Veget Hist Archaeobot (2003) 12:233–247 DOI 10.1007/s00334-003-0021-8 ORIGINAL ARTICLE Magdalena Ralska-Jasiewiczowa · Dorota Nalepka · Tomasz Goslar Some problems of forest transformation at the transition to the oligocratic/Homo sapiens phase of the Holocene interglacial in northern lowlands of central Europe Received: 17 March 2003 / Accepted: 25 September 2003 / Published online: 20 November 2003 Springer-Verlag 2003 Abstract The paper discusses the main changes in the composition of mixed deciduous forests which occurred mostly between 5,000 and 2,000 b.p., based on selected pollen diagrams from the lowlands of Germany, Denmark and Poland, and including two pollen diagrams from varved sediments, used as reference sites, and on isopollen maps for Poland. The Ulmus retreat is shown on maps, and additional data for its pathogenic origin are presented. Corylus declines at ca. 3,500 b.p. at both reference sites, and its connection with Fagus expansion in the west and Carpinus expansion in the east is discussed. The nature of post-Atlantic transitory shrub– forest communities with dominant Corylus and Quercus is presented. Relationships between the history of Fagus and Carpinus and the development of human settlements are shown. Human impact has been admitted as one of the most important driving forces determining vegetational development since the time of fully developed Neolithic cultures. Other very important abiotic factors were the climate (particularly after 2,500 b.p.), and soil degradation. of climatic and ecological processes can be divided into four main phases (Fig. 1): 1. The first cryocratic phase corresponds to the transition from the late glacial to the early interglacial, when the formation of primitive soils, and their colonisation by pioneer herb communities begins. 2. The subsequent protocratic phase is a period of ameliorating climate, with soils still poor in humus. Formation of grasslands, shrubs and pioneer forests is in progress. 3. The mesocratic phase is a long, warm period when many mesotrophic trees end their migrations, multispecies mixed forests are being formed and settle on fertile, often brown soils. Keywords Oligocratic stage · Synchrony · Corylus decline · Fagus/Carpinus relationships · Human impact Introduction The Holocene is the most recent, still progressing interglacial of the Quaternary. According to the classical model proposed by Iversen (1958), any interglacial cycle M. Ralska-Jasiewiczowa ()) · D. Nalepka W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakw, Poland e-mail: [email protected] Fax: +48-12-4219790 T. Goslar Poznań Radiocarbon Laboratory, Rubież 46, 61-612 Poznań, Poland Fig. 1 Interglacial cycle in western Europe (Iversen 1958, modified by Andersen 1964, 1966). The scheme presents the main changes in regional vegetation and soils, as well as the inferred regional changes of climate (after Birks 1986) 234 Fig. 2 Pollen diagram of varved sediments from Lake Meerfelder Maar, Eifel, W Germany (selected pollen curves, after Kubitz 2000), shown to illustrate the synchrony of some main changes in forest composition around the transition to the oligocratic stage of the Holocene, with those recorded in the pollen diagram from Lake Gościa˛ż, central Poland (Fig. 3) 4. The deteriorating, or retrogressive telocratic phase has been further subdivided by Andersen (1964, 1966) into two stages—in the first, soils becoming impoverished (oligocratic stage) and, in the following stage, major climate cooling (telocratic stage s. str.). Both are accompanied by corresponding vegetational changes, characterised by the expansion of species which can tolerate leached soils and a cooler, more humid climate. first—between the protocratic and mesocratic phases— was characterised by intensive tree migrations and forestformation processes, with strong competition between tree species. The second—between the mesocratic and oligocratic phases—began after a long period of relative stabilisation of mixed, species-rich forests associated with the climatic optimum of the Holocene. Soils and later also climate deteriorated progressively, giving rise to a transformation of vegetation communities. In this paper we focus on the second transitional period, because by then the impact of human populations on the natural environment had already reached a rather advanced stage, and we attempt to determine how far it is possible to identify relationships between the natural and anthropogenic influences in this process. Recent precise investigations on sedimentary archives, yielding series of multi-proxy data based on fine-resolution dating, help to present some of those problems. Sequences of undisturbed series of varved (annually laminated) sediments come from key sites, such as in the Eifel region, W Germany, the lakes Holzmaar and Meerfelder Maar (Fig. 2) of volcanic origin (Zolitschka 1989, 1992, 1998; Litt