Late Quaternary history of spruce in southern Europe

Review of Palaeobotany and Palynology 120 (2002) 131^177 www.elsevier.com/locate/revpalbo

Late Quaternary history of spruce in southern Europe Cesare Ravazzi C.N.R. ^ Istituto per la Dinamica dei Processi Ambientali, Via Mangiagalli 34, 20133 Milan, Italy Received 19 September 2001; accepted 29 November 2001

Abstract The late Quaternary history of fossil spruces in southern Europe (Picea abies (L.) Karsten and Picea omorika (Pancic) Purkyne) is based on 163 selected pollen, charcoal and macrofossil records. The timing of immigration of P. abies is estimated from data where the Picea curve passed the threshold value of 4%. P. abies occupied the southern European mountain ranges ^ excluding the Pyrenees ^ during the middle part of the last interglacial. Spruce reached its late Quaternary maximum expansion during the early Weichselian, after which it retired from central Europe and expanded in southern Europe during the middle Weichselian interstadials. A general decline in geographical distribution occurred during the last glacial maximum, and populations were most restricted during the Alpine deglaciation. The concept of ‘glacial refugia’ does not apply to residual populations because current climatic reconstructions relate periods of maximum spruce decline to maximum continental dryness during the growth season, rather than to full glacial conditions. Spruce took part in late glacial and early Holocene tree expansions in the eastern Alps and Carpathian, but failed to spread from residual populations in the Apennines and the Pirin^Rila^Rhodopes Mountains. These differences are explained by the influence of oceanic air masses on upper forest belts with relation to geographic location and maximum elevation of mountain ranges. Late glacial spruce expansion in the Alps coincides with the abrupt warming at 14 700^14 500 yr cal BP. High migration rates were reached in the upper forest belts (e.g. 1500^2300 masl) during the early Holocene, and decreased since about 6 kyr cal BP, as a result of climatic cooling during the Neoglaciation (treeline depression), ecological competition with other tree species (Abies alba), climatic and physical setting of the highest ranges in western Alps, and human impact. The long late Quaternary fossil history of presently isolated spruce stands from the Apennines accounts for their state of genetic differentiation, which could not be fully understood from the shorter time interval of postglacial events. : 2002 Elsevier Science B.V. All rights reserved. Keywords: southern Europe; plant migration; Quaternary refugia; genetic diversity; Picea abies

1. Introduction Norway spruce (Picea abies Karst.) is one of the most widespread tree species of Europe, and it forms an important component of boreal and mixed conifer ^ broad-leaved forests. Because of

E-mail address: [email protected] (C. Ravazzi).

its wide utilization in forestry, there is much interest in its ecological requirements and genetic structure at the population level. Climatic change and increased air pollution have drawn special attention to the spruce decline in central Europe (Schulze et al., 1989) and to rapid shifts in the position of treeline at high latitudes (Kullmann, 1986, 1993) and in the Alps (Holtmeier, 1993). Knowledge of the late Quaternary history of this

0034-6667 / 02 / $ ^ see front matter : 2002 Elsevier Science B.V. All rights reserved. PII: S 0 0 3 4 - 6 6 6 7 ( 0 1 ) 0 0 1 4 9 - X

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species provides a basis for comparing historical provenances and late Quaternary migration pathways with patterns of genetic variation among living populations. Indeed the genetic variability is structured by variations in geography, ecology, and reproductive systems but also by historical events (Lagercrantz and Ryman, 1990; Taberlet et al., 1998). In the present work, spruce is chosen in an investigation of paleoecological problems of past biome reconstruction, tree migration, refugial patterns induced by climate change, and the in£uence of history on genetic variation among living populations. Several studies have described the late glacial and Holocene history of spruce and the impact of climatic change and human activity on forest history in the Alps (Markgraf, 1970; Tallentire, 1973; Burga, 1988; de Beaulieu et al., 1993; Lang, 1994). Late glacial timing and patterns of migration also depend on the location and extent of tree-survival areas during the last glacial, a question hampered by the di⁄culty of detecting small tree populations by paleobotanical methods. Detection of spruce at particular sites bene¢ts from the moderately high production of pollen (Markgraf, 1980; Hicks, 1994), its anemophilous pollen+seed dispersal, and the easy identi¢cation of the pollen (Beug, 1961). Huntley and Birks (1983) suggest that the area where a tree taxon ¢rst appears in the postglacial corresponds with its survival area during the full glacial, i.e. a ‘glacial refugium’ according to Huntley and Birks (1983) and Bennett et al. (1991). These authors used informally the term ‘glacial refugium’ to refer to an area of any size in which a taxon persisted, during a cold phase. In the present work the term refugium is used in a stricter sense as an area in which a taxon persists during a disturbance event that caused its extinction in contiguous areas where it previously occurred (see also Lynch, 1988). A survival area is here de¢ned as the region to which a taxon is reduced during unfavorable periods. The identi¢cation of refugial and survival areas requires evaluating the history prior to the full glacial, using the last interglacial as a starting point to highlight processes induced by the late Quaternary phases of coldest climate on tree dynamics and migration.

This synthesis has been aided by several new paleobotanical investigations, including radiocarbon-dated sites from the Alps, the Apennines, and the Balkan peninsula, and recent progress in re¢ning the late Pleistocene stratigraphy (e.g. van Kolfschoten and Gibbard, 2000). In addition, late glacial spruce pollen and macrofossils have recently been described from the eastern Italian Pre-Alps (Avigliano et al., 2000), a poorly studied area where spruce had not been reported, and late Pleistocene spruce records are available from maar deposits in central Italy (Allen et al., 2000; Magri, 1999; Magri and Sadori, 1999). Possible centers of survival in the eastern Po Plain, the Adriatic depression, the eastern Alpine and Slovenian ranges, and the Pannonian plain during the last glacial maximum (LGM) can also be discussed thanks to information provided by Sercelj (1996), Schmidt et al. (1998, 2000), CombourieuNebout et al. (1999) and Willis et al. (2000).

2. Materials and methods 163 late Pleistocene sites with Picea abies pollen and/or charcoal and macrofossil records were selected from the Massif Central, Alps, Po Plain, Adriatic Basin, Apennine Mountains, Balkans, Pannonian plains, and central Europe (Figs. 1 to 3 and Table 1). Sites are referred in the text by their identi¢cation code listed in Table 1. In the time span of radiocarbon dating (e.g. the last 35 000 yr BP), sites without 14 C ages were generally avoided, except in poorly investigated regions (e.g. the Apennine Mountains and the southern Pre-Alps). A simpli¢ed pollen diagram has been provided for a few important sites that needed stratigraphic revision (Figs. 7, 10, 11 and 13). 2.1. Methods for determining ¢rst spruce occurrence Although the occurrence of a conifer species in a mountainous area would best be estimated from pollen in£ux values (Hicks, 1994) or macrofossils, especially cones and needles (Wick, 1996), such data are only available at a few sites. A further problem is the similarity of Larix and Picea wood

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133

Fig. 1. The main geographical regions mentioned in the present paper. Grey shading indicates elevation s 500 masl (the same in Figs. 2, 4^6, 8 and 9). Codes: Pa, Paris; Ly, Lyon; Br, Bruxelles; Am, Amsterdam; Be, Bern; Mi, Milano; Ro, Roma; Ko, KObenhavn; Be, Berlin; Pr, Praha; Lj, Ljubljana; Wi, Wien; Za, Zagreb; Ba, Bratislava; Sa, Sarajevo; Bu, Budapest; Ti, Tirane; Wa, Warsawa; Be, Beograd; Sk, Skopje; At, Athina; So, So¢ja; Vi, Vilnius; Min, Minsk; Buc, Bucuresti; Ki, Chisinau; Kiy, Kiev.

anatomy (Bartholin, 1979), the reason why charcoal is commonly identi¢ed as ‘Picea/Larix’. The only diagnostic character, i.e. the shape of exterior borders of pits in ray tracheids (Bartholin, 1979; Schweingruber, 1990), is hardly detectable in fossil material and only recently its taxonomical value has been con¢rmed and applied to charcoali¢ed fragments (Anagnost et al., 1994; Talon, 1997; Jagels et al., 2001). This di⁄culty has hampered the reconstruction of spruce history during the last glacial, when charcoal fragments are often

the only preserved plant material (e.g. in loess). In this paper, the identi¢cation of Picea from wood charcoal without speci¢c diagnosis will be questioned. Picea pollen curves in the Alps normally show a long tail or low values before a distinct rise. At many sites this step in the pollen percentage curve is accompanied by the ¢rst occurrence of stomata (Ammann and Wick, 1992). Stomata are often interpreted as sign of local presence rather than long-distance transport (Welten, 1982; Gaillard,

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Fig. 2. Location of sites mentioned in the present paper. Number and country codes refer to Table 1 in which site information is reported. Symbols: F Spruce pre-LGM record. Selected Picea records from LGM to present time: R A fossil record outside the present distribution range (e.g. extinct population) with 14 C dates. O A fossil record outside the present distribution range, without 14 C dates. b A fossil record within the present distribution range, with 14 C dates. a A fossil record within the present distribution range, without 14 C dates. * A fossil site with no Picea record.

1984; Ammann, 1989), and therefore this step in pollen abundance is taken as evidence of Picea occurrence. However, at some sites this pollen percentage pattern is unclear (e.g. A11, I7, I26, Fig. 15), and therefore I have preferred to establish a threshold value, which is discussed below. 2.2. Basis for threshold value of continental sites Ammann and Wick (1992) found that in the Alpine pollen records Picea values of 3% occur

in samples with stomata. At many sites, this value also approximates the distinct step in the pollen curve as already mentioned. Interestingly, modern pollen samples for forest types in Finland that include Picea abies show spruce values exceeding 4% (Hicks, 1986, 1994). In boreal forests, however, spruce pollen percentages are depressed by the over-representation of Pinus sylvestris (Markgraf, 1980; Hicks, 1986). In the Alps, spruce stands stressed by severe climate at the treeline £ower only periodically, every 4^10 yr (Bor-

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135

alpine sites and to the low-altitude Pinus^Picea parkland that were present in the late Pleistocene. Caution is required to evaluate the spruce-rich communities present in the Apennines during the late Pleistocene from pollen data, because these forests have no modern analogues. Nevertheless, 5% pollen values are considered evidence of spruce occurrence in central Italy (Magri, 1999; Magri and Sadori, 1999). 2.3. Marine and £uviatile sites Large rivers commonly carry reworked pollen, and pollen buoyancy commonly produces a marked over-representation of saccate grains in coastal sediments fed by rivers (Bernard and Pons, 1985; Heusser, 1988). The source of spruce pollen in these deposits is unclear. In this paper, marine sites will be considered without reference to pollen percentages. 2.4. Compilation of present and past distribution maps Fig. 3. Main sites discussed in the Venetian Pre-Alps. Grey shading indicates the maximum glacier extent during the LGM (Avigliano et al., 2000).

tenschlager, 1970). This irregular pollination strongly reduces the average pollen in£ux. Pollen assemblages from glacier ice in an Alpine valley covered by spruce-dominated forests register less than 10% Picea on average (Bortenschlager, 1970). These data, in agreement with Markgraf (1980), mean that long-distance pollen transport of Picea would not exceed 5%. Huntley and Birks (1983, p. 285) reached the same conclusion by comparing a recent surface sample map of Picea pollen percentage with the modern distribution of the tree, arguing that Picea pollen percentages higher than 5% indicate local presence. These pollen^vegetation relationships consistently suggest that the timing of late glacial and Holocene immigration of Picea abies in a region can be derived by estimating the radiocarbon age of the depth where the Picea curve passed a threshold value (T in Table 1) of 4 (3^5)%. This value applies to most of late glacial and Holocene

The available data from macrofossils and pollen sites that matched the above pollen criteria (Picea v 4%) served for compilation of past distribution maps (Figs. 5, 6, 8, 9, 14 and 16). The number of available sites is inadequate to establish spruce limits for di¡erent time periods, especially in case of the LGM map. The reconstructions are therefore tentative, and several question marks remain. The map of the present-day natural range of spruce (Fig. 4) was reconstructed by pollen data from historical times and from information on plant ecology (references in caption). Some modern spruce stands reported for central Europe in the Atlas Florae Europeae (Jalas and Suominen, 1988) do not belong to its natural area (Holzer and Philippi, written communication) and were omitted in Fig. 4. The Balkanic range is from Fukarek (1970).

