13
basin characteristics of the sampling site, analyses in coastal plains must take into account other more specific factors, such as fluvial transport and deposition of pollen grains coming from the entire drainage basin, and marine transport and sedimentation processes. The topic of fluvial transport of pollen and its implica- tions for palaeoenvironmental reconstruction has been addressed in various studies in the European context (Brown et al., 2007), including Italy (Collins et al., 2001), and in tropical environments (Moss et al., 2005). Since coastal plains represent the terminal tracts of watercourses, where the water energy significantly decreases, fluvial deposition of pollen grains is accentuated with respect to upstream tracts (Chmura et al., 1999; Roe and van de Plassche, 2005). Therefore, analyses of coastal sediments must consider the pollen input deriving from the vegetation of the entire river catchment, mixed together with the ‘local’ component coming from the littoral vegetation. The Mediterranean coasts have been significantly colonized and exploited by man for several millennia and therefore exhibit a modified (‘disturbed’) state that seems less favourable for palaeoecological research aiming to reconstruct past ‘natural’ environments. Instead it is exactly this aspect of coastal areas that is particularly interesting for environmental archaeology and Introduction Mediterranean palaeoecological studies have focused mostly on the mountain belt, where small depositional basins conserve con- tinuous biostratigraphies and allow the evolution of the mountain vegetation in response to the main climate changes and human activity during the Holocene to be determined (for the NW Apennines see Braggio Morucchio et al., 1989, 1991; Lowe and Watson, 1993; Bertolani Marchetti et al., 1994; Lowe et al., 1994; Mori Secci, 1996; Watson, 1996; Branch, 2004; Guido et al., 2004b; Menozzi et al., 2007). On the contrary, palaeoecological studies along the Mediterranean coasts are insufficient, probably because of the scarce presence of swampy areas that could allow the extraction of long, continuous sedimentary sequences poten- tially rich in micro- and macroremains. Moreover, in coastal areas, the difficulty of interpreting the origin of the pollen in the sedi- ments is higher: the taphonomy of the pollen grains deserves par- ticular mention. Apart from the usual factors influencing the relevant source area of pollen (RSAP), such as vegetation and Abstract: Cores from four coastal plains of the Mar Ligure Sea in N Tuscany and E Liguria (Italy) were inves- tigated by means of pollen analysis to delineate the Holocene landscape history of the NW Italian coasts. In the first half of the Holocene (c. 9800–7000 cal. yr BP) all four sites show elevated percentages of Abies pollen which suggest the local presence of fir woods (with Ulmus, Tilia, etc.). In the second half of the Holocene (from 7000 cal. yr BP), Abies becomes locally extinct along the coasts leaving space for the development of mosaic landscapes formed by open meso-thermophilous woods (with deciduous Quercus, Alnus, Corylus) and Mediterranean maquis (with Erica cf. arborea). The new data represent a significant contribution to the recon- struction of the landscape history of the NW Italian coasts and of the history of fir in Italy. Along all examined cores discontinuous pollen records show that initially the coastal areas were characterized by retrodunal wet- lands; after c. 6000 cal. yr BP only the larger plains in N Tuscany remained extensively damp while the smaller plains in E Liguria were buried (and/or drained). Thus, these buried deposits of ‘fossil’ coastal wetlands proved to be only partially useful for high-resolution environmental archaeology and history studies. Nonetheless they are unique traces of ecosystems that provided important local economic resources for millennia and formed elements of the coastal cultural landscapes which have almost totally disappeared today. Key words: Palynology, vegetation history, Abies, NW Mediterranean, coastal plains, ‘fossil wetlands’. The Holocene 19,8 (2009) pp. 1161–1172 © The Author(s), 2009. Reprints and permissions: http://www.sagepub. co.uk/journalsPermissions.nav 10.1177/0959683609345077 *Author for correspondence (e-mail: [email protected]) The Holocene landscape history of the NW Italian coasts Cristina Bellini, 1,2 * Marta Mariotti-Lippi 2 and Carlo Montanari 3 ( 1 Laboratorio di Archeologia e Storia Ambientale, Università di Genova,Via Balbi 6, I-16126 Genova, Italy; 2 Dipartimento di BiologiaVegetale, Università di Firenze,Via La Pira 4, I-50121 Firenze, Italy; 3 Dipartimento per lo studio delTerritorio e delle sue Risorse, Università di Genova, Corso Dogali, 1/ M, I-16136 Genova, Italy) Received 17 November 2008; revised manuscript accepted 11 June 2009

The Holocene landscape history of the NW Italian coasts

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basin characteristics of the sampling site, analyses in coastalplains must take into account other more specific factors, such asfluvial transport and deposition of pollen grains coming from theentire drainage basin, and marine transport and sedimentationprocesses. The topic of fluvial transport of pollen and its implica-tions for palaeoenvironmental reconstruction has been addressedin various studies in the European context (Brown et al., 2007),including Italy (Collins et al., 2001), and in tropical environments(Moss et al., 2005). Since coastal plains represent the terminaltracts of watercourses, where the water energy significantlydecreases, fluvial deposition of pollen grains is accentuated withrespect to upstream tracts (Chmura et al., 1999; Roe and van dePlassche, 2005). Therefore, analyses of coastal sediments mustconsider the pollen input deriving from the vegetation of the entireriver catchment, mixed together with the ‘local’ component comingfrom the littoral vegetation.The Mediterranean coasts have been significantly colonized

and exploited by man for several millennia and therefore exhibita modified (‘disturbed’) state that seems less favourable forpalaeoecological research aiming to reconstruct past ‘natural’environments. Instead it is exactly this aspect of coastal areas thatis particularly interesting for environmental archaeology and

Introduction

Mediterranean palaeoecological studies have focused mostly onthe mountain belt, where small depositional basins conserve con-tinuous biostratigraphies and allow the evolution of the mountainvegetation in response to the main climate changes and humanactivity during the Holocene to be determined (for the NWApennines see Braggio Morucchio et al., 1989, 1991; Lowe andWatson, 1993; Bertolani Marchetti et al., 1994; Lowe et al., 1994;Mori Secci, 1996; Watson, 1996; Branch, 2004; Guido et al.,2004b; Menozzi et al., 2007). On the contrary, palaeoecologicalstudies along the Mediterranean coasts are insufficient, probablybecause of the scarce presence of swampy areas that could allowthe extraction of long, continuous sedimentary sequences poten-tially rich in micro- and macroremains. Moreover, in coastal areas,the difficulty of interpreting the origin of the pollen in the sedi-ments is higher: the taphonomy of the pollen grains deserves par-ticular mention. Apart from the usual factors influencing therelevant source area of pollen (RSAP), such as vegetation and

