History of Larix decidua Mill. (European larch) since 130 ka

Rémy Petit

Retrospective studies focussing on forest dynamics using fossil and genetic data can provide important keys to prepare forests for the future. In this study we analyse the impact of past climate and anthropogenic changes on Larix decidua Mill. (European larch) populations based on a new range-wide fossil compilation encompassing the last 130 ka and on recently produced genetic data (nuclear, mitochondrial). Results demonstrate that during the last 130 ka L. decidua persisted close to its current distribution range and colonized vast areas outside this range during the first two early Weichselian interstadials (c. 87-109 ka and c. 83-78 ka), reaching a distributional maxima in the north-central European lowlands. Some fossil sites point to notably rapid responses to some abrupt climate events DansgaardeOeschger cycles and Heinrich Events). Combined fossil and genetic data identify at least six MIS 2 refuges and postglacial recolonization pathways. The establishment of extant L. decidua forests dates back to the first two millennia of the Holocene (c. 11.5-9.5 ka) and the onset of anthropogenic impact was inferred since the late Neolithic (c. 6 ka), with major changes occurring since the Bronze Age (c. 4 ka). During the last 300 years human-induced translocations resulted in recent admixture of populations originating from separate refuges. Altogether, the results of this study provide valuable clues for developing sustainable conservation and management strategies targeting ancient genetic lineages and for studying evolutionary issues.

Quaternary Science Reviews 124 (2015) 224e247 Contents lists available at ScienceDirect Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev History of Larix decidua Mill. (European larch) since 130 ka nchez-Gon ~ i d, Re my J. Petit b, a Stefanie Wagner b, a, *, Thomas Litt c, Maria-Fernanda Sa a INRA, UMR1202 BIOGECO, F-33610 Cestas, France Univ. Bordeaux, UMR1202 BIOGECO, F-33615 Pessac, France c Univ. Bonn, Steinmann Institute of Geology, Mineralogy and Paleontology, D-53115 Bonn, Germany d Ecole Pratique des Hautes Etudes, UMR 5805 EPOC, CNRS, Univ. Bordeaux, F-33615 Pessac, France b a r t i c l e i n f o a b s t r a c t Article history: Received 27 April 2015 Received in revised form 29 June 2015 Accepted 1 July 2015 Available online xxx Retrospective studies focussing on forest dynamics using fossil and genetic data can provide important keys to prepare forests for the future. In this study we analyse the impact of past climate and anthropogenic changes on Larix decidua Mill. (European larch) populations based on a new range-wide fossil compilation encompassing the last 130 ka and on recently produced genetic data (nuclear, mitochondrial). Results demonstrate that during the last 130 ka L. decidua persisted close to its current distribution range and colonized vast areas outside this range during the ﬁrst two early Weichselian interstadials (c. 87e109 ka and c. 83e78 ka), reaching a distributional maxima in the northecentral European lowlands. Some fossil sites point to notably rapid responses to some abrupt climate events (DansgaardeOeschger cycles and Heinrich Events). Combined fossil and genetic data identify at least six MIS 2 refuges and postglacial recolonization pathways. The establishment of extant L. decidua forests dates back to the ﬁrst two millennia of the Holocene (c. 11.5e9.5 ka) and the onset of anthropogenic impact was inferred since the late Neolithic (c. 6 ka), with major changes occurring since the Bronze Age (c. 4 ka). During the last 300 years human-induced translocations resulted in recent admixture of populations originating from separate refuges. Altogether, the results of this study provide valuable clues for developing sustainable conservation and management strategies targeting ancient genetic lineages and for studying evolutionary issues. © 2015 Elsevier Ltd. All rights reserved. Keywords: Fossil compilation Genetics Last interglacialeglacial cycle Range-wide Refuges European larch 1. Introduction Knowledge of long-term forest history against the background of past climate and anthropogenic impact provides a valuable basis to prepare forests for the future (Millar and Brubaker, 2006; Petit et al., 2008). Recent advances in historical vegetation and climate research open new perspectives for reconstructing individual tree histories in the context of long-term and short-term climate changes of the last interglacialeglacial cycle (Fletcher et al., 2010; ~ i, 2010; Wolff et al., 2010). Similarly, Harrison and Sanchez Gon recent progress in population genetics helps increase the precision of historical inferences that can be derived from extant populations thanks to the improved quality of genetic datasets and the improved performance of genetic assignment methods (e.g. De Carvalho et al., 2010; Tollefsrud et al., 2009). Altogether, these * Corresponding author. INRA-Univ. Bordeaux, UMR BIOGECO, 69 route d'Arcachon, 33610 Cestas, France. E-mail address: stefanie.wagner@pierroton.inra.fr (S. Wagner). http://dx.doi.org/10.1016/j.quascirev.2015.07.002 0277-3791/© 2015 Elsevier Ltd. All rights reserved. advances encourage new steps in reconstructing individual tree histories using fossil and genetic data, which should eventually help understand the consequences of ongoing changes on extant tree populations and help develop appropriate forest conservation and management strategies (Hu et al., 2008; Millar and Brubaker, 2006; Petit et al., 2008). A precise documentation of long-term climate variability of the last interglacialeglacial cycle (c. 130,000 years ¼ 130 ka) is provided by marine sedimentary records documenting three major periods of different ice volume in the high latitudes of the northern hemisphere, each of them lasting several millennia. The ﬁrst one corresponds to Marine Isotope Stage (MIS) 5, the penultimate warm period, the second one to MIS 4, 3 and 2 deﬁning the last glacial, and the third one to MIS 1, the present-day interglacial. Greenland ice core archives (North GRIP Members, 2004; Wolff et al., 2010) identiﬁed a series of rapid warming and cooling events taking place within less than 100 years in superposition to this long-term climate variability: the DansgaardeOeschger (DeO) cycles, that triggered warm and cold phases called Greenland Interstadials (GI) and Greenland Stadials (GS), respectively. These cycles have S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 counterparts in North Atlantic marine sedimentary records and some of the rapid cooling events are associated with iceberg discharges in the North Atlantic (Heinrich events, HE) (Bond and Lotti, 1995; Heinrich, 1988). The impact of such strong releases of freshwater in the ocean produced cold phases called Heinrich nchez Gon ~ i and Harrison, 2010). GI, GS and HS Stadials (HS) (Sa lasted between 500 and 3000 years. Direct comparison between North Atlantic climate records and pollen sequences obtained from the same marine sediments have shown that forests responded rapidly (in c. 100 years) and synchronously to long-term and shortterm North Atlantic and Greenland climate changes (Fletcher et al., ~ i et al., 2008). In contrast, responses of individual 2010; S anchez Gon tree species have so far been neglected albeit this should be very valuable for developing ﬂexible conservation and management strategies for future forests (Millar and Brubaker, 2006). Existing historical tree studies focus on the late- and postglacial period. Some of them rely on combined fossil and genetic data to identify MIS 2 refuges and postglacial recolonization pathways, e.g. in Europe for Quercus, Fagus sylvatica, Pinus sylvestris, Abies alba and Picea abies (Cheddadi et al., 2006; Liepelt et al., 2009; Magri et al., 2006; Petit et al., 2002; Tollefsrud et al., 2008). Other paleobotanical investigations evaluate the importance of climatic and anthropogenic factors during postglacial recolonizations, e.g. for European beech (F. sylvatica) climate turned out to be the most important factor in general (Huntley et al., 1989), but in some areas close to the northern range limit anthropogenic factors were predominant (e.g. Bradshaw and Lindbladh, 2005; Tinner and Lotter, 2006). In fact, human-induced translocations have the potential to signiﬁcantly alter the original genetic composition of pop€ nig et al., 2002; Lowe et al., ulations (Deguilloux et al., 2003; Ko 2004). Altogether, these examples illustrate how precise historical reconstructions beneﬁt from the combined use of fossil and genetic data. In this study we endeavour a new step in reconstructing a tree species history by taking advantage of recent advances made in paleoecology and population genetics. We chose the conifer Larix decidua as a model offering many advantages and having not yet been subject to a combined range-wide paleoecological-genetic study. Owing to its pioneer character, responses to climate and anthropogenic changes can be expected to be particularly rapid. Its current distribution range is rather small and well covered by fossil sites. L. decidua is found at high-altitudes in the Alps and in other Central European mountains (Carpathians, Sudetes), as well as in the Polish lowlands (see Fig. S1, Supporting information). Overall its distribution is centred in the subalpine vegetation belt of the central Alps characterized by continental climate conditions. In the other range parts altitudinal distribution and climate vary widely. Among European conifers L. decidua is unusual as it is a deciduous species, thereby increasing the chance to ﬁnd macrofossil evidence (especially needles and stomata). Another advantage is that its pollen productivity and dispersal ability are very limited, leading to , 2007; Pela nkova and Chytrý, very local pollen signal (Jankovska €gren et al. 2008a, 2008b). A recent genetic study has 2009; Sjo uncovered the most recent part of its history (last 300 years), which was marked by human-induced translocations (i.e. movements of seed or seedlings across the range), followed by artiﬁcial mixture of geographically distant gene pools (Wagner et al., 2015). It also recovered the ancestral genetic structure at nuclear and mitochondrial markers. This information can now be used together with fossil data to reconstruct the species more ancient history. In this study we focus on the range-wide history of L. decidua during the last interglacialeglacial cycle (130 ka) by compiling fossils and by interpreting them together with the recently produced nuclear and mitochondrial data. The issues we address are (1) the chronological documentation of range changes since the last interglacial (130 ka), 225 (2) the identiﬁcation of some consequences of abrupt climate events, (3) the identiﬁcation of MIS 2 (27.8e14.7 ka) refuges and postglacial (14.7 kaepresent) recolonization pathways, and (4) the importance of past climate versus anthropogenic impact. 2. Material and methods 2.1. Fossil data compilation and mapping Pollen, macrofossil, stomata and charcoal data for 130 ka to 19 ka corresponding to the last interglacial until the end of the Last ~ i et al., 2012) Glacial Maximum (LGM, Mix et al., 2001; S anchez Gon was extracted from the original literature. For the last 19 ka, data was compiled from the Alpine Palynological Database (ALPADABA), the Czech Quaternary Palynological database (PALYCZ, http:// botany.natur.cuni.cz/palycz, Kunes et al., 2009), the European pollen database (EPD, Davis et al., 2013; Giesecke et al., 2013) and the original literature, in this sequence. After compilation, data was assigned to consecutive, stratigraphically meaningful time intervals: MIS 5, which comprises the last interglacial sensu stricto nchez Gon ~ i et al., corresponding to the Eemian (c. 130e112 ka, Sa 2012), the ﬁrst early Weichselian interstadial corresponding to nchez Gon ~ i, the St. Germain 1/Brørup (c. 109e87 ka, Müller and Sa 2007), the second early Weichselian interstadial corresponding to the St. Germain 2/Odderade (c. 83e78.2 ka, Sanchez Goni et al., 2013) and the third early Weichselian interstadial corresponding to the Ognon 1 (DO 20, c. 76.4e75.5 ka, Sanchez Goni et al., 2013; nchez Gon ~ i and Wolff et al., 2010), MIS 4 (c. 73.5e59.4 ka, Sa ~ i and Harrison, 2010), MIS 3 (c. 59.4e27.8 ka, S anchez Gon Harrison, 2010), MIS 2 (c. 27.8e14.7 ka) subdivided into HS 2 (c. 26.5e24.3 ka), LGM (c. 23e19 ka) and HS 1 (c. 19e15 ka, Mix et al., nchez Gon ~ i and Harrison, 2010; Stanford et al., 2011), MIS 1 2001; Sa subdivided in Late-glacial interstadial (Bølling/Allerød, c. 14.5e12.8 ka, Ammann et al., 2006; Litt et al., 2001; Rasmussen et al., 2006), Younger Dryas (c. 12.8e11.7 ka, Ammann et al., 2006; Litt et al., 2001; Rasmussen et al., 2006) and consecutive 1000 year intervals for the Holocene (c. 11.5 ka until presenteday). For each time interval and each site, we mapped mean Larix pollen percentages and presence/absence of macrofossils, stomata and charcoals using ArcGIS 9.3 (ESRI 2009). As previous studies focussing on pollen abundance have revealed that sometimes only few or no pollen grains of Larix are detected even when the species is , 2007; Pela nkova and growing nearby the study site (e.g. Jankovska € gren et al., 2008a, 2008b), we also took into acChytrý, 2009; Sjo count single pollen ﬁndings, while keeping in mind that scarce pollen ﬁndings can also be of non-local origin (e.g. Ortu, 2002; € gren et al., 2008a, 2008b). Sjo 2.2. Genetic data The genetic study of Wagner et al. (2015) shows that Larix decidua is characterized by an exceptionally strong genetic structure of both biparentally inherited nuclear markers and maternally inherited mitochondrial markers (Figs. 1 and 2). Speciﬁcally, the genetic differentiation at both marker classes is particularly strong between the Alpine and the eastern range part, suggesting that the genetic lineages from these two regions have become separated a long time ago. Overall, seven nuclear genetic groups, so-called genetic clusters, and four major mitochondrial variants, so-called haplotypes, have been identiﬁed. In the Alps, four nuclear clusters (clusters 1e4) and two of the most common mitochondrial haplotypes (haplotypes 16 and 18) are distributed longitudinally. In most cases, nuclear clusters are associated with one frequent mitochondrial haplotype (nuclear