et al. 1997, 1998; Kubitz 2000), and in central Poland, Lake Gościa˛ż (Fig. 3) of glacial origin (Ralska-Jasiewiczowa et al. 1998, 2001). These According to this scheme, the present state is the oligocratic phase of the Holocene interglacial. However, it refers to natural processes forming an interglacial cycle, whereas the Holocene interglacial differs essentially from the preceding ones, due to the presence of Homo sapiens. Human populations, active from the beginning of the recent interglacial, expand very fast, transforming the environment according to their needs. In the oligocratic/ telocratic phase of the Holocene, the impact of man on the environment increases rapidly, influencing vegetation, soils and finally also the global climate. The consequences of human activities and natural changes can hardly be distinguished from each other. The richest and most interesting periods of interglacial environmental change are two transitional periods. The 235 Fig. 3 Pollen diagram of varved sediments from Lake Gościa˛ż, central Poland (selected pollen curves, after Ralska-Jasiewiczowa et al. 1998), shown to illustrate the synchrony of some main changes in forest composition from around the transition to the oligocratic stage of the Holocene, with those recorded in the pollen diagram from Lake Meerfelder Maar (Fig. 2) sequences seem to provide signals of events referring to the whole area of northern central European lowlands, as shown by selected pollen diagrams (cf. below). The first version of this paper, which discussed only the pollen diagrams and did not include the isopollen maps, was presented at the ELDP (European Lake Drilling Project) meeting in Potsdam in May 2001. Discussion Material and methods The majority of pollen diagrams used in this article were digitised from the original papers. The only diagram from Flgelner Holz was plotted using the raw pollen data. We are very grateful to K.E. Behre for rendering them accessible to us. The Meerfelder Maar diagram was digitised from a copy obtained from the author, B. Kubitz, to whom we would like to express our thanks. In all, 188 sites have been compiled to trace the processes of forest transformation during the Late Glacial and the Holocene, and new isopollen maps have been produced by a group of Polish palynologists as a joint project. A set of maps illustrating the transition to the oligocratic phase of the Holocene is presented in this paper (cf. Appendix). The elm decline It has generally been accepted that the post-optimum transformation of deciduous woods began around 5,000 b.p., at the time of the so-called Ulmus decline (elm decline), broadly recorded in the European pollen diagrams. This was the period when the development of Neolithic settlements, which started in northern central Europe around 6,500 b.p., accelerated through the decreasing density of woodlands brought about by a large reduction in elm trees. The problem of the Ulmus decline in Holocene pollen diagrams was the subject of heated discussion for tens of years—different hypotheses were offered, e.g. change of climate, human impact, and finally the spread of the socalled “Dutch” elm disease caused by an ascomycete fungus (Ceratocystis ulmi), and carried over by bark beetles of the genus Scotylus (Rackham 1980). The majority of Holocene palynologists accepted the argu- 236 ments and evidence of elm disease as the triggering factor for elm retreat and, in consequence, for the acceleration of Neolithic land occupation (e.g. Groenman-van Waateringe 1983; Girling and Greig 1985; Birks 1986; Molloy and O’Connell 1991; Peglar 1993). There are isolated reports of the fall in Ulmus being recorded as 700– 800 years earlier, e.g. at Wolin Island at the NW Baltic coast (Latałowa 1992a, 1992b), or as recurring several times between ca. 6,500 and 5,400 cal b.p. in NW Denmark (Andersen and Rasmussen 1993). The pattern of Ulmus distribution on Polish isopollen maps shows the process of Ulmus decline as being more differentiated in time and space than previously accepted, but consistent within areas of varying geomorphology and landscapes (cf. Appendix). The distribution of Ulmus in the mesocratic deciduous forests of around 6,000 b.p. and somewhat after was fairly stable. Higher pollen values were occurring in the south, in uplands (Mamakowa 1961; Ralska-Jasiewiczowa 1977; Bałaga 1998), and in the mountains (Gil et al. 1974; Ralska-Jasiewiczowa 1980; Obidowicz 1996), reaching values of up to 10% or more, where U. glabra must have been the dominant elm species. During the Atlantic period, mountain elm was probably an essential element of the lower montane forest zone in the whole of the Polish Carpathians ( Ralska-Jasiewiczowa 1980). Today it still occurs above 1,200 m a.s.l. in the Tatras, and ca. 1,150 m a.s.l. in the Bieszczady Mts. (Zieliński 1979). The isopollen maps (cf. Appendix) show that Ulmus pollen was