Abstract: Cores from four coastal plains of the Mar Ligure Sea in N Tuscany and E Liguria (Italy) were inves-tigated by means of pollen analysis to delineate the Holocene landscape history of the NW Italian coasts. In thefirst half of the Holocene (c. 9800–7000 cal. yr BP) all four sites show elevated percentages of Abies pollenwhich suggest the local presence of fir woods (with Ulmus, Tilia, etc.). In the second half of the Holocene (from7000 cal. yr BP), Abies becomes locally extinct along the coasts leaving space for the development of mosaiclandscapes formed by open meso-thermophilous woods (with deciduous Quercus, Alnus, Corylus) andMediterranean maquis (with Erica cf. arborea). The new data represent a significant contribution to the recon-struction of the landscape history of the NW Italian coasts and of the history of fir in Italy. Along all examinedcores discontinuous pollen records show that initially the coastal areas were characterized by retrodunal wet-lands; after c. 6000 cal. yr BP only the larger plains in N Tuscany remained extensively damp while the smallerplains in E Liguria were buried (and/or drained). Thus, these buried deposits of ‘fossil’ coastal wetlands provedto be only partially useful for high-resolution environmental archaeology and history studies. Nonetheless theyare unique traces of ecosystems that provided important local economic resources for millennia and formedelements of the coastal cultural landscapes which have almost totally disappeared today.

Key words: Palynology, vegetation history, Abies, NW Mediterranean, coastal plains, ‘fossil wetlands’.

The Holocene 19,8 (2009) pp. 1161–1172

© The Author(s), 2009. Reprints and permissions: http://www.sagepub.co.uk/journalsPermissions.nav

10.1177/0959683609345077

*Author for correspondence (e-mail: [email protected])

The Holocene landscape history of theNW Italian coastsCristina Bellini,1,2* Marta Mariotti-Lippi2 and Carlo Montanari3

( 1Laboratorio di Archeologia e Storia Ambientale, Università di Genova, Via Balbi 6, I-16126 Genova,

Italy; 2Dipartimento di Biologia Vegetale, Università di Firenze, Via La Pira 4, I-50121 Firenze, Italy;3Dipartimento per lo studio del Territorio e delle sue Risorse, Università di Genova, Corso Dogali,

1/ M, I-16136 Genova, Italy)

Received 17 November 2008; revised manuscript accepted 11 June 2009

landscape history studies. These rely on the possibility of inte-grating stratigraphical and biological data with the informationderiving from archaeological and historical sources to focus thediverse aspects of the physical, economical and social environ-ments of the past. For the prehistorical (Palaeolithic–Iron Age)and historical (Iron Age–present) periods, several sources areavailable in NW Italy and interdisciplinary work of this type hasrecently been carried out (see eg, Maggi and Pearce, 2005;Benvenuti et al., 2006; De Pascale et al., 2006; Menozzi et al.,2007). Human influence in coastal plains is often precociouslysignalled by water-management practices and for this reasonmany wetlands have slowly disappeared from the Mediterraneancoasts and with them the typical vegetation (Grove and Rackham,2001). The almost complete absence of ‘active’ wetlands alongthe coasts of the Mar Ligure Sea (and N Tyrrhenian Sea) today,drew attention to the possibility of using buried deposits ofcoastal ‘fossil wetlands’ as biostratigraphical archives (ie, sourcesfor landscape history studies), and to the opportunity of exploit-ing available deep cores for a preliminary evaluation of theseburied archives. The concept of ‘fossil wetlands’ seems to berather new in this sense, since it has previously exclusivelyreferred to fossil beds preserving mineralized fossil remains(Holman, 1968; Mead et al., 1998). Furthermore, archaeologicaland historical research shows that for millennia the coastal wet-lands of NW Italy formed landscape elements which were impor-tant for the economy of the local human populations in terms ofwater resources, space for settlement, animal husbandry, farmingpractices, commercial ports, etc. (Maggi, 2004). These ‘fossilwetlands’ should therefore be considered as part of the cultural-environmental heritage to be protected and valued in the sameway as all other wetland types (a similar environmental archaeol-ogy and history approach to wetland studies is described inLagerås, 2003). Therefore, the multiproxy study of long and short

drillings (off-site data) integrated by levels from archaeologicalexcavations (on-site data) can help reconstruct the pressure ofhuman populations on a territory, particularly on the vegetation,which contributed to the formation of the cultural landscapes. Aswe are learning to evaluate only in recent times, the history ofthese landscapes is characterized by the interplay of the ‘con-scious’ activities, generally testified by the archaeological anddocumentary archives, and the ‘unconscious’ activities recordedin the biostratigraphical archives. The success of pollen analysisin landscape history studies certainly resides in the potential ofrevealing information on both the environmental (local andregional levels) and cultural contexts of the past.Recently, new pollen studies from coastal lake sites in S Italy

(Di Rita and Magri, 2009; Noti et al., 2009) are beginning toenrich the available data on the Holocene vegetation history ofthe Italian coastal belt. Along the coasts of NW Italy (Figure 1)there are many coastal plains of different origin and extent, lyingon marine and continental deposits. Sedimentary deposits of sig-nificant thickness occur near the mouths of the major watercourses and in a lacustrine basin (the Massaciuccoli lake), andcontain the information for the reconstruction of the natural andcultural history of the coastal areas. In this paper, the vegetationhistory of four coastal sites by the Mar Ligure Sea is recon-structed through pollen analysis of ten cores and an attempt totrace the Holocene landscape history of the NW Italian coasts ismade by delineating typological and chronological correlationsbetween the various pollen sequences. Palynological research onsedimentary series of the NW Italian coasts has been sporadicallyundertaken in the past (Galletti Fancelli, 1971, 1979; Montanariet al., 1998; Arobba et al., 2001; Mariotti-Lippi et al., 2007a, b).Correlating this data with new research could allow a broader pic-ture of the history of these coasts and of the man–environmentrelationship to be revealed.