cluster 1 with haplotype 16, cluster 2 with haplotype 18, cluster 3 with haplotype 16, cluster 4 226 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Fig. 1. Nuclear genetic structure of Larix decidua (modiﬁed after Wagner et al., 2015). Pie charts represent cluster composition of 40 native populations. Dotted circles represent population considered to have been introduced. Fig. 2. Mitochondrial genetic structure of Larix decidua (modiﬁed after Wagner et al., 2015). Pie charts represent haplotype (H) composition of 40 native populations. The other ones represent other sequence variants. with haplotype 18). However, there are two populations from the central Alps (Swiss region of Valais) that depart from this pattern (populations 27 and 83, Figs. 1 and 2, Wagner et al., 2015). In these populations, cluster 1 is associated with haplotype 18. Furthermore, a rare haplotype (haplotype 22) is only detected in the southwestern extremity of the Western Alps: it is ﬁxed in a population from the Maritime Alps (population 81) and mixed with haplotype 16 in a population from further north (population 77). In the eastern part of the range, one nuclear cluster was found in the Sudetes Mountains (cluster 5), one in the western Carpathians (cluster 6) and one in the eastern and southern Carpathians (cluster 7). In the Polish lowlands nuclear assignments were less clear and clusters from the Sudetes and the western Carpathians co-occurred. An additional analysis of nuclear data focussing on Central Europe S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 populations has uncovered an additional sub-cluster in the southeastern Carpathians (Wagner, unpublished data). The original genetic structure has been altered by recent human-induced translocations (last 300 years) involving artiﬁcial mixing of geographically distinct populations (Wagner et al., 2015). 3. Results There were 355 sites with Larix fossils found among 1026 sites compiled from across Europe (Fig. S2, Table S1eS8, List S1 and List S2, Supporting information). Speciﬁcally, Larix fossils were reported in 105 out of 156 sites from the Alpine Palynological Database, 30 out of 114 sites from the Czech Quaternary Palynological database, 51 out of 549 sites from the European pollen database and 169 out of 207 sites from the literature. 3.1. Last interglacial sensu stricto (c. 130e112 ka) For the last interglacial, data compilation yielded 41 fossil sites, with 14 sites documenting Larix pollen or macrofossils (Fig. 3, Table S1, Supporting information). These Larix sites are located in the Alpine region and in northecentral Europe. In the Alps, Larix fossils occur in the northern (e.g. Mondsee, 540 m a.s.l., DrescherSchneider, 2000a) as well as in the southern Alpine foreland (e.g. Lake Fimon, 23 m a.s.l., Pini et al. 2010). In northecentral Europe, they occur in the Polish and German lowlands (e.g. Kittlitz, 170 m a.s.l., Horoszki Duze, 180 m a.s.l., Rederstall, <100 m. a.s.l.; Erd, 1973; Granoszewski, 2003; Menke and Tynni, 1984). All evidence for Larix is restricted to the transitions from the bracketing glacial periods and disappears in the course of the interglacial. During the lisey 1/Herning), scattered Larix evidence is subsequent stadial (Me still found in some sites (Behre et al., 2005; Behre and Lade, 1986; Erd, 1973; Granoszewski, 2003; Menke and Tynni, 1984), but this stadial is clearly dominated by non-arboreal cold steppic taxa. 227 3.2. First early Weichselian interstadial (c. 109e87 ka) For the ﬁrst early Weichselian interstadial (St. Germain 1/ Brørup), we compiled 42 fossil sites, including 28 Larix sites (Fig. 4, Table S2, Supporting information). In the northecentral European lowlands, Larix fossil distribution reaches a maximum since the last 130 ka. Pollen percentages of lowland sites reach up to 18% and are corroborated by numerous macrofossil ﬁndings (e.g. Osterwanna, Oerel and Keller, Behre, 1974; Behre et al., 2005; Behre and Lade, 1986; Menke, 1970). Further Larix pollen evidence from outside the current distribution range exists from the Vosges Mountains (La Grande Pile, de Beaulieu and Reille, 1992). Pollen and macrofossils within or close to the current range are documented in the northern and southern Alpine foreland (e.g. Füramoos, Mondsee, Lake Fimon; Drescher-Schneider, 2000a; Müller, 2001; Müller et al., 2003; Pini et al., 2010). Altogether, fossil representation largely peaks in the second half of this interstadial. The observed increase of larch fossils has been correlated to an abrupt cold event termed “WFII kryomer” (Caspers and Freund, 1997) in northecentral Europe (e.g. Caspers, 1997; Caspers and Freund, 1997; Litt et al., 1996; Menke and Tynni, 1984; Müller, 2001; Müller et al., 2003; Pini et al., 2010) and “Montaigu” (Woillard, 1978) in the French Massif lisey 2/Rederstall), scattered Central. In the subsequent stadial (Me Larix evidence still exists, though non-arboreal cold steppic taxa are predominant (e.g. Drescher-Schneider, 2000a; Erd, 1973; Hahne et al., 1994; Müller, 2000, 2001). 3.3. Second and third early Weichselian interstadial (c. 83e78.2 ka and c. 76.4e75.5 ka) For the second early Weichselian interstadial (St. Germain 2/ Odderade), compilation yielded 34 sites, 20 of which document Larix (Fig. 5, Table S3, Supporting information). The fossil distribution range is similar to the one observed during the ﬁrst early Fig. 3. Fossil distribution of larch from 130 to 112 ka and sites cited in the text for this period: (01) Mondsee (02) Lake Fimon (03) Kittlitz (04) Horoszki Duze (05) Rederstall. 228 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Fig. 4. Fossil distribution of larch from 109 to 87 ka and sites cited in the text for this period: (01) Mondsee (02) Lake Fimon (06) Keller (07) Osterwanna (08) Oerel (09) Füramoos (10) La Grande Pile. Fig. 5. Fossil distribution of larch from 83.0 to 78.2 ka and sites cited in the text: (01) Mondsee (02) Lake Fimon (06) Keller (07) Osterwanna (08) Oerel (09) Füramoos (10) La Grande Pile. S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Weichselian interstadial, though in northecentral Europe fossil representation is reduced compared to ﬁrst early Weichselian interstadial (Behre et al., 2005; Behre and Lade, 1986; Caspers and Freund, 1997; Litt et al., 1996; Menke and Tynni, 1984). In northcentral Europe, Larix fossils disappear after this interstadial, whereas in the Alps they reoccur during the third early Weichselian interstadial (Ognon 1/Oerel), in some sites with exceptionally high pollen percentages (up to 20%) (e.g. Drescher-Schneider, 2000b; Grüger, 1979; Grüger and Schreiner, 1993; Müller, 2001; Müller et al., 2003; Wegmüller, 1992; Welten, 1982a). After this interstadial Larix fossils disappear from the Alpine region and fossils of non-arboreal cold steppic taxa become predominant. 