Even the occurrence of reddeer and wild boars suggests the presence of forestedpatches [38, 39].A steppe environment is mostly characterized by low pre-cipitation levels. In Sicily such a characteristic is furthersupported by overall negative correlation of oxygen iso-topic data with the arboreal pollen (and positive corre-lation to the abundance of Artemisia grains) in Pergusa

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A multidisciplinary approach to reveal the Sicily Climate and Environment over the last 20 000 years

Figure 3. Down-core variations of climatic/environmental proxies for the last 20 kyr BP, each plotted with its own age model. From the left: δ18Orecord of Greenland ice cores NGRIP, a proxy for air temperature in that region [23]; Paleoclimatic curves of planktonic foraminiferaand calcareous nannofossils at ODP Site 963, Sicily Channel [13, 18]; Abundance variations of K+ and Na+ in the Greenland GISP2ice core [22, 25]; Relative abundance changes (%) in arboreal pollen grains of Lago Grande di Monticchio sediments [34]. The leftcolumn indicates boundaries among last glacial period: Bølling-Allerød, Younger Dryas and Holocene.

Lake sediments. In fact, the highest δ18O values of the lastglacial are interpreted as phases of much reduced rainfalland high evaporation in a dry environment [33, 40]The framework traced out above is fully confirmed by aGeneral Circulation Model carried out on the last glacialin Europe. Severe climate and aridity conditions were es-tablished even south of 45°N. In the simulation, tempera-tures were lower than 2-7°C, with strong westerly activ-ity, while precipitation accounted only for 1.5-5 mm day−1.Finally, the dominant vegetation type in southern Europewas constituted from temperate grassland, reflecting thesubstantial decrease in precipitation [41].

3. Deglaciation

The transition between the last glacial period and theHolocene took 3 kyr (from 14.7 to 11.7 kyr BP) and wasthe result of higher insulation energy received in responseto orbital variations. It was a re-organization phase ofthe climate system which involved changes in ice sheetvolume, Atlantic overturning meridional circulation, at-mospheric greenhouse gases and global mean tempera-ture [42–46].In the Northern Hemisphere, the warming was not grad-ual but occurred through two suborbital climatic fluctua-

tions: Bølling-Allerød (warm) and Younger Dryas (cold),first recognized in Greenland ice cores and North Atlanticsedimentary records [21, 47–49]. Recent studies demon-strated that suborbital-scale climatic fluctuations can berecognized even in the Mediterranean Sea. These eventsseem to be simultaneous to those in the Greenland icecores and North Atlantic sediments, possibly because ofteleconnession phenomena [16, 17, 28, 29, 31, 32, 50–57]. In particular, in the Sicily Channel (southern Sicilycoast), geochemical and micropaleontological analyses al-lowed the whole sequence of suborbital climatic fluctua-tions across the last 115 kyr to be recognized [13, 14, 58].The warming in the lower part of the Bølling-Allerød canbe evaluated in about 5°C in the southern Tyrrhenian Sea,on the basis of alkenones [16] and in about 3-4°C in theSicily Channel, on the basis of planktonic foraminiferaassemblages (Figure 3) [13, 14]. Just after about 1,000years, in coincidence of the Younger Dryas, the TyrrhenianSea and Sicily Channel SST would again be lowered toglacial values [13, 16]. The Bølling-Allerød warm phaseand the Younger Dryas cold spell can be interpreted asrespectively wetter and drier, following the interpretationof the whole stadial-interstadial sequence in the SicilyChannel [13, 14, 18, 58].Uncertainty in the chronology of Pergusa Lake sedi-ments does not show the vegetation pattern transforma-

