tree species composition and historic chang- es of … · tree species composition and historic...

61Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe

Tree species composition and historic chang-es of the Central European oak/beech region

Carsten Rüther 1 and Helge Walentowski 2

1 Lorettoplatz 8, 72072 Tübingen, Germany 2 Bavarian Forest Institute (LWF), Am Hochanger 11, 85354 Freising, Germany

Corresponding author: Carsten Rüther, email: [email protected]

Citation: Rüther, C. and Walentowski, H., 2008. Tree species composition and historic changes of the Central European oak/beech region. In: Floren, A. and Schmidl, J. (eds): Canopy arthropod research in Europe, pp. 61-88, bioform entomology, Nuremberg.

Abstract

After the ending of the � nal glaciations, tree species re-immigrated to Central Europe from their southern refuges. Fagus sylvatica (beech) was the last principal tree species to arrive at the northernmost areas of the Central European region, viz. north-western Germany, Denmark and southern Sweden. Due to their high competitive ability, F. sylvatica plays a predominant role in Central European Fagion-forest communities, whose native ranges are lowlands and low mountain ranges. Depending on local soil and climatic conditions, other tree genera are included such as Quercus (oak), Picea (spruce) and Abies (� r) (both Pinaceae). Indeed, only under exceptional environmental conditions is the competitive ability of F. sylvatica limited in such a man-ner that other tree species may predominate. Even rare occurrences of one associated tree species within beech forests may assure a long habitat tradition for specialised biocoenoses. Since the Neolithic period, human activities interfered with the forest-covered regions of Central Europe. The forests were changed sig-ni� cantly by multiple historical uses and management techniques. Anthropogenic transformation of the forests was driven by wood pasture, pollarding techniques, litter utilisation, coppicing, coppicing with standards and high-forest systems (selective and clear-cutting). These had crucial effects on tree species composition as well as the stand climate of the forests. They changed the spatial and temporal structures of the stands as well as the cycling of soil matter. Another effect of human activities was drastic reduction of forest cover. The unwitting transition of tree species composition � nished in the Middle Ages. Since then, human interference has concentrated on targeted timber selection. These alterations make clear that structural components and site conditions of commercial forests deviate considerably from those of the original natural forests. Modern management methods have to consider biodiversity and the degree of naturalness with regard to the tradition of old-growth trees, natural tree species combination and natural ground layer vegetation. Important stan-dards for a sustainable forestry combined with conservation of biodiversity are heterogeneity, complexity and scale.

Zusammenfassung

Nach der letzten Eiszeit wanderten die Baumarten aus ihren südlichen Refugien wieder nach Mitteleuropa ein. Als letzte der Hauptbaumarten erreichte die Buche (Fagus sylvatica) die nördlichen Gebiete Mitteleu-ropas: Nordwestdeutschland, Dänemark, Südschweden. Aufgrund ihrer ausgeprägten Konkurrenzkraft spielt sie in den mitteleuropäischen Fagion-Gesellschaften eine vorherrschende Rolle; derartige buchen-reiche Wälder würden natürlicherweise weite Teile der niederen Lagen und der unteren und mittleren Mit-telgebirgsstufe bedecken. Andere Baumarten wie Eiche, Fichte und Tanne sind in Abhängigkeit von den lokalen Boden- und Klimabedingungen am Aufbau der Baumschicht mehr oder weniger stark beteiligt. Nur

62 Rüther and Walentowski: Tree species composition and historic changes…

Eemian periods (128000-115000 B.P.), large herbi vores were still present. These periods showed a very similar development of for-est reestablishment after glaciation in com-parison with the early Holocene (10300 B.P.). Therefore, the stated signi� cant impact of the large herbivores, which became extinct at the start of the Holocene on the vegetation has to be rejected (LWF 2000).

Conversely, Central Europe is also a zone of most intensive anthropogenic use. Since at least the second half of the sixth mille-nium before Christ (B.C.), forests of natural origin were transformed progressively by human in� uence (Lang 1994, Küster 1996, 1998a). In north-western Germany as well as in some loess regions of central and southern Germany, the � rst farm cultiva-tions were established. The regions’ inhab-itants farmed � elds and bred cattle, pigs, sheep and goats. Cattle pastured freely in the surrounding forests, a method still prac-tised locally in extensively-managed areas in the Balkan Peninsula and Carpathians. In addition to the production of timber, � re-wood, lumber and leaf litter, the early farm-ers also took fruit for their diet, to extract oil or as a winter supply for their domestic animals. Further forest uses followed, such as the production of charcoal or ash burning

Introduction

Naturally, the temperate forest zone of Central Europe would be a mostly uniform forest area (forest cover 95%), dominated by deciduous trees. Only dunes, marshes, bogs, rocky outcrops and few alpine regions above the upper timberline were once un-wooded (Ellenberg 1996). The vast progress made in palaeobotany and the amount of data available prove conclusively that Europe was natu rally forested, either with or without mega-herbivores (Litt 2000). Pollen diagrams from many localities across Central Europe show a clear pattern of post-glaciation refor-estation, its tree species combinations and also the degree of openness of European forests, both during Eemian inter-glacial pe-riod and the beginning of the Holocene prior to human intervention. Pollen samples show exactly the same result: measured by the ratio of non-tree pollen to tree-pollen, the European landscape was nearly covered by woodland which formed forests in the true sense and without resembling a park land-scape. This is true for the entirety of Cen-tral Europe, since none of the known pollen pro� les deviate from this picture by showing signi� cant amounts of non-tree pollen. In both the Holstein (347000-362000 B.P.) and the

auf Sonderstandorten ist die Konkurrenzkraft der Buche derart eingeschränkt, dass andere Baumarten zur Vorherrschaft gelangen können: daher kann gerade das seltene Vorkommen einer Begleitbaumart innerhalb von Buchenwäldern als Beweis für eine lange Habitattradition des Bestandes herangezogen werden. Seit dem Neolithikum be ein� usst der Mensch die Waldlandschaft Mitteleuropas. Die Wälder wurden aufgrund der vielfältigen histo rischen Nutzungen und Bewirtschaftungstechniken erheblich verändert. Für die Veränderun-gen waren in erster Linie die Waldweide, die Schneitelwirtschaft, die Streunutzung, die Nieder- und Mittel-waldwirtschaft sowie die Hochwaldwirtschaft (Kahlschlag, Plenterwaldwirtschaft) verantwortlich. Diese hatten entscheidenden Ein� uss auf die Baumartenzusammensetzung und das Bestandsklima der Wälder. Darüber hinaus veränderten sich die räumliche Struktur und die zeitliche Entwicklung der Bestände sowie die Stoff-kreisläufe im Boden. Parallel dazu führten die anthropogenen Eingriffe zu einer drastischen Verringerung der Wald� äche. Bis ins Mittelalter erfolgten die Veränderungen eher unbewusst; seit dem Spätmittelalter hat der Mensch durch die Auswahl bestimmter Baumarten bewusst in die Baumartenzusammensetzung eingegriffen. Die Veränderungen machen deutlich, dass die strukturellen Eigenschaften und die Standortbedingungen der bewirtschafteten Wälder deutlich von denen natürlicher Wälder abweichen. Um den weiteren Verlust an Bio-diversität aufzuhalten, muss die moderne nachhaltige Forstwirtschaft klar de� nierten Mindestanforderungen an Biodiversität und Naturnähe gerecht werden, d.h. in Bezug auf Strukturparameter (v.a. Alt- und Totholz), standortscharakteristische Arten (Baumartenzusammensetzung, Bodenvegetation) und Funktionalität (räum-liche und zeitliche Heterogenität und Komplexität).