also then fairly abundant up to the Wielkopolska (Poznań-Gniezno region) in W Poland, there spreading over fertile soils, and also over the loess uplands. The central and western lowlands had less than 5% Ulmus pollen. This was the Holocene optimum of elm distribution. Around 5,200 b.p. Ulmus retreated from its upland distribution southwards, towards the mountains. At 5,000 b.p. a decline in its pollen values down to 2% or less occurred in a W–E area in N Poland, where U. laevis seems to have dominated over U. glabra (RalskaJasiewiczowa 1983). Subsequently (4,800 b.p.), this area broadened, covering the northern half of Poland, with pollen values of 3–5% remaining only as little islands in a few places in northern lake districts. The southern area of higher values (<5%) retreated to the Western Carpathians. The development of such a pattern of Ulmus reduction corresponds quite logically with the pathogenic hypothesis, and it is not concordant with the migration routes of successive Neolithic cultures. However, it seems convincing, as already supposed (Girling and Greig 1985; Birks 1986; Peglar 1993; Ralska-Jasiewiczowa et al. 1998), that Neolithic people took advantage of elm disease to clear the deciduous woods on more fertile soils faster and more easily. The following maps show the progressing, southward shift of reduced (1–3%) Ulmus values. At 3,000 b.p. the 5% isoline is not exceeded anywhere in Poland. At the time when late Neolithic cultures were declining, the mature soils were already subject to the leaching and podsolisation processes of the meso- to oligocratic stages Fig. 4 The plant communities forming the Quercion petraeae alliance in Poland (after Matuszkiewicz 1981) of the interglacial. After a temporary stage of pioneer copses composed of Betula, Populus tremula or Alnus incana depending on habitat, the invasion of many abandoned areas by wood species caused the development of forest–shrub vegetation of different composition. A rather open, woodland type with dominant hazel shrubs and free-standing, abundantly flowering oak trees must have been quite common. Oak–hazel vegetation types There are no strict modern analogues for such communities, but similarities can be found in the Quercion petraeae alliance (Fig. 4), such as the Lithospermo– Quercetum subboreale, a xerothermic scrub-wood of oceanic-Mediterranean affinities, now a relict in Poland, or the more shrubby Peucedano cervariae–Coryletum. Finally, we may also mention the Potentillo albae– Quercetum with a rich herb layer, open and rather heliophilous, but with little Corylus. Peucedano cervariae–Coryletum and Potentillo albae–Quercetum are thought to be closely connected with the vegetation of forest edges and marginal communities of anthropogenic origin (Rhamno–Prunetea, Trifolio–Geranietea), or as directly forming in woodland habitats after earlier, extensive long-term cattle grazing (Matuszkiewicz 1981). Between 4,000 and 3,400 b.p., a very distinct fall in Corylus pollen is indicated not only in Polish diagrams, but also rather frequently in those from the adjacent countries (e.g. Eifel region, Brauer et al. 2000, and Kubitz 2000; SE Westfalen, Pott 1986; see Fig. 2 and below). In the pollen profiles from both reference areas (Figs. 2, 3), this retreat is even more drastically indicated in the influx than in the percentage diagrams. The onset of this process was fixed at around 3,400 b.p. (=3,700 cal b.p.) in Lake Gościa˛ż, and at ca. 3,490 b.p. (=3,810 cal b.p.) in both lakes from the Eifel region. It is very clear that the effective migration and expansion of Fagus coming from the SW, and of Carpinus from the SE into central Europe, started at nearly the same time (Fagus—Eifel at 3,510 b.p.=3,840 cal b.p., and Carpinus—Lake Gościa˛ż at 3,420 b.p.=3,750 cal b.p.). In the case of Corylus, climatic changes seem to be the least probable reason for its decline. There are some hints of both thermal oscillation towards climate cooling (varve evidence from the Eifel region synchronous with the Lobben glacier advance in the Alps; Zolitschka 1992) and a decrease in climate humidity (a long period of decreased, although oscillating, humidity is recorded in 237 the lake levels of S Sweden between ca. 5,500 and ca. 3,000 b.p., Gaillard and Digerfeldt 1991; evidence of a drier period in the lakes of N Poland between 4,000 and 3,000 b.p., Ralska-Jasiewiczowa and Starkel 1988). If these climatic oscillations had any impact on vegetation changes, they would tend to affect the spread of latemigrating trees like Carpinus and Fagus. Corylus has a wide temperature tolerance and is also very adaptable to different soil types. In the Polish isopollen map for 4,300 b.p. (cf. Appendix), nearly all Poland is covered with Corylus pollen values <10%; in western-central Poland, in the zone around Gdańsk Bay, and at the south-eastern margin of the country those values reach 20%. At 