3. Chrono-, climatic, and biostratigraphy The late Pleistocene reference chronostratigraphy for Central and southern Europe is presented

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136

Table 1 Location and site information of paleobotanical records selected in the present paper Site code

Site name

Geographical district

Altitude

Site type

Length of Evidence total record of spruce

Age of ¢rst spruce occurrence or occurrence intervals

Reference

Finsinger, in press Brugiapaglia, 1997 Brugiapaglia, 1997 Brugiapaglia, 1997 Schneider, 1978 Schneider, 1978 Wick, 1996; Wick Olatunbosi, 1997 Wick, 1994a,b Orombelli and Ravazzi, 1995 Zoller et al., 1977; Pini, submitted Gehrig, 1997 Gehrig, 1997 Baroni and Ravazzi in preparation Speranza et al., 1996 Beug, 1964 Gru«ger, 1968 Gru«ger, 1968; Ko£er, 1994 Lona and Torriani, 1944 Lona, 1941 Seiwald, 1980 Kral, 1983 Kral, 1980 Lona, 1962 Lona, 1957 Kral, 1991 Kral, 1986 Casadoro et al., 1976 Avigliano et al., 2000 Bondesan, 1999 Bortolami et al., 1977 Mu«llenders et al., 1996 Bertolani Marchetti, 1967 Fuchs, 1969 Kral, 1982

(masl) code: I) western Alps 2054 western Alps 1460 western Alps 820 western Alps 2305 southern Alpine fringe 580 southern Alpine fringe 240 southern Alpine fringe 230

l/p l l p l l/p l

lg-H H H LH lg-H lg-H lg-H

P s 1% T T T T P s 2% P s 2%

ca5 4.5 1.5 2.0 6 5.7 8.0 7.4

7 8 9

Lago Basso Cerete Pian di Gembro

central Alps southern Pre-Alps central Alps

2250 450 1350

l p(l) l/p

H lg-EH lg-H

T T T

9.3 = 9.4 8.0 8.2

10 11 11

Col di Val Bighera Passo del Tonale near Passo del Tonale

central Alps central Alps central Alps

2087 1883 1920

p(l) p p

lg-H lg-H MW

T T T

9.4 9.4 46^40

12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 28 29 30 31

Pian Venezia Lago di Ledro Fiave' Bondone Vegiose Laghestel Dura Moos Alpe Siusi Forcellona Lago di Fimon Arqua' Petrarca Wieser-Werfer Comelico ^ Lago S. Anna Fornaci di Revine Palughetto di Cansiglio Colle Umberto Venezia, several cores Venezia, several cores Motte di Volpedo Val Caltea Malga Varmost

eastern Alps southern Pre-Alps southern Pre-Alps southern Pre-Alps eastern Alps eastern Alps eastern Alps eastern Alps southern Pre-Alps Berici hills Euganei hills eastern Alps eastern Alps southern Pre-Alps southern Pre-Alps southern Alpine fringe Venetian Lagoon Venetian Lagoon Venetian Lagoon southern Pre-Alps southern Pre-Alps

2270 655 650 1550 1250 900 2080 1880 1330 26 100 2075 1420 260 1040 145 0/3 0 0 900 1480

p l l/p p p l/p p p p l l p p l l/p g m m m l p

H lg-H lg-H lg-H H H H H lg-H lg-H lg-H H H lg lg-EH LGM MW-H s MP-H LGM-H MW H

T T T P s 2% T T T T T T T T T T T W T T T WT T

10.2 8.4? 10.2 10.3 10.2 = 10.7 10.3 10.1 10.2? 10.2 = 10.7 lg lg 9.4 = 10.2 s 10.2 17.2 14.1 21.0 LGM s 45^40; 30?^25 s 27^? 34.2 /

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Italy (Alps and Po Plain) (country 0 Lago Vei del Bouc 1 Lago di Lod 2 Lago di Villa 3 Torbiera di Sant’Anna 4 Lago di Alice 5 Lago di Biandronno 6 Lago di Annone

Table 1 (Continued). Site code

Site name

Geographical district

Altitude

Site type

Length of Evidence total record of spruce

Age of ¢rst spruce occurrence or occurrence intervals

Reference

Kral, 1982 Kral, 1982

(masl) 1440 920

l l

H lg-H

T T

13 = 14.1 s 13

987

p

LH

*

/

35

Agoraie

northern Apennines

1330

p

H

*

36 37 38 39 40 41 42

Casanova Lagdei Prato Spilla Forl|' core 1934 Forl|' core 1952 Massaciuccoli Col¢orito

northern Apennines northern Apennines northern Apennines Po Plain Po Plain Tyrrhenian coast central Apennines

1055 1255 1350 25 29 10 750

p p p p p p p

MH-LH * MW?-H T lg-H * MW-LGM? T MW-LGM? T LGM? C lg-H T

43 44 45 45

Lago di Mezzano Lagaccione Lago di Vico (central) Lago di Vico (marginal)

central central central central

455 355 505 505

l l l l

LGM-lg EW-H EW-H EW-H

* T *P s 1.5% T

46 47 48 49 50

central Apennines central Apennines central Apennines central Apennines southern Apennines

220 45 650 1600 655

l l l g l

MW-H s Em-H s Em-H LGM-lg EW-H

T *P s 2% * * *

51 52

Stracciacappa Valle di Castiglione Piana del Fucino Campo Imperatore Lago Grande di Monticchio Ca'nolo Nuovo core RF/93-77

Braggio Morucchio et al., 1978 / Braggio Morucchio and Guido, 1975; Cruise, 1990 / Cruise, 1990 s 19.1? Bertoldi, 1981 / Lowe, 1992 LGM? Firbas and Zangheri, 1934 LGM? Firbas and Zangheri, 1954 LGM? Marchetti and Tongiorgi, 1936 / Brugiapaglia and de Beaulieu, 1995 / Ramrath et al., 1999 ca. 75; 50^40 Magri, 1999 EW; 25^50 (max 1.9%) Magri and Sadori, 1999 EW, 25^50 (max 16%) Francus et al., 1993; Leroy et al., 1996 50^40 Follieri et al., 1998 EW; MW (max 2.7%) Follieri et al., 1988 / Narcisi, 1995 / Giraudi and Frezzotti, 1997 / Watts et al., 2000

l/p m

MW; lg EW-H

* P s 5%

/ ( s 45^40)

Gru«ger, 1977 Lowe et al., 1996

53

core MD 90-917

m

LGM-H

P s 3%

s 20^18.7

54

core 309

m

lg-H

P s 3%

ca. 7

Combourieu-Nebout et al., 1999; Siani, 1999 Gru«ger, 1975

55

core 353

m

lg-H

P s 3%

ca. 7

Gru«ger, 1975

ls ls

MW/LGM Ch LGM Ch

37.7 24.9

Geyh et al., 1969 Pe¤csi, 1977

Apennines Apennines Apennines Apennines

southern Apennines central Adriatic Sea

900 3152 water depth southern Adriatic Sea 31010 water depth southern Adriatic Sea 3929 water depth southern Adriatic Sea 31207 water depth

Hungary (H) 1 Budapest Bakony foothills 2 site 17 in Willis et al., 2000 Donau valley

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32 Laghetto di Somodogna southern Pre-Alps 33 Fusine/Weissenfels southern Pre-Alps Italy (Apennines, central and southern Italy) 34 Lajone northern Apennines

137

138

Table 1 (Continued). Site code

Site name

Geographical district

Altitude

Site type

Length of Evidence total record of spruce

Age of ¢rst spruce occurrence or occurrence intervals

Reference

ls

LGM

Ch

24

Geyh et al., 1969

a

LGM

Ch

32.4

Willis et al., 2000

ls

LGM

Ch

31.8^27.6

Willis et al., 2000

310

l/p

lg-H

T

s 15^10.5

Willis et al., 1995

(masl) 3

310

l/p

lg-H

T

s 14.6^10.1

Willis et al., 1997

1740 1900

p p l

lg-H H lg-H

P s 2% T T

13-x MH-LH 3.8 ca

4 5 6

Sedmo Rilsko Besbog Kupena

2095 1050 1300

l l l/p

lg-H LH lg-H

T T T

4.5 4.0^1.3 5.5 ca

Filipovitch, 1985 Filipovitch, 1982 Bozilova and Smith, 1979; Bozilova, 1995 Bozilova and Tonkov, 2000 Stefanova and Bozilova, 1995 Huttunen et al., 1992

7

Varna

50

l

Em (?)

T

early Em (?)

Bozilova and Djankova, 1976

Bauerochse and Katenhusen, 1997 Bortenschlager, 1984 Bortenschlager, 1984 Bortenschlager, 1984 Bortenschlager, 1976; Bortenschlager and Bortenschlager, 1981 Oeggl and Wahlmu«ller, 1992, 1994; Oeggl, 1994 Klaus, 1987; DrescherSchneider and Papesch, 1998; Drescher-Schneider, 2000 Draxler and van Husen, 1978 Brosch, 2000 Draxler, 1977 Schmidt et al., 1998

Rila Mountain Pirin Mountain western Rhodopes Mountains Stara Planina Mountain

Austria (A) 1 Las Gondas

northern Tirol

2160

l/p

H

T

9.3

2 3 4 5

Wildmoos Gerlos Lindenmoos Giering

northern northern northern northern

1435 1590 640 820

p l/p l/p l/p

H lg-H lg-H lg-H

T T T T

9.5 10.3 10.5 10.2

6

Hirschbichl

Eastern Tirol

2140

l/p

lg-H

T

10.6

7

Mondsee

northern Alpine fringe 540

l

Em-EW

T

zone E2 (end)-EW

8 9 10 11

Ramsau Seetaler See Ro«dschitz La«ngsee

Austrian Alps Austrian Alps Styria Carinthia

fp l l l

MW lg-H lg-H LGM-H

T T T T

35.7 10.7 10.5 = 10.2 LGM?; ca. 21^19.0; ca. 14

Tirol Tirol Tirol Tirol

1225 790 548

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site 31 in Willis et al., 2000 central Hungarian Plain 4 site 7 (unnamed) in Willis eastern Hungarian et al., 2000 Plain 5 group of sites unnamed in eastern Hungarian Willis et al., 2000 Plain 6 Batorliget eastern Hungarian Plain 7 Kis-Mohos To¤ eastern Hungarian Plain Bulgaria (BG) 1 Manitsa Vitosha Mountains 2 Kumata Vitosha Mountain 3 Sucho Ezero Rila Mountain

Table 1 (Continued). Site code

Site name

Geographical district

Altitude

Site type

Length of Evidence total record of spruce

Age of ¢rst spruce occurrence or occurrence intervals

Reference

Drescher-Schneider, in press Fritz, 1965 van Husen, 1989 Drescher-Schneider, unpublished Drescher-Schneider, unpublished Haesaerts et al., 1996; Damblon, 1997b Haesaerts et al., 1996; Damblon, 1997b

(masl) Kohltrattenmoor Lengholz Hohentauern Leopoldsteiner See

Styria Carinthia Steiermark Alps Styria

880 570 1260 628

l/p l/p fp l/p

lg-H lg-H MW lg-H

T T T T

10.7 s 12.3 35.0 11.2

16

Rohr

Burgenland

248

l/p

lg-H

T

10.8 = 11.1

17

Willendorf II

lower Austria

230

ls

MW-LW

Ch

ca. 48^27

18

Schwallenbach

lower Austria

230

ls

MW-LW

Ch

ca. 45^34.2

ls

MW

Ch

35^30

l/p

MW

T

ca. 29^?

ls

MW

Ch

31.5^30

Behre, 1989

Gru«ger and Schreiner, 1993

Czech Republic (CZ) 1 Pavlov II

Moravia

2

Bulhary

Moravia

3

Dolni Vestonice

Moravia

Germany (D) 1 Oerel

160

Haesaerts et al., 1996; Damblon, 1997a Rybnickova and Rybnicek, 1991 Kneblova, 1954; Damblon, 1997b

lower Saxony

12

p

Em-MW

T

northern Alpine foreland northern Alpine foreland Leipzig Basin Leipzig Basin Leipzig Basin

578

l

Em-EW

T

Em (zone E4^E6); EW part Em (zone E2)-EW

650

l

MP-EW

T

Em-EW

100 107 98

l l l

Em Em MP-EW

T T T

northern Alpine foreland

605

l

MP-MW

T

Em (zone Em (zone Em (zone EW part Em (zone (MW)

Great Britain (GB) 1 Chelford

Cheshire

20

p

EW

T

EW

France 1 2 3

northern France Massif Central Massif Central

10 1080 1200

l/f l/p l

EW LGM-H MP-H

T T T

EW Emontspohl, 1995 reworked Reille and de Beaulieu, 1988 EM (E4)-St. Germain Pleniglacial; MW-36 1c; St. Germain 2-early

2

Jammertal

3

Wurzach

4 5 6

Neumark Grabschu«tz Gro«bern

7

Samerberg

(F) Fampoux Limagne Le Bouchet

Mu«ller, 2000

E4^E6) E4^E6) E4^E6);

Litt, 1994; Litt et al., 1996 Litt et al., 1996 Litt, 1994

E1)-EW;

Gru«ger, 1979a,b

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12 13 14 15

Whitehead, 1977; Holyoak, 1983

139

140

Table 1 (Continued). Site code

Site name

Geographical district

Altitude

Site type

Length of Evidence total record of spruce

Age of ¢rst spruce occurrence or occurrence intervals

Reference

Em (E4)-St. Germain de Beaulieu and Reille, 1992b 1c; St. Germain 2-MW Em (E4^E6); St. de Beaulieu and Reille, 1984 Germain 1b^c; St. Germain 2; MW (zone J2) 1.1 Wegmu«ller, 1977 0.4 Wegmu«ller, 1977 1.7 Ponel et al., 1992; Brugiapaglia and Barbero, 1994 2.7 David, 1997 1.0 Wegmu«ller, 1977 4.5 de Beaulieu et al., 1993 3.2 de Beaulieu et al., 1993 4.2 de Beaulieu, 1977; de Beaulieu et al., 1994 / Kharbouch, 2000 Woillard, 1978; Woillard and Em (E4)-EW (excl. Montaigu, Melisey II); Mook, 1982; de Beaulieu and Reille, 1992a,b MW (Pile) MP Bazile et al., 1977

(masl) Ribains

Massif Central

1190

l/p

MP-MW

T

5

Les Echets

Rhone valley

267

l/p

Em-H

T

6 7 8

Tourbie're de Chirens Le Lauza Canard (Taillefer Massif)