Anthropogenic impact on Central Euro-pean forests has differed both regionally and temporally. Thus, the original conditions of these forests can only be estimated. Never-theless, some quantitative changes can be detected and qualitative effects assessed. These include:

(1) diminishment of welfare and protective functions;

(2) alteration of tree species composition (including direct and indirect in� uence on the role of indigenous species for the vegetation and introduction of for-eign species);

(3) modi� cation of forest structure;(4) impact on forest dynamics/cyclic phas-

es;(4) modi� cation of micro-climate;(5) change to substratum conditions, e.g.

nutrient supply;(6) alteration of ground cover species

(abundance and species combination) and

(7) shift of competition equilibrium; devel-opment of secondary vegetation forma-tion and plant communities of varying degrees of naturalness.

In order to understand the radical altera-tion of the character of forest vegetation, an investigation of the ecological properties of the tree species (e.g. geobotanical site fac-tors, shade tolerance, coppice sprouting, dis-semination, reproduction) is crucial. Abrupt spatial changes in species composition of Central European forests re� ect small-scale patterns in topographical and geological site factors, as well as management in� uences. Therefore, it is appropriate to de� ne discrete forest units, i.e. plant associations or vegeta-tion units generated by clustering, rather than arrange plots along underlying environmental gradients. Well-de� ned forest units are es-sential for mapping projects which are intend-ed for management strategies. For assess-ment of the transformation caused by human inference to forests with reference to former vegetation conditions, the post-glacial forest history is outlined on the basis of palynologi-cal data.

and subsidiary uses such as bee keeping or tan-bark peeling.

These forest uses effected a complex change of structural and ecological conditions in the forests, alterations of species abun-dance and species combination as well as a clear reduction of the forest area. Decisive factors for the degree of anthropogenic trans-formation were duration, intensity and nature of the impacts and the resultant ecological site conditions. In extensively-grazed areas, non-natural open pastures with individual groups of bushes as well as park-like stages and groves, alternating with closed forest stands, were formed (Pott 1983, Ellenberg 1996). Instead of natural forests, numerous substitute communities emerged, so that a di-verse mosaic of secondary forests, pastures, meadows, heaths and � elds is found today. Fragmentation, soil degradation and isolation of forests contributed to these effects.

Present-day forests in Central Europe have been modi� ed completely by man. In order to obtain an impression of the forests before their anthropogenic transformation, we must visit inaccessible and high-altitude forest areas. Some relics of virgin forests worth mentioning are Rothwald (Northern Limestone Alps, Austria) and Kubany (Krušné hory and Šumava, respectively; both Czech Republic) (Zukrigl 1978, Leibundgut 1982, Zukrigl 1984). Due to previous and current human impact, forest reserves and forest na-tional parks provide only a reduced picture of original forests. For numerous forest areas, the label virgin forest (e.g. Neuenburger Ur-wald, Nitzschke 1932, see also Rüther and Peppler-Lisbach 2007) is misleading. Most of these citations concern rare and irregularly-used (e.g. as forest pasture or for browsing) or otherwise modi� ed natural forests, but by no means original virgin forests.

Available data on forest dynamics, struc-ture, species composition, species propaga-tion and matter cycles and their interaction with soil and vegetation are derived, almost exclusively, from investigations of secondary forests. However, these do not permit extra-polation for forests of natural origin, such as those encountered by the � rst Central Euro-pean farmers.


Site conditions

Geology and morphology

In terms of geology and morphology, Cen-tral Europe may be divided into two principal zones (Walter 1992):

(1) Most of the Central European lowland, situated in the region’s north, is below 50m a.s.l. and attains elevations of 200m a.s.l. in few places only. Its tallest peak is Wie�yca (329m a.s.l.), which is part of the Pomeranian young moraine in Poland. Both the geologi-cal material and the shape of the surface are comparatively young. Vast quantities of sedi-ment were deposited during the Pleistocene ice ages and formed the surface texture. Dur-ing the inter-glacial and the Holocene periods, further relocating and modelling processes occurred. The base saturation of the soils dif-fers depending on the age of the deposit: Old moraine landscapes are strongly decalci� ed, whereas young ones are rich in bases. The pre-Quaternary underground is only revealed in few places (e.g. shelly limestone occur-rences at Rüdersdorf, Berlin).

(2) The low mountain range with heights between 200 and 1000m a.s.l. follows south-wards. Individual summits [e.g. Feldberg (Black Forest), Großer Arber (Bavarian For-est), ] attain elevations between 1,500 and 1,600m a.s.l. Mountain ranges and basin-like landscapes appear as mosaics adjacent to each other. The low mountain ranges are older than the lowlands. Their basement is formed by eroded moun-tain trunks of the Variscian period (age 420 to 250 million years). These carry, with some ex-ceptions, the layers of the Mesozoic (Triassic, Jurassic, Cretaceous) as well as Caenozoic (Tertiary period, Quaternary) overlying strata. The orientation of the mountain ranges and axial depressions are caused by tectonic rea-sons. Frequent orientations are NNE-SSW (Rhenish), SE-NW (Hercynian) and SW-NE (from the Krušné hory). The northern Alpine foothills are added to the central mountain area in this concept, composed of Caenozoic sediments and with a surface texture with the regional character of a hill country.

Outline of the area

The boundaries of Central Europe vary signi� cantly depending on their historical, cultural, political, geological, climatic or geo-botanical de� nition. The simple geographical term concentrates on the central part of the European continent and, from this, Ellenberg (1996) gave a combined climatic-geobotani-cal de� nition with Central Europe lying be-tween 47o and 53oN. Essentially, this covers Germany, Poland, the Czech Republic, Slo-vakia, Austria, Switzerland, Luxembourg and parts of the adjacent states.

In our study we follow the boundaries of Rubner and and Reinhold (1953) and Mayer (1984), which are de� ned by natural forest re-gions. The Central European oak/beech for-est region covers Germany (excluding East Frisia and the East Frisian islands), Denmark (excluding western Jutland), the southern part of Norway as well as the southern point of Sweden, the northern part of Poland (in-cluding the north-west and south-west re-gions and the Silesian basin), the Czech Republic (excluding the territory east of the River Morava), as well as parts of non-Alpine Austria and Switzerland. Its western bound-ary approximates the frontiers of Germany, France and the Benelux countries (Fig. 1).