3,800 b.p. Corylus has rather high, evenly distributed values above 10% in western, central and northern Poland, and also in the Carpathians and Roztocze Mts. The values over 20% occur in the peri-Baltic areas. The lower values (below 10%) cover the eastern part of the country up to the Podlasie border and the pre-Carpathian depression with the Małopolska Upland. Up to 3,300 b.p., Corylus percentages have reduced across the whole country except for the lake districts, the coastal zone, and the other limited areas of higher humidity in the mountains and eastern foothills. The further decrease to below 5% proceeds again from SE–E, following a similar trend as on the 3,800 b.p. map. As continental climate cannot be the explanation for this, we have to consider other reasons for the phenomenon. The development of Early Bronze Age cultures, like the Mierzanowicka culture which spread in the Małopolska region, practising rather primitive arable agriculture but broadly dependant on cattle grazing, may have contributed to the shrinking hazel distribution. The migration and expansion of Fagus and Carpinus from the SE–E and SW is synchronous with the Corylus fall, and possibly this migration pressure was one factor in the reduction of Corylus-dominated open shrub–woods. In the model of the interglacial cycle (Fig. 1), the Corylus–Quercus zone falls at the transitional time between the mesocratic and oligocratic phases. In the older interglacials, however, particularly in the Eemian with respect to the duration and character of vegetation cycles, no distinct corresponding phase has been observed. It can be concluded that the Holocene Corylus– Quercus period was a transitional one, without ecological equilibrium, and was merely a response to the cyclic abandonment of lands used at different times by different Neolithic populations. Various successional processes may have developed in such areas, leading to the formation of a forest cover different from the mixed deciduous forests of Atlantic type, which could not reexpand due to changed soil conditions and changing climate. Thus, Corylus–Quercus-dominated vegetation may have persisted for some time, being perhaps the oldest anthropogenic scrub–forest community. Beech and hornbeam expansion Around 3,500 b.p. (ca. 3,800 cal b.p.), the later-immigrating trees Carpinus and Fagus started to spread from their previous habitats to the south of the area discussed towards the central Europe lowlands. Obviously these two tree genera, having different ecological requirements, started their migrations in different landscapes and climatic conditions at a similar time. Fagus sylvatica, a tree of a cooler and more humid, rather oceanic climate, has heavy fruits and slow migration rates. However, it can easily colonise Corylus shrubbery, as its seeds are mostly distributed by animals and, at the seedling and juvenile stages, it has the best shade tolerance of all deciduous trees (Dzwonko 1990). Carpinus betulus, preferring more continental conditions, produces wind-distributed seeds. Its seedlings can hardly penetrate the rich herb layer, and high percentages are lost. Moreover, during juvenile growth the seedlings need 2–4 times more light than most trees of deciduous forests (Faliński and Pawlaczyk 1993). So, a high density of Corylus, together with a thick ground layer of already stabilised open vegetation, hampers its reproduction. However, once established after the pioneer tree stages with their decreased density of herb layer, it will very easily reproduce by suckering, colonising any forest gaps (Vera 2000). Also, it is very resistant to grazing and any other form of mechanical damage. In this way both trees have strong, but different potentials for expansion into areas occupied by the unstable Corylus–Quercus communities. In general, archaeological periodisation schemes for central Europe place the ending of the Neolithic period at 3,800 b.p. Kubitz (2000) marks the upper boundary of the Neolithic in the Eifel area at 3,800 cal b.p. (=3,490€20 b.p.). In the region of central Poland at Lake Gościa˛ż, the Late Neolithic Comb-Pitted Pottery Culture remained in situ, according to archaeologists, until ca. 3,860 cal b.p. (=3,580 b.p.), although its distinct traces in the pollen diagram end much earlier (before 4,000 b.p.; Ralska-Jasiewiczowa et al.1998). W to NW Germany Tracing the expansion of Fagus and Carpinus in the pollen diagrams from the central European lowlands (Fig. 5), one should begin in W Germany and continue in an easterly direction (Fig. 5 and below). The pollen diagram from Meerfelder Maar (336 m a.s.l.; Litt et al. 1997; Kubitz 2000) is shown in Figs. 2 and 6. The huge maxima in Fagus developing from around 3,500 b.p. (3,700–3,800 cal b.p.) mirror the decline in Corylus and are accompanied by a slight rise in anthropogenic herb indicators, connected undoubtedly with some Early Bronze settlement activities. A small presence of Carpinus pollen begins in the later part of the Fagus culmination, slightly before 2,900 b.p. (ca. 3,000 cal b.p.). The following long-term depression of the Fagus pollen 238 Fig. 5 Map showing the geographical location of mentioned sites in N Germany, Denmark and N Poland. 