3.4. MIS 4 (c. 73.5e59.4 ka) For MIS 4, compilation resulted in 25 sites, only nine of them located in central Europe and only three of them exhibiting very scarce and discontinuous Larix pollen evidence (Fig. 6, Table S4, Supporting information). In Füramoos (662 m a.s.l.) documenting MIS 5eMIS 2, Larix evidence is missing (Müller, 2001; Müller et al., 2003). Altogether, pollen assemblages are dominated by nonarboreal cold-steppic taxa. 3.5. MIS 3 (c. 59e27.8 ka) For MIS 3, 50 sites were compiled, 25 of which include Larix fossils (Fig. 7, Table S5, Supporting information). These Larix sites are located in the northern and southern Alpine foreland and the western Carpathians with adjacent areas (Bohemia, Moravia and the Pannonian Plain). In the western Carpathians, Larix pollen percentages from Saf arka and Jabl unka are conspicuously high (up , 2003; to 20%) and corroborated by macrofossils (Jankovska et al., 2002; Jankovska and Pokorny, 2008). These Jankovska 229 pollen sequences document a predominance of Larix, Betula and Pinus cembra until approximately 30 ka when Picea becomes predominant (Kunes et al., 2008). In Moravia and the Pannonian Plain, Larix is present in several single-dated samples with pollen, macrofossils and charcoals from between 55 ka and 28 ka (e.g. Damblon and Haesaerts, 1997; Damblon et al., 1996; Erd, 1973; Geyh et al., 1969; Komar et al., 2009; Krolopp, 1977; Mamakowa and Starkel, and Rybnícek, 1991; 1974; Musil, 2003; Opravil, 1994; Rybní ckova Willis and van Andel, 2004). In the northern Alpine foreland, Larix pollen is found again in Füramoos (662 m a.s.l.) together with Betula pollen at approximately 54 ka (LPAZ B1) and at 45 ka (LPAZ B2). This has been interpreted as a response to the Dansgaard Oeschger cycles 14 and 12 (Fletcher et al., 2010; Müller, 2001; Müller et al., 2003). South to the Alps, Larix is documented by pollen and macrofossils in Lake Fimon (23 m a.s.l.) and Lago della Costa (7 m a.s.l.) (Kaltenrieder et al., 2009; Pini et al., 2010; Wick, 2006). The pollen sequence of Lake Fimon documents an increase of Larix and other arboreal taxa at approximately 50 ka (LPAZ FPD15) after a period of predominating non-arboreal cold steppic taxa (LPAZ FPD14) that has also been interpreted as a response to DeO 12 and DeO 14, although in the pollen of Lake Fimon these two events are documented as a cluster, i.e. their individual impact cannot be discerned (Pini et al. 2010). In addition, Larix charcoals are detected at the southern Alpine margin in northern Italy dated between 46 ka and 36 ka (Maspero, 1996). The prevalence of Larix and cold steppic taxa in Lake Fimon prior to c. 31 ka (LPAZ 17c) nchez Gon ~ i and could correlate to HS 3 (c. 32.7e31.3 ka) (Sa Harrison, 2010) but resolution and dating will have to be improved to corroborate this hypothesis. The Lago della Costa sequence starting at c. 32.5 ka also documents cold steppic communities with Larix contribution (pollen and macrofossils evidence) at that time (Kaltenrieder et al. 2009). Fig. 6. Fossil distribution of larch from 73.5 to 59.4 ka and the site cited in the text for this period: (09) Füramoos. 230 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 a rka (12) Jabl Fig. 7. Fossil distribution of larch from 59.4 to 27.8 ka and sites cited in the text for this period: (02) Lake Fimon (09) Füramoos (11) Saf unka (13) Lago della Costa. * DeO cycles 12 and 14: b Larix. 3.6. MIS 2 (c. 27.8e14.7 ka) 3.6.1. HS 2 (c. 26.5e24.3 ka) and LGM (c. 23e19 ka) For this period, 40 sites were compiled, eight of them documenting Larix fossils (Figs. 8 and 9, Table S6, Supporting information). There is continuous evidence of Larix fossils from the Alpine and Carpathian region, pointing to its persistence in each of these two regions. South to the Alps Larix is documented in Lago della Costa and Lake Fimon (Kaltenrieder et al., 2009; Pini et al., 2010). In Lago della Costa, Larix pollen and macrofossils slightly increase between 27 ka and 23 ka (LPAZ APG-3) together with fossils of other light demanding woody and herbaceous taxa, which correlates well with HS 2 (c. 26.5e24.3 ka). A similar tendency, though less precisely dated, is documented in Lake Fimon (LPAZ 19). In the Hungarian Plain, detailed inferences can be based on a r (86 m a.s.l.) (Sümegi et al., 2013). recent investigation of Lake Fehe As in Lago della Costa, pollen evidence of Larix and of cold-steppic taxa increase prior to the LGM (c. 26.4e24.9 ka, FT-2), at a time r corresponding to HS 2 (Sümegi et al., 2013). In addition, Lake Fehe documents a decrease of Larix and cold-steppic taxa concomitant to an increase of Picea in the two bracketing intervals (c. 28e26.4 ka, FT1 and c. 24.9e23.2 ka, FT-3), which correlates to DeO 3 and DeO r and in Nagymohos, located in 2 (Sümegi et al., 2013). In Lake Fehe the north-eastern Hungarian Mountains, Larix is present during the LGM (Magyari, 2002; Magyari et al., 1999; Sümegi et al., 2013). In addition, Larix and Picea-Larix charcoals dated to the HS 2 and the nk, 1960; Geyh LGM are found in the Pannonian Plain (Gaborí-Csa et al., 1969; Vogel and Waterbolk, 1964; Willis and van Andel, arka, pollen and macro2004). In the western Carpathian site Saf a, 2003; Jankovska fossils are found throughout the LGM (Jankovsk and Pokorny, 2008). A single sample et al., 2002; Jankovska including pollen and macrofossils dated to c. 20 ka is also reported from Smerek (600 m a.s.l.) in the south-eastern Polish Bieszczady Mountains (Ralska-Jasiewiczowa, 1980; Wacnik et al., 2004). Pollen evidence dated to c. 20 ka also exists from Labský d ul in the Sudetes , 2004), though dating Mountains (Engel et al., 2010; Jankovska deduced by interpolation will have to be conﬁrmed. 3.6.2. HS 1 (c. 19e15 ka) For this period, compilation resulted in 208 sites, 88 of which include Larix (Fig. 10, Table S7, List S1 and List S2, Supporting information). Strongest evidence exists from low altitudinal sites (<500 m a.s.l.) located in the southern Alpine foothills and areas nearby the Carpathians and the Sudetes. In the south-western Alps pollen, stomata, and macrofossil evidence exists from the province of Turin, e.g. fossil wood dated to 18.3 ka and even older needle and pollen ﬁndings from Lago Piccolo di Avigliana (353 m a.s.l.) (Finsinger and Tinner, 2006; Finsinger et al., 2006; Vescovi et al., 2007), and pollen and stomata evidence assigned to the Oldest Dryas (e.g. Lago di Viverone, 220 m a.s.l., Torfsee, 270 m a.s.l., Schneider, 1978), the terrestrial counterpart of HS 1 (Naughton et al., 2007). Further eastwards, in the foothills of the central Alps (c. 8.5e10 E, provinces of Lombardy and Ticino), Larix evidence is weaker. Scarce records exist for instance from Lago di Biandronno (239 m a.s.l.), Lago di Origlio (416 m a.s.l.) and Lago di Gaiano (334 m a.s.l.) (Gehring, 1997; Schneider, 1978; Tinner et al., 1999; Vescovi et al., 2007). Further east, at longitudes >10 E (provinces of Trentino and Venetia), evidence gets stronger again, e.g. there are tree trunks and pollen percentages up to 8% dated to 18.1e17.0 ka from Fornaci di Revine (224 m a.s.l.) (Casadoro et al., 1976; Friedrich et al., 1999; Kromer et al., 1998), macrofossils dated between 18.5 ka and 17.5 ka from Lago Lucone (249 m a.s.l.) (Valsecchi et al., 2006), and macrofossils, stomata and pollen since c. 17 ka from Lago di Ragogna (188 m a.s.l.) (Monegato et al., 2007). In the eastern Alps >14 E, less fossil sites are available. Early pollen evidence dated to c. €ngsee (548 m a.s.l., Austrian state of Carinthia) 18.7 ka exists from