Fig. 1: Natural forest regions of Europe (according to Rubner and Reinhold 1953, Mayer 1984).


as well as the windward and leeward situation (Fig. 3). Increasing altitude above sea level is linked to a thermal and hygric gradient. For the Bavarian Forest, Baumgartner (1970) in-dicated a decrease of 0.5°C per 100m differ-ence in altitude. This gradient is overlain by a north-south decline. From north to south, the altitude belts shift upwards because an-nual average temperatures on the same al-titude increase and the vegetation period is prolonged. Therefore, the upland parts of the Black Forest with high annual temperatures have milder winters in comparison with the Harz Mountains. However, with increasing continental conditions (i.e. eastwards) the vegetation levels are shifted vertically, due, primarily, to lower occurence of cloud cover and more intensive thermal radiation.

With increasing altitude, precipitation in-creases and reaches a maximum on the

Climate

Climatically, Central Europe is affected by both atlantic and continental climates. In the west, the atlantic character predominates with a balanced cool-humid climate, without tem-perature extremes and rainfall in all seasons (oceanic conditions). Eastwards, the conti-nental character increases with warmer sum-mers and colder winters; annual precipitation with brief and heavy rainfall concentrates on the summer months (continental conditions). From north to south, annual precipitation in-creases notably towards the Alps. The rain catcher effect is pronounced in summer. It is caused by the humid Atlantic airstreams form the north-west meeting on the mountains (Fig. 2).

In the low mountain ranges, the macro-climate is modi� ed notably by the elevation

Fig. 2: Climatic diagrams from the planar-colline to sub-montane level of Central Europe. Data base: www.klimadiagramme.de.

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(e.g. the dry and hot summer 2003). Temper-ate, cool-humid climate favours tree growth, in particular summer-green hardwoods such as beech and oak.

Late glacial and post-glacial immigration of the tree species

In the glacial stages of the Pleistocene epoch, an open peri-glacial dwarf shrub- and grassland-tundra prevailed in Central Europe. In addition to species of present-day Nordic tundra [e.g. Betula nana L. (Betulaceae) and Dryas octopetala L. (Rosaceae)], species of

mountain crests. Additionally, the amount of rainfall is in� uenced by the windward or lee-ward position (Fig. 3). This contrast is particu-larly pronounced when comparing the rain-catching Harz Mountains and the adjacent leeward central German dry region.

Altogether, the sub-oceanic to sub-conti-nental transition in climate in Central Europe can be described as follows: the temperature extremes are not pronounced (summers are rarely more than 30°C and winters rarely are below -20°C). Expanded springs and au-tumns extend the vegetation period. Precipi-tation can occur throughout the year. Longer hot or cold periods only arise exceptionally

Fig. 3: Climatic diagrams from low mountain ranges in Central Europe. Data base: Schirmer (1969), Bay-FORKLIM (1996) and www.klimadiagramme.de.

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The post-glacial (Holocene) period began with a distinct warming. In the pre-boreal pe-riod (10300-8800 years ago) birch and pine expanded their ranges again. In the follow-ing boreal climatic epoch (8800-7500 years ago), there was a major expansion of Cory-lus L. (Betulaceae). In birch-dominated land-scapes, pines spread out considerably for a short time. At the same time, species of mixed oak forests [Ulmus L. (Ulmaceae), Quercus L. (Fagaceae), Acer L. (Aceraceae), Tilia L. (Tiliaceae), Fraxinus excelsior L. (Oleaceae)] expanded their ranges and rapidly replaced the light-demanding pioneers. In the south-eastern low mountain ranges, Picea abies A. Dietr. (Pinaceae) appeared for the � rst time. During the Atlantic period (7500-4500 years ago), species of mixed oak forests became dominant. In the valleys of the northern low-lands and in other smaller areas, a mass expansion of Alnus Miller (Betulaceae) oc-curred. Pinus was the only genus to persist as a domi nant taxon in more continental areas of the eastern lowlands. Picea covered the higher elevations of the eastern low mountain ranges to the Harz Mountains. In the second half of this period (from 6200 years ago) Fagus L. [principally F. sylvatica L. (Fagaceae)] and Abies alba Miller (Pinaceae) began to spread out in the southern low mountain ranges of the area. In loess regions, the � rst Neolithic farmers interferred with the natural succes-sion (as shown by palynological evidence). The sub-boreal period (4500-2800 years ago) was characterised by a signi� cant cooling and increasing precipitation. The species of mixed oak forests were displaced gradually by the progressive propagation of beech and � r as well as of Carpinus betulus L. (Corylaceae) and Picea in the north-east. The propagation of beech may have been accelerated by the anthropogenic opening up of the mixed oak forests in the Bronze Age (e.g. Küster 1996, 1998a, Pott 1997a, 2000).

In the older sub-Atlantic epoch (2800-1000 years ago) the competition balance shifted in favour of the beech (the so-called beech period). This was caused by cool, humid, sub-oceanic climatic conditions, similar to the present-day climate, which permitted beech-es to colonise lowland and upland forests

the dry steppes such as Artemisia spp. (Aster-aceae) and Helianthemum spp. (Cistaceae), as well as species of Chenopodiaceae, most of them common in eastern Europe (Firbas 1949/52, Lang 1994) are found. Only at its northern margin the Central European area was covered by nordic inland ice. The last glacial period is therefore called the Weichse-lian glaciation. Further to the south, the Alps were covered by a massive ice sheet, extend-ing southwards to the northern Alpine foothills (with its � nal period being the Würm glacia-tion). Around some peaks of the low mountain ranges (e.g. Black Forest, Bavarian Forest, Krkonošé), there were local glaciations (Er-genzinger 1967, Liedtke 1975, Walter 1992). With the gradual warming in the last glacia-tion (up to 10000 years ago) the ice sheet slowly melted. However, the post-glacial re-forestation proceeding from south to north did not occur immediately. Theories about post-glacial migration differ signi� cantly: trees and shrubs spread from their Pleistocene glacial refuges in south-eastern and south-western Europe via different routes and with different speed, so that they reached dominance suc-cessively or at the same time. The character-istic succession of the vegetation, depending on the climatic changes has been de� ned as the so-called Mitteleuropäische Grundfolge (Central European basic sequence) (Firbas 1949/52).

The � rst tree species, i.e. those resistant to frost, but also light demanding pioneer spe-cies such as those of Betula L. and Pinus L. (Pinaceae), along with Salix L. and Populus tremula L. (both Salicaceae) colonised the steppe areas. Forest succession started in the north-west and the west with high percentages of Betula, whilst in the south-west, south-east and the east Pinus dominated. During the so-called Alleröd interstadial (11900-10800 years ago) the greatest forest density and cover of the last glaciation was reached. At the end of this period a new relocation of the forests took place. It was caused by a climatic setback. This � nal glaciation (10800-10300 years ago) was characterised by an open landscape with tundras poor in trees (birch and pine) and ar-eas devoid of forest (low mountain ranges).


northward from one inter-glacial period to the next.