1 Meerfelder Maar (Kubitz 2000), 2 Erndtebrck (Pott 1985, 1986), 3 Rehhornsmoor (Drfler 1989), 4 Flgelner Holz I (Behre and Kučan 1994), 5 Holmegaard (Aaby 1986), 6 Felchowsee (Jahns 2000), 7 Kołczewo (Latałowa 1989, 1992a, 1992b; Ralska-Jasiewiczowa and Latałowa 1996), 8 Żurawiec (Latałowa 1989), 9 Puszcza Darżlubska (Latałowa 1982, 1989), 10 Lake Mały Suszek (Miotk-Szpiganowicz 1992), 11 Lake Skrzetuszewskie (Tobolski 1990, 1991), 12 Lake Gościa˛ż (RalskaJasiewiczowa et al. 1998), 13 Woryty (Pawlikowski et al. 1982; Ralska-Jasiewiczowa and Latałowa 1996) curve (ca. 2,650–1,640 b.p.=ca. 2,700–1,500 cal b.p.) finds its explanation in the high maximum of herbs, dominated by human indicators, which covers a long series of subsequent cultural phases from the Hallstatt C, through the La Tne and Roman times, up to the beginning of the Migration period and the onset of the next Fagus rise. This second, shorter Fagus maximum is synchronous with the Carpinus rise, culminating during the early stage of woodland re-expansion. The distinct Corylus-pollen peak from an early phase of forest regeneration shows the relation of hazel expansion to settlement history. The deep depression of the NAP curve coincides with this shrub- and tree-regeneration period which commonly occurs in central European pollen diagrams in connection with the Migration period. The pollen diagram from the mire at Erndtebrck, 470 m a.s.l. (Pott 1985, 1986; Fig. 6), located in the lowmountain area of Siegerland, SE Westfalen, is shown here for comparison. This only documents the forest history from around 3,500 b.p. (ca. 3,800 cal b.p.). It also reveals a Fagus rise from ca. 3,300 b.p. (ca. 3,500 cal b.p.), very similar to and only slightly later than that in the Eifel diagrams. Fagus regression starts from ca. 2,400 b.p., possibly due to human activities from early Roman (preRoman?) times, and then, after a short regeneration phase, from the Roman settlement period, lasting until ca. 1,700– 1,600 b.p. It seems that also here the decline of the Roman period and the beginning of the Migration times provoke forest regeneration also enabling Carpinus to expand. Such a pattern is rather typical for foothill and low montane landscapes. The record of this site is not complete and not fully comparable, because the NAP raw data were not available and so the diagram is based on the AP sum. However, the curves of human indicators in the complete diagram (Pott 1985) clearly show the pattern described above; the settlement phases are also Fig. 6 Age correlation of selected pollen curves (Carpinus, Fagus, Corylus, NAP) from five pollen diagrams from N Germany and one diagram from Zealand, Denmark, showing the age relationships between the Corylus fall and Carpinus and/or Fagus rise, and the inverse correlation between the maxima of both trees and maxima of NAP (for authors of pollen diagrams see Fig. 5 caption) 239 Fig. 7 Age correlation of selected pollen curves (Carpinus, Fagus, Corylus, NAP) from seven pollen diagrams from N Poland (for further explanation see Figs. 5, 6) 240 characterised by rises in Corylus values. In the central Mnsterland, the oak–hornbeam region of the low NW part of Westfalen, Carpinus begins to expand from ca. 2,700 b.p., reaching values up to 30% AP around the time of the birth of Christ (Pott 1985). The following German sites illustrate the history of woodland in the North Sea coastal zone. At Rehhornsmoor in the Cuxhaven region, Lower Saxony (Drfler 1989; Fig. 6), the pollen record represents the woods of fresh to humid soils formed on loam beds where, after ca. 5,000 b.p., oak was the main wood-forming tree together with hazel. The first slight rise in the Fagus curve began here rather late, after 2,500 b.p., clearly induced by the change of climate. However, it coincides with the Corylus fall, and also with the beginning of a distinct settlement phase. Probably in this area influenced directly by maritime climate, the rise of humidity in the earlier Subatlantic period triggered the Fagus migration in spite of land use, but only the events connected with the Migration period around 1,600 b.p. enabled its vigorous expansion. Then the wood regeneration phase also starts with a short episode of Corylus development. Synchronously, a slow spread of Carpinus develops gradually, up to the small culmination closely preceding the high Fagus expansion which rapidly ousted the hornbeam during the economic breakdown ca. 600–400 years ago. For the duration of its Holocene history in central Europe, a distinctly decreased vitality of hornbeam in