La S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 231 a rka (14) Lake Fehe r. x Heinrich events 2 Fig. 8. Fossil distribution of larch from 27.8 to 23.5 ka and sites cited in the text for this period: (02) Lake Fimon (13) Lago della Costa (11) Saf or 3: b Larix: * DeO cycles 3 and 2: a Larix. arka (13) Lago della Costa (14) Lake Fehe r (15) Smerek (16) Nagymohos (17) Labský d Fig. 9. Fossil distribution of larch from 23.5 to 19 ka and sites cited for this period: (11) Saf ul. 232 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 arka (16) Nagymohos (17) Labský d Fig. 10. Fossil distribution of larch from 19 to 15 ka and important sites cited for this period: (11) Saf ul (18) Piccolo di Avigl. (19) Lago di Origlio €ngsee (21) Selle di Carnino (22) Lake Galeş (23) Lake Brazi. (20) Fornaci di Revine (21) La (Schmidt et al., 1998). Evidence from mid- and high-altitudes largely consists in exceptional single pollen grains in pollen assemblages dominated by non-arboreal cold steppic components. For instance, in the Alps, evidence exists in Lac de Villa (820 m a.s.l., Aosta), ZeneggenHellelen (1520 m a.s.l., Valais), Krotenweiher (1310 m a.s.l., Tirol), Grosses Überling Schattseit Moor (1750 m a.s.l., Carinthia) (ALPADABA, EPD; Bortenschlager, 1984; Brugiapaglia, 2007; Kirsai et al., 1989; Welten, 1982b). A remarkable early single pollen ﬁnding dated to 16.2 ka originates from Selle di Carnino (1905 m a.s.l.) located in the southern extremity of the Western Alps (Maritime Alps) (ALPADABA; de Beaulieu, 1977), an area where fossil sites of this period are very sparse. In the area of the Carpathians and the Sudetes, evidence con a rka, tinues in sites also covering the LGM (e.g. Labský d ul, Saf , 1984, 1991, 2004, 2008; Nagymohos; Engel et al., 2010; Jankovska Kunes et al., 2008; Magyari, 2002; Magyari et al., 1999). Other sites located in the forelands of the Sudetes and western Carpathians provide additional evidence: Vlcí rokle (583 m a.s.l.), Wolbrom (375 m a.s.l.) and Jaslo (250 m a.s.l.) (EPD, PalyCZ). In the southern Carpathians (Retezat Mountains), two high-altitudinal sequences document Larix pollen and stomata since their respective onset at about 15 ka (Lake Brazi, 1740 m a.s.l.; Lake Galeş, 1990 m a.s.l., Magyari et al., 2012). 3.7. MIS 1 (since c. 14.5 ka) 3.7.1. Late-glacial interstadial (c. 14.5e12.8 ka) During the Late-glacial interstadial (Bølling/Allerød), Larix fossil evidence from across the range becomes stronger as compared to before and occurs from low- to mid-altitudes (Fig. 11, Table S8, List S1 and List S2, Supporting information). In the south-western Alps at low altitudes Larix ﬁrst increases together with Betula and Pinus and then decreases with the increase of thermophilous taxa, whereas it becomes more abundant at mid-altitudes (e.g. Lago Piccolo di Avigliana, 353 m a.s.l; Lago di Viverone, 220 m a.s.l.; Torfsee, 270 m a.s.l.; Finsinger et al., 2006; Schneider, 1978). For instance, there are needles reported from Lac de Villa (820 m a.s.l.) re de Pilaz (1460 m a.s.l.) located in the Aosta valley. In and Tourbie contrast, in the same valley, macrofossils are still missing at the re de Champlong (2320 m a.s.l.) high-altitudinal site Tourbie (Brugiapaglia, 1997, 2001, 2007). It is notable that another pollen record is reported from Selle di Carnino. Further eastwards, in the provinces of Lombardy and Ticino, Larix evidence stays weak, due to the dominance of Pinus, Betula and Picea (e.g. Ilyashuk et al., 2009; Schneider, 1978; Tinner et al., 1999; Vescovi et al., 2007; Zoller and Kleiber, 1971). At longitudes >10 E, fossils increase at mid-altitudes and decrease at low-altitudes: in Pian di Gembro (1350 m a.s.l.) and Palughetto (1040 m a.s.l.), pollen and macrofossils increase whereas in Lago di Ragogna (188 m a.s.l.) they decrease concomitant to the increase of thermophilous taxa (Monegato et al., 2007; Pini, 2002; Vescovi et al., 2007). At highaltitudes evidence remains scarce, as for instance in Passo del Tonale (1883 m a.s.l.) and Col di Val Bighera (2087 m a.s.l.) (Gehring, 1997; Vescovi et al., 2007). Evidence from the Adige-Inn valley is abundant, e.g. Zotensenk (560 m a.s.l.), Sommersüss (870 m a.s.l.), Lanser Moor I/III (840 m a.s.l.), Totenmoos (1718 m a.s.l.) (ALPADABA; Bortenschlager, 1984; Heiss et al., 2005; Seiwald, 1980). In Totenmoos macrofossils, stomata and pollen are detected regularly since c. 14.2 ka (Heiss et al., 2005). At the south-eastern Alpine €ngsee (Schmidt et al., margin, Larix pollen is still found in the La 1998), and there is additional macrofossil and pollen evidence from the Slovenian Lake Bled (475 m a.s.l.) (Andri c et al., 2009). In the Carpathians, evidence is stronger than before in the two southern Carpathian high-altitudinal sites (Magyari et al., 2012) as well as in mid- altitudinal sites of the north-eastern and western S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 233 re de Pilaz (25) Fig. 11. Fossil distribution of larch from 14.5 to 12.8 ka and important sites cited for this period: (18) Piccolo di Avigl. (21) Selle di Carnino (23) Lake Brazi (24) Tourbie Majola Pass (26) Palughetto (27) Lago di Ragogna (28) Totenmoos (29) Lake Bled (30) Zarnowiec. Carpathians (Feurdean and Bennike, 2004; Jankovsk a, 1984, 1991; Ralska-Jasiewiczowa, 1980; Wohlfarth et al., 2001). Additional evidence comes from the Pannonian Plain (Zsombo Swamp, jezero, 1105 m a.s.l.; PALYCZ, 92 m a.s.l.; EPD), Bohemia (Plesne , 2006) and the Polish lowlands (Niechorze, 5 m a.s.l., Jankovska Zarnowiec Peat Bog, 5 m a.s.l. and Woryty, 105 m a.s.l.; EPD). 3.7.2. Younger Dryas (c. 12.8e11.7 ka) During the Younger Dryas, the longitudinal and latitudinal extent of the fossil distribution range does not change importantly (Fig. 12, Table S8, List S1 and List S2, Supporting information). However, altitudinal changes can be noticed. At some low- to midaltitudinal sites, Larix fossils increase concomitantly to the decrease of less cold tolerant tree taxa, e.g. Lago Piccolo di Avigliana (353 m a.s.l.), Lago di Origlio (416 m a.s.l.), Lago di Ledro c et al., 2009; (652 m a.s.l.) and Lake Bled (475 m a.s.l.) (Andri Finsinger et al., 2006; Joannin et al., 2013; Tinner et al., 1999; Vescovi et al., 2007) whereas at some high-altitudinal sites fossils decrease, e.g. Pian di Gembro (1350 m a.s.l.), Lago di Lova c et al., 2009; (1299 m a.s.l.) and Totenmoos (1718 m a.s.l.) (Andri Heiss et al., 2005; Pini, 2002). Similar altitudinal changes are , 1984, 1991; documented in the Carpathians (e.g. Jankovska Koperowa, 1962). 