Dauphine¤ Dauphine¤ Ise're

480 1130 2055

l/p l/p p

lg-H H MH-LH

T T T

9 10 11 12 13

Plan des Mains Le Besset Prarion ^ Servoz La Flatie're Lac Long Infe¤rieur

Savoie Savoie northern Pre-Alps northern Pre-Alps Alpes Maritimes

2050 1835 1850 1430 2090

p p l/p l/p l/p

lg-H LH lg-H H lg-H

T T T T T

14 15

Lac des Grenouilles La Grande Pile

Alpes Maritimes Vosges

1993 330

l l/p

MH-LH Em-H

* T

Causses

380

t

MP

N, C

Jylland

20

p

BrOrup T interstadial

second part of interstadial

Andersen, 1961

0

l

Em

T

zones E5^E6

van Leeuwen et al., 2000

0

l

Em-EW

T

Em E5-EW

Zagwijn, 1961; Cleveringa et al., 2000

Carpathian Mountains 1840 Carpathian Mountains 1650 Prut River basin 140

l/p l/p ls

lg-H lg-H MW-LW

T T, S Ch

13.1 ca. 18.0^17.5; 11.1 37^24

Pop, 1971; Farcas et al., 1999 Farcas et al., 1999 Haesaerts et al., 1996; Damblon, 1997b

Dnestr River basin

150

ls

lg-H

Ch

22.9^19.7

Haesaerts et al., 1998

southern Balkans

285

l

lg-H

P s 1%

/

Willis, 1992a, 1994

16 Peyre Denmark (DK) 1 BrOrup Hotel ^ Moor The Netherlands (NL) 1 Amsterdam 2

Amersfoort

Romania (RO) 1 Taul Zanogutii 2 Iezerul Calimani 3 Mitoc Malu Galben Moldova (Mo.) 1 Cosautsi Greece (GR) 1 Gramousti

central coastal central coastal

European plain European plain

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4

Table 1 (Continued). Site code

Site name

Geographical district

Altitude

Site type

Length of Evidence total record of spruce

Age of ¢rst spruce occurrence or occurrence intervals

Reference

Willis 1992b, 1994 Bottema, 1974; Tzedakis, 1994 Bottema, 1974 Bottema, 1974 Tzedakis, 1999 Wijmstra, 1969; Wijmstra and Smith, 1976

(masl) southern Balkans southern Balkans

1800 500

l/p l/p

H MP-H

P s 1% P s 1%

/ /

4 5 6 7

Khimaditis Edessa Kopais Tenaghi Philippon

southern Balkans southern Balkans southern Balkans Aegean coast

560 120 100 40

l/p l/p l/p l/p

H H Em-H EP-H

P s 1% P s 1% P s 1% P s 1%

/ / / 40.2^40.2

northern Balkans southern Pre-Alps eastern Alps southern Pre-Alps southern Pre-Alps Ljubliana Barje Ljubljana Barje eastern Alps

1 100 1170 1120 430 300 300

f/m l/f p l/p l/f l l a

lg-H LGM lg-H lg-H lg-EH MW-H lg-EH MW

T W T T T T T Ch

10.2 = 10.4 22.5 13.4 s 13 s 11.2 10.7 14.1 = 14.4 (44 = 33)

Ogorelec et al., 1981 Sercelj, 1981 Sercelj, 1971 Culiberg et al., 1981 Culiberg, 1991 Culiberg and Sercelj, 1980 Culiberg, 1991 Brodar and Brodar, 1983

Dalmatian Islands

m/l

H

*

/

Beug, 1967

Dalmatia Dalmatian Islands Dalmatian Islands

329 water depth 3 0 15

l/p l-m l

H lg-H lg-H

T T *P s 2%

10^8.0 12.7 16^14

Brande, 1973 Schmidt et al., 2001 Schmidt et al., 2000

southwestern Jura southwestern Jura central Jura Swiss Plateau Swiss Plateau Swiss Plateau Swiss Plateau northern Pre-Alps northern Pre-Alps western Alps western Alps Swiss Plateau western Alps central Alps Swiss Plateau

1300 1035 1050 514 639 460 476 1000 1260 2095 2010 425 1940 1880 434

l/p l/p p l/p l/p l/p l/p l/p l/p l/p p l/p p l/p l/p

lg-H lg-H H lg-H MP-MW lg-H H H MH-LH lg-H MH-LH lg-H MH-LH H lg-H

T T T T T T T T T T T T T T T

5.7 6.6 5.6 2.8 Em (E3)-MW 5.0 3.4 7.3 5.9 6.3 5.6 = 5.7 5.3 6.3 7.0 = 7.2 0.6

Wegmu«ller, 1966 Wegmu«ller, 1966 Matthey, 1971 Ammann, 1989 Wegmu«ller, 1992 Welten, 1947 Zoller, 1962 Welten, 1952, 1982 Heeb and Welten, 1972 Markgraf, 1969 Welten, 1958 Lotter, 1988 Zoller et al., 1966 Mu«ller, 1972 Ro«sch, 1985

Slovenja (SLO) 1 Secovlje 2 Anhovo 3 Barje Sijec na Pokljuki 4 Ledine 5 Selca 6 Notranje Gorice 7 Zamedvejca 8 Potocka Zijalka Croatia (HR) 1 Malo Jezero 2 Vid I Neretva 3 Valun Bay 4 Lake Vrana Switzerland (CH) 1 Le Marais des Amburnex 2 Les Cruilles 3 Le Cachot 4 Lobsigensee 5 Gondiswil 6 Burgmoos 7 Fa«tzholz 8 Egelsee bei Diemtigen 9 Ha«ngstli 10 Bo«hnigsee 11 Aletschwald 12 Rotsee 13 Go«scheneralp 14 Segnes 15 Nussbaumer See

141

Rezina Ioannina

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2 3

8.4 9.0 9.9 = 10.2 T T T H lg-H H l/p l/p l/p 2220 1440 1546

8.8 = 9.0 8.6 EW T T T H lg-H EW p l/p l/f 2010 1020 445

Faninpass Selva Lai Nair 19 20 21

central Alps central Alps northern Pre-Alps Alp Marschol Crapteig Walensee

(masl)

16 17 18

central Alps central Alps central Alps

Reference Age of ¢rst spruce occurrence or occurrence intervals Length of Evidence total record of spruce Site type Altitude Geographical district Site name Site code

Table 1 (Continued).

Site code: country abbreviation+site number (see also Fig. 2). Site type: l = limnic deposits; p = peat; f = £uviatile deposits; g = till; m = deposits of marine and transitional environments; ls = loess; t = travertines; a = archeological site. Length of total record: P = Pleistocene (E = early, M = middle); Em = Eemian; W = Weichselian (divided into E, M, L); LGM = last glacial maximum; lg = late glacial; H = Holocene (divided into E, M, L). Evidence of spruce: T = pollen 4% threshold (cf. text); P = pollen par; N = needles; S = stomata; C = cones; W = wood; Ch = charcoal; * = negative evidence (e.g. no spruce record or pollen percentages 6 4%). A pollen par P s 1% was represented for some negative sites and P s 3/5% for marine sites. Age of ¢rst spruce occurrence (in kyr cal BP) or occurrence intervals: age of ¢rst signi¢cant value of spruce, based on 4% threshold or other evidence mentioned above. Ages separated by a hyphen are lower and upper boundary of an interval of signi¢cant occurrence of spruce. Ages separated by = are calibration extremes of a single 14 C age. Di¡erent intervals of spruce occurrence are separated by a semicolon.

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Burga, 1980 Burga 1980 Ammann and Tobolski in Schindler et al., 1985 Wegmu«ller, 1976 Burga, 1987 Welten, 1962

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in Table 2. Radiocarbon ages have been calibrated with CALIB 4.0 (Stuiver et al., 1998) and reported as yr cal BP. Out of the CALIB range (e.g. 24 000 yr cal BP: Stuiver et al., 1998) radiocarbon ages have been calibrated using a secondorder polynomial equation based on U/Th-dated corals (Bard et al., 1998). Calendar ages provided by varves and ice cores are reported as yr cal. Time intervals are normally expressed in kyr cal BP. Climatic transitions are used in the present work to subdivide the late Pleistocene into stratigraphic units. The resulting periods of climatic signature are reported as ‘climatic phases’. The term LGM is used with reference to the last maximum glacial advance in the Alps and the Apennines, with several culminations from about 27 to 17.5 kyr cal BP (Alessio et al., 1978; Orombelli, 1983; Niessen and Kelts, 1989; van Husen, 1989; Florineth, 1998; Giraudi and Frezzotti, 1997; Bondesan, 1999). The subdivision and boundary ages of Eemian and Weichselian (Table 2) are from Martinson et al. (1987), Tzedakis et al. (1997) and Kukla et al. (1997). Eemian pollen zonation in central Europe (zones E1^E6) is from Zagwijn (1961). Local stages are used in Table 1. Although several interstades preceding the LGM have been recognized in southern Europe from loess sequences (Haesaerts et al., 1996), karstic rock-shelters (Broglio, 1999), marine cores (Lowe et al., 1996), and lacustrine sequences (Watts et al., 2000; Follieri et al., 1998; Ramrath et al., 1999; Allen et al., 2000), these episodes are not easy correlated. Therefore many authors refer to ‘middle pleniglacial interstadials’ between 55 and 33 kyr cal BP. These are characterized either in the Mediterranean and in continental Europe by distinct peaks of arboreal pollen and marked soil horizons between loess sequences (Cremaschi et al., 1990; Paepe et al., 1990; Gerasimenko, 2000; Haesaerts et al., 1996). A sharp interstadial episode from 43 to 39 kyr cal BP evident in rockshelter and marine successions from northern Italy and from the Monticchio pollen record, southern Italy, has been tentatively correlated with the Hengelo interstadial in The Netherlands (Bortolami et al., 1977; Mu«llenders et al., 1996; Broglio and Improta, 1995; Allen and Huntley, 2000).

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The ‘late glacial interstadial’ (a term coined by Pennington) refers to the interval between the ¢rst marked increase in temperature after the LGM and the beginning of Younger Dryas (YD) (de Beaulieu et al., 1994b; Lowe et al., 1996). The subdivision and terminology of the late glacial interstadial in central Europe have recently been revised (Litt and Stebich, 1999; Litt et al., 2001). The following concepts are useful in subdividing the late glacial period in southern Europe too. The ‘Meiendorf pollen stage’ has been assigned to the ¢rst part of late glacial interstadial, with onset at 14 450 varve yr at Meerferlder Maar (Brauer et al., 1999) and at 14 700 yr cal in the GISP2 ice record (Meese et al., 1997). This results in a new subdivision of late glacial interstadial into biozones (Meiendorf, Oldest Dryas, BOlling,

143

Older Dryas, AllerOd) (Litt and Stebich, 1999). A substantial di¡erence from previous schemes used in central Europe, the Alps, and Greenland (Mangerud et al., 1974; Welten, 1979; Ammann and Lotter, 1989; Stuiver et al., 1995; Orombelli and Ravazzi, 1996) is the incorporation of Oldest Dryas in the late glacial interstadial. The warming at 14.7 kyr cal in the Greenland record is commonly taken as the end of pleniglacial (beginning of Last Termination). However, the Alps and Apennines were deglaciated well before the Last Termination as de¢ned in GRIP core (Niessen and Kelts, 1989; Giraudi and Frezzotti, 1997). Thus, the interval between the initial deglaciation of pre-alpine lakes (e.g. about 17.5 kyr cal BP based on Niessen and Kelts, 1989) and the onset of the late glacial interstadial (e.g. 14.45 kyr cal

Fig. 4. Present geographical distribution of Picea omorika (in black, from Fukarek, 1975) and of Picea abies (stripped pattern, mainly from Jalas and Suominen, 1988; Schmidt-Vogt, 1977). Distribution information for French Alps is from Collignon Trontin (2000), for Bosnia and Hercegovina from Fukarek (1970), and for Black Forest and central Europe from Holzer and Philippi (personal communication). Roman numbers mark the districts referred in the text.

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Table 2 Chronostratigraphy, climatic and biostratigraphy used in the present paper

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varve ages) in the Alps and the Apennines will be informally referred as ‘Alpine deglaciation’ (Table 2).