maritime climate areas is obvious. In the adjacent region of Lower Saxony, several pollen diagrams were presented by Behre and Kučan (1994) in the context of a large project on the history of the Flgeln “Siedlungskammer”. The diagram from the kettle-hole mire Flgelner Holz I is reproduced here (Fig. 6). After the elm decline around 4,400 b.p., forest cover was composed of birch–oak and alder woods, similar to those described from the environs of Rehhornsmoor in the adjacent Cuxhaven region. It was inhabited in the early Neolithic by the tribes of the “Trichterbecher” culture which was based mostly on animal husbandry. The appearance of Secale cereale starts from ca. 2,370 b.p., and is coincident with the slight fall in the Corylus curve, synchronous with the first significant rise in Fagus. Since that time indicators of agricultural activities occur regularly. Excavation from the pre-Roman period showed a Celtic field-pattern where probably a slow rotation system was used until the 1st to 2nd century a.d. Fagus, and from ca. 1,840 b.p. also Carpinus percentages were rising slowly but continuously up to ca. 1,640 b.p. when a distinct rise to a peak in Fagus began, together with a depression in herb values. The deepest depression of these coincides with the Fagus maximum at 1,100 b.p. (880– 990 b.c.) and lasts until 1480–1950 a.d. Denmark In the pollen diagram from Holmegaard (Fig. 6) in Zealand, Denmark (Aaby 1986), Fagus and Carpinus appear synchronously, around ca. 3,100 b.p. (3,350 cal b.p.), but Fagus expands rapidly and forms a huge maximum within ca. 200 years, reflecting closely the Corylus fall. Carpinus is present only scarcely. These events clearly precede the development of settlements around 2,400–2,300 b.p. which, during its first stage, resulted in very strong devastation of Fagus woods. It permitted hornbeam to expand a little, and later Corylus shrubs regenerated on gradually exploited and abandoned fields and pastures. This phase was followed by the second Fagus expansion during the Migration period. Carpinus remained at very low values all the time, most probably for climatic reasons. E Germany and Poland At Felchowsee in NE Brandenburg (Germany, Fig. 6), close to the Lower Odra River Valley (Jahns 2000), Fagus and Carpinus appear simultaneously with a very slow Corylus decline. This is still happening during the Neolithic settlement period clearly recorded in the diagram between ca. 5,279 b.p. and ca. 3,700 b.p. The woods of the region were clearly dominated by oak and pine, with a very small contribution from other trees, apart from birch. The vitality and expansion potential of both species, Fagus and Carpinus, seem to be nearly equal. Carpinus starts spreading around 2,400 b.p., during the recessional settlement phase between the La Tne/preRoman and Roman periods, and Fagus follows by the end of this recession phase, from ca. 2,100 b.p. However, again both trees reach their maximum values during the Migration period. Those analyses of Fagus/Carpinus competitive relationships within areas influenced by the maritime climate of the North Sea and SW parts of the Baltic Sea point to human impact as an important factor in the process of vegetation transformation, while the regional climate type and climatic changes were also contributing in an essential way to these processes. Analyses of pollen data from the Polish lowlands show the eastward continuation of the Fagus versus Carpinus history. The pollen diagram from Kołczewo on Wolin Island (Fig. 7), at the outlet of the Odra River into the Baltic Sea (Latałowa 1989, 1992b; Ralska-Jasiewiczowa and Latałowa 1996), reveals the occurrence of deciduous forests with abundant oak and hazel up to ca. 2,900 b.p. However, the hazel already shows its first distinct depression after 3,300 b.p. The Early Bronze, and then the Late Bronze (Lusatian) peoples settled in the area, impeding the spread of Carpinus and Fagus, both already present in the region—Carpinus from before 3,000 b.p., and Fagus appearing slightly later (ca. 2,900 b.p.). The Subatlantic change of climate at ca. 2,500 b.p. accelerated the expansion of Fagus which lasted through the Migration period until Medieval times. Carpinus never played any significant role in the region. The successive pollen diagrams from sites distributed to the east show the gradual change in the Fagus/ 241 Carpinus relationship: the pollen diagram from Żurawiec (Fig. 7) situated in the middle Baltic coastal zone (Latałowa 1989, 1995; Ralska-Jasiewiczowa and Latałowa 1996) documents the rapid expansion of Carpinus during the Migration period, after the devastation of forests by cultures previously settled there ceased. This very rapid expansion closely preceded the invasion of Fagus. Such a sequence of events was most probably caused by the gradual extension of the cool, humid climate eastwards, as suggested by Latałowa (1995). At the