3.7.3. Holocene (since c. 11.7 ka) In the course of the Holocene, Larix fossils become increasingly restricted to mid- and high-altitudinal sites, corresponding to its current distribution range (Figs. 13e24). At low altitudes, competing tree taxa become predominant. As early as between 10.5 ka and 9.5 ka fossil distribution range largely reﬂects the modern distribution range, with the only difference that some lowaltitudinal occurrences disappear around 5 ka. For the ﬁrst two millennia of the Holocene, fossils indicate the establishment of extant L. decidua forests (Table 1, Figs. 13 and 14). In the central Swiss Alps (Valais, Engadine) and the central Italian Alps (Lombardy, Ticino), forest establishment has been dated between 11.5 and 10.5 ka based on high resolution macrofossil and pollen records (e.g. Gobet et al., 2005; Kaltenrieder et al., 2005; Lang and Tobolski, 1985; Tinner and Ammann, 1996; Wick and Tinner, 1997). This can be corroborated by additional sites documenting stomata and pollen (e.g. Maloja Pass, 1865 m a.s.l., Aletschwald, 2017 m a.s.l., Ilyashuk et al., 2009; Welten, 1982b). In the south-western Alps (province of Turin) continuous pollen curves as well as macrofossils from the Aosta Valley also indicate an establishment between 11.5 and 10 ka (Brugiapaglia, 2007). In the southern extremity of the western Alps, forest establishment cannot be dated conclusively. Based on pollen data (de Beaulieu, 1977; Ortu, 2002), a preliminary estimation would be c. 10 ka. In the Inner French Alps, macrofossils, charcoals and pollen records (Ali et al., 2005; Blarquez et al., 2009; Ponel et al., 2011) indicate an establishment between 9 ka and 8 ka, though earlier scattered macrofossils point to an earlier presence of Larix in the Vanoise Massif (Lac du Loup, 2035 m a.s.l., Blarquez et al. 2009). In the central Eastern Alps, records covering the early Holocene are scarce. Two combined macrofossil-pollen records from the Tyrol indicate that forests got established at c. 11.4 ka (Totenmoos, 1718 m a.s.l., Hirchbichl, 2140 m a.s.l.; Heiss et al., 2005; Oeggl and Wahlmüller, 1994). In the southern Carpathians, the two high-altitudinal records from Lake Brazi and Lake Gales indicate forest establishment at around 11.5 ka (Magyari et al., 2012). For the Western Carpathian chronologies are insufﬁciently resolved to provide an accurate estimation of larch forest establishment. However, fossils point to forest dynamics comparable to those observed in the Alps. Two sites investigated for macrofossils and pollen (Tarnowiec, 220 m a.s.l., Tranawa Wynza, 670 m a.s.l., Harmata, 1987; Koperowa, 1962) and further pollen sites (e.g. Bobrov, 620 m a.s.l., 234 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Fig. 12. Fossil distribution of larch from 12.8 to 11.7 ka, sites cited in the text for this period and Larix fossil tendencies: (18) Piccolo di Avigl. (19) Lago di Origlio (28) Totenmoos (29) Lake Bled (31) Lago di Ledro (32) Pian di Gembro. Fig. 13. Fossil distribution of larch from 11.5 to 10.5 ka, important sites and dates of extant forest establishment: (23) Lake Brazi (25) Majola Pass (28) Totenmoos (33) Champlong Rion (38) Lago Basso (39) Hirchbichl. Lake (34) Sant Anna peat bog (35) Pilaz peat bog (36) Simplon (37) Guille S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 235 Fig. 14. Fossil distribution of larch from 10.5 to 9.5 ka, important sites and dates of extant forest establishment: (40) Clapeyret (41) Biecai peat bog (42) Laghi dell'Orgials. Fig. 15. Fossil distribution of larch from 9.5 to 8.5 ka. pleso, 1494 m a.s.l., PALYCZ) show Hozelec, 685 m a. s. l., Popradske for instance that the species shifted from low to mid- and highaltitudes at the onset of the Holocene. During the late Neolithic (c. 6 ka) Larix was detected at higher altitudes than at present-day. In the central Swiss Alps it has been shown that since c. 6 ka, the timberline (limit of closed forests) and 236 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Fig. 16. Fossil distribution of larch from 8.5 to 7.5 ka. Fig. 17. Fossil distribution of larch from 7.5 to 6.5 ka. the treeline (tree limit) for L. decidua were lowered by about 300 m and 180 m, respectively (Kaltenrieder et al., 2005; Tinner, 2007; Tinner and Theurillat, 2003). These studies also showed that the demise at the timberline was primarily human-induced (e.g. caused by wood cutting and burning) whereas the demise of the treeline was primarily climate-induced. Such kind of diebacks of S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 237 Fig. 18. Fossil distribution of larch from 6.5 to 5.5 ka. Fig. 19. Fossil distribution of larch from 5.5 to 4.5 ka. Larix at the treeline have also been observed during cold-humid periods (CE-1eCE-8) including the 8.2 ka event (Haas et al., 1998; Tinner, 2007). Investigations of numerous fossil sites from across the Alps show that since c. 4 ka (Bronze Age) anthropogenic activities, in particular grazing after ﬁre, elicited the development of “L€ archenwiesen” (engl. ¼ larch meadows) corresponding to L. 238 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Fig. 20. Fossil distribution of larch from 4.5 to 3.5 ka. Fig. 21. Fossil distribution of larch from 3.5 to 2.5 ka. decidua stands with herbaceous under-storey currently representing an abundant vegetational type in the Alpine region (e.g. Blarquez et al., 2009; Brugiapaglia, 2007; Gehring, 1997; Gobet et al., 2003; Kral, 1979, 1980, 1982; Muller et al., 2000; Ortu et al., 2003; Pini, 2002; Talon, 2010; Wick, 1996). These studies also document alternation of intensiﬁed land-use (Bronze Age, Iron Age, S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Fig. 22. Fossil distribution of larch from 2.5 to 1.5 ka. Fig. 23. Fossil distribution of larch from 1.5 to 0.5 ka. 239 240 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Fig. 24. Fossil distribution of larch during the last 500 years. * Modern distribution data compiled by E. Welk, AG Chorology, Geobotany Department, University of Halle, based on map 21b in Meusel et al. (1965), and ourselves. Map produced using ArcGIS 9.3 (ESRI 2009). ** Ice sheet extent after Ehlers and Gibbard (2004). Roman Period, Middle Ages) and land abandonment involving Rion, Kaltenrieder et al., 2005) populational declines (e.g. Guille and expansions (e.g. Lac du Loup, Blarquez et al., 2009). Data from the Carpathian region suggest a similar interplay between climatic and anthropogenic factors since the Bronze Age (e.g. Feurdean and , 2008; Willis, 2008; Harmata, 1995; Rybní cek and Rybníckova , 1974; Speranza et al., 2000; Wacnik et al., 2004). Rybní ckova Data for the last 500 years indicate an increase of occurrences beyond the previous natural distribution range, in particular in the Czech Republic and in Slovakia (Fig. 24). Such rapid expansion is likely of anthropogenic origin, as shown by genetic analyses (Wagner et al., 2015). 