1162 The Holocene 19,8 (2009)

Figure 1 Map of the coastal strip of the Mar Ligure Sea in N Tuscany and E Liguria (Italy) with the location of the four coring sites

Materials and methods

The study areaThe coastal strip of the Mar Ligure Sea considered here liesbetween the Arno river mouth to the south and the promontory ofPortofino to the north (Figure 1). The cores come from fourcoastal plains: two wider plains in N Tuscany, the Pisa and theVersilia Plains, and two narrow plains in E Liguria, the SestriLevante and Rapallo Plains. The geological setting of this coastalarea (c. 140 km long) is described considering two separate units:the Pisa-Versilia Plain and the Tigullio gulf.The Pisa-Versilia Plain is a triangular-shaped area delimited

from NW to SE by the NApennines, extending from the mouth ofthe Magra River at the Liguria/Tuscany boarder to the mouth ofthe Arno River near Pisa. From inland to the coast, the structureof this coastal plain is characterized by a succession of alluvialfans followed by swampy depressions alternated with a series ofsandbars running parallel to the coast and forming dry dune anddamp interdune systems. The upper part of the basin fill is madeup of Upper Pleistocene and Holocene fluvial and marine sedi-ments following sea-level change in connection with glacio-eustaticoscillations (Mazzanti, 2002; Aguzzi et al., 2007); in particular,the so-called ‘Versilia transgression’ between 10 400 and 6600 BP(Federici, 1993; Antonioli et al., 2000; Lambeck et al., 2004).The Tigullio gulf represents the eastern part of the wider Genoa

gulf whose arched-shape is a consequence of the Alpine–Apennineaxis curve forming a chain of coastal hills and mountains. Similarlyto the rest of the Riviera Ligure, the Tigullio gulf is characterizedby narrow coastal plains (Sestri Levante, Chiavari-Lavagna,Rapallo) drained by small streams (Gromolo and Petronio in SestriLevante and Boate and S. Francesco in Rapallo) unable to formtrue deltas, alternated by steep rocky slopes and small gravellybeaches. The subsoil sediments of the coastal plains referred to asthe Holocene fill of the Tigullio gulf are also associated with the‘Versilia transgression’ and delineate the widespread swampyretrodunal areas separated from the sea by systems of sandbars(Fanucci et al., 1987; Carobene and Firpo, 2004; Ottomano, 2004).In the case of the NW Italian coastal landscapes presented here,

the taphonomic considerations mentioned above imply that thepalaeovegetational reconstruction based on the pollen record fromcoastal cores will be relative not only to lowland areas, ie, theplains where the coring sites are located, but also to the uplandareas, ie, the relieves (hills and/or mountains) lying immediatelybehind them (Figure 1). The coastal plains considered here, how-ever, are on the whole rather narrow and dominated by ‘coastal’relieves: in E Liguria, for example, 700–800 m a.s.l. are reachedin 3–5 km inland from the coast. Also, the watercourses that drainthe E Liguria area are characterized by small extensions and low-flow regimes. In this study area only theArno River may representa significant source of sediment load in a coastal plain, ie, the Pisaarea; however, the Arno River delta is relatively young since itwas formed only in the Etruscan-Roman period, c. 3000–2500 yrago (Caporali et al., 2005), therefore, before that period the sedi-ment transport of the river was not sufficient to form a delta.Moreover, the cuspate shape of its delta today suggests a low riverinput and a low mean annual sediment budget (Pranzini, 2001).Along the coast the climate is essentially Mediterranean/sub-

Mediterranean with 15°C mean temperature and 900 mm mean pre-cipitation. Coastal E Liguria represents the northernmost limit of theMediterranean bioclimate but it is also under the influence of theAtlantic bioclimate (Barberis et al., 1992; Vagge, 1999). As a conse-quence this littoral area is characterized by the presence of meso-Mediterranean vegetation which can be referred to phytosociologicalliances such as Quercion ilicis and Quercion pubescenti-petraeaeand the absence of the thermo-Mediterranean Oleo-Ceratonion

alliance (Gentile, 1986). In general, a patchwork of Mediterraneanmaquis (with Quercus ilex L., Juniperus oxycedrus L., J. communisL., Erica arborea L., etc.) and open mesophilous woodlands (withAlnus glutinosa L. Gaertner, Fraxinus oxycarpa Bieb., Populus albaL. and Ulmus minor Miller) represent what survives of the MarLigure Sea coastal vegetation, while Quercus spp., Carpinus betulusL. and Ostrya carpinifolia Scop. forests occur on the nearby slopes(Mariotti, 1993; Tomei and Zocco Pisana, 1994, Tomei et al., 1995).The mosaic (wet/dry) coastal landscapes are highly endangered

ecosystems at present, and the wetlands characterized by Phragmitesaustralis (Cav.) Trin. ex Steudel, Cladium mariscus L., Typhaangustifolia L., Carex elata All., etc. (Tomei et al., 1995) are themost vulnerable. Among the coastal plains examined here, onlythe Massaciuccoli Lake in the Versilia Plain is still partiallyformed by extensive wetland ecosystems (even though the corestudied comes from an area of the basin which at present is totallydrained and extensively cultivated). All the other coring sites arenow located in highly urbanized areas where almost no superficialtrace remains of the past retrodunal wetlands.

The coresThe examined cores were extracted between 1998 and 2004 bydifferent geological teams for different investigation purposes. Thecorings in the Massaciuccoli Lake basin and in Pisa were under-taken for palaeoenvironmental reconstruction purposes, the focus ofthe stratigraphical and sedimentological studies being, respectively,sea-level change in the NW Mediterranean (Antonioli et al., 2000;Lambeck et al., 2004) and the Arno River floodplain evolution(Benvenuti et al., 2006). The cores from Sestri Levante and Rapallowere extracted for hydrogeological risk assessment and mitigationduring construction works in these cities. Ten cores were subjectedto pollen analysis: six from Sestri Levante (S3–8), two from Pisa(PSR4 and PSR5) and one from Massaciuccoli (ENEA) andRapallo (RES); the depths reached by the corings are variable: theENEAcore reaches 90 m (the upper 34 m are presented here), whilethe other cores range between 15 and 30 m. All cores are character-ized by alternations of gravelly, sandy, silty and clayey sediments,including peat layers, corresponding to alternations of marine, tran-sitional brackish-lagoonal and terrestrial environments in the fourcoastal plains. In agreement with sandy and/or gravelly levels in thecores the organic component of the biostratigraphical archives isscarce and poorly preserved; furthermore, these sediments arerelated either to full marine, transitional or highly dynamic fluvialenvironments, all of which are generally unsuited for pollen analy-sis. The result is a fragmented pollen record made up of sequenceswith interruptions and/or short sequences.