following sites, the first from the eastern part of the Baltic coastal zone (Puszcza Darżlubska, Latałowa 1982, 1989; Ralska-Jasiewiczowa and Latałowa 1996), and the next located to the south in the West-Pomeranian Lake Districts (Lake Mały Suszek, Miotk-Szpiganowicz 1992; Ralska-Jasiewiczowa and Latałowa 1996; Fig. 7), the influence of marine climate is distinctly weaker. In both cases Carpinus begins to rise around 3,000 b.p., preceding the development of the Lusatian culture. At Puszcza Darżlubska Carpinus increases slightly, together with a slight rise in very low Fagus values at the breakdown of this culture after 2,500 b.p., and then reaches a maximum at both sites during the Migration period. Fagus increases a little again, but never assumes a major role in these regions. In areas of stronger continental climate influence, such as the southern lake districts of the Wielkopolska region (Poznań-Gniezno region, Lake Skrzetuszewskie, Tobolski and Okuniewska-Nowaczyk 1989; Tobolski 1990, 1991), the south-eastern margins of the lakelands at Płock Basin in central Poland (Lake Gościa˛ż, Ralska-Jasiewiczowa et al. 1998), and the western part of the Mazurian Lake Districts (Woryty, Pawlikowski et al. 1982; RalskaJasiewiczowa and Latałowa 1996; Fig. 7), Fagus completely loses its forest-forming ability. The pollen data from the latter three sites will be described together, because the changes they demonstrate are very similar. The first Carpinus expansion develops in all the three profiles between ca. 3,700 and 3,400 b.p., and coincides with the Corylus fall at the end of the Neolithic. At Lake Skrzetuszewskie the Carpinus maximum is enormous (up to ca. 40% of total pollen), and lasts several hundreds of years. In the Wielkopolska Lake District area, a region of very fertile soils, Carpinus became the most common tree (Tobolski 1990, 1991). Settlement activities at that time, as shown by the anthropogenic pollen indicators, were not very intensive in the region in question. At Lake Gościa˛ż, the first Carpinus expansion developed at a similar time as at the Skrzetuszewskie Lake (3,700–3,500 b.p.), but its maximum spread was less marked (slightly exceeding 20%) and lasted no more than 200–300 years. The Early to Late Bronze (Lusatian) cultures had already started to clear the hornbeam-dominated woods there by about 3,200 b.p., and these clearances lasted, as illustrated by the record from the varve sequence, for ca. 1,000 years overall. At Woryty the first rise in Carpinus pollen values was slightly delayed compared with the latter two lakeland sites, and was still smaller (ca. 10%). The young hornbeam stands must have been cleared quite early by the Late Bronze populations arriving and settling in the lake’s immediate surroundings (Da˛browski 1981). The following, rather short woodland regeneration phase occurred at Lake Gościa˛ż and at Woryty between 2,300 and 2,100 b.p. This was the consequence of the successional processes which followed the retreat of the Lusatian culture, through lake water-level rise and paludification of the lake surroundings which started after 2,500 b.p. In both cases the restoration of hornbeam woods was preceded by the spread of pioneer birch stands. At Lake Skrzetuszewskie the maximum in settlement indicators following the first huge culmination of Carpinus is dated at 2,700 b.p., and the second Carpinus maximum at 2,460 b.p. Thus, the expansion of Lusatian settlers seems to have been much shorter here (if these dates are correct). The next deforestation of the regenerated hornbeam woods seems to have been by the populations of the Roman period after 2,000 b.p. The wood devastation, although lasting a much shorter time, must have been intense. The last expansion of hornbeam-dominated woods in our area very clearly resulted from events connected with the Migration period. The dates of ca. 1,600–1,500 b.p. nearly everywhere mark the beginning of hornbeam spread (at Lake Skrzetuszewskie, it is dated at 1,800 b.p.), but the duration of the spread is highly dependent on the local history of human settlement and economy. The Fagus and Carpinus successions illustrated on the isopollen maps for Poland (cf. Appendix) reveal successional distribution on a broader scale, including the southern areas of the country and tree migration routes. Fagus maps demonstrate a very consistent migration pattern (cf. Appendix). At around 4,000 b.p., beech entered the Polish mountains from the SE and SW, spreading gradually over the uplands. Its later migration from the NW along the seacoast evidently developed in close connection with the Subatlantic climate change, only after 2,500 b.p. The maximum Fagus distribution had been reached between 1,700 and 1,400 b.p. Around 1,000 b.p., beech representation in the forests of NW Poland decreased substantially. Carpinus is the most problematic taxon for mapping (cf. Appendix) because