4. Discussion 4.1. Responses to long-term and short-term climate variability MIS 5eMIS 2 This study provides a detailed fossil-based account of the history of Larix decidua of the last 130,000 years. Long-term and short-term climate changes documented between MIS 5 and MIS 2 elicited multiple range changes of Larix decidua in areas close to its current distribution as well as in areas lying far outside of its current distribution. During the last interglacial Larix occurred in the Alpine and east-central European region. It expanded early at the Table 1 Establishment of extant Larix decidua forests. Locality Region Altitude [m a.s.l.] Reference Continuous fossils [ka] Fossil type Clapeyret Biecai peat bog Laghi dell’Orgials Sant Anna peat bog Pilaz peat bog Aigue Agnelle Lac du Loup Lac des Lauzon Simplon Hobschensee Rion Guille South-western Alps South-western Alps South-western Alps Western Alps Western Alps Central Western Alps Central Western Alps Central Western Alps Central Alps Central Alps 2260 1920 2243 2304 1900 2300 2035 2180 2017 2343 10.2 10.0 9.8 11.0 11.5 8.3 8.3 8.0 11.4 11.1 Pollen Pollen Pollen Pollen, macrofossils Pollen, macrofossils Charcoals Macrofossils Pollen Macrofossils, pollen Macrofossils, pollen Lej da San Murezzan Majola Pass Lago Basso Totenmoos Hirschbichl Lake Brazi Central Alps Central Alps Central Alps Eastern Alps Eastern Alps Southern Carpathians 1768 1865 2250 1718 2140 1740 de Beaulieu 1977 Ortu 2002 Ortu 2002 Brugiapaglia 2007 Brugiapaglia 2007 Ali et al., 2005 Blarquez et al., 2009 Ponel et al., 2011 Lang and Tobolski 1985 Wick and Tinner 1997, Kaltenrieder et al., 2005 Gobet et al., 2005 Ilyashuk et al., 2009 Wick and Tinner 1997 Heiss et al., 2005 Oeggl and Wahlmüller 1994 Magyari et al., 2012 10.7 11.2 11.0 11.5 11.4 11.5 Macrofossils, pollen Stomata, pollen Macrofossils, pollen Macrofossils, stomata, pollen Macrofossils, pollen Macrofossils, stomata, pollen S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 interglacial onset, highlighting its role as a pioneer in formerly glaciated areas. In analogy to what happened during the current interglacial, we inferred that Larix became restricted to high-altitudes during the temperate stages of the last interglacial due its low competitive ability leading to its replacement by more competitive thermophilous species at lower altitudes. However, Larix reoccurred at lower altitudes during climatic deterioration at the transition from the last interglacial to the last glacial when thermophilous species regressed. During the ﬁrst two early Weichselian interstadials (c. 87e109 ka and c. 83e78.2 ka) Larix reached distributional maxima in the northecentral European lowlands where it built up boreal forests under relatively continental climate conditions. Both the method of climate indicator species based on macrofossils as well as the probability density function method based on pollen and € bern in central Germany (Hoffmann et al., macrofossils from Gro 1998; Kühl et al., 2007) indicate mean July temperatures of about 15e16 C whereas the reconstructed mean January temperatures range between 12 and 14 C and the annual mean precipitation between 500 and 600 mm (Kühl et al., 2007). During the ﬁrst early Weichselian interglacial Larix exceptional expansion was prompted by an abrupt cold event (WFII kryomer/Montaigu) stressing the important impact of short term climatic episodes on tree species distribution. The distributional maxima of the two early Weichselian interstadials highlight the great potential of Larix to rapidly colonize the European continent if concurrence is low due to climate conditions prohibiting the expansion of more temperate trees. In north-central Europe, Larix forests were replaced by cold steppic communities after the second early Weichselian interstadial whereas in the Alpine region there was another important advance of Larix during the third Early Weichselian interstadial. This can be explained by the strong climate gradient covering Europe at that time, induced by icesheet growth, implying that continentality in northecentral anchez Europe was higher than in the Alpine region (Müller and S ~ i, 2007; Sa nchez Gon ~ i et al., 2005). Though poorly docuGon mented by fossil sites, it becomes clear that during the extremely severe cold stage of MIS 4 Europe was dominated by cold steppic environments (Fletcher et al., 2010). Scarce Larix fossil evidence was found in each of the two main regions (Alps, east Central Europe). This evidence together with the particularly deep genetic split between extant populations of these two regions (Wagner et al., 2015) leads us to assume that Larix persisted during MIS 4 in each of the regions. During the climatically more favourable MIS 3, marked by intermediate ice volume, Larix recolonized low- and mid-altitudinal ranges of both regions. Two Alpine sites demonstrate the positive inﬂuence of DeO cycles 14 (c. 54.2 ka) and 12 (c.46.8 ka) favouring the early spread of Larix among other pioneers. This contrasts with earlier DeO cycles (DeO 17: c. 59.4 ka and DeO 16: c. 58.2 ka) that did not trigger such a spread (dates of DeO cycles after Wolff et al. 2010), an observation that is in agreement with a study demonstrating that at latitudes above 40 N DeO 12 and 14 triggered the most prominent forest adnchez Gon ~ i et al., 2008). Yet, it can be hypothesized that vances (Sa DeO 12 and 14 triggered range changes of Larix at a broader scale, which will have to be conﬁrmed by additional evidence. A pollen r) documented the sequence from the Hungarian Plain (Lake Fehe consequences of alternating warming and cooling events during the last interval of maximum ice volume: Larix declined during GI 3 and GI 2 and expanded during HS 2 and HS 3. Similar changes during HS 2 and HS 3 were observed in the southern Alpine region (Lago della Costa, Lake Fimon). These examples illustrate the great potential of Larix to rapidly respond to climate changes taking place at the millennial scale. As other studies demonstrated that HS 2 resulted in the demise of temperate forests and 241 desertiﬁcation across Europe (e.g. Fletcher et al., 2010; Naughton et al., 2007), it can be expected that at least this stadial impacted Larix at a broader scale. 4.2. MIS 2 refuges and ﬁrst recolonizations Refuges of MIS 2 can be identiﬁed based on fossils occurring during the LGM (maximal ice volume) and the subsequent HS 1 (exceptional drought) together with the ancient genetic structure (Fig. 25). Fossils found between 23 ka and 16 ka and genetic data point to six distinct refuges: three are located south of the Alps (Turin, Veneto region and Carinthia) and another three are located in east central Europe (Sudetes, western Carpathian and southern Carpathians). These six locations correspond well to six of the seven nuclear clusters identiﬁed by genetic assignment of multilocus nuclear genotypes from a range-wide sample of extant populations, indicating that theses six clusters originated from the refuges identiﬁed by fossils: cluster 1 from the Turin area, cluster 2 from the Veneto region, cluster 3 from Carinthia, cluster 5 from the Sudetes, cluster 6 from the western Carpathians and cluster 7 from the southern Carpathians. Cluster 4 is found at the eastern Alpine margin that is poorly covered by fossil sites for that time. However, another study has shown that the eastern Alpine margin provided favourable environments allowing the persistence of endemic herbaceous taxa associated to upper mountainous forests (Tribsch € nswetter, 2003). This suggests that cluster 4 originated and Scho from another refuge situated at the eastern Alpine margin that could not be detected by paleontological data due to insufﬁcient fossil site coverage (“cryptic refuge”), a hypothesis to be conﬁrmed by additional data. Finally, considering the occurrence of early preliminary fossil evidence (Selle di Carnino, c. 16.2 ka) in the south-western Alps together with the occurrence of the rare mitochondrial haplotype 22 only found in this part of the range, we also suggest the existence of another refuge in this region that was less glaciated than other parts of the Alps (de Beaulieu, 1977). First recolonizations leading to the establishment of Larix pioneer forests at low-altitudes (<500 m) in formerly cold-steppic environments occurred since c. 16 ka (Vescovi et al., 2007). More prominent advances occurred between 14.5 ka and 12.8 ka in response to the abrupt warming of the late-glacial interstadial corresponding to DeO 1. At that time Larix shifted to higher altitudes in the Alps and the Carpathians as well as across the Polish Plain. These changes, documented by multiple fossil sites, demonstrate the important range-wide impact of DeO 1. However, at that time treeline formed by Larix and other species remained about 600 m below the current level (e.g. Gobet et al., 2005; Tinner, 2007; Tobolski and Ammann, 2000). The subsequent Younger Dryas cold episode (c. 12.8e11.7 ka) did not cause major longitudinal or latitudinal range changes, but elicited altitudinal shifts (mainly declines close to the timberline and expansions at lower altitudes). 