(4) The west-east oriented Alps represent-ed migration barriers both for the retreat into the glacial refuges, and for the inter-glacial expansion. The Mediterranean can be inter-preted as an additional barrier.

(5) In the glacial refuges, the climatic con-ditions were so extremely unfavourable that none of the forest communities occurring in Central Europe could survive there (Beug 1977). These facts can be summarised as fol-lows:

(1) Since arboreal taxa became extinct north of the Alps, e.g. during the Würm gla-ciation, several thousand years elapsed until plant and animal species re-immigrated from refuges in regions south of the Alps. This was determined by climatic and biotic conditions but also by historical events.

(2) Glacial refuges are thought to have contained a large share of the intra-speci� c biodiversity of the temperate biota (e.g. hap-lotype diversity in Fagus has a strong latitudi-nal gradient).

(3) Modern forest communities started to develop in the Holocene during re-immigra-tion of tree species. Beech forests were as-sessed to be rather young in comparison with other Central European forest communities, since Fagus re-immigrated as one of the last tree species. In the north of Central Europe, they are not older than approximately 3000 to 4000 years, which is not more than 30 to 60 tree generations (Pott 1997a, 2000).

(4) Less than 40 tree species (30 decidu-ous and 7 coniferous) occur in extant forest communities of Central Europe. 90% of the forest communities investigated were domi-nated by one or two principal tree species only (Fig. 4).

(5) Some relict communities which re-semble tree species combinations of earlier Holocene periods

- The Betula pubescens Ehrh./Sorbus au-cuparia L. (Rosaceae) community, colonising nutrient-poor boulder beds with considerable accumulation of humus, represents an exam-ple from the early Holocene, similar to some relict pine forests (Ellenberg 1996, Rüther 2003).

(demontane spreading), in part accompanied by � r and, in higher altitudes, by spruce. The north-western lowlands were still dominated by oak forests but, in the north-eastern low-lands, Carpinus was evident. The more re-cent sub-Atlantic period (starting from 1000 years ago) was shaped by a growing anthro-pogenic impact. The cultural epoch of the Iron Age was characterised by increasing agricul-tural land-use together with decreasing forest cover and an economical promotion of certain timber species. Initially, pioneer species such as birch and pine and then species that could be coppiced, such as hazel, elm, hornbeam and oak, pro� ted from man-made clearings and exploitation of forests. Finally, the signi� -cance of broad-leaved trees decreased, whilst spruce and pine gained in importance. Initially this transition occured more or less unwitting-ly by man. Later, this became active, caused by the conversion of coppice (with standards) systems to high forest systems, which were harvested by clear-cutting.

Potentially natural forest vegetation of today

Tree species and forest communities

The Central European � ora of the Quater-nary was characterised by a massive spe-cies depletion, in particular with respect to the arboreal taxa (the Pleistocene extinc-tion). The pre-Pleistocene dendro� ora, once rich in species, decreased to approximately 50 genera (Walter and Straka 1970, Ellen-berg 1996). Several factors may have con-tributed to the extent of these extinctions (Lang 1992).

(1) At least � ve prolonged cold glacial peri-ods alternated with shorter warm inter-glacial periods.

(2) Climate changes between glacial and inter-glacial periods occurred very rapidly.

(3) The glacial periods became progres-sively colder. None of today’s dominating broad-leaved trees could have survived the � nal Würm glaciation north of the Alps. Gradually, re-immigration distances became greater, so that the species migrated less far


- oxygen on waterlogged soils;- nutrients on acid podsols;- iron and phosphorus on calcareous sites

and- stability on unconsolidated landslides.Many Central European tree species were

able to occupy similar habitats, i.e. their phys-iological amplitude overlapped within a wide range (Ellenberg 1996). The ecological ampli-tude refers only to those habitats where a spe-cies could grow and reproduce successfully in competition with others. In Central Europe, Fagus is a particularly competitive and shade-tolerant tree species, driving other species out. Its ecological amplitude is nearly identical to its physiological behaviour (Fig. 5).

The competitiveness of beech had a great effect on the number of tree species in Cen-tral European forests. Fig. 6 shows that beech dominated upland forests both on acidic and on calcareous sites. Other tree species occur-ring on nutrient-rich beech forest habitats are classi� ed as associated trees (see Hordely-mo-Fagetum in Fig. 4), which dominated only temporarily in certain dynamic phases or at the ecological margins of a forest community. The in� uence of beech in reducing species diversity is illustrated in Fig. 6. A high diversity of tree species can only be expected in nu-tritious habitats outside natural beech domi-nance. In particular, these characteristics ap-plied to:

(1) lowland riparian forests (Ulmenion mi-noris; Fig. 4, no. 35) and ravine forests (Tilio Acerion: Fig. 4, nos. 16 and 22) on sites with high dynamics which provided a large pool of temporary niches and spatial micro-habitats and

(2) warm-dry, isolated oak/shrub forests (Quercion pubescentis-petraeae; Fig. 4, no. 25) and oak/hornbeam forest habitats (Carpinion; Fig. 4, no. 23).

- Tilio-Acerion communities are examples for vegetation pictures during the oak/mixed woodland time (Atlantic period) (Müller in Oberdorfer 1992) on unstable and nutrient-rich skeletal soils.

Tree species composition related to site fac-tors and competition

The rate of plant growth is limited by the environmental resource which is in least sup-ply. On a regional or greater scale, the range of a tree species is mainly determined by the climate. For example, temperature may be a limiting factor in locations of higher altitude. According the Map of the Natural Vegeta-tion of Europe (Bohn 2000/2003), beech is common from planar-colline up to sub-alpine levels, with an emphasis on sub-montane to montane regions. In the montane to high montane levels, beech is accompanied by � r and spruce. However, a climatically-caused drought limit could not be identi� ed for beech in the area. Even in the central German dry region east of the Harz Mountains, beech forests prosper at an annual precipitation of 500mm or less (i.e. large areas in the forest of Ziegelroda between Querfurt and Artern; Leuschner 1998). In sub-continental lowlands of eastern Central Europe with warm sum-mers, the vegetation consists of oak/horn-beam forests. On over-exploited, poor and dry sandy soils in this region, pine prevails. In the lowlands, particularly in the west, aci-dophilous oak/mixed forests are widespread, but their degree of naturalness is question-able due to their former utilisation in coppice and incomplete post-glacial re-immigration of beech. In higher low mountain ranges (e.g. Harz Mountains, Black Forest, Bavarian For-est, Krušné hory, Sudetic and Carpathian Mountain ranges) coniferous forests with spruce and � r form the upper forest belt. With continental climate increasing, the proportion of spruce increases, whilst the west (e.g. the Black Forest) is dominated by � r.