Pollen analysis, microcharcoal count andradiocarbon datesAs a general rule, for the preliminary assessment of these littoralcores, subsampling for pollen analysis was undertaken every 10cm, where possible. During sampling coarse sands and gravelswere avoided or, when tentatively sampled, generally were barrenof pollen. Chemical treatment of samples followed standardmethodologies (Fægri and Iversen, 1989) and pollen concentra-tions (Absolute Pollen Frequency, APF) were calculated as pollengrains/gram of known sediment volume (2 ml). Pollen analysis wascarried out by LM at 400–1000×. Identification follows literature(Punt et al., 1976–1996; Moore et al., 1991; Reille, 1992–1998;Chester and Raine, 2001) and reference collections of the Palynologylaboratories of the University of Genoa and Florence, Italy. Pollenpercentages were calculated on the total land plant pollen sum(TLP, excluding wetland herbs and Pteridophyta). Only sampleswith TLP≥250 grains are reported in the pollen diagrams whichwere made with TILIA2.0 and zoned with CONISS (Grimm, 1994,2004); the histogram form was chosen to better underline the

Cristina Bellini et al.: Holocene landscape history of the NW Italian coasts 1163

Table 1 Radiocarbon dates from the cores

Site Core Depth (cm) Sediment Material AMS radiocarbon Cal. ages (2σ)date (yr BP) (yr BP)

Pisa PSR4 570 Clay Cerastoderma glaucum 5800 ± 80 6430–6045Massaciuccoli ENEA 100 Peaty clay Marsh 2206 ± 52a 2326–2128

210 Clayey silt Marsh 2716 ± 37a 2850–2775260 Clayey silt Wood 3827 ± 40 4344–4150750 Sandy silt Marine shell 5471 ± 41 5896–58302930 Sandy silt Wood 8419 ± 48 9513–93303210 Sandy-clayey silt Cerastoderma glaucum 8970 ± 50 9800–94453400 Sandy-clayey silt Cerastoderma glaucum 9590 ± 60 10 236–10 056

Sestri Levante S3 1420 Sandy silt and peat Wood 5894 ± 45 6850–6560S4 1800 Peat Wood 5161 ± 60 6180–5740

2490 Sandy silt Wood 7213 ± 65 8180–7930Rapallo RES 910 Clayey silt Charcoal 4563 ± 50 5330–5040

1660 Sandy silt Charcoal 7175 ± 45 8060–7930

a Conventional date.

1164 The Holocene 19,8 (2009)

discontinuities of the pollen records along the cores. Some pollentaxa (eg, Alnus, Corylus) sometimes occur with very high values:in order to better evaluate the trends of the other taxa they wereexcluded from the TLP sum.Microcharcoal analysis follows Sadori and Giardini (2007):

charcoal particles between 10 and 50 µm in diameter were countedon pollen slides and their concentrations (microcharcoals/gram)were calculated with the same method as pollen grains.The scarcity of organic material along the cores is reflected by

the overall low number of radiocarbon dates in this study (Table 1).All 14C dates are AMS except for two from the ENEA core inMassaciuccoli as reported in Lambeck et al. (2004); the date fromthe PSR4 core in Pisa is as reported in Benvenuti et al. (2006). Thedates from the S3 and S4 cores in Sestri Levante and the RES corein Rapallo are calibrated with CALIB Rev. 5.0 (Stuiver andReimer, 1993; Reimer et al., 2004).

Results

Overall, considering the sandy component forming most sedi-ments along the examined cores, pollen concentrations weresufficiently high, even though quite variable more or lessaccording to the stratigraphic alternations along the cores. Forexample, in the ENEA core from Massaciuccoli, TLP concen-trations are generally > 10 000 grains/g (max. 24 000 grains/g)in the lower sandy-clayey silts, while in the upper peaty levelspollen concentrations reach c. 400 000 grains/g. In the RES corefrom Rapallo, instead, TLP concentrations are much less vari-able (averaging around 16 000 grains/g) and seem not to clearlydepend on the stratigraphy; here, pollen concentrations andpercentages follow similar trends.

Pisa coresThe corings were undertaken in the context of the archaeologicalsite of Pisa San Rossore, where Roman ships were found, with theaim of obtaining geo-stratigraphical information on the LowerArno river Valley (Benvenuti et al., 2006). Pollen analysis wascarried out on cores PSR4 and PSR5 (43°43′N,10°23′E) located c.20 m apart; core PSR5 has no 14C dates available but thegeo-stratigraphical correlation with core PSR4 allows some broadchronological attributions.

PSR4 (Figure 2A)The bottom of the sequence (14–6 m) is characterized by high per-centages of Abies pollen (oscillating between 20% and 60%,

reaching a peak of 70% at 8.5 m) and significant percentages ofPinus (attributed mainly to P. sylvestris/mugo). Around 6400 cal.yr BP Abies percentages become very low while deciduousQuercus and Quercus ilex t. dominate. The top two samples showhigh percentages of Fagus (25–27%).

PSR5 (Figure 2B)Along the sequence Urticaceae (cf. Urtica) pollen is over-repre-sented and was excluded from the TLP sum. The lower part of thesequence records high Abies percentages (20–40%) with somepresence of deciduous Quercus, Alnus and Corylus. At the top,Abies percentages drop while deciduous Quercus, Alnus, Corylusand Quercus ilex t. increase.

Massaciuccoli coreThe results of pollen analyses of the upper 34 m of the ENEA core(43°48′N,10°19′E) from the Massaciuccoli Lake Basin in theVersilia Plain are reported in Mariotti-Lippi et al. (2007b). Herethe pollen record is reinterpreted and the diagram redrawn anddivided in two (ENEA1–2), following the important interruptionin the pollen sequence caused by a thick sand layer in the corebetween c. 8 and 28 m (c. 6000–9500 cal. yr BP).

ENEA1 (Figure 3A)Around 9800 cal. yr BP Abies pollen percentages are high(20–40%). Numerous oscillations of Abies and Poaceae percent-ages occur before the interruption.

ENEA2 (Figure 3B)The pollen sequence restarts at c. 6000 cal. yr BP with still signif-icant percentages of Abies (c. 20%), but deciduous Quercus is alsohigh (up to 20%). Corylus, Alnus and other riverine trees andshrubs dominate after the decline of Abies. The top of the sequence,around 2300 cal. yr BP, is characterized by high percentages ofdeciduous Quercus, Quercus ilex t. and Erica t.