of its very fast reaction to changes in settlement, which are often non-synchronous. Its ability to spread rapidly gives a chaotic impression on the maps but, in fact, there is some consistency in the pattern formed: coming from the SE (Bieszczady Mts.) around 4,000 b.p., Carpinus migrates very fast in a NW direction and around 3,300 b.p., just before the expansion of Lusatian culture, establishes its first, very broad centre in central-west Poland (Wielkopolska). The only areas with a less-intense spread of hornbeam remain the peri-Baltic and northern lake district zones. This migration and expansion pattern is in contrast to that of well-known, older reconstructions, e.g. Firbas (1949, p. 263, Fig. 36). Between 2,900 and 2,500 b.p., the areas dominated by hornbeam shrink gradually due to the development of Lusatian settlement. The decline of this culture after 242 Fig. 8 Isopollen maps for Ulmus 2,500 b.p. is reflected in some renewal of Carpinus woods, at first rather patchy (2,400 b.p.), but gradually covering nearly the whole country at values of 5–7%. In the south-western areas of Dolny Śla˛sk (Lower Silesia) up to the Wielkopolska lake districts and lowlands, the hornbeam percentages are still higher at that time. A brief setback caused by the increase of economic activities during the Roman period (1,800 b.p.) is recorded in eastern Poland, partly in the Carpathians, their foothills, the uplands and in the central coastal zone. On the other hand, the Migration period (1,400 b.p.) was the most advantageous time for Carpinus. Its expansion was very strong in most parts of the country, decreasing gradually from the uplands of S Poland and Polesie Lubelskie towards the SE, to values below 5% in the central Carpathians and their foothills. The last map shows the decrease in Carpinus pollen percentages dating back to the earliest period of the Polish State. 243 Fig. 9 Isopollen maps for Corylus Conclusions The transformation of forest cover during the transition between the mesocratic and oligocratic phases of the Holocene (ca. 5,000–3,000 b.p.) was driven to a great extent by man. However, the later general change towards a more maritime climate (from ca. 2,500 b.p.) with higher humidity, lower temperatures, and milder seasonal contrasts made an essential contribution to the histories of late-migrating trees, especially to the extension of Fagus in the NW territories of Poland. Regional settlement activities were combined with, and strongly dependent on, the regional climate and its changes and, in this respect, the climate was also indirectly responsible for successive human population expansions, contractions and changes in activity, influencing the development and reduction of woodland cover and its composition. Appendix The Appendix shows isopollen maps for Poland: Fig. 8, Ulmus; Fig. 9, Corylus; Fig. 10, Fagus; and Fig. 11, Carpinus, at individually selected time slices. These maps have been based on data of the following authors: M. Ralska-Jasiewiczowa1, Dorota Nalepka1, Zofia Balwierz2, Krystyna Bałaga3, Anna Filbrandt-Czaja4, Wojciech Granoszewski1, Krystyna Harmata5, Krzysztof Krupiński6, Mirosława Kupryjanowicz7, Małgorzata Latałowa8, Jacek Madeja5, Ewa Madeyska1, Mirosław Makohonienko9, Kazimiera Mamakowa1, Krystyna Milecka9, Grażyna Miotk-Szpiganowicz10, Agnieszka Noryśkiewicz11, Bożena Noryśkiewicz12, Małgorzata Nita13, Andrzej Obidowicz1, Iwona Okuniewska-Nowaczyk14, Kazimierz Szczepanek5, Kazimierz Tobolski9, Agnieszka Wacnik1, Adam Walanus15, Krystyna Wasylikowa1, Joanna Zachowicz10 1. W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakw, Poland 2. Department of Geomorphology, Łdź University, Narulowiera 88, 90-139 Łdź, Poland 3. Department of Physical Geography and Palaeogeography, Maria Curie-Skłodowska University, Akademicka 19, 20-033 Lublin, Poland 4. Institute of Biology and Environment Protection Nicholas Copernicus University, Gagarina 9, 87-100 Toruń, Poland 5. Institute of Botany, Jagiellonian University, Lubicz 46, 31-512 Krakw, Poland 244 Fig. 10 Isopollen maps for Fagus 6. Polish Geological Institute, Rakowiecka 4, 00-975 Warszawa, Poland 7. Institute of Biology, Białystok University, Świerkowa 20b, Białystok, Poland 8. Laboratory of Palaeoecology and Archaeobotany, Department of Plant Ecology and Nature Protection, Gdańsk University, Legionw 9, 80-441 Gdańsk, Poland 9. Quaternary Research Institute, Adam Mickiewicz University, Fredry 10, 61-701 Poznań, Poland 10. Polish Geological Institute, Sea Geology Branch, Kościerska 5, 80-328 Gdańsk, Poland 11. Institute of Archaeology and Ethnology, Nicholas Copernicus University, Podmurna 9/11, 87-100 Toruń, Poland 12. Institute of Geography, Nicholas Copernicus University, Fredry 6/8, 87-100 Toruń, Poland 13. Earth Science Faculty, University of Silesia, Be˛dzińska 60, 41200 Sosnowiec, Poland 14. 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