4. The Picea abies late Quaternary history in southern Europe Two species of spruce occur in Europe at present: Picea abies (L.) Karst. (Norway spruce) and Picea omorika (Pancic) Purkyne (Serbian spruce). The latter is restricted to the Drina Basin Mountains (Fukarek, 1975), on the present boundary of Bosnia^Hercegovina and Jugoslavija (Fig. 4). The present range of Norway spruce can be divided into three loosely connected parts (I^ III) and some fragmented spots (IV^VI, Fig. 4): (I) the Hemiboreal zone of Fennoscandia and the Baltic and Russian regions, mainly occupied by the subsp. obovata (Ledeb.) Domin. ; (II) the Hercynic^Carpathian area; (III) the Alps, central Europe, and the northern Dinarid Alps, mainly occupied by the subsp. europaea Jurkev. et Parf.; (IV) the Western Balkan Mountains in Bosnia^ Hercegovina, Montenegro, Serbija, Kosovo, northern Albania, northern Macedonia (Fukarek, 1970); (V) the Rila^Pirin^Rhodopes Mountains (Fukarek, 1970; Schmidt-Vogt, 1977; Jalas and Suominen, 1988; Lang, 1994) and (VI) the northern Apennines of Parma and Tuscany (Chiarugi, 1936, 1958). In zones II^VI the subsp. abies is prevalent. The present paper focuses on the late Quaternary history of spruce in zones III and VI, but neighboring areas are also discussed. 4.1. From Eemian to LGM During the middle Eemian interglacial (e.g. pollen zone E3, Zagwijn, 1961) Picea abies grew in a mountain belt in the Alps, the Massif Central, and the Carpathians (Fig. 5) and extended to a Scandinavian^Ukrainan zonal range (Zagwijn,

145

1989a,b). At that time, Picea-dominated pollen pro¢les from sites above 550 m altitude in the northeastern Alps (sites CH5, D3, D7, A7, Fig. 2) and Carpinus were abundant at lower elevations (site D2; Mu«ller, 2000). Drescher-Schneider (2000) found little or no spruce in western localities, and suggested that a migration from the east and southeast took place in the second part of the last interglacial. A coeval expansion of Picea from mountain ranges towards lower altitudes is also documented in pollen diagrams from the Alps and northern Alpine foreland (DrescherSchneider, 2000; Mu«ller, 2000), the Vosges foothills and the Carpathians (e.g. Imbramovice, Poland, Mamakova, 1976; La Grande Pile, F14, Woillard, 1978; Les Echets, F5, de Beaulieu and Reille, 1984) leading to the invasion of the central European plains. An earlier Eemian migration in Germany is suggested at the Neumark site (D4 in Fig. 2, Seifert, 1990; Zagwijn, 1996, p. 463). The southern limit of spruce in Europe is di⁄cult to determine precisely. No pollen sites of Eemian age are known from the Italian Alps (Coltorti and Ravazzi, 2000; Pinti et al., 2001). Spruce was present in the northern Apennine and the Balkans during the Eemian, but, judging from Sercelj (1966, 1996), Wijmstra (1969) and Follieri et al. (1988, 1998) it was con¢ned to mountain belts at the end of the interglacial. A temperate phase of possible Eemian age from the eastern Bulgarian plain (Varna, BG7, Fig. 2) contains 10^20% Picea (Bozilova and Djankova, 1976). The Eemian record of Picea omoricoides Weber (a close fossil relative of Picea omorika) is still unclear, as macrofossils of this age are very rare (sporadic needles at D7: Gru«ger, 1979b) and its pollen cannot be identi¢ed to the species level (Zagwijn, 1961 and personal communication; Andersen, 1965; West, 1980). During the early Weichselian stadial/warm^ temperate alternations referred to MIS 5a^d (interstadials St. Germain 1/VdC 12 and St. Ger-

MIS = marine oxygen isotope stage. References to local stages and estimated ages of boundaries: northern Germany from Behre (1989) and Behre and van der Plicht (1992); Alps from Wegmu«ller (1992); E-France (La Grande Pile) from de Beaulieu and Reille (1992a); central Italy from Follieri et al. (1998). Chronology and correlation of early Weichselian local stages follow the scheme proposed by Allen and Huntley (2000).

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Fig. 5. Tentative map of spruces (Picea abies+Picea omoricoides) distribution in central and southern Europe during the ¢rst part of the Eemian interglacial (Eemian zone E3/IV, cf. Turner, 2000). Data from sites reported in Fig. 1 and from Zagwijn, 1989a, 1996.

main 2/VdC 14, 107^75 kyr BP, Table 2), pollen data indicate oscillations in forest taxa, suggesting abrupt vegetational changes. Temperate species better adapted to cool-wet climates, like Picea and Abies, expanded in the northern foothills of the Alps and in the plains of west-central Europe (Gru«ger, 1979a,b; Welten, 1982; Behre, 1989; Zagwijn, 1989a,b; Gru«ger and Schreiner, 1993; Emontspohl, 1995; Caspers and Freund, 2001). Picea omoricoides was present in a distinct Picea zone in central Europe during the BrOrup interstadial, 91^105 kyr BP (Behre, 1974, 1989). Near Walensee Lake (CH18, Swiss northern Pre-Alps) deposits representing part of the early Wu«rm (early Weichselian) are rich in both Picea abies and P. omoricoides, and the needles of the latter are abundant (Ammann and Tobolski, in Schindler et al., 1985). At Samerberg (D7) Picea omorika-

type pollen (sensu Beug, 1961) was recognized in all temperate phases of early Wu«rm (Gru«ger, 1979b). These occurrences of P. omoricoides, far west and north of its present Balkanic range, might indicate that it occupied most of the spruce range reported in Fig. 6. Macrofossils of P. abies subsp. obovata (Siberian spruce) have been found at Chelford in England during this interstadial (GB1, Phillips, 1976; Whitehead, 1977; Holyoak, 1983). On the basis of comparative pollen morphology of recent and past spruce populations, Birks (1978) was able to distinguish a pollen type characteristic of the recent Hemiboreal populations of spruce (zone I in Fig. 4, dominated by P. abies subsp. obovata (Ledeb.) Domin.), including the Chelford fossil pollen, from recent populations of the Alps and southern Europe. This suggests that early Weichselian spruce from cen-

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147

Fig. 6. Maximum spruce expansion (Picea abies+Picea omoricoides) during the early Weichselian (pollen zone Saint Germain 1c, and BrOrup WFII, i.e. MIS 5b). Data from sites reported in Table 1 and from Zagwijn, 1989a,b, 1996. A glacio-eustatic sea level of 315 m (not shown) has been adopted for the Saint Germain 1c (Richards et al., 1994).

tral and western Europe was best related to the lineage of P. abies subsp. obovata rather than subsp. europaea. The former also occurred in the Alps and the Massif Central during older Quaternary times, as shown by fossil cones (Bazile et al., 1977; Ravazzi et al., in press), in areas today occupied by the subsp. europaea. Southward, Picea reached the central Italy foothills, as documented by its presence in pollen records of s 4% on the Tyrrhenian side of the Apennine chain (Lagaccione, I44, and Lago di Vico, I45, pollen records: Francus et al., 1993; Leroy et al., 1996; Magri and Sadori, 1999; Magri, 1999). In the case of Lago di Vico, a late Pleistocene Picea maximum at the very end of the early Weichselian is documented by two pollen records, one from outside the present lake (Francus et al., 1993; Leroy et al., 1996) and

the other in a water depth of over 20 m (Magri and Sadori, 1999), with the former showing a higher Picea peak (7% versus 2%), that suggests local spruce in the riparian forest. Picea also expanded in Rhodopes Mountains (Wijmstra, 1969) and in southeastern France during the second part of interstadials St. Germain 1 and 2, both in the Rhone valley (Les Echets, de Beaulieu and Reille, 1984) and in the mountain belt of Massif Central (Reille and de Beaulieu, 1988; de Beaulieu and Reille, 1992a,b) (Fig. 6). According to paleoclimatic reconstructions, these phases of maximum spruce expansion follow the warm climatic optimum of temperate episodes (St. Germain 1 and 2) and mark a moderate decrease in annual temperature and a stable to slight increase in growing season precipitation (Guiot et al., 1989, 1992). Oceanic conditions for the maximum

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northern expansion of spruce during the Eemian are inferred by Andersen (1965, 1966) and Zagwijn (1996), who note the co-occurrence of Ilex aquifolium (a tree requiring oceanic climates and mild winters) and spruce in several sites. The early pleniglacial culmination eliminated spruce from most of Central and southern Europe. Picea omoricoides was probably restricted to the Balkans, as suggested by the absence of younger macrofossil and pollen ¢nds. Picea abies maintained continuous pollen curves with 1^5% values at several sites in the northern alpine foreland (Seilern phase in Gondiswil, CH5, Wegmu«ller, 1992; Samerberg, D7, Gru«ger, 1979b) and central Italy (Lagaccione, I44, Magri, 1999). Marine deposits in central Adriatic (I52, Fig. 7, Lowe et al., 1996) also show a continuous Picea curve after the transition from early Weichselian to early pleniglacial. In Fig. 7 this transition is marked by the Quercus drop at end zone I. In southern France and Massif Central spruce pollen

nearly disappears. South of the Alps and in the Balkans this interval is poorly documented, and the idea of an early pleniglacial withdrawal cannot be demonstrated. Subsequently, a moderate expansion took place during the middle pleniglacial interstadials at several sites in southern Europe. From the northern alpine foreland, spruce moved to central Europe during early interstadials of the middle pleniglacial, as shown at La Grande Pile, F15 (Pile interstadial), but it did not reach northern Germany. This absence might suggest that time allowed for expansion northward during interstadials was too short (Behre, 1989), or that climate was unfavorable. Conversely, south of the Alps in the Adriatic Basin the record of spruce is continuous up to the latest middle pleniglacial interstadials, 40/33 kyr cal BP (Bortolami et al., 1977; Mu«llenders et al., 1996; Lowe et al., 1996, Fig. 7). During the interstadials between 50 and ca. 34 kyr cal BP, spruce reached its late Pleistocene maximum in

Fig. 7. Simpli¢ed pollen diagram from site I52 (core RF/93-77), central Adriatic Sea (from Lowe et al., 1996).

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the central Apennines (16% in the marginal core at Lago di Vico, I45, Leroy et al., 1996 ; 5.4% at 28.1 m depth at Lagaccione, I44, Magri, 1999), being present on foothills around Rome (Magri, 1999), but did not reach the southern Apennines (I51, Gru«ger, 1977; I50, Allen and Huntley, 2000). It occurred in the Po Plain and the Adriatic Basin (Bortolami et al., 1977; Mu«llenders et al., 1996; Lowe et al., 1996). The middle Wu«rmian Picea cones at Massaciuccoli (I41, Fig. 2) on the Tyrrhenian coast (Marchetti and Tongiorgi, 1936), though undated, might have been deposited during this middle pleniglacial interstadial interval. About 34 770 [ 580 yr 14 C BP (40 200 yr cal BP or 45^50 kyr using orbital chronology, Mommersteeg et al., 1995), spruce pollen appeared in the long succession of Tenagi Philippon (GR7 in Fig. 2, 12 km from the Aegean coast;

149

Wijmstra, 1969). This record includes four samples with 3.5% Picea according to the published pollen diagram (1% according to the spreadsheet data, Mommersteeg, personal communication), suggesting that spruce was part of an open Pinus^Picea^Juniperus forest surrounding the Tenagi Basin (Wijmstra, 1969). Whether this pollen belongs to Picea omoricoides (as suggested by Wijmstra) or to Picea abies (as suggested by present distribution of this species in the Rhodopes range, see Fig. 4) remains unclear, but this is the only evidence of a Weichselian preLGM spruce range in the Balkan Mountains. On the other hand, in the entire upper Pleistocene succession of Ioannina (Tzedakis, 1994; Frogley et al., 1999) and Kopais, GR7, western Greece (Tzedakis, 1999), Picea is absent (Tzedakis, personal communication, 2000).

Fig. 8. Tentative map of maximum expansion of Picea abies in southern Europe during middle pleniglacial interstadials (55^35 kyr cal BP). Data from sites reported in Table 1. A glacio-eustatic sea level of 320 m has been adopted, which corresponds to the highstand related to MIS 3 (Lundberg and Ford, 1994; Antonioli and Ferranti, 1996).

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During middle pleniglacial interstadials, spruce is distributed both in the lowlands and in the mountain belts of southern Europe. Between ca. 46 and 40 kyr cal BP, Picea abies^Pinus cembra forests occupied a vegetation belt in the inner central Alps below 1800 m altitude (Passo del Tonale, I11, Baroni and Ravazzi, in preparation). In the Massif Central, Picea was present near Le Bouchet Lake (F3, 1200 m altitude), between s 40 and ca. 36 kyr cal BP (pollen zones 26^24, Reille and de Beaulieu, 1990). The occurrence of charcoal identi¢ed as Picea in the Hungarian lowland since 37 kyr cal BP (U4 and U5, Willis et al., 2000 ; see 4.2. Onset of the LGM) and in the lower Austria since 46 kyr cal BP (sites A17 and A18, Damblon et al., 1996; Haesaerts et al., 1996; Damblon, 1997a,b) suggests it already occupied this region during earlier interstadials. Thus, the range of Picea abies retreated from central Europe and expanded in southern Europe in the early pleniglacial. Maximum development was reached during the interstadials between 50 and about 38 kyr cal BP (Fig. 8). This middle pleniglacial pattern of Picea is the starting point for evaluating its LGM range. 4.2. Onset of the LGM In the westernmost part of its distribution, spruce makes its last regional appearance about 35 kyr cal BP before its local extinction from the Massif Central (Reille and de Beaulieu, 1990) and the Lyonnais (de Beaulieu and Reille, 1984). The few 14 C-dated records extending back before the LGM from the Adriatic Basin show a spruce decline between 45 and 41 kyr cal BP and discontinuous occurrence after 39^37 kyr cal BP (Bortolami et al., 1977; Mu«llenders et al., 1996). A similar pattern is evident in Slovenian pollen records (Culiberg, 1991; Sercelj, 1996), but there the lack of radiocarbon dates allows only a rough estimate of the time of spruce decline. In the Notranje Gorice record (SLO6, Slovenian lowland, Culiberg and Sercelj, 1980), the decline occurred between 40 and 29 kyr cal BP (Fig. 15). The highresolution records available from the volcanic region in central Italy (Follieri et al., 1998) also show a general decrease at 35^30 kyr cal BP,

modulated by several stadial^interstadial episodes (framed in the ‘Lazio Complex’). Spruce maxima follow periods of oak and mark the latter part of temperate episodes, as observed during St. Germain episodes. Some of these temperate phases still supported spruce in a low-elevation belt of the eastern Alps between 35 and 29 kyr cal BP. In the Italian PreAlps, spruce wood and pollen dated 29 350 [ 460 14 C yr BP and calibrated 34 250 yr cal BP was found in Val Caltea (I30, 900 m altitude, Fig. 3; Fuchs, 1969). In the Austrian Alps a Pinus^Picea parkland is reconstructed and dated between 35.7 and 35.1 kyr cal BP (Ramsau, A8, and Hohentauern, A14, Fig. 2): it occupied the valley bottom in the basin of the Enns River, but not the upper slopes (Draxler and van Husen, 1978; van Husen, 1989). In the Slovenian Alps, wood charcoal identi¢ed as Picea has been found in an archeological site of Aurignacian culture (Potocka Zijalka, SLO8, Brodar and Brodar, 1983). The Aurignacian cultural complex in eastern Alps is now dated between 42 and 33 kyr cal BP (Broglio, 1997). Later on, the spruce record was restricted to the lowlands and low foothills. In the Venetian Lagoon are remarkable spruce-pollen peaks (20^ 25%) in peat deposits dated 23 450 [ 500 14 C yr BP (27 600 yr cal BP; Bonatti, 1968; Bertolani Marchetti, 1967) and 21 750 [ 730 14 C yr BP (25 600 yr cal BP; Bortolami et al., 1977; Mu«llenders et al., 1996). These organic deposits formed in a closed environment, where reworking of sediments was minimal. These pollen values therefore demonstrate dense spruce stands during the ¢rst part of the LGM in the margin of the Adriatic depression, dried up by the glacio-eustatic regression (Fig. 9). In the Hungarian Plain, macroscopic charcoal ( s 0.2 cm3 ) identi¢ed as Picea is found up to 27.6 kyr cal BP (Willis et al., 2000). In lower Austria and south Moravia, Picea dominates microscopic charcoal assemblages from humic and gley layers, embedded in loess sequences up to 27 kyr cal BP. Several hundred of microscopic charcoal fragments were identi¢ed, and some of them dated by AMS (Willendorf II, A17; Schwallenbach, A18; Pavlov, CZ1 ; Dolni Vestonice, CZ3 ; Kneblova, V., 1954; Haesaerts