Sestri Levante coresOf the six cores examined only the two inland ones by the Gromolostream, S3 and S4 (44°16′N,9°24′E), located c. 700 m apart, gaveuseful results; in both cores the results are limited to sandy silt andpeat levels deposited between c. 8000 and 6000 cal. yr BP.

S3 (Figure 4A)The sequence shows the almost complete dominance of Erica cf.arborea pollen, never falling below 20%. Also noteworthy are the

Figure 2 Pisa Plain cores. (A) Pollen diagram of PSR4 (selected taxa). (B) Pollen diagram of PSR5 (selected taxa)

Cristina Bellini et al.: Holocene landscape history of the NW Italian coasts 1165

deciduous Quercus pollen percentages that oscillate mostly inopposition with Erica percentages. The macroremains analysis ofthe core (8.8 m) has shown small wood and charcoal fragments ofAbies, Pinus, Alnus, Corylus, Quercus dec., Acer together withRhamnus/Phillyrea and Buxus. Moreover, a nearby stratigraphyrevealed mainly charred fruits and seeds of Corylus, Rubus andVitis at 13–20 m (c. 6000 uncal. BP; Campana et al., 2008).

S4 (Figure 4B)In this very short sequence Alnus pollen was over-represented andwas excluded from the TLP sum. The deepest sample (c. 8000 cal.yr BP) of the sequence is peculiar as it is formed by low tree(< 50%) and high terrestrial herbs (32%) percentages. Interestingtaxa occur in the pollen assemblage, eg,Olea (3%), Juniperus (3%),Erica cf. arborea (10%), Cerealia (7%, mainly Hordeum group),wild grasses (4%), Urticaceae (cf. Urtica, 10%). Except for thisdeepest sample, the bottom of the sequence shows high Abies per-centages (20–40%), while at the top Abies percentages are low anddeciduous Quercus and Corylus dominate.

Rapallo coreThe coring was undertaken in the city centre in 2004. Theupper 9 m of the RES core (44°21′N, 9°13′E) are formed bysandy silt sediments which resulted generally barren of pollengrains, as did the incompact gravelly sands of the lowest part(c. 17.5–20 m). The pollen sequence is therefore limited to thesandy-clayey silt and peat levels deposited between 8000 and5300 cal. yr BP.

RES (Figure 5A)Corylus pollen is over-represented in the sequence and wasexcluded from the TLP sum. Around 8000 cal. yr BP very highpercentages of Abies (40–60%, with a peak of 76% at 15.7 m)are recorded and are accompanied by noteworthy percentages ofUlmus (up to 21% at 16.1 m) and Tilia (up to 12% at 16.3 m).With Abies pollen, often found clumped and in an excellentstate of preservation, numerous coniferous wood fragmentsreferable to Abies are recorded. In the middle–upper part of thecore these taxa decline while deciduous Quercus and Alnusincrease. Quercus ilex t., Erica cf. arborea and terrestrial herbs(eg, Poaceae, Chenopodiaceae, Rumex) also attain significantpercentages around 5000 cal. yr BP in agreement with highmicrocharcoal and Trilete spore (mostly attributed to Pteridium)concentrations (Figure 5B).

Discussion

Pisa PlainThe results of pollen analysis show that in the early Holocene thePisa area was densely forested by fir (Abies) woods with pine(mostly Pinus sylvestris/mugo) and some oak (Quercus decidu-ous). Damp and salt-prairies were widespread along the coast.Later, the diffusion of fir decreased and at c. 6400 cal. yr BP firwas no longer part of the vegetation of the coast and perhaps alsoinland; open deciduous and evergreen oak (Quercus ilex t.) woodsbecame the dominant vegetation in the mid to late Holocene.

Figure 3 Versilia Plain core. (A) Pollen diagram from section ENEA1 (selected taxa). (B) Pollen diagram from section ENEA2 (selected taxa)

1166 The Holocene 19,8 (2009)

The upper metres of the pollen sequences are geostratigraphicallyand geoarchaeologically attributed to the pre-Roman, Roman andpost-Roman periods (Benvenuti et al., 2006), but the results ofpollen analysis are very fragmentary in connection with the occur-rence of sandy sediments; however, it is interesting to note theappearance of beech (Fagus) pollen. Signals of local presence ofbeech in the plain were also recorded in the oldest levels of the on-site pollen sequences from the Roman Ship site of Pisa SanRossore (Mariotti-Lippi et al., 2007a), and confirmed by the abun-dance of beech macroremains recovered in the same site (Bertacchiet al., 2008). In general the on-site pollen sequences, relative topre-Roman and Roman times, show the alternation of woody veg-etation during intervals of soil and climate stability, and shrubbyand/or open vegetation during intervals of deforestation andswamping and, therefore, of high environmental instability in thePisa Plain. The mid- to late-Holocene environmental instability,caused by the Arno River channel instability and by the frequencyof floodings, was connected with climate change (higher temper-atures and precipitation) and sea-level fall between 6400 and 2000BP (Benvenuti et al., 2006).

Versilia PlainPollen analyses of the upper 34 m of the ENEA core from theMassaciuccoli Lake Basin in the Versilia Plain resulted in thelongest and most continuous sequence of the four sites, eventhough a significant interruption is present here as well. Also, theavailability of numerous 14C dates makes the study of the Holocenevegetation history of the Versilia Plain the one with the bestchronology. In the early Holocene (9800–9500 cal. yr BP) fir-dominated woods with deciduous oak, alder (Alnus) and hazel(Corylus) were widespread in the plain and surrounding area;

numerous fluctuations of the trees+shrubs/terrestrial herbs ratiohint at cycles of retreat and recovery of these woods in the plain(Mariotti-Lippi et al., 2007b). The fluctuations of the woody veg-etation indicate environmental instability together with oscilla-tions in the water-table level possibly due to marine waterinfiltrations in the Versilia Plain. The stratigraphical and sedimen-tological analyses performed on the core showed that the ‘VersiliaTransgression’ was in the rate of 7 mm/yr between 10 400 and6600 BP (Antonioli et al., 2000; Lambeck et al., 2004). The sedi-ments in this part of the core, mainly ascribable to full marineenvironments, are generally unsuited for pollen conservation: theresult is an interruption of the pollen sequence between c. 9500and 6000 cal. yr BP. At c. 6000 cal. yr BP the pollen record restartsin the ENEA core and suggests that mesophilous woods domi-nated by deciduous oak were widespread in the Versilia Plain andinland areas, even though some presence of fir was stillrecorded. In the mid to late Holocene the sequence shows theretreat of fir and the expansion of alder, hazel and willow (Salix)thickets which were enriched by grapevine (Vitis) and ivy(Hedera) between 4500 and 2500 cal. yr BP. An expansion ofevergreen oak and tree-heath (Erica cf. arborea) is recorded inthe late Holocene and could be a consequence of the canalizationand reclamation works in these coastal swamps in the Romanperiod. In fact, the Fossae Papirianae mentioned in the TabulaPeutingeriana of Imperial Age evoke the presence in theMassaciuccoli Lake Basin of navigable canals for run-off to thesea (Mazzanti and Pasquinucci, 1983).

Sestri Levante PlainThe subsoil sediments of the city of Sestri Levante in E Liguriaproved to be suitable for pollen analyses only in the levels

Figure 4 Sestri Levante Plain cores. (A) Pollen diagram of S3 (selected taxa). (B) Pollen diagram of S4 (selected taxa)

Cristina Bellini et al.: Holocene landscape history of the NW Italian coasts 1167

deposited between c. 8000 and 6000 cal. yr BP and referable toswampy retro-dunal environments. The results of pollen analysesshow that in the early to mid Holocene the Sestri Plain and nearbyrelieves were characterized by a mosaic landscape with dry envi-ronments formed by sandy soils where Mediterranean maquis wasestablished, and damper environments characterized bymesophilouswoods with some presence of fir, perhaps restricted to the foothillsof the Apennines. The high environmental biodiversity is con-firmed by the occurrence of macroremains which, as a rule, have amore local significance than pollen. This patchy landscape isascribable to the presence of swampy depressions created by a sys-tem of sandbars within the coastal plain and probably also to theconfluence of its two main streams. Such a landscape is still pre-served in the Pisa-Versilia Plain today.The analyses at Sestri Levante showed that a tree-heath-domi-

nated Mediterranean maquis was present in the plain and pollenanalyses in the nearby Chiavari Plain testify that this maquis waswidespread in E Liguria in the early to mid Holocene (Guido etal., 2004a). The available data do not allow us to formulatehypotheses on the origin and history of this local high tree-heathmaquis and on which factors controlled its diffusion as done in theW coast of Corsica (Reille, 1992; Reille et al., 1997). TheNeolithic subsistence activities in the area, especially clearingwith fire and pasture (Maggi, 2004; De Pascale et al., 2006) couldhave favoured an expansion of tree-heath which previously mighthave been limited to the drier areas of the coasts. Starting with theEneolithic, the Italian Copper Age, the well-documented quarry-ing and mining activities in the Sestri Levante area (Maggi andPearce, 2005) probably maintained the diffusion of tree-heath,similarly to what emerged from archaeobotanical studies along the

coasts of S Tuscany in the Etruscan period (sixth century BC,Mariotti-Lippi et al., 2002).

Rapallo PlainThe pollen record for Rapallo, as for Sestri Levante, is availableonly for a determined time period, between c. 8000 and 5500 cal.yr BP, when the sediments of the buried deposit are ascribable toretro-dunal wetlands. Along the entire sequence the strong signalof hazel is evident and suggests the presence of hazel woods in theRapallo Plain in the early and mid Holocene. Fir-dominatedwoods with elm (Ulmus), lime (Tilia) and deciduous oak werewidespread in the early Holocene. Regarding the fir micro- andmacroremains recovered in the core, it is not possible to evaluatethe importance of their fluvial transport to the coast; however, asalready mentioned, the distance covered by the watercourses isbrief (Figure 1). A correlated signal of presence of fir woods in theearly to mid Holocene comes from pollen analysis of cores fromthe city of Genoa (Brandolini et al., 1997; Montanari et al., 1998).These fir woods have totally disappeared today from this plain aswell as from all plains and relieves of coastal NW Italy and as aconsequence there are no modern analogues of this type of vege-tation for comparison.In the middle part of the Rapallo sequence, supposedly around

7000 cal. yr BP, the signal of fir woods starts decreasing and at thetop, 5500 cal. yr BP, there is an expansion of open meso-ther-mophilous woods with oak, hazel and alder. In Rapallo thedecreasing trend of fir pollen is coincident with an increasingtrend of charcoal particles along the core; a similar opposite trendwas observed in a pollen and microcharcoal study from theMassaciuccoli Lake by Colombaroli et al., (2007) who hint at a

Figure 5 Rapallo Plain core. (A) Percentage diagram of RES (selected taxa). (B) Concentration diagram of RES (selected taxa): ×100 pollengrains/g and microcharcoals/g

1168 The Holocene 19,8 (2009)

possible correlation between the retreat of fir-dominated woodsand an increase in the regional fire regime in the mid Holocene.

The vegetation history of the NW Italiancoastal landscapesThe time-period subdivision attempted in Table 2, and on whichthe landscape history study of the NW Italian coasts is based,should only be considered as a broad reference outline since thelimited number of 14C dates from the cores hinders the possibilityof establishing accurate intra-site chronological attributions and,thus, inter-site chronological correlations are especially difficultand perhaps misleading.In general, in the early Holocene, these coastal landscapes were

prevalently woody, while in the mid and late Holocene, more openand/or mosaic landscapes prevailed. Elevated percentages of firpollen (>20% TLP, often reaching 40% and 60%), are a commonfeature of the oldest parts of the sequences from the cores.Threshold pollen percentages for fir are not well-defined; how-ever, 10% has been used to mark the Holocene population expan-sion in the Alps (Burga and Perret, 1998). Also common to allcores is the decline of fir and its disappearance in the most recentparts of the sequences and, therefore, the local extinction of firwoods from these coastal plains and relieves in the mid to lateHolocene. From the pollen record presented here it is evident thatin the early Holocene other trees such as pine, hazel, alder, elm,lime and deciduous oak were widespread along the NW Italiancoasts. The data also show that in the same period Mediterraneanevergreen elements were scarce or at least scarcely represented inthe pollen spectra, apart from high (pollen) productive speciessuch as evergreen oak and tree-heath, significantly represented in

all pollen sequences. Perhaps Mediterranean elements were lim-ited to the drier, sandy areas of the coasts. An exception to thisgeneral scheme seems to be the Sestri Levante record whichshows widespread Mediterranean maquis, within a mosaic land-scape scenario, starting from the early Holocene and continuing inthe mid Holocene. It is possible that both environmental andanthropogenic factors are responsible for the different palaeoveg-etational reconstruction in Sestri Levante.Regarding the chronology of the vegetation dynamic of the coastal

forested landscapes of NW Italy, the oldest Holocene data comingfrom the Massaciuccoli lake inform us that fir was already dominantin the woods starting from 9800 cal. yr BP and it is continuouslyrecorded there since at least 130 000 yr BP (Menozzi et al., 2003).The data coming from Pisa, Sestri and Rapallo show that fir woodswere still widespread from 8000 to 7000 cal. yr BP (Table 2). Theretreat of the fir woods is placed between 7000 and 6000 cal. yr BPin Pisa, Sestri and Rapallo, while in Massaciuccoli their presence isstill recorded after 6000 cal. yr BP. At 5000 cal. yr BP fir disappearsalso from the Versilia Plain and nearby slopes and open meso-thermophilous woods become established (Table 2).Therefore, in the first part of the Holocene fir woods were wide-

spread along the NW Italian coasts. This statement needs proofthat the pollen in the sediments of the cores reflects the local veg-etation and is not the result of fluvial transport accumulation in thecoastal plains. In order to discern the waterborne from the localpollen component in the record, Brown et al. (2007) suggest con-sidering the general preservation state of the pollen grains. An in-depth study of this type was not carried out here; however, theaverage percentage of indeterminate (reworked) pollen grains forthe entire pollen record does not exceed 2%, which suggests an

Table 2 The Holocene vegetation history of the NW Italian coasts

Chronology (cal. yr BP) Site Sediment Principal taxa Vegetation

Late Holocene 4000–2000 Massaciuccoli Peaty clay dec. Quercus – Open thermophilous woodsQ. ilex – Erica

5000–4000 Massaciuccoli Peaty clay Vitis – Alnus – Salix Damp thickets

Mid Holocene 6000–5000 Rapallo Sandy clay and peat Corylus – Alnus – Open meso-thermophilous woodsdec. Q. – Q. ilex

Sestri Sandy silt Erica Mediterranean maquisMassaciuccoli Sandy silt dec. Q. – Abies – Mesophilous woods

Alnus – Corylus with firPisa Clay dec. Q. – Q. ilex Open oak woods

7000–6000 Rapallo Sandy silt Corylus – Abies – Mesophilous woodsdec. Quercus with fir

Sestri Sandy silt and peat Erica – Mediterranean maquisCorylus – Alnus Damp thickets

Pisa Clay dec. Q. – Q. ilex Open oak woods

Early Holocene 8000–7000 Rapallo Sandy silt Abies – Corylus – Fir-dominated woodsUlmus – Tilia

Sestri Sandy silt and peat Erica – Mediterranean maquisCorylus – Alnus – Mesophilous woods with firAbies – dec. Quercus

Pisa Clay Abies – Pinus – Fir woodsdec. Quercus –Chenopodiaceae Salt prairies

10 000–9500 Massaciuccoli Sandy-clayey silt Abies – dec. Q. – Fir-dominated woodsAlnus – Corylus –Poaceae Damp meadows

Cristina Bellini et al.: Holocene landscape history of the NW Italian coasts 1169

overall excellent state of preservation of the pollen grains andrules out a strong influence of long-distance transport. Anotherelement in favour of a local input of fir pollen in these coastalcores comes from the nature of the sediments: in fact, high per-centages of fir pollen were recorded not only in silts and clays butalso in peat levels, which give a more trustworthy indication of thelocal character of these pollen assemblages.Signals of the presence of fir woods in the first part of the

Holocene also come from other coastal areas of the Mar LigureSea, eg, Genoa (Montanari et al., 1998) and the Albenga Plain(Arobba et al., 2001) in W Liguria, and in the coasts of SE France(Nicol-Pichard and Dubar, 1998). RegardingMediterranean France,however, Muller et al. (2007) suggest that the postglacial pollenrecord of (silver) fir from coastal cores should not be interpretedin terms of local presence but rather of reworking and/or long-distance pollen transport. The early Holocene presence of fir atlow altitude is also signalled at Lago dell’Accesa, central Tuscany(Drescher-Schneider et al., 2007), even if new studies have addedsome doubt on the actual local presence of fir around the lake(Colombaroli et al., 2008). Interestingly, in S Tuscany pollenanalysis in the coastal plain of the Ombrone River shows thatthroughout the Holocene fir was never an element of the vegeta-tion of the Grosseto Plain (Biserni and van Geel, 2005); this sug-gests that the southern limit of the early to middle Holocenediffusion of fir woods along the coastal plains and relieves of NWItaly could be located between the LivornoHills and the ‘Metallifere’Hills, in north-central Tuscany.In the N Apennines, numerous studies indicate the dominance

of fir in the mountain woods of the early Holocene (eg, Watson,1996; Branch, 2004; Guido et al., 2004b; Terhürne-Berson et al.,2004; Menozzi et al., 2007). In particular, in the LigurianApennines fir appears together with deciduous oak and elm start-ing from 10 600 yr ago, and becomes the dominant element of themountain woods between 9000 and 7000 yr ago (Branch, 2004).Finally, in the Alps, fir population expansion takes place from9000 to 5000 cal. yr BP in connection with soil maturation devel-opment (van der Knaap et al., 2005).

Therefore, the data presented here represent a significantcontribution to the hypothesis that from the early to midHolocene fir was widespread in the woodlands from the coaststo the mountains of Liguria and N Tuscany and that, togetherwith the mountain areas, the coastal plains and hills of NW Italywere also possible glacial refugia for fir. Besides the citedPleistocene data from Massaciuccoli (Menozzi et al., 2003),abundant 40 000-yr-old fir pollen was recorded in the coastalmountains near Sestri Levante (Molinari et al., 2008), and anew pollen study from the Arno River coastal plain in Tirrenia(Tuscany) shows an expansion of fir during a late phase of thelast interglacial and suggests the presence of glacial refugiaalong these coasts (Ricci Lucchi, 2008). Besides, Massaciuccoliis currently a well-known area of subtropical, glacial andAtlantic relicts in Italy, which denotes a strong conservativecharacter of this environment (Menozzi et al., 2003).Some doubt remains on the fir species’ attribution since

much morphological variability was observed during the analyses(although mainly falling within the range of the Abies alba pollenmorphotype) and pollen identification alone proved to be insuffi-cient for unambiguous determinations. One hypothesis regards thepossibility that the fir species occurring in the coastal forests of theNW Mediterranean area in the early to mid Holocene was not A.alba but rather another Mediterranean fir species similar to A. pin-sapo, A. marocana, A. numidica and/or A. nebrodensis (Andrieu-Ponel et al., 2000). Another hypothesis is that it was indeed A.alba populating lowland and coastal areas of NW Italy, but per-haps it was an extinct and different ecotype from the presentmountainous species. Today, in fact, silver fir grows only in themountain forests of the Italian Apennines, the closest occurrenceto the Mar Ligure coasts being in the Apuan Alps (Rovelli, 1995).An important contribution to this debate in Italy could certainlyderive from the study of fir macrofossils (charcoal, wood, needles,cones, etc.) combined with ancient and modern DNA, as has beendone for beech (Magri et al., 2006), and for silver fir in Switzerland(Burga and Hüssendörfer, 2001) and throughout Europe (Liepeltet al., 2009).

1170 The Holocene 19,8 (2009)

Since there are no modern analogues of fir-dominated woodsalong these coasts today, this research confirms that the presentdiffusion of this type of vegetation is strongly reduced in respect tothe past. In fact, this reduction has been observed throughout theNW Mediterranean area by many studies which have ascribed itmainly to human impact, particularly deforestation in terms of fireactivity (Branch, 2004; Carcaillet and Muller, 2005; de Beaulieu etal., 2005; Colombaroli et al,. 2007; Drescher-Schneider et al.,2007; Menozzi et al., 2007). The microcharcoal count carried outin this study was useful in Rapallo for detecting a possible corre-lation between the retreat of fir-dominated woods and an increas-ing regional fire-regime trend. In the S Alps, the mid-Holocenedecline of fir was certainly connected to forest fires, but also toother human impact processes, climatic changes, and to a lowgenetic variability of its populations (Wick and Möhl, 2006).According to the data presented here, the retreat of fir woods in

NW Italy began firstly along the coastal plains and relieves in themid Holocene, and only secondarily in the inland mountains.In fact, in the N Apennines the pollen records indicate the fir

decline in the late Holocene when beech became dominant in theAbies-Fagus mountain woods (Lowe et al., 1994; Mori Secci,1996; Branch, 2004; Menozzi et al., 2007). Even though at pres-ent the data are not sufficient to completely rule out the possibil-ity of reworking and/or long-distance pollen transport (especiallyfluvial transport), an element in favour of the local input of firpollen derives from the trend of its percentages and the timing ofthis trend along the sequences. If the fir pollen in these coastal sed-iments was entirely fluvially transported from upland areas, thenthe percentage trends on the coasts would follow those comingfrom the numerous pollen studies in the mountain areas of Liguriaand Tuscany (Apennines), which show the persistent dominanceof fir in the woods at least till the Roman Period, or even up to theMiddle Ages, ie, long after the decline emerging from thesecoastal data at c. 6000 cal. yr BP.In the smaller plains of Eastern Liguria the sediments deposited

in the late Holocene (after c. 6000 cal. yr BP) are formed by sandylayers, generally unsuited for pollen analysis, related to highlydynamic fluvial environments. Repeated flooding episodes, con-nected to increasing environmental instability and human pres-sure, caused the final burial of the wetlands and drainage of thesenarrow coastal plains by the Mar Ligure Sea.

Conclusion

The pollen data presented here give strong evidence for a regional-scale vegetation shift in the NW Italian coasts in the mid Holocene.In fact, this period was characterized by climatic change and envi-ronmental instability in connection with sea-level fluctuations. Oneof the main changes in the environmental features of NW Italy is thedisappearance of the fir-dominated woodlands spreading from thecoasts up to the mountains. This successional substitution of pioneerconifer woods by broadleaved forests is a common event during theearly Holocene. In the Ligurian Apennines, however, it is evidentonly from the Roman to Medieval periods when human impact wasstronger (Lowe et al., 1994; Branch, 2004; Menozzi et al., 2007).The conifer/broadleaf substitution could have been the consequenceof natural soil development at the end of the postglacial coloniza-tion processes, even if widespread erosion is often suggested (and itwould be confirmed by the progradation of the coastal plains) as aconsequence of deforestation which started at least from theNeolithic. In fact, archaeobotanical research (for a general review inLiguria, see Arobba and Caramiello, 2006; for Tuscany, Mariotti-Lippi and Mori-Secci, 2007 and Bellini et al., 2008) suggests thathuman activities during the Neolithic, especially clearing practicesfor agriculture and pasture, represented an important factor in the

acceleration of soil erosion and transport of fine material to the sea.From the CopperAge, quarrying and mining activities further inten-sified erosion and flooding, and the progradation of the coastalplains caused the definitive burial and drainage of the smallercoastal wetlands and the spread of mosaic landscapes formed byopen meso-thermophilous woods and Mediterranean maquis.Overall, the new data represent a significant input to the recon-

struction of the landscape history of the NW Italian coasts and of thehistory of fir in Italy. This research also assesses the validity of lit-toral biostratigraphical archives as sources for palaeovegetationalstudies. Even if these ‘fossil wetlands’ furnished discontinuouspollen records, they provided an important contribution to environ-mental archaeology and history studies and they often are the onlytraces remaining of ecosystems which represented important localeconomic resources for millennia. Today, wetlands form elements ofthe Italian coastal landscapes which are almost totally disappearing.

Acknowledgments

This PhD research was partially funded by the Culture 2000 ECLProject ‘Our Common European Cultural Landscape Heritage’(2005–2007) coordinated by Professor Krzywinski,Universityof Bergen (Norway). Special thanks to the SoprintendenzaArcheologica della Toscana, particularly to Dr P. Pallecchi, for thestudy opportunity in Pisa; Professor F. Garbari and Professor F.P.Bonadonna (University of Pisa) for the Massaciuccoli core;Professor M.G. Mariotti (University of Genova); Dr N. Campana(Museo archeologico di Chiavari), Dr A. De Stefanis and Dr P. DeStefanis (Geosarc) for the Sestri Levante cores. We are grateful toDr M.A. Guido and Dr B.I. Menozzi (University of Genova) foraid in pollen analysis and useful suggestions; S. Placereani(University of Genova) and T. Gonnelli (University of Florence)for technical assistance; E. Menicagli (University of Florence) formap design. We would also like to thank Professor D. Magri(University of Rome, La Sapienza) and an anonymous reviewerfor their comments which improved the paper.

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