The occurrence of a species on local scale is determined by the substrate characteristics. Examples for limited resources are:

- moisture on dry and shallow rendzina soils;


high-altitudes from lowlands up to mountains

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1 Alnetum viridis 2 Rhododendro-Pinetum mughisubalpine 3 Vaccinio-Pinetum cembrae

4 Adenostylo glabrae- / Homogyne-Piceetum5 Calamagrostio villosae-Piceetum6 Aceri-Fagetum7 Ulmo glabrae-Aceretum pseudoplatani8 Sorbo ariae-Aceretum

9 Luzulo luzuloidis-Fagetum, montanemontane 10 Galio-Fagetum, montan

11 Galio rotundifolii-Abietetum 12 Aposerido-Fagetum

13 Seslerio-Fagetum14 Calamagrostio variae-Pinetum15 Fraxino-Aceretum16 Adoxo-Aceretum17 Vaccinio vitis-idaeae-Abietetum

submontane 18 Hordelymo-Fagetum19 Luzulo luzuloidis-Fagetum, colline-submontane

20 Galio-Fagetum, colline-submontane21 Carici-Fagetum22 Aceri platanoidis-Tilietum platyphylli23 Galio sylvatici-Carpinetum24 Stellario holosteae-Carpinetum

colline 25 Potentillo albae- und Cytiso-Quercetum 26 Calamagrostio arundinaceae-Quercetum

27 Luzulo luzuloidis-Quercetum28 Leucobryo-Pinetum

planar 29 baltic Melica uni� ora-beech forest 30 Deschampsia � exuosa-beech forest

31 Betulo-Quercetumriparian 32 Carici remotae-Fraxinetumforests 33 Pruno padis-Fraxinetum

34 Alnetum incanae35 Querco roboris-Ulmetum minoris36 Salicetum albae

alder carr 37 Carici elongatae-Alnetum glutinosae38 Vaccinio uliginosi-Betuletum pubescentis

peatland 39 Vaccinio uliginosi-Pinetum sylvestrisforests 40 Vaccinio-Pinetum rotundatae

41 Bazzanio trilobatae-Piceetum

categories: principal tree species (pt) (predominant in the upper canopy layer) obligate abundant

Fig. 4: Phytocoenological behaviour and signi� cance of tree species in forests of Central Europe.


lowlands and uplands pioneer character riparian forests

Tilia

cord

ata

Carp

inus b

etulu

s

Quer

cus r

obur

Quer

cus p

etra

ea

Acer

cam

pestr

e

Prun

us a

vium

Tilia

platyp

hyllo

s

Acer

plat

anoid

es

Sorb

us to

rmina

lis

Betu

la pe

ndula

Popu

lus tr

emula

Sorb

us a

ria

Pinu

s sylv

estri

s

Malu

s, Py

rus

Betu

la pu

besc

ens a

gg.

Prun

us p

adus

Ulm

us ca

rpini

folia

Ulm

us la

evis

Alnu

s inc

ana

Popu

lus n

igra

Salix

alba

Salix

div.

spec

.

15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 pt st ct pt n1 2 2 - 51 1 5 - 72 1 2 - 51 - 6 3 101 1 4 - 62 1 3 2 81 3 3 1 82 1 4 - 71 2 2 2 71 2 4 - 72 1 6 1 101 2 4 1 81 1 7 - 91 - 7 - 84 1 5 - 102 1 11 2 162 1 5 2 101 - 14 2 171 1 5 3 101 1 11 2 151 - 11 1 133 1 11 - 153 3 6 3 152 2 7 3 142 1 11 142 3 3 1 91 2 5 3 111 - 6 1 81 1 7 2 111 1 3 5 101 1 4 1 72 4 6 4 162 2 8 4 161 2 9 - 123 3 10 1 171 1 5 - 71 2 3 2 81 1 1 2 51 2 4 - 71 3 1 - 51 - 6 1 8

secondary tree species (st) companions (st) � pioneers (pt) (subdominant in the (no distinct pioneer (intial phases of succession upper canopy layer) character) after disturbance) obligate abundant accessory abundant accessory abundant


cies typical for the region and adapted to the site (such as Quercus, Fraxinus or Acer) may react rapidly and become established as ser-al species. In mixed montane forests, beech has a shorter average life expectancy (200-300 years) in comparison with spruce (300-400 years) and � r (400-500 years). Hence the � r becomes dominant particularly in late development phases, i.e. late over-mature phase and regeneration phase.

The opening up in the over-mature phase is crucial (e.g. in terms of gap- and patch-dynamics). Leibundgut (1982) proved that in beech-dominated mixed montane forests, most collapse and rejuvenation took place on small-scale by loss of individual old trees. Usually, on these sites the next beech gen-eration followed directly, either evenly spread or in clusters. It may be concluded that only in larger gaps or patches, rapidly-growing species, such as Fraxinus excelsior L. and Acer pseudoplatanus L., reached a tempo-rary dominance, which contributed to an in-creased species diversity. Comparable tree

Tree species composition related to time

During a development cycle of up to 600 years, a virgin forest passes through different phases (cyclic changes, Fig. 7). These include growing, ageing and dying of tree individuals and show completely different structures. The succession of a complex virgin forest system is undetermined and subject to random � uc-tuation (factors from outside the forest which affect growth, e.g. insect attacks, wind and snow damage and others).

Most different micro-climates, light regimes and niches provide a temporally-graduated and species-rich rejuvenation. In particular, temporal heterogeneity in virgin forests caus-es large structural variety and creates spatial patterns of different dimensions and ecologi-cal niches. Due to random � uctuation, some light-demanding and semi-tolerant tree spe-

Fig. 5: Ecogram of tree species which form forests at sub-montane levels of Central Europe (Leu-schner 1998). The shaded part marks site condi-tions where Fagus can reach dominance without human interference. In transition to thermophilous oak/mixed woodlands and sub-continental Quer-cus/Carpinus forests (range a), dominance of Fagus is adapted to closed stands (a favourable micro-climate) and intact humus cover. In nutrient-poor and strongly acidic habitats (range b) Fagus can predominate if intact humus cover guarantees a comparatively favourable nutrient supply. Moist soil locations (range c) can favour Fagus, partic-ularly on sandy substrates and under an oceanic climate (lower drought endangerment of the upper soil), contrasting Quercus and Carpinus.

Fig. 6: Approximate tree species numbers in the tree layer of important alliances and sub-alliances of Central European forest communities (Leusch ner 1998). The numbers apply to well-developed (i.e. comparatively species-rich) stands and are based, mainly, on relevés in Oberdorfer (1992). In these tables, only those tree species which achieved an abundance of at least 25% constancy in the rele-vant associations were considered.


between 4th. and 6th. centuries anno domini, A.D.), forest percentage increased distinctly (Andersen and Berglund 1994). Similarly, in the late Middle Ages (with the occurrence of agrarian crises and the plague), as well as in the Thirty Years War (1618-1648), numer-ous forest areas could regenerate (Hasel and Schwartz 2002).