4.3. Early Holocene recolonization and establishment of extant forest The abrupt temperature rise (4e6 C) at the beginning of the Holocene (c. 11.5 ka) triggered the establishment of extant L. decidua forests and involved treeline shifts over as much as 800 m within 200e300 years (Tinner, 2007). Our range-wide fossil compilation reveals that a major recolonization took place during the ﬁrst two millennia of the Holocene. The corresponding migration pathways can be inferred based on combined fossil and genetic data (Figs. 25 and 26). In the Alps, L. decidua likely expanded out of the four Alpine refuges using different Alpine valleys as corridors (Fig. 27). In addition, some limited recolonization might have 242 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 Fig. 25. Early fossil evidence (23.5e16 ka) and nuclear genetic clusters (after Wagner et al., 2015). Fig. 26. Early fossil evidence (23.5e16 ka) and mitochondrial haplotypes, haplotypes with an asterisk are microsatellite variants, the other ones simple sequence variants (after Wagner et al., 2015). S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 243 Fig. 27. Postglacial recolonization of the Alpine range. Dashed lines symbolize introgression of mtDNA. occurred from the potential south-western refuge, a hypothesis to be conﬁrmed by additional evidence. Recolonization of the Central Alps (<13 E) mostly started from sources in the Veneto region as most extant populations are assigned to nuclear cluster 2 and mitochondrial haplotype 18. The exceptional occurrence of two populations assigned to cluster 1 and haplotype 18 (c. 8 E, 46 N, region of Valais) might indicate that in these cases populations originating from the Turin refuge (nuclear cluster 1) swamped out populations from the Veneto refuge (mitochondrial haplotype 18) by pollen mediated gene ﬂow (e.g. Du et al., 2011). If this was the case, it seems likely that genetic exchanges took place before populations became restricted to high-altitudes as the Aosta zone separating extant cluster 1 populations and cluster 2 populations represents a particularly strong biogeographical barrier (e.g. ThielEgenter et al., 2011). Further eastwards, the Adige-Inn valley should have constituted a major corridor of migration. In the area of Carinthia data were scarce but migration in different directions seems likely. In east central Europe, extant populations should originate from geographically close-by refuges (Sudetes, western Carpathians and southern Carpathians). The additional nuclear subcluster detected in the south-eastern Carpathians may signify that there had been several small scattered refuges there, rather than a single large one, though this needs to be conﬁrmed by further investigations. 4.4. Past climate impacts versus past anthropogenic impacts After the ﬁrst two millennia of the Holocene L. decidua became largely restricted to high-altitudes due to the climate driven expansion of more competitive thermophilous taxa replacing it at low-altitudes. Anthropogenic changes occurred since the Late Neolithic (c. 6 ka) and became more important since the Bronze Age (c. 4 ka) when they started to provoke major populational declines and expansions. Major anthropogenic changes altering the ancient genetic structure started with the onset of translocations of forest reproductive material (seed or seedlings) across different range parts about 300 years ago (Wagner et al., 2015). Seen in the light of this study, translocations and artiﬁcial admixtures identiﬁed by Wagner et al. (2015) have reached such an extent that the original refugial gene pools have become hard to identify (in particular in the cases of the Sudetes and western Carpathians). Combined results of this study and of the genetic survey can now be taken as a guideline for establishing conservation strategies targeting the ancient genetic lineages. 5. Conclusions Our fossil compilation precisely documents range-changes of L. decidua over the last interglacialeglacial cycle in the context of long-term and short-term climate variability and anthropogenic changes. It uncovered distributional maxima of the species in northecentral Europe during the ﬁrst two early Weichselian interstadials. Several sites illustrate the species rapid response, within a few centuries, to millennial scale warming and cooling events (DeO cycles and HE). Seven (or possibly eight) last glacial refuges were identiﬁed with genetic data, six of which were corroborated by fossils, demonstrating the quality of the fossil compilation. Anthropogenic impact started at the end of the Neolithic and became more important since the Bronze Age, with important consequences for extant L. decidua populations. The genetic structure of the populations is determined by climate (ancient genetic structure) but also by human-induced translocations that changed the genetic make-up of extant populations, raising conservation concerns. The detailed historical inferences provide important keys for investigating adaptation and selection of L. decidua populations under ongoing climate change, for forthcoming palaeogenetic studies and for establishing management strategies for future forests, including plantations. Author contributions This work was part of a bi-national PhD thesis of SW, co- 244 S. Wagner et al. / Quaternary Science Reviews 124 (2015) 224e247 supervised by TL and RJP in Germany and France, respectively. SW conceived the study with TL and RJP and got additional advice from MFSG. SW performed the compilation and the mapping. SW analysed the data and wrote the paper with TL, RJP and MFSG. All authors have revised the manuscript and approved the ﬁnal version. Acknowledgements This project was funded by the German Research Foundation (DFG) (DFG LI 582/18-1) and the German Academic Exchange Service (DAAD). Data was compiled and mapped at the Steinmann Institute of the University of Bonn. We thank Thomas Giesecke and collaborators, W.O. van der Knaap, Petr Kunes, Rachid Cheddadi for pollen database access. For advices on the use of vegetation models we thank Rachid Cheddadi, Manuel Chevalier, Christian Ohlwein and Sophie Stolzenberg. 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