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Fig. 9. Tentative range map of Picea abies in central and southern Europe during the LGM (25^17.5 kyr cal BP). Glaciers and ice sheets maximum expansion (shaded) from Habbe (1995) and Giraudi and Frezzotti (1997). The Adriatic lowstand coast line is from Correggiari et al. (1996). Data from sites reported in Table 1.

et al., 1996; Damblon, 1997a,b). However, Willis’ and Haesaerts’ identi¢cations lack a detailed diagnosis to distinguish Picea from Larix. The local occurrence of Picea in south Moravia at 30 100 yr cal BP is supported by pollen s 5% in a reconstruction of a Larix-dominated parkland, including Picea, Pinus sylvestris and Pinus cembra (Bulhary, CZ2, Rybnickova and Rybnicek, 1991). 4.3. Spruce survival at the LGM culmination The LGM began a phase of maximal reduction of the spruce range, with extinction in some districts. The spruce decline continued up to the late glacial period. 4.3.1. The Adriatic Basin and the Apennines Marine cores from central Adriatic provide a

record of the LGM, although the origin of spruce pollen is unclear. Core MD 90-917 from the southern Adriatic (Combourieu-Nebout et al., 1999) shows continuous low values of spruce pollen from s 20 to 18.7 kyr cal BP. The overlying interval lacks spruce pollen (520^400 cm) and marks a period when other conifers (Abies, Pinus) declined and Artemisia^Ephedra^Chenopodiaceae steppe elements expanded. Conifers may have occupied refuges, and semideserts may have reached their maximum expansion in the Adriatic Basin. By correlation of cores MD 90-917 and KET 8216 (200 km NE, Fig. 2), Combourieu-Nebout et al. (1999) provided age boundaries of 16.8 and 14.1 kyr cal BP and related the interval 520^400 cm to the Oldest Dryas or to Heinrich event H1. Accordingly, the retreat of conifer trees to their minimum range in the LGM in southern Europe

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would have lasted up to the late glacial. The chronology of core MD 90-917 has been thoroughly investigated by Siani (1999) and Siani et al. (2000), who provided a set of 12 AMS radiocarbon ages from the MD 90-917 core and discussed in detail the e¡ect of the reservoir 14 C correction. The last occurrence of Picea is framed between the (reservoir-corrected) ages 18 300 [ 150 and 16 020 [ 130 yr 14 C BP, which are calibrated 21 800 and 19 170 yr cal BP. The subsequent late glacial spread of conifers is re-dated 14 100 yr cal BP. Therefore, the phase of conifer contraction spanned a time interval longer than that estimated by Combourieu-Nebout et al. (1999), between the end of the LGM culmination and the late glacial interstadial onset. Other pollen records from Adriatic cores (I52 ^ Fig. 7 ^ and I53, Lowe et al., 1996, taken farther to the north in the Central Adriatic, and I54^I55, Gru«ger, 1975, taken to the south) show a similar pattern of conifer contraction during the LGM and alpine deglaciation. Unfortunately, no ages are available for the LGM interval and the beginning of late glacial. The LGM decline of conifer forests in central Italy is also shown in pollen records from volcanic lakes. Among them, the record at Lagaccione shows the highest spruce pollen percentage in zones LGC-11 and -13 (lowermost Lazio Complex in Table 2). There the Picea curve ends between 24.3 and 19.2 kyr cal BP (Magri, 1999). On the whole, spruce withdrawal from central Italy and the Adriatic Basin towards the present-day relict area in northern Apennines took place about 23 kyr cal BP (based on Follieri et al., 1998). Spruce probably survived the LGM in the (northern) Apennines. This hypothesis was proposed by Chiarugi (1936) but ignored by Huntley and Birks (1983) and Lang (1994) in their studies of late glacial and Holocene plant distribution of Europe. Several arguments support the spruce refugia in the northern Apennines. Bertoldi (1981) reports a continuous record of spruce pollen between late glacial and early Holocene from a mountain lake in the northern Apennines (Lagdei site, I37, 1250 masl, Fig. 10). Pollen maxima (9%) occur in zone A of Lagdei record, referred to BOlling/Older Dryas pollen zones by Bertoldi.

Lowe (1992) revised the stratigraphy of this record and attributed zone A to the late glacial interstadial (13^11 kyr 14 C BP according to Lowe, 1992), as found in the Prato Spilla records (I38, 1250^1550 masl, 10 km from Lagdei site). However, Prato Spilla registers only sporadic spruce pollen. Actually, the Lagdei pollen zone A does not resemble any of the zones described by Lowe (1992). AP/NAP and Artemisia curves suggest that Bertoldi’s zone A pre-dates local reforestation, which in turn is older than the base of the Prato Spilla record, dated 12 360 [ 55 yr 14 C BP (Lowe, 1992), i.e. 14.35 kyr cal BP. Radiocarbon ages from Prato Spilla sections are probably affected by reservoir e¡ects, as evidenced by the fact that the 14 C boundary ages of YD are 500^600 14 C yr systematically older than expected. A third interpretation is proposed for Lagdei zone A in Fig. 10, where it is included in the Alpine deglaciation phase. Accordingly, it is concluded that autochthonous spruce was present in the region (the northern Apennines of Parma) shortly after the LGM culmination and before 15 kyr cal BP. The depletion of spruce pollen at time of reforestation suggests that tree expansion favored mixed Abies ^ broad-leaved forests (Lowe, 1992), and not spruce. The data suggest a reduction of the Apenninic spruce range from the LGM up to the late glacial interstadial. This reduction may lead one to think that living spruce stands (VI in Fig. 4), con¢ned on the highest massifs of northern Apennine (Ferrarini, 1962, 1977; Magini, 1972), represent a ‘glacial relict’ (Magini et al., 1980). In fact, they have a direct phylogeographic linkage with late glacial populations (Giannini et al., 1994). Nevertheless, as shown by Chiarugi (1936, Fig. 13), the present development of spruce in the northern Apennines is the result of the middle to late Holocene spread of forest in an oceanic climate instead of a progressive relictual process. The Holocene expansion followed unfavorable dry periods during YD and early Holocene (Watson et al., 1994) responsible for a substantial contraction of the widespread LGM range throughout the northern Apennine. A controversial question is whether this Holocene expansion included regional migrations. This point is discussed in 5.2. Comparing

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Fig. 10. Simpli¢ed pollen diagram from Lagdei, I37, northern Apennine Mountain (redrawn from Bertoldi, 1981).

paleobotanical and phylomolecular evidences for alpine migration routes. 4.3.2. The Italian Alps and the Po Plain It is commonly agreed that spruce did not maintain LGM refugia in the Italian Alps, because north Italian spruce populations did not take part in the late glacial reforestation (Kral, 1979). New data from the Italian border of PreAlps (NE Italy) suggest spruce occurrence at the end of the LGM. A trunk recently found in £uvioglacial deposits covered by till on top of the outermost moraine of LGM age in the piedmont glacial amphitheatre of the Piave glacier (Colle Umberto site, I27, Figs. 2 and 3) is dated 17 670 [ 320 yr 14 C BP (Bondesan, 1999), i.e. 21 000 yr cal BP. This individual settled in the £uvioglacial plain and was transported by the gla-

cier. As already pointed out, the distinction of Picea from Larix wood from microscopic characters (Bartholin, 1979) is not easy, but a Picea identi¢cation of this trunk is probable, according to ray tracheids bordered pit features (Pignatelli et al., in preparation). No pollen record is available to document the extent of spruce stands in this proglacial area, but the paleobotanical site of Fornaci di Revine (I25), only 5 km from Colle Umberto (Figs. 2 and 3), may help. The Revine clay pit was open to the inner slope of a moraine related to the LGM (Casadoro et al., 1976). It provided a series of Larix trunks, dated between 18.1 and 17.1 kyr cal BP (Casadoro et al., 1976; Friedrich et al., 1999), on which a tree-ring chronology for the alpine deglaciation is currently in progress at the Hohenheim laboratory. At Revine, macroscopic remains of Picea are absent (Frie-

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drich et al., 1999; Martinelli, personal communication, 2000). A pollen record coeval to larch trunks (Casadoro et al., 1976) shows a Pinus, Larix, Betula, Salix, Artemisia, and Gramineae assemblage, characteristic of a taiga parkland. Picea pollen has a discontinuous record, suggesting long-distance transport. However, in the upper part, dated 17.2 kyr cal BP, it increases to 3^ 4%. As already discussed, these pollen values in-

dicate its local occurrence in the region (Bortenschlager, 1970; Hicks, 1986, 1994), and are similar to those from the base of the Palughetto di Cansiglio record, a neighboring site (I26, Figs. 3 and 11) that spans the subsequent time interval (since 15.4 kyr cal BP). Cansiglio is a large mountain karstic plateau (25 km2 , 1000^1400 masl) rising above the Venetian^Friulian plain. During the LGM, the glacier dammed a small intermorainic

Fig. 11. Simpli¢ed pollen diagram from Palughetto di Cansiglio, I26, southern Italian Pre-Alps, and radiocarbon time control.

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lake ^ the Palughetto ^ at the northern border of the plateau (Avigliano et al., 2000; Peresani et al., 2000). The Palughetto ¢lling succession consists of ¢ne-detrital deposits overlain by peat. The basal peat is a moderately humi¢ed litter made up of needles, cones, and branchlets of Larix decidua, twigs and wood of Betula and Alnus catkin scales, as well as mosses. Cones and other remains of spruce are also present. Upward in the stratigraphical section, vegetative parts of spruce and larch are dominant (e.g. branches, stumps in situ and crushed trunks). This deposit is an accumulation of trees in situ, likely killed during waterlogging phases (Avigliano et al., 2000). Fig. 11 shows a simpli¢ed pollen record from the interval 40^210 cm depth, including peat initiation (116^ 114 cm, 14 362^14 030 yr cal BP). From the base of the diagram upwards, Picea has a continuous curve with low values. The establishment of Larix^Picea forest is marked by an important step at 122 cm depth, e.g. 6^8 cm below peat initiation (interpolated age: 14.9 kyr cal BP), when spruce pollen reaches 4%, very close to the ¢rst spruce cone at 119 cm. A cone collected 5 cm above the basal peat contact provided an age of 14.0^14.3 kyr cal BP. This is the oldest late glacial cone of Picea abies Karst. in southern Europe. The macrofossil record of spruce extends up to the beginning of the Holocene. More than 50 cones have been analyzed from late glacial levels (Fig. 12) and identi¢ed as Picea abies var. europaea Jurkev. et Parf. (Avigliano et al., 2000), according to the

155

cone-scale shape and cone dimensions (SchmidtVogt, 1977; Schmidt, 1989). The succession of sites in the eastern Italian Pre-Alps (Val Caltea, Venetian Laguna, Colle Umberto, Revine, and Palughetto di Cansiglio) allows one to trace a possible history of Picea between 34 and 11 kyr cal BP at the southern Alpine border. Picea stands populated the lower mountain belt of the Pre-Alps at 34 kyr cal BP. Between 27 and 25 kyr cal BP, spruce was present in the eastern Po Plain and still at the alpine border 21.0 kyr cal BP. After leaving the Piave glaciated area between 21.0 and 18.5 kyr cal BP, it did not re-immigrate soon after deglaciation (as Larix and Pinus sylvestris did) but maintained a signi¢cant in£ux in the pollen rain. In the £uvioglacial plain a continuous curve is re-established about 17 kyr cal BP. Colonization of spruce and larch on the mountains (Cansiglio) surrounding the Piave glacial amphitheatre occurred at about 14.9 kyr cal BP, and forest settlement was coincident to the time of peat initiation, and close to the beginning of the late glacial interstadial, 14.3^14.0 kyr cal BP. Spruce was present in this mountain area throughout the late glacial interstadial and the YD and became the dominant forest tree during early Holocene (Fig. 11). The hypothesis that spruce survived during the last glaciation in the eastern Po Plain was already raised by Lona (1957), who reported a continuous record of spruce pollen during the ‘Late Wu«rm’ ( = pleniglacial and late glacial) in a basin close to

Fig. 12. Spruce cone at 80 cm depth of Palughetto di Cansiglio record (cf. Fig. 11). Three cones from this level were dated 11 605 [ 85 yr 14 C BP.

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the Euganei hills (Arqua' Petrarca, I22). However, pollen levels dating to the Alpine deglaciation displayed low spruce percentages, derived either from small populations of spruce locally or from long-distance transport. 4.3.3. Slovenja, Carinthia and the northern Balkans Spruce is commonly thought to have survived the LGM in the Slovenian foothills (Huntley and Birks, 1983; Culiberg and Sercelj, 1980; Sercelj, 1996). Available pollen records in the lowland show spruce decline before the LGM onset and no continuous record up to the late glacial interstadial (Notranje Gorice, SLO6, Culiberg and Sercelj, 1980). A log, identi¢ed as Picea, was found in the foothill region near the Italian^Slovenian border and dated at 18 790 [ 300 yr 14 C BP (Anhovo, SLO2, Sercelj, 1981), i.e. 22.3 kyr cal BP. This might represent either a survival spot or the latest relict population subsequently extinct at 22^19 kyr cal BP. Evidence of local spruce during the LGM in southeastern Alps is provided by the La«ngsee pollen record, in the Drau valley in Carinthia (A11, 540 masl, Schmidt et al., 1998). During the pleniglacial, the area was covered by the Drau glacier that formed a moraine-dammed lake (Van Husen, 1976). The sediment record spans the late pleniglacial and the alpine deglaciation. Before the de¢nitive spruce occurrence at the site at about 14 kyr cal BP (Fig. 15), a Picea pollen of s 4% is found in zone PZ1/3, older than 15 535 [ 160 yr 14 C BP, i.e. 18.6 kyr cal BP. This zone may tentatively be framed between about 21 and 19 kyr cal BP. In this interval, the reconstructed landscape is a Pinus^Larix^Picea taiga with widespread Juniperus understory. This record suggests local survival of spruce during the late Pleiniglacial in parkland formations in the lower mountain belts of eastern Alps. The basal zone of the La«ngsee record ( s 21 kyr BP) shows further, remarkable Picea peaks (up to 22%). Because of low pollen concentration and prevalent detrital input by glacial melting, this pollen may originate from reworking interstadial deposits (Schmidt et al., 1998), as argued in other situations in Carinthia (Fritz, 1978). The Picea records in La«ngsee need further investigation, currently in progress.

Early appearance of spruce in the Ljubljana basin at 14.5^14 kyr cal BP (SLO7, Culiberg, 1991 ; Jerai, personal communication, 2000), in the mountain belt of the Julian Alps and Carinthia during the late glacial interstadial (Ledine, SLO4, Culiberg et al., 1981; Lengholz, A13, Fritz, 1965) suggest it survived in several areas during LGM. Farther south in Croatia and Bosnia^Hercegovina, available data are scarce. The existence of survival areas in the northern Balkans is suggested by the early continuous appearance of Picea pollen in late glacial and early Holocene pollen records from near the Dalmatian coast (Fig. 2): Neretva valley (HR2, Brande, 1973), Valun Bay (HR3, Schmidt et al., in press), and Lake Vrana (HR4, Schmidt et al., 2000), very close to mountain ranges. Although late glacial Picea pollen representation remains 6 4% in these coastal sites (2^3% in the late glacial interstadial at Lake Vrana), and local spruce presence is not documented, spruce populations in the mountains are probable. 4.3.4. The Hungarian Plain, Romanian Carpathians, lower Austria, Moravia Lang (1994) and Willis et al. (1995, 1997, 2000) suggest that Picea survived the full glacial maximum in the Hungarian Plain. The pollen records of Batorliget and Kis-Mohos To¤ in the eastern plain (Fig. 2) provide unequivocal evidence of Picea^Pinus forests before 14.6 kyr cal BP (Willis et al., 1995, 1997). The occurrence of widespread spruce in the Hungarian Plain between 35 and 27 kyr cal BP is shown by identi¢cation of charcoal in buried paleosols in loess (Willis et al., 2000). Although older authors mention Picea^ Larix wood and charcoal from the intermediate period between 27 and 19 kyr cal BP (Ve¤rtes, 1964; Dobosi, 1967; Geyh et al., 1969), these non-speci¢c identi¢cations do not eliminate the doubt of prevalent or exclusive Larix occurrence at the LGM in the Hungarian Plain instead of spruce. A pollen record through the LGM is not available in this area, and the hypothesis of Willis et al. (2000) for widespread spruce in ‘the Hungarian landscape during the last full glaciation’ is limited to the interval before 27 kyr cal BP.

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Fig. 13. Composite pollen diagram from Lago di Greppo, 1442 masl, and Lago di Braccioli, 1295 masl, northern Apennine Mountain (from Chiarugi, 1936). The pollen sum includes arboreal pollen only. Quercetum mixtum = sum of Quercus, Tilia, Ulmus.

Spruce persistence through the LGM in the eastern Hungarian Plain is demonstrated by the continuous high percentages (5^20%) of spruce pollen at sites of Batorliget (Fig. 2, H6, Willis et al., 1995) and Kis-Mohos To¤ (Fig. 2, H7, Willis et al., 1997) between 17 and 11 kyr cal BP. From these records it appears that Picea grew in the eastern part of the Hungarian Plain during the LGM and late glacial. Spruce was then replaced in the early Holocene by the spread of broadleaved forests. In the neighboring Carpathian Mountains, Picea immigrated during the late glacial interstadial but its expansion dates to early or middle Holocene (Pop, 1971; Ralska-Jasiewiczova, 1980; Farcas et al., 1999). At Iezerul Calimani (RO2, 1650 masl), spruce pollen is recorded between 18 and 17.5 kyr cal BP (2^4%), then it declines in abundance before increasing again in the Holocene (Farcas et al., 1999). This pattern may suggest that thinned spruce populations survived the pleniglacial in some mountain districts of the Romanian Carpathians, but did not withstand the late glacial. Several loess sequences in lower Austria and in

Moravia, mentioned in the previous section, are interrupted at the beginning of the LGM, thus they provide only limited insight about tree persistence during LGM in these areas. The upper cultural layer at sites of Pavlov II (site CZ1) and Willendorf II (site A17) contain microscopic charcoal mostly assigned to Picea, dated between 25.8 and 23.2 kyr 14 C BP, i.e. 30.2 and 27 kyr cal BP (Damblon et al., 1996; Haesaerts et al., 1996; Damblon, 1997b). 4.3.5. The southwestern Balkans and Rila^Pirin^ Rhodopes Mountains The history of spruce range in Bosnia^Hercegovina, Montenegro, Albania, and Macedonia (V in Fig. 4) is poorly known. Surprisingly, the several Pleistocene and Holocene pollen records from northwestern Greece (Bottema, 1974; Willis, 1992a,b,c, 1994; Tzedakis, 1994, 1999; Gerasimidis and Athanasiadis, 1995) do not report any spruce pollen, even though they are close to the present southern limit of spruce. As discussed by Willis (1992c), the present spruce range in these regions is better explained as the result of conifer expansion at the beginning of the Atlantic Period

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(between 8.2 and 7.4 kyr cal BP), instead of a relict from pre-Holocene time. A similar hypothesis has been raised for the Bulgarian mountain ranges (Bozilova and Tonkov, 2000). However, the Tenaghi Philippon pollen diagram (see 4.1. From Eemian to LGM) supports the occurrence of Picea sp. during early and middle Weichselian in the Rhodopes^Rila^Pirin Mountains. Although no records extending through the LGM are available, the occurrence of spruce in the Vitosha Mountain during the late glacial interstadial is shown by Filipovitch (1982). In the Pirin Mountain, spruce is absent until the beginning of the Atlantic Period (Stefanova and Bozilova, 1995). This early postglacial spruce record suggests either autochthonous stands (as assumed in Fig. 9) or immigration from the northern Siroka and Vracanska ranges. As observed in the northern Apennine and in western Balkans, the presentday distribution of spruce in this district (region V in Fig. 4) results from the mountain expansion of conifer forests (Bozilova, 1975a, 1995; Bozilova and Tonkov, 2000; Filipovitch, 1982; Huttunen et al., 1992) in an oceanic climate, starting about 8 kyr cal BP.

4.4. An updated outline of late glacial and Holocene spruce migration in the Alps The calibrated 14 C chronology concerning late glacial and Holocene spruce immigration in the Alps is presented in Figs. 14^16. The method used for recognition of levels marking the immigration of Picea from percentage pollen records is presented and discussed in 2. Materials and methods. Information about the sites selected for this ¢gure is in Fig. 2 and Table 1. Several authors (Markgraf, 1970; Kral, 1979; Huntley and Birks, 1983; Lang, 1994) have proposed a general trend from east to west to south in the western Alps, i.e. the general pattern of spruce migration followed the main structural units of the Alpine chain. Reconstructed timing and pathways shown in Figs. 14^16 show considerable departure from this picture. Eastern Alps were already occupied by spruce at the beginning of the late glacial interstadial (Fig. 16), i.e. well before the beginning of the Holocene warming (Kral, 1980, 1982, 1989; Huntley and Birks, 1983). This early establishment was already suspected by Bortenschlager (1970), considering that

Fig. 14. Calibrated 14 C chronology of the late glacial and Holocene spruce migration in the Alps and Alpine forelands (symbols as in Fig. 2). The map reports the age of the stratigraphical depth where the Picea pollen curve passes the value of 4% (site information in Table 1). Dashed line: present distribution boundary (district III in Fig. 4). Grey shading indicates elevation s 200 masl south of the Alps and s 500 masl north of the Alps (in dark gray: elevation s 3000 masl). Symbols as in Fig. 2. An arrow indicates a small spot in the Apennine mountain belonging to district VI in Fig. 4.

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Fig. 15. The Picea percentage curve plotted against age from the pertinent levels of selected diagrams from the Alpine region and Alpine forelands, ordered according to an east^west gradient. The westernmost site (Le Lac Long) belongs to the Maritime Alps. An arrow shows the level where the 4% threshold is reached. Scissors indicate truncations in pollen diagrams. Sites are listed in Table 1.

even spruce pollen 6 5%, observed at several lateglacial records in the eastern Alps, still indicates local populations (cf. 2. Materials and methods). Recent macrofossil ¢nds from the Cansiglio Plateau provide support for this hypothesis. A late glacial migration pathway from Slovenia and Dinaric Alps, previously proposed (Schmidt-Vogt, 1977; Kral, 1982; Huntley and Birks, 1983; Lang, 1994), is contradicted by spruce survival during the phase of Alpine deglaciation at the border of the Italian Pre-Alps. Moreover, the Cansiglio record shows that spruce immigration in the Venetian Pre-Alps is coincident with that of other species or even somewhat earlier. An abrupt late glacial rise in arboreal pollen to over

80% is common in all pollen records from the Alps (Schneider and Tobolski, 1983; Ammann, 1989; Ammann and Wick, 1994; Orombelli and Ravazzi, 1995; Wick, 1996; Gehrig, 1997) and the northern Apennines (de Beaulieu et al., 1994b). This increase suggests that mass expansion of trees occurred forming forests. The radiocarbon ages for this event (12.2^12.5 kyr 14 C BP calibrated 14 700^14 050 yr cal BP) are close or slightly younger than the age of the ¢rst late glacial abrupt warming as estimated by varves in central Europe (14 450 yr cal BP, Brauer et al., 1999; Litt et al., 2001) and by annual ice layers from Greenland (14 700 yr cal BP, Meese et al., 1997). Therefore, it is proposed that mass expan-

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sion of trees, including spruce in eastern Pre-Alps, represents a direct response to the late glacial warming. The available data do not make clear the range of Picea at the end of the late glacial and fail to document the e¡ects of YD on Picea dynamics in the Alps. However, in the several pollen records from the eastern Pre-Alps (Cansiglio, I26; Fusine, I33; Somodogna, I32; Lengholz, A13; La«ngsee, A11), Picea immigration during the YD is not evident, and in sites where spruce was present decreases little (Bortenschlager, 1970). Likely, the YD resulted in a standstill phase. At the end of the YD, spruce was still absent west of longitude 12‡E on both side of the Alps. Between 10.8 and 9.5 kyr cal BP Picea rapidly expanded throughout the eastern and central Alps. An average migration rate of 100 m/yr can be estimated for this time interval (Ravazzi and Pini, submitted), a value consistent with the maximum (250 m/yr) reported by Lang (1994). Zoller (1960) suggested a delay in the immigration along the southern border of the Alps and concluded that the invasion of the Swiss Plateau involved populations following a northern route through the Inn valley. This view has been supported by Kral (1979), Welten (1982) and Burga (1982, 1988), who provide detailed discussion about the time when the treeline altitude was crossed at the alpine pass. Yet recent investigations on the Ital-

ian side of the central and eastern Alps have excluded important time lags related to a barrier produced by the Alpine ridge. Instead, the migration front appears to have been compressed in longitude, and no time lag is evident in the southern Alps (Figs. 14 and 16). In the Carinthian Plain and eastern Pre-Alps, Picea migration precedes the spread of broad-leaved trees and shrubs (e.g. La«ngsee, A11; Rohr, A16). Contrary, in central Alps, Quercus, Ulmus, Tilia and Corylus expand earlier than Picea. There, spruce migration took place in a subalpine belt (e.g. 1500^2300 m altitude), because lower belts supported dense broad-leaved forests (Tallentire, 1973; Lang, 1994; Oeggl and Wahlmu«ller, 1994). Moreover, because of its upper thermal limit (0‡ mean January, 18‡C mean July; Bernetti, 1995; Zagwijn, 1996), spruce could not expand downward during the early Holocene warm period. The evidence of a compact front is consistent with a high-altitude migration pattern and renders the hypothesis of distinct pathways through the northern and southern Alpine borders invalid. Most alpine passes of the inner eastern and central Alps are set in the subalpine^lower alpine belt, and during the hypsithermal interval (e.g. 10^7 kyr cal BP) seed and pollen dispersal outcrossing was easy for spruce. However, during the middle and late Holocene the migration pattern changes radically. In some parts of the Swiss and western Alps, the

Fig. 16. Western boundary distribution of spruce in the Alps at 13.5, 9.0, 4.5 kyr cal BP (coarse lines) and present distribution (dotted pattern). Data from Fig. 14. Symbols as in Fig. 2. Grey shading indicates elevation s 200 masl.

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range of spruce seems to have stabilized for several thousand years, but sudden invasions also occurred (e.g. the Swiss Plateau: Markgraf, 1970). The causes for these reduced or unstable migration rates have been widely debated. Several factors seem to have been involved: the competition with other tree species, notably Abies (de Beaulieu et al., 1993), and anthropogenic deforestation coincident with agricultural activities in the last 5000 yr (Markgraf, 1970), and climatic change. These latter two combined e¡ects are probably responsible for the expansion of conifers during the last 8000 yr in the central Alps and in Switzerland (Markgraf, 1970; Tallentire, 1973 ; Pini, submitted, see Fig. 15). The dynamics of altitudinal belts along a north/south transect through central Alps are of special interest in this respect. During early Holocene, Picea abies and/or Abies alba forests were con¢ned to the upper altitudes (e.g. above 1300 masl) of inner valleys (I9^11, Gehrig, 1997 ; Pini, submitted), but in the southern Pre-Alps they grew over 200 masl (I4^6, I31^33, Kral, 1980; Wick, 1996; Tinner et al., 1999; see also Fig. 11). Rainfall regimes may account for these di¡erences, because the Pre-Alps are under oceanic in£uence, and inner valleys experience continental conditions (Ozenda, 1985). The expansion of conifer forests to the inner valleys £oors at about 8.2^8 kyr cal BP may be related to an increasing oceanic in£uence at the beginning of the Atlantic Period (Tinner et al., 1999) or at the 8.2-kyr cal BP event (von Grafenstein et al., 1998). A similar climatic control on Picea expansion has been observed in the Apennines and Balkans during the Holocene (see previous section). The combined physical and climatic e¡ects of the highest reliefs might be envisaged in Valle d’Aosta. A striking delay of spruce immigration there (Brugiapaglia, 1997) may be related to the position of the Mischabel^Monte Rosa^Monte Bianco massifs (Pennine Alps), reaching the highest altitudes in the Alps and forming a continuous range over 3000 m altitude (Fig. 14, in dark gray). This ridge probably acted as a physical and climatic barrier to migration southward and westward because of high elevations of passes and of the continental climate of inner valleys, where a

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spruce belt is nowadays missing (Ozenda, 1985). A well-documented migration route took place on the north side of the Pennine Alps, through the Jura and northern French Alps (Wegmu«ller, 1977; de Beaulieu et al., 1993). A possible second route through the Italian side of southwest Alps is poorly documented by paleobotanical records. Sites at the southwestern border of the Alps (Maritime Alps) display continuous spruce pollen representation earlier than expected from the timing of a migration from the north (5.3 kyr cal BP at F14, Kharbouch, 2000; about 5 kyr BP at I0, Finsinger, in press ; 4.2 kyr cal BP at F13, de Beaulieu, 1977 ^ Figs. 14 and 16). The genetic structure of recent spruce populations from this area would also exclude relationships with the northern provenances (see 5.2. Comparing paleobotanical and phylomolecular evidences for alpine migration routes).

5. Discussion 5.1. Population dynamics, ‘glacial refugia’, and climate The extent of ice and permafrost cover in Europe during the late Pleistocene cold phases limited the suitable areas for thermophilous trees to southern Europe (Bennett et al., 1991; Willis, 1996). It is currently believed that temperate tree species persisted in small areas of refugial character surrounded by regions with unfavorable climate. The term ‘glacial refugia’ is commonly used to indicate survival areas during the entire time span of the last glacial period (e.g. from about 70 to 10 kyr). ‘Glacial refugia’ of temperate trees occurred in the mountains of the Balkan, Italian and Iberian peninsulas at higher altitudes, because of higher moisture levels (Beug, 1967; Bottema, 1974; Huntley and Birks, 1983; Bennett et al., 1991). In southern Europe dryness would have limited the distribution of cold^temperate conifer trees (Abies, Picea, Pinus pro parte), in spite of their resistance to low temperatures. Thus some authors have ¢gured a ‘glacial refugial’ pattern even for cold-resistant conifer trees (Huntley and Birks, 1983; Lang, 1994; Combou-

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rieu-Nebout et al., 1999). The concept of glacial refugium as drawn by some authors (Willis, 1996) assumes a simple pacemaker-like response of tree species to glacial^interglacial cycles, i.e. expansion towards northern latitudes occurs during interglacials, but no inverse way back to ‘glacial refugia’ occurs at their end. At the end interglacial populations became cut o¡ and died in situ, as no mechanisms for driving back thermophilous £ora from the north can be traced (Bennett et al., 1991). The data presented in this paper show that Picea abies distribution experienced a complex history during the last glacial period. The maximum expansion is not of interglacial age, and the maximum contraction does not coincide with full glacial conditions. In order to assess the mobility of this species through time, it is helpful to provide a brief summary on its speci¢c ecological boundaries in cold environments. Norway spruce withstands 1^1.5‡C annual average, and a frost resistance till to 336/ 338‡C, with a minimum annual rainfall of 450 mm in coldest climates (Schmidt-Vogt, 1977; Tranquillini, 1979). These limits are higher compared to boreal species from North America (P. mariana, P. glauca) that occur in dry and cold climates (Thompson et al., 1999). The upper alpine altitudinal limit of spruce is 2100^2350 masl (Tranquillini, 1976, 1979; Schmidt-Vogt, 1977; Holtmeier, 1993; Andreis et al., 1996; Rossi et al., 2001). The lower temperature limit for viable seeds in cones is 32‡C at £owering time (Folladori, 2000). The behavior of spruce in the recent Holocene speaks to its pioneering ability in cold and open environments. At the present alpine treeline, the reproductive success of Picea abies and Larix decidua is similar, and both act as major pioneers on the deglaciated terrain after the Little Ice Age (Schmidt-Vogt, 1977; Matthews, 1992; Rossi et al., 2001; Folladori, 2000). Conversely, in forest environments, competition with other trees, notably Abies, is a main factor accounting for reduced migration rates (de Beaulieu et al., 1994; Brugiapaglia, 1997; Burga, 1988). In open environments, the reduced competition among forest species favors the population dynamics and migration of pioneering species, in-

cluding spruce. Open vegetation also increases the average limits of pollen and seed dispersal of wind-pollinated and anemochorous species (Van der Pijl, 1982). Therefore, a reduction in density of individuals may enhance long-distance pollination and outcrossing. These mechanisms may limit the loss of genetic variability (richness in alleles) within residual populations of restricted size. All together, these facts suggest that cold periods were favorable for dispersal, migration, and adaptation of spruce populations. However, compared to Pinus and Larix, Picea is more sensitive to extreme continental climate and particularly to winter desiccation (Tranquillini, 1979; Sakai and Larcher, 1987). Reduced snow cover and strong winter winds eliminate spruce from the timberline ecotone in the Alps (Tranquillini, 1979). Also, spruce does not take part in the forest/steppe ecotone in the Eurasiatic boreal zone (Komarov, 1968; Walter and Breckle, 1986). Thus, the match of ecological requirements to habitats reached by diaspores is a main factor controlling species distribution. It is therefore appropriate to assess the main distribution changes by analyzing associated habitat changes. The paleobotanical records presented in previous sections of this paper show that the spruce range expanded in southern Europe during the middle pleniglacial interstadials. There is no reason to exclude regional and southward migration at that time (e.g. from the western Alps to the Adriatic Basin and to the Rhone valley, or from the northern to central Apennines to the coastal region, or from the Carpathians to the Pannonian plains). This south-centered maximum distribution (including Alps, northern and central Apennines, Po Plain, terrestrialized part of the Adriatic Basin, northern Dinaric Alps, Hungarian Plain, Carpathians) was longitudinally continuous, and probably no geographic breaks existed eastward to the range of spruce in Russia (Fig. 8). Plains were occupied by open boreal forests, e.g. a taiga parkland, dominated by Pinus sylvestris, Pinus cembra, Betula sspp., Juniperus sp., Picea abies and Larix decidua. Sedimentary processes, soil development, and the archeological record suggest that south and east of the Alps the middle pleniglacial interstadials experienced cool to cold^tem-

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perate climate, allowing soil development under forest cover (al¢sols) and no loess accumulation (Cremaschi, 1990; Cremaschi et al., 1990; Gerasimenko, 2000). In the Alps, there is evidence of peat accumulation at 1900 m altitude (Baroni and Ravazzi, in preparation). Conifer forests occupied a mountain belt up to over 1000 m altitude. Cool^ temperate conditions provided suitable habitats for enlarging conifer forests in the lowlands, building up corridors between the mountain regions of southern Europe: one of them is the northern Adriatic, partially dried up by glacio-eustatic regression, a second one was the lower Austria and Hungarian Plain. The later interval from 40 to 23 kyr cal BP is marked by declining spruce. New establishments are not recorded during this phase. In the Alps, the spruce decline was linked to timberline depression that occurred between 35 and 30 kyr, as suggested by comparing the Tonale (Baroni and Ravazzi, in preparation) and Enns valley records (van Husen, 1989). This decline was not progressive but modulated by stadial^interstadial events, as shown by oscillating pollen values in several records in central Italy (Follieri et al., 1998; Magri, 1999). Therefore it is suggested that the remaining populations were not strictly refugial in character: part of them may have resulted from new establishments under pressure of the £uctuating climate, during this time interval in Europe (Dansgaard et al., 1993; Watts et al., 2000). The density of populations thinned; however, several records considered suggest that a continuity of distribution along a longitudinal transect between the Alps, the Carpathians and Moldavia was maintained, at least up to 23 kyr BP. The Adriatic corridor was probably also still maintained. Of interest is the question raised by Combourieu-Nebout et al. (1999), suggesting an extreme reduction of conifer development in the Adriatic Basin during Alpine deglaciation phase due to extreme dryness. Because of spruce sensitivity to dry, continental climate, this may be regarded as the most critical period for its survival. Detailed paleoclimatic records available in central and southern Italy from varves, SST, pollen, lake levels, lake-sedimentation regimes, and glacial advances (Giraudi and Frezzotti, 1997; Kallel et

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al., 1997; Combourieu-Nebout et al., 1999; Huntley et al., 1999; Magri, 1999; Magri and Sadori, 1999; Ramrath et al., 1999; Cacho et al., 2001) suggest phases in which the climate was drier than and as cold as the LGM occurred between 23 and 14 kyr cal BP. Moderate glacial advances in the central Apennines are found to coincide with phases of increased winter runo¡ and decreasing arboreal pollen percentage in lakes between 21^18 and 14^12.8 kyr cal BP (Ramrath et al., 1999). This juxtaposition is explained by a combination of very dry summers and abundant winter precipitation. An extreme dryness during the growing season under semiarid climate (e.g. annual precipitation (cm)/mean annual temperature (‡C) 6 2, according to Barry and Chorley, 1992) probably contributed to spruce decline from the Apennine chain, the Adriatic Basin, the Po Plain, Slovenja, and the Romanian Carpathians during the Alpine deglaciation. This situation may have led to late glacial refugia. The location of residual populations during late glacial is discussed by Combourieu-Nebout et al. (1999). They suggest conifer refugia (Abies, Picea, and probably Pinus) on top of a Balkanic mountain belt of broad-leaved species occupying middle altitudes. Spruce pollen found in the Adriatic Sea deposits would originate from there, with the Adriatic depression being like a semidesert. Actually, the source of spruce pollen may be from the Po River discharge, the main river feeding the Adriatic. In fact, the conifer pollen from Adriatic Sea cores does not prove a refugial conifer belt at high altitudes in the Balkans, although survival there is plausible. Moreover, the data presented in previous sections are consistent with the survival of spruce populations in the Po Plain and in the northern Apennines during the Alpine deglaciation phase. The widespread loess cover between the Po Plain and the Dalmatian border at the LGM (Cremaschi et al., 1990; Cremaschi, 1990) along with arboreal pollen percentages from pollen records from Adriatic continental lowlands that are similar to levels recorded in pollen rain from present forest/steppe (Peterson, 1983), suggest that the forest/steppe ecotone had a wide extent in northern Italy at the end of LGM and the late glacial. Indeed the abundance of Picea char-