In the lowlands, in particular in the loess regions of Central Europe, in early prehistori-cal periods (e.g. Mesolithic), settlements and their areas of arable land were used for some decades only. Primitive land use, mainly pro-ducing annual crops, effected pronounced dynamics (secondary successions). Farm-ing practices depended on traditional know-ledge. Natural forest vegetation was cleared by burning without removing the roots. Land use was rarely permanent, since it was aban-doned for recuperation when yields declined markedly (e.g. Kossack 1982, 1995, Lüning et al. 1997). Under this in� uence of shifting cultivation, the proportion of the forest area remained relatively high. From the Neolithic period, human settlement behaviour changed. Instead of a nomadic existence, peoples be-came increasingly settled and established permanent dwellings. These profound altera-tions instigated by the beginning of agricul-ture are called the Neolithic Revolution (e.g. Küster 1998a, 1998b).

In the Bronze and Iron Ages (2200-750 B.C. and 750-15 B.C., respectively) and early Antiquity, human impact on soil and vegeta-tion became greater with increasing popula-tion, with need for fodder (due to the unfa-vourable winter temperatures) and with new cultivation techniques on arable land (Lang 1994). Initially, human disturbance concen-trated, almost exclusively, on lowlands and uplands that were easy to cultivate.

From the early Middle Ages, unaffected forest regions of the low mountain ranges, such as the Bavarian Forest, became more heavily settled and cultivated (Stalling 1987, Nelle 2002, Rüther and Nelle 2006). Due to adverse climatic conditions unfavourable for sustained agriculture in many places, the set-tlements were later abandoned (Fehn 1963) and forest cover remained comparatively high.

species successions were also observed in the former woodland pastures of north-wes-tern Germany (Koop 1982).

Anthropogenic changes: transition of the forest vegetation

Duration and intensity as well as effects of anthropogenic in� uence vary within differ-ent growth ranges of Central European for-ests. The comparison of palyonogical data indicates temporal and regional differences between coastal, lowland, upland, and low mountain regions (Pott 1997b). The degree of landscape changes were determined by not only (i) the varying intensity of human activi-ties on a regional scale but also (ii) by the lo-cal site conditions or (iii) the regenerative abil-ity of the habitats (see the overview in Pott 1988). As a result, an intensely structured landscape with a great number of vegetation types developed during the period of exten-sive management (Burrichter 1977).

Forest area

Under anthropogenic in� uence, the ex-pansion of the forest area was subjected to permanent change. Most spatial alterations occurred in times of upheaval or major crises. Palynological data show that, for example, during the Migration of Nations (its culmination

Fig. 7: Cyclic changes in mixed mountain forests, il-lustrated by the example of virgin forest relics of the lower Austrian limestone Alps (according to Zukrigl et al. 1963).


Wood pasture is an ancient type of multi-functional use. The resulting structure was a mixture of woodland and grassland that developed through a long history of grazing by domestic herbivores under gnarled trees (often pollarded, see below). Wood pastures were widespread in lowland and upland land-scapes through the Middle Ages and until the 19th century. The resulting timber was small in diameter and was harvested for fuel, con-struction of wooden items and possibly char-coal. The effects of wood pasture were:

(1) opening up of the forest landscape;(2) encouragement of the thicket stage of

trees and ground layer species;(3) promotion of tree species with vigorous

sprouting and regeneration capacity (e.g. Carpinus, Fraxinus, Corylus and Ulmus).

(4) selection of woody species which bore spines or thorns [e.g. Juniperus L. (Cu-pressaceae), Rosa L., Prunus L. and Crategus L. (all Rosaceae)] or toxic for the cattle (e.g. Alnus).

(5) suppression of shade tolerant, but less regenerative climax trees like beech.

Similar species selections were caused by pollarding techniques. Pruning was performed for winter feeding and bedding (e.g. Pott 1983, Haas and Rasmussen 1993). Many of the pol-larded trees are now ancient. Some of these relicts of both wood-pasture and pollarding remain in parts of Central Europe (e.g. Neu-enburger Urwald, Nitzschke 1932, see also the overview for north-western Germany in Pott and Hüppe 1991). The complex structure of the gnarled old trees provided a number of micro-habitats, such as bark crevices and tree holes, which are not found in high for-est and this made them suitable for species of fungi, invertebrates and bats. The effects of the wood-pasture on the growth habit and the phenotype of beech were shown for the Bavarian Forest and the Black Forest (Sey-fert 1975, Schwabe and Kratochwil 1987, respectively).Today, the ground vegetation of such in� uenced forest stands is dominated by Agrostis L. and/or Holcus lanatus L., with Fes-tuca rubra L., Deschampsia � exuosa (L.) Trin. (all Poaceae) and herbs, such as Potentilla erecta (L.) Raeusch. (Rosaceae) and Galium

In the late Middle Ages and in the begin-ning of modernity, reduction of the forest area reached its greatest extent. Decisive factors were industry and mining, in particular the iron and glass industries, as well as salt mines, which required much wood or wood products (i.e. charcoal and potash). Major substantial disturbances were found in partial regions of the Harz Mountains (Hillebrecht 1982, Bartels 1996), the Siegerland (Pott and Speier 1993), the Black Forest (Brückner 1981, Golden-berg 1996), the Upper Palatinate (Lutz 1941, Vangerow 1987) as well as the Bavarian Al-pine Uplands (Bülow 1962, Knott 1988, Zi-erhut 2003). At the same time, reforestation measures took place in many deserted areas; for example, these have been documented for the Nuremberg Reichswald in 1368 (Man-tel 1968). In the course of modern forestry, at the end of 18th century, reforestation had become mandatory. Since this period, the for-est area in different regions of Central Europe has increased again.

Today, approximately 30% of Central Eu-rope are covered by forest (Mayer 1984). The largest connected forest areas can be found in the central mountain regions (e.g. the Pfälzer Wald, the Odenwald-Spessart-Rhön hills and the Bavarian Forest). However, the forest ar-eas are reduced greatly and have become isolated and fragmentated, particularly in the loess-covered lowland and upland regions (e.g. Franconian plateau, northern Alpine foothills), where cultivation was easy. This also applied to offshore areas [e.g. Schles-wig-Holstein (northern Germany), Denmark and Skåne (southern Sweden)].

Tree species composition

Due to human activity and usage, the tree species composition of the original forests was changed markedly. Until the Middle Ages these changes occurred more or less unwit-tingly by man. Commencing in the late Middle Ages and becoming more widespread in the modern era, organised forestry was under-taken in western Europe (Köstler 1956) with the intention of promoting certain tree species (e.g. 1529: Waldpuech für das Reichenhalle-sche Waldwesen).