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Alessandro Incarbona, Giuseppe Zarcone, Mauro Agate, Sergio Bonomo,Enrico Di Stefano, Federico Masini, Fabio Russo, Luca Sineo

tion through the deglaciation [33, 59]. However, a firmchronology was established for the last 12.8 kyr and high-lights the transition towards a much wetter climate and thedevelopment of forests and woods, between the YoungerDryas and the Holocene [33, 40].High-resolution pollen data able to illustrate vegetationchanges across the deglaciation in the Mediterraneanarea have been collected from several marine cores. Theyreport the passage from a semi-desert open environmentof the last glacial to an arboreal-rich environment typicalof the early phase of the Bølling-Allerød [36, 60–62].More information is available from Lago Grande di Mon-ticchio (Basilicata, southern Italy). Even if this record isnot from Sicily, we argue that it belongs to the same cli-matic system, today under the influence of the North At-lantic Ocean SST, and is characterized by the same sea-sonality, with winter atmospheric perturbation and sum-mer high pressure establishment. As a consequence, evenif the vegetation pattern of this part of southern Italy isdifferent from that of Sicily, it possibly reacted in a similarway during the last deglaciation period.One of the most striking characteristics of the LagoGrande di Monticchio record is associated with very rapidand sharp vegetation changes, occurring at a century-scale along the last 130 kyr [34, 63, 64]. The terminal partof the last glacial period was characterized by a steppe-like vegetation, similar to that already discussed for sedi-ments of Pergusa Lake (see Section 2). Starting from theBølling-Allerød base, a gradual increase in tree-pollengrains testifies to the development of a wooded steppe,culminating later in a temperate forest (Figure 3). TheYounger Dryas cooling led the environment to revert to awooded steppe environment [34].Among vertebrates, the possible occurrence of dormouseand Arvicola, at the transition between the Castello faunalComplex and Holocene assemblages, suggests an increaseof precipitation and the development of a suitable arborealcover. Even Equus hydruntinus might have disappearedin this interval from Sicily and south Italy, because of thereduction of open landscapes [9, 39].About 14.5 kyr BP, at the beginning of the last deglacia-tion, there was a low stand of about -90 m [6]. Sicilywas still wider than today, at about 33 000 km2, but theconnection with Malta island was likely submerged (Fig-ure 2b). In the Messina Strait exists a possible corridorlocated at 72 m below sea level, between Punta Pezzoand Ganzirri. Several studies provided estimates of re-gional uplift rates in the Calabrian-Peloritan Arc, usuallybetween 0.5 and 2.5 mm yr−1 [1, 65–67]. More recently, anestimate of 1.07 and 6 0.95 mm yr−1 has been reportedfor Northeastern Sicily (S. Alessio) and southern Calabria(Scilla) respectively [2]. An important co-seismic contribu-

tion might have been added, due to local fault actions. Inparticular, in the Messina Strait, the Scilla Fault activityin the late Holocene would have increased the uplift rateup to 2.1 mm yr−1 [68, 69]. Thus, a shallow seaway mighthave formed between 15.9 and 11.6 kyr BP, when sea levelrose from -89 to -50 m [6].Occasional evidence of fossil man in Sicily, for instancefrom Riparo di Fontana Nuova (Ragusa), can be ascribedto the Aurignatian culture (late Paleolithic), to about30 000 years ago. Apparently, Sicily territories wereabandoned after 30 000 years and remained empty ofhuman settlement for several thousand years [70]. Thefirst demographic boom was found during the Epigravet-tian [71–75], at the transition between the last glacialperiod and the Bølling-Allerød. The best evidence canbe seen at Grotta di San Teodoro (Acquedolci, Messina),where well-preserved skeletal remains of seven individu-als were found. An accelerator mass spectrometry radio-carbon analysis on the skeleton St1 was carried out, andgave a calibrated age of 14 750 years BP [76]. The ancienthistory of human dispersion in Sicily might therefore betied to the crossing of the Messina Strait barrier whosehistory, as discussed above, is not still fully understood.

4. Holocene

Starting from the Holocene, Sicily assumed a modernphysiographic feature, especially since 7.4 kyr BP whensea level rose up to -13 m at (Figure 2c).Holocene climate was thought to be steady, with a uniquecooling episode occurring at about 8.2 kyr BP [77]. How-ever, advance and retreat phases in European glacierswere reported by [78]. More recently, such an instabil-ity has been proven by Ice Rafted Detritus (IRD) levels inthe northern North Atlantic (Figure 4) [79, 80]. Holoceneclimatic anomalies are now recognized in several recordsof both hemispheres [81].Holocene climatic anomalies have been identified in sed-imentary cores retrieved from the southern TyrrhenianSea and northern Sicily Channel [16, 19]. These stud-ies, based on geochemical and micropaleontological data,depict a series of cooling (2-4°C) and productivity increaseepisodes that match with the IRD discharge episodes ofthe North Atlantic (Figure 4). They are interpreted as aresult of stronger northerly wind action, which might haveprolonged even into the summer season. The Holoceneclimatic instability of the area is further supported by fourepisodes of brief cooling, recorded in the Sicilian-TunisianChannel on the basis of dinoflagellate cyst and planktonicforaminifera assemblages [82].Climatic anomalies in the Holocene record of the Sicily

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A multidisciplinary approach to reveal the Sicily Climate and Environment over the last 20 000 years

Figure 4. Down-core variations of climatic/environmental proxies for the last 12 kyr BP, each plotted with its own age model. From the left:Abundance variations of Ice Rafted Detritus lithics from North Atlantic Ocean [79, 80]. The solid line shows Holocene changes onan expanded scale. Labels B1-B8 indicate the so-called “Bond cycles”, used as a master record for Holocene climatic anomalies;percentage variations of the calcareous nannofossils species F. profunda at ODP Site 963, Sicily Channel, which are inverselyrelated to primary productivity [19]; Sea Surface Temperature variations, based on alkenones ratio, at core BS 79-38, southernTyrrhenian Sea [16]. C1-C5 indicate episodes of cooling. In gray, Holocene climatic anomaly intervals reported from multiple marineand continental records [81].

Channel seem to be repeated every 1 500 years [19],like in the North Atlantic, Pacific Ocean and Green-land [25, 79, 80, 83–87], reflecting a common response ofdifferent regions to climate forcing whose origin is still notwell understood. The last climatic anomaly is known asthe Little Ice Age (LIA) and its main phase spans from 1550to 1850 AD, when many glaciers of the Northern Hemi-sphere had the most extensive advance since the YoungerDryas [88–92]. Severe winters during the LIA, with frozenlakes and rivers and icy canals, for instance in Italy, theNetherlands and England, are reported from historicalchronicles. Different temperature reconstructions carriedout on Northern Hemisphere records envisage drops be-tween 0.5°C and 1°C [93–95].The impact of the LIA in Sicily is witnessed by two stud-ies. A 2°C SST decrease was reported by geochemicalanalysis on the Vermetid Reefs along the northern Si-

cilian coast [96]. A prolonged drought was suggestedat Erice (Trapani) by the statistical study of the numberof religious processions ‘ad petendam pluviam’ (to invokerain) [97]. These studies highlight the cooling and droughtthat characterized Sicily during the LIA.The Sicily vegetation pattern underwent abrupt modifica-tions starting from the Holocene, in response to increasedvalues of insolation, temperature and precipitation. Thesedimentary record of Pergusa lake (667 metres abovesea level), representative of the inner part of the island,shows that a forest environment developed gradually andculminated at 10 kyr BP [33, 59, 98], while in mountainregions, such as Nebrodi and Madonie (over 1 200 metresabove sea level), vegetation consisted of beech, oak, andfir forests [99]. Since 10 kyr BP, there was a significantvegetation cover loss which followed a general aridifica-tion trend.

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The pollen record of southern coastal regions, such as‘Biviere di Gela’ and ‘Gorghi Tondi’ coastal lakes (Fig-ure 1) indicates that the most significant change occurredat 10 kyr BP when Mediterranean maquis grew (for in-stance Pistacia shrubland). Since 7 kyr BP, a forest en-vironment of evergreen broad-leaved trees developed andresisted up to 2.7 kyr BP, when the environment opened(open maquis, garrigue and grassland-prairie) [99, 100].The decrease in precipitation levels since the middle-late Holocene has been also reported by geochemicaldata of lacustrine sediments and of Carburangeli Cavespeleothems [33, 40, 101, 102]. This phenomenon seems tobe tied to the southward shift of the Intertropical Conver-gence Zone and to the monsoon activity decrease occur-ring at 5.5 kyr BP that caused desertification and droughtin North Africa [103, 104]. Possibly, this phenomenon af-fected the central Mediterranean region and precipita-tion levels in Sicily, even if the link between the Africanmonsoon and the activity of high pressure cells over theMediterranean region is not understood [105].The precipitation reduction trend had an impact on thevegetation pattern. Even more important might have beenthe impact of human activities, such as agriculture andbreeding. Peaks of micro-charcoal grains suggest thatpeople might have begun soil exploitation in the middleHolocene [33, 99]. However, only with Greek colonization,and especially under Roman domination, the landscapewas intensively modelled, leading to a significant vegeta-tion cover loss. Under natural or near-natural conditions,vegetation cover would be far more important than it istoday, with Quercus and Olea forests and Mediterraneanmaquis restricted to drier situations [99, 100].The Holocene anthropogenic impact is also evident fromthe extinction of numerous animals on the island. Hunter-gatherer populations decimated several large mammals,such as wild boars and red deer [106], and also provokedthe extinction of endemites indirectly, through the intro-duction of domestic species and alteration of the vegeta-tion cover [9].

5. Conclusion

During the Last Glacial Maximum, about 20 kyr ago, sealevel was at about 110 m below the present one, [5–8] allowing the mammalian assemblage of Sicily to un-dergo a large turnover as well as a complete rearrange-ment [9]. Air temperature reconstruction envisages tem-perature lower than 7-8°C, with respect to today [10],while SSTs were possibly colder than 2-6°C [13, 15].A significantly strengthened atmospheric circulation wasdue to stronger and more prolonged northwesterly ac-

tion [19, 20, 28–30, 32]. Vegetation was characterized bya steppe or semi-steppe environment [33], but some sur-viving mesophilous and thermophilous species could havefavored the rapid central Mediterranean re-colonizationonce climate amelioration happened [33].The transition between the last glacial period and theHolocene occurred through two suborbital climatic fluctu-ations: Bølling-Allerød (warm) and Younger Dryas (cold).The warming in the lower part of the Bølling-Allerød canbe evaluated in about 5°C in the southern Tyrrhenian Seaand in about 3-4°C in the Sicily Channel. Just after about1 000 years, in coincidence of the Younger Dryas, SSTwould again be lowered to glacial values [13, 16].Holocene climatic anomalies have been identified in sedi-mentary cores retrieved from the southern Tyrrhenian Seaand northern Sicily Channel [16, 19]. The last climaticanomaly, known as the Little Ice Age, is witnessed by a2°C SST cooling along the northern Sicilian coast [96]and by a prolonged drought [97].A forest environment developed gradually and culminatedat 10 kyr BP in the inner part of the island and moun-tain regions [33, 59, 98, 99]. Since 10 kyr BP, therewas a significant vegetation cover loss which followeda general aridification trend. The decrease in precip-itation levels, since the middle-late Holocene, is alsoreported by geochemical data from lacustrine sedimentsand speleothems [33, 40, 102]. In southern coastal re-gions, Mediterranean maquis expanded about 10 kyr ago.Since 7 kyr BP, a forest environment of evergreen broad-leaved trees developed and resisted up to 2.7 kyr BP, whenthe environment became open (open maquis, garrigue andgrassland-prairie), due to intensive anthropogenic land-use [99, 100].

Acknowledgments

We are grateful to two anonymous reviewers who providedvaluable comments and suggestions. This study was sup-ported by MURST ex 60% grants E. Di Stefano.

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