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coal in humic layers embedded in loess sequences from lower Austria, Moravia, Hungary, Moldavia and Siberia may suggest either that spruce took part to the forest/steppe ecotone (e.g. tree stands con¢ned to favorable micro-environmental pockets under semiarid conditions) or that marked £uctuations in wetness produced biome oscillations from tundra and steppe to boreal forest (e.g. open boreal forests extensively covering stable areas). A critical moisture balance for Picea can be suggested either in the Adriatic depression, because of its negative altitude at present sea level, or in the Pannonian basin, because of its higher continentality. Moister conditions, partially edaphic in nature, occurred along the river beds in the highest plains of the Po River basin and on cool slopes on the Venetian and Slovenian foothills at the border of eastern Pre-Alps. According to these interpretations, the minimum range reached by spruce in southern Europe was not during full glacial time, but rather during maximum continental dryness during the growing season. In this respect, it is helpful to consider the situation of spruce during the LGM in the Alps, where glacial advances were much more pronounced than in the Apennines. Some authors (Bortenschlager, 1970; Paganelli, 1996) postulated ‘glacial refugia’ of Picea abies in the Pre-Alps, particularly on south-facing slopes. As already shown, the paleobotanical record from the Italian Pre-Alps supports a LGM survival of Larix, Pinus sylvestris, and Pinus cembra (Maspero, 1996). Picea probably survived also in the Po Plain and in the eastern and Slovenian Pre-Alps. According to climatic reconstructions based on glaciological methods, low mountain belts of Pre-Alps could have o¡ered habitats compatible with spruce survival during the LGM. In the central Alps, the present spruce treeline lies about 800 m below the snowline (based on Pel¢ni, 1994; Folladori, 2000). Current climatic reconstructions suggest that the LGM in the Alps and the Apennines occurred under a cold and wet climate, especially south of the Alps (Giraudi and Frezzotti, 1997; Florineth, 1998). Assuming the LGM precipitation regime to be similar to present, the estimated ELA altitude during the LGM culmination in southern Italian Alps (1400^1650 m altitude,

von Klebelsberg, 1953; Fuchs, 1969; Carraro and Sauro, 1979) and in the central Apennines (1750 m altitude, Giraudi and Frezzotti, 1997) would have allowed Larix^Picea^P. sylvestris^P. cembra stands up to 600^800 m altitude in the Alps and 900 m in the Apennines. Under such a climate, there is no reason to assume a refugial character for spruce populations living during the LGM culmination at the border of the Alps and in the northern Apennines. 5.2. Comparing paleobotanical and phylomolecular evidences for alpine migration routes Lagercrantz and Ryman (1990) described the genetic structure of Picea abies populations from northern and central Europe by studying polymorphism and morphological variations in proteins. The observed pro¢le of genetic di¡erentiation displays a good correspondence with the postglacial history of migrating populations from the Carpathians and from Russian ancestors. The central European provenances display low levels of heterozygosity attributed to bottleneck e¡ects, that is, the genetic variability was lost due to restriction in population size in the Balkanic refugia (Lewandowski et al., 1997). Giannini et al. (1994) studied isozyme systems from several populations in the Alps and found levels of heterozygosity higher than those reported by Lagercrantz and Ryman (1990). The Italian alpine populations were found to show a higher degree of within-population variation than the populations from the northern side of the Alps, even if both groups were derived from the same ancestral genetic pool in the Balkans, as suggested by Huntley and Birks (1983). This assumption is retained by Vendramin et al. (1999) and Sperisen et al. (2001) interpreting the spruce mitochondrial DNA variation from natural populations of spruce in southern Europe. However, the present study has shown that the Italian Alpine populations probably derive from an original pool set in the Po Plain and the Adriatic Basin, which might have kept peculiar alleles distinct from those originated from the Pannonian region and the Balkans. The pattern found by Giannini et al. (1994) might therefore re£ect the contribution of several

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ancestral pools to the reforestation of di¡erent areas of the Alps. Giannini et al. (1994) also studied the population living in the Apennines and found it to be markedly di¡erentiated and provided with some unique alleles. The long paleobotanical history of the Apennine populations, extending back at least to the beginning of early Weichselian, allowed time to accumulate the observed amount of genetic di¡erentiation. According to Borghetti et al. (1988) and Giannini et al. (1994), the distribution of some alleles and cone variability points to a phylogeographic relation among Apennines and western Alpine populations. This hypothesis was recently corroborated by Collignon and Favre (2000), who studied the genetic structure of French spruces (French Alps, Jura, Vosges) using random ampli¢ed polymorphic DNA analysis. Populations from the French Alps show a latitudinal structuring, the northernmost provenances maintaining close molecular relationships to the Jura and Swiss spruces. Instead, the southernmost ones di¡er so much that they could not originate from the same genetic pool. Collignon and Favre (2000) suggest a postglacial migration route from the Apennines. Contrary to these results, Scotti et al. (2000), using sequencecharacterized ampli¢ed region markers, found an isolated population in the southwestern Italian Alps that could not be grouped with the Apennine genetic pool. They also observed a closer genetic relationship between the Apennine stand and the northeastern Alpine pools, although the former has features that could not be found in the Alpine populations. They concluded that the Apennine stand is a marginal population of the eastern Alpine lineages and rejected the hypothesis of a glacial refugium. Either Collignon and Favre (2000) and Scotti et al. (2000) limited their historical analysis to the postglacial history or overlooked previous events that could be re£ected in the present genetic structure of spruce populations. As shown in 4.3. Spruce survival at the LGM culmination, Holocene migration patterns in the Apennines are not documented. Since the LGM, spruce declined and did not play a role in the late glacial reforestation. Some pollen diagrams at high altitude in the Ligurian Apennines, where spruce is presently absent, report a middle^

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late Holocene continuous spruce-pollen curve (Agoraie, Braggio Morucchio and Guido, 1975; Lajone, Braggio Morucchio et al., 1978). Recent investigations of the same sites (Cruise, 1990; Guarisco and Ravazzi, unpublished data), however, lack spruce pollen. Therefore, a postglacial pathway from the northern Apennines towards the western Alps is not supported by the paleoecological record. On the contrary, the paleobotanical data favor an old (pre-LGM) relation between the Apennine, southern French, and the Po Plain spruces. As discussed in 4.1. From Eemian to LGM, 4.2. Onset of the LGM maximum and 4.3. Spruce survival at the LGM culmination, and shown in Figs. 10 and 12, the middle Weichselian spruce range was initially continuous across the Po Plain and the western Alps and the connection with residual Apennine populations was probably interrupted after the LGM. Even later, the reproductive interaction among spruces living in the northern Apennines and the northeast of Italy was probably relevant, because of the extreme size reduction of the Apennine stand at the late glacial end (see 4.3. Spruce survival at the LGM culmination) that enhanced the probability of gene £ow through pollen. Consequently the present genetic a⁄nities of spruce in the northern Apennine do not contradict its LGM survival there. Finally, the question raised with the status of genetic isolation observed in southeastern Alpine populations points to the need for further paleobotanical investigations in the Maritimes Alps, in the Ligurian Alps and in the Haute Provence to look for a possible center of survival so far neglected. Otherwise, the hypothesis of human introduction should be explored.

6. Conclusions The late Quaternary distribution of spruce in central and southern Europe has a complex history linked to environmental and climatic change. The history of Picea omorika, one of the two species presently living in Europe, could not be traced, because it is only sporadically represented in paleobotanical records (by its fossil relative Pi-

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cea omoricoides). The record of Picea abies is adequately documented by continuous paleobotanical records in some regions (central Europe, Massif Central, the Alps) since the late glacial. Elsewhere, and for older time intervals, the reconstruction is tentative. After a maximum expansion towards the Nordic regions during the early Weichselian, the western European range of P. abies was reduced in southern regions at the onset of the early pleniglacial. A new expansion occurred in southern Europe during middle Weichselian interstadials. In this time interval, spruce formed extensive forests in a continuous range in the lower mountain belts and in the plains between the Alps, the Carpathians, and Ukraine. The Apennines and Balkans were probably connected by a corridor through the Adriatic Basin. Starting from this middle Weichselian pattern, a general decline occurred during the LGM. However, it is suggested that a longitudinally continuous distribution of spruce between the Alps and the Carpathians was maintained at least up to 23 kyr BP. This distribution argues against a marked restriction of population sizes during most of LGM. The hypothesis of restricted ‘glacial refugia’ in the northern Balkans and Carpathians, previously proposed, is inappropriate. Phases of marked conifer withdrawal seem to be unrelated to full glacial conditions in southern Europe, because the maximum glacier extent is not related to maximum continental dryness during the growing season, which is the most critical period for spruce survival. Dry maxima probably occurred during the alpine deglaciation, between the end of the LGM and the onset of the late glacial interstadial. Spruce took part in the late glacial treemass expansion in the eastern Alps and Carpathians, whereas it failed to spread in the Apennines and in the Pirin^Rila^Rhodopes Mountains. This pattern might re£ect the distribution of wet climates, which in turn is related to the in£uence of oceanic air masses and to altitudinal variations. In the southernmost mountains of Europe, the expansion of the Abies^Picea forest is limited to the middle^late Holocene, due to increased climatic humidity. Climate and location of residual populations in£uenced the migration rates and directions of

postglacial dispersal of spruce over the alpine region. The late glacial spruce expansion in the Alps is coincident with or very close to the abrupt warming that occurred at 14 700^14 500 yr cal BP in the northern hemisphere. During the early Holocene warm interval, spruce expansion reached maximal migration rates at high altitudes (e.g. 2000 masl) in the Alps. In the second part of the Holocene, its migration rates decreased, in response to the climatic e¡ects of the Neoglaciation and to ecological competition with other tree species (notably Abies) encountered along way. The middle^late Holocene spruce expansion in the Alps also points to the role of people and of the local orographies and climates, as shown in the region of highest elevations of the Mischabel^ Mt. Bianco^Mt. Rosa Massifs. Because of the fragmentary nature of the fossil record and of the low pollen representation of spruce, its past distribution has often been underestimated. Some of its past range has been ignored, like the populations living in the Apennines. Spruce has a long history (pre-LGM) in the Apennines: this explains the status of marked genetic di¡erentiation of the living populations. Because of the low number of well-dated paleobotanical records in northern Italy, the late glacial expansion from the border of the Po Plain through the eastern Italian Pre-Alps was unknown to previous authors. At some places in the Alps, the age of immigration occurred earlier than previously estimated, so that a supposed delay in the immigration along the southern border of the Alps is no longer supported. Comparing late Quaternary history of spruce with patterns of genetic variation among living populations is a di⁄cult task because of the low number of populations genetically investigated and because of the technical re¢nements that in the last years have disproved the results of previous work. The time window used by most researchers (the postglacial) to evaluate the in£uence of historical events on genetic structure of European tree populations seems too short compared to the amount of genetic variability that was probably not eliminated by the restriction of populations during the LGM. This is especially important in the case of spruce, which was not

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reduced to a con¢ned refugium but survived in large regions in southern Europe. Hope is now placed in fossil DNA extraction and identi¢cation, because this will allow a real integration of paleobotanical and molecular methods.

Acknowledgements This work is part of the project ‘Environmental evolution and human impact after the last glacial time in the Alps’ promoted by the C.N.R. ^ Istituto per la Dinamica dei Processi Ambientali, Milan. An early version of the manuscript bene¢ted from the revisions by Brigitta Ammann, Lucia Wick (University of Bern), Jacques Louis de Beaulieu (University St. Je¤rome Marseille), Ruth Drescher-Schneider (Graz), and Waldo Zagwijn (Santpoort), who provided suggestions and recommendations. Ruth Drescher-Schneider and Walter Finsinger made available unpublished pollen records from southeastern Austria and the Maritime Alps. The revisions by Cathy Whitlock (University of Oregon) and Henri Hooghiemstra (University of Amsterdam) greatly improved the quality of the manuscript. Special thanks are due to Herbert Wright (University of Minnesota) for the hard work revising the English form. However, I am fully responsible for mistakes and further alterations. I am indebted to many others who helped with special questions raised in the paper: Remo Bertoldi (University of Parma) contributed with information on the Lagdei record; Giuseppe Siani (University of Paris Sud) provided his expertise on the late glacial chronology in the Adriatic Basin; Chronis Tzedakis (University of Cambridge) and Herman Mommersteeg (University of Amsterdam) assisted me with the Greek records; Donatella Magri (University of Roma I) with maar lakes in central Italy and Freddy Damblon (Institut Royal des Sciences Naturelles de Belgique, Bruxelles) with loess sequences; Roberto Avigliano (University of Udine) helped ¢nding the outcrops of Colle Umberto; Nicoletta Martinelli and Olivia Pignatelli (Dendrodata) were involved with the antrachological diagnosis of spruce wood; Adam Holzer and Georg Philippi (Staatlisches Museum fu«r Naturkunde Karlsruhe)

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and Ivo Trinajstic (Zagreb) helped with problems concerning the present-day natural range of spruce. Without the invitation by Marco Peresani (University of Ferrara) to study peat bogs on Venetian Pre-Alps, I could not have found the fossil spruce remains from which the present research originates. Roberta Pini (CNR, Milan) and Verushka Valsecchi (University of Milan) gave an important contribution to the paleobotanical investigation at Cansiglio. Lastly, I must thank Renata Perego who did much work compiling most ¢gures. This research is supported by the projects of the C.N.R. ^ Istituto per la Dinamica dei Processi Ambientali, Milan, by C.N.R. ^ P.F. ‘Beni Culturali’ (U.O. Universita' di Ferrara, CT no. 97.00597 PF 36) and by the Azienda Regionale Veneto Agricoltura.

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