� rst plantings of Pinus were documented in the14th century for the Nuremberg Reichswald (Mantel 1968). This introduced a gradual change from broad-leaved to conifer-domi-nated forests (Fig. 8). As a pioneer with few demands, Pinus established itself spontane-ously on strongly degraded soils (e.g. in the Upper Palatinate, Lutz 1941, Diepold 1945). Pinus bene� tted from anthropogenic exploi-tation and deforestation of the landscape, as did Picea in the low mountain regions. Both genera expanded outside of their native ar-eas (e.g. Black Forest, Harz Mountains, lower Bavarian Forest; Brückner 1970, Schubart 1978, Rüther 2003, respectively).

Finally, from the middle of the 18th and until the beginning of 19th century, organised for-estry promoted high forest systems (Hartig 1791, Cotta 1817). Silviculture (as a theory and practice of controlling forest establish-ment, composition and growth) was devel-oped as discipline for the manipulation of for-ests in order to enhance certain forest values or products. Due to previous forest devasta-tion and exploitation of the natural resources, a lack of timber, perhaps inspired further by the idea of a rapid economic pro� t (theory of � nancial rotation), resulted in the planting of age-class forests, dominated by Picea, Pinus and Pseudotsuga menziesii (Mirb.) Franco (Pinaceae) (introduced from Paci� c North America). Table 1 shows that the relationship between broad-leaved trees and conifers has become reversed in comparison with the original situation. An additional and signi� cant factor was that considerable areas of native broad-leaved forests were cleared for farm-ing. The proportion of Picea increased notably, whilst the percentage of Abies declined over the last two hundred years. A well-known ex-ample of the decline of Abies is from the Fran-conian Forest, where, in the Middle Ages, a � r-promoting selection system was operated. It was based on the export of raft-wood along the rivers Main and Rhine to the Netherlands (Fig. 9). Selective timber production resulted in a change of tree species combination: prior to the 16th Century, we estimate a ratio of 40% of broad-leaved trees and 60% of Abies. An initial consequence was the elimination, by creation of forest pasture, of rare high-quality

saxatile L. (Rubiaceae). It contained some woodland edge species, such as Teucrium scorodonia L. (Lamiaceae), but few typical woodland species.

Coppicing is the oldest form of forestry; woven hazel screens used for � shing have been dated back to 5000 B.C. (in the Neolith-ic period). Much of the ancient semi-natural woodland in Central Europe developed un-der the coppice management system. Many broad-leaved trees, including Corylus, can be cut down to the stumps, which re-grow producing multiple stems called poles. Usu-ally, the poles were harvested every 15 to 25 years as fuel, the poles of hazel were har-vested approximately every eight years and converted into a wide range of products. Tra-ditionally, willow [Salix L. (Salicaceae)] poles were used in hurdles. In addition to Corylus and Salix, Carpinus, Tilia, Acer, Fraxinus and Alnus were promoted. Quercus, Ulmus and Populus (Salicaceae) are also adapted to coppicing, but have less regeneration poten-tial. Like wood pasturing, pollarding coppicing suppressed not only Fagus, but also conifers such as Abies, Picea and Pinus. Requiring longer rotation periods, beech-dominated coppices were limited to some regions with milder winters, such as the margins of the low mountain ranges in north-western Germany (e.g. Pott 1981) .

Coppicing with standards, a variation of traditional coppicing, was established in the Middle Ages. Individual trees were excluded from short-term activities. Quercus, in particu-lar, was promoted, since it served in pig-keep-ing (e.g. Hesmer and Schröder 1963, Schöller 2001). This form of management was particu-larly common in densely-settled loess regions of Central Europe. This sometimes led to shortages of wood (Ellenberg 1996).

This preservation of standards and the oak mast for pig-keeping promoted a con-scious tree species for the � rst time. Also in low mountain regions, human activities in-� uenced the tree species composition of the forests to a greater extent [e.g. plenter forests (i.e. individual tree selection, group selection system) promoting Abies directly, Wirth 1956, Rüther 2005)]. At the same time, reforesta-tions of many deserted sites took place. The


Fig. 8: Forest development in Central Europe. All illustrations simpli� ed from Walter (1986). Top: earlier sub-Atlantic epoch from 2800-1000 years ago). Center: prevailing tree species of forests in the Middle Ages. Bot-tom: prevailing tree species around 1900 A.D..


withered before the roots had reached min-eral soil. In 1830, productive forests were formed exclusively by Abies and Picea, but, after 1850 the former was � nally displaced by the latter. Reforestations often succeeded with Pinus and Picea only, and the decline of Abies was accelerated by large-scale open-ings caused by undifferentiated management methods. Another important reason for the dramatic decline of Abies was intensive man-agement of game. Comparable simultaneous decreases and increases of Abies and Picea, respectively, are known for example from south-western Moravia (Malek 1980) and eastern Bavaria (Eichenseer 1998).

Fig. 10 compares natural and current stocking in relation to the present total forest area in Bavaria. Due to centuries of human settlement and exploitation, light-demanding and partially shade-tolerant tree species were favoured (Pinus; in former times Quercus and later Picea), whereas shade-tolerant cli-max trees [Fagus, Abies and Taxus L. (Tax-aceae)] were replaced. The proportion of Fa-gus amounted to 12% and Abies 2%, which, in both cases, is approximately 20% of the natural cover. Conversely, Picea and Pinus have bene� ted from their utilisation with the cover of the Picea, which is natively limited to mountains, bogs, boulder beds and podsols, having quintupled.

Contemporary forestry faces great chal-lenges, similar to those of 250 years ago. Conversion of unstable non-natural conifer forests (e.g. Olsthoorn et al. 1999, Klimo et al. 2000, Zerbe 2002), increasing demands for its social function, climatic change/global warming, intensi� ed wind damage and insect pests have been accompanied by a glut in the timber trade and rapid staff reduction within in forest administration. Nevertheless, sustaina-

hardwood. This was followed by disorganised use, including selective removal of timber (e.g. material for joinery). Fagus, unwieldy in growth habit and incapable of � otation, was used less. In 1665 the share of broad-leaved trees (exclusively Fagus) was not more than 10%, whilst Abies and Picea covered 80% and 10%, respectively (Wirth 1956).

Exploitation and the breaking up of the canopy led to an increasing cover of Picea. Within the raw humus cover and dwarf shrub blankets with intensively-matted roots, Fagus could either not germinate or its seedlings

Tab. 1: Comparison of original (i.e. early sub-Atlantic) and current tree species composition in forested regions of Central Europe (after Mayer 1984).

broad–leaved trees

conifers beech oak others pine spruce � r

original 66 34 38 20 8 20 7 7current 29 71 14 10 5 44 25 2

Fig. 9: Changes of tree species combination in the Franconian forest due to human impact.


gramme operating in Lower Saxony. Accord-ing to national forest laws, site or vegetation classi� cations are mandatory declarations as a basis for all forest projects. An evaluation of the forest resource inventories showed that, in regenerating forests, broad-leaved trees are increasing gradually. During the previous 20 years, the proportion of deciduous trees in national forests in Germany increased by nearly 10% within 8 years. This was caused by the direct promotion of deciduous trees. Other key factors are:

(1) reduced game densities;(2) rapidly regenerating humus forms due

to atmospheric nitrogen immissions up to 30 kg/ha/a and

(3) global warming, as it supports decidu-ous trees of temperate zones, with conifers dominating boreal, sub-boreal or montane zones.

However, in some landscapes of Central Europe, Fagus or Quercus disappeared en-tirely or became so rare that rejuvenation on suitable habitats did not occur spontaneously. The lack of nursery trees, selective game damage, unfavourable micro-climate, frag-mentation and isolation of forests or dense/impenetrable ground layer vegetation compli-cated resettlement.

Forest structures

The structure of the stand, in particular the complexity and dynamics of the forest cano-py, is of fundamental importance to the for-est dynamics, e.g. spatial heterogeneity and temporal change in understorey vegetation, patterns in regeneration mosaics, and micro-climatic variation (Norman and Campbell 1989, Song et al. 1997). The structure of the forests was modi� ed by different forms and intensities of utilisation. The wide range of cli-matic, geological and geobotanical conditions in Central Europe produced a differentiated settlement history and speci� c management techniques, resulting in regionally different forest structure types. Fig. 11 shows the pos-sible modi� cations of the natural forest in the hill and mountain country depending on the use. The dynamics between utilisation and abandonment produced numerous transition

ble forest management is increasingly orient-ed towards the natural biological resources of a landscape (by taking advantage of primary production, thus minimising the need for in-tervention).

The following factors require special con-sideration:

(1) ecology (soil conditions, indicator spe-cies groups and vegetation units/natu-ral forest communities with their princi-pal, secondary and pioneer trees) and

(2) forest genetics (sources, genetic vari-ability of tree populations).

Individual stand characteristics (stand his-tory, genetics, current and natural vegetation, succession stage, ecological site character-istics) are analysed for the suitability of tree species. Recommendations are given for dif-ferentiated, economically rational and ecologi-cally sound sylvicultural options. Site-adapted biodiversity and rich structural heterogeneity (chapter “forest structures”, see below) of the stands are considered as indicators for a sus-tainable forest management under the risks of climatic change/global warming (e.g. Ferris and Humphrey 1999). The maintenance or re-establishment of forests in as natural a state as possible is a strategy for sustainability [e.g. as shown by LÖWE (=langfristige ökologische Waldentwicklung, Otto 1989), a forestry pro-

Fig. 10: Natural and current stocking related to the total forest area of Bavaria. The natural stocking was projected approximately over the area of the natural forest communities. The current stocking data is based on the national forest inventory.


can affect substantially the adaptive abilities of tree stands and genetic resources of tree species (Müller-Starck 1996), and which may be occupied by the entire range of regionally typical admixed/secondary tree species, in-cluding rare, sensitive and endemic species (e.g. Otto 1989, Schölch et al. 2003).

Stages of forest development

Each kind of forest utilisation disrupts the development cycle of woodlands, in particu-lar with regard to old-growth and over-mature stages (chapter “tree species composition related to time”, see above). In prehistoric times, the human and animal impacts oc-curred temporarily only and were limited spa-tially (chapter “forest area”, see above). After utilisation, the sites were able to re-grow, so that, apart from site changes, these impacts could be compared with some natural events like forest � re or wind throw.

From the Neolithic period onwards human impact became more intense, due to new

stages between the individual structure types, leading to a considerable structural diversity even on small sites.

There are very little data on the structure of original virgin forests in Central Europe (chapter “tree species composition related to time”, see above). Relicts of virgin forests re-mained small-scale in climatically rough and inaccessible locations within montane and sub-alpine zones. Observations within these locations cannot be applied by inference to lowland forests. The principal differences be-tween commercial forest and virgin forest are the high degree of order (regular structure), the limited heterogeneity, breaks in continu-ity of old-growth-habitats and the quantity and quality of deadwood (Walter 1986).

With reference to stand structure, (i) the size and shape of gaps and patches, (ii) the uneven cutting of individual trees (resulting in small and large groups) and (iii) selective working resemble natural forest conditions. In addition, nature-oriented forestry practice tries to create niches of different size, which

Fig. 11: Possible modi� cations depending on the use of the natural forest (e.g. Luzulo-Fagetum) in hill (left) and mountain country (right). There are numerous transitions between the forest structure types represented (from Walentowski et al. 2004).

Forest structures lowand and upland regions Forest structures mountainous regions


ground layer was modi� ed profoundly. Forest litter utilisation and removal of sods displaced the original forest � oor species and supported the invasion of indicators of soil degradation such as Calluna vulgaris (L.) Hull (Ericace-ae). Particularly large areas of heathland de-veloped on the sandy soils (on old moraine landscapes) in north-western Germany and in Denmark (Ellenberg 1996). Heather also spread in the forests of low mountain ranges such as the upper Bavarian Forest (Rüther 2003).

The forests opened up due to woodland pasturing, so that the stand climate changed. Speci� c shade-tolerant species on the forest � oor disappeared and were replaced by light-demanding herbs and shrubs. Additionally, some grass species, adapted to grazing but unknown from forests, were able to invade. In addition to tree species, grazing animals also enabled the spread of numerous herbs and shrubs, which possessed toxic ingredients or thorns (this is dependent on the natural zone, e.g. Pteridium aquilinum (L.) Kuhn (Denns-taedtiaceae), J. communis L., Ilex aquifolium L. (Aquifoliaceae), Prunus spinosa L., Cra-taegus spp., Rosa spp., (all three Rosaceae), Genista spp. (Fabaceae), Thymus spp. (La-miaceae).

All of (i) the method, frequency and inten-sity of utilisation, (ii) vertical and horizontal structure of the stand, (iii) canopy closure, (iv) mixture of tree species, (v) isolation, fragmen-tation and partition of the forest area, (vi) the history of the forest stand and (vii) the stage of forest development are potentially deter-minant factors of the species composition of the ground layer in today’s commercial for-ests. Man-made forest edges or openings are habitats for some light-demanding species which are absent in natural forests or occur in natural open stages of forest development only. These species immigrate from either ad-jacent agricultural areas or from natural open-ings such as rocks, stone � elds, debris cones and riparian areas. In forests with a young habitat tradition, some relicts of former land use (e.g. dry grasslands, alpine pastures, lit-ter meadows, vineyards and orchards) may be present.

With reference to the composition of the

cultivation systems. Forests and � elds were cultivated permanently and systematically, so that the natural development of forests was affected profoundly. Forest pasture caused different vegetation types, with degeneration and regeneration complexes occurring simul-taneously (Pott 1988). With the decrease of pasturing, regressive reforestation processes occurred. Due to enduring extensive pasture, thorn bushes played a decisive role in forest regeneration, due to their protective function for the natural thicket stage. The intensity of the pasture instigated many dynamic, spa-tially and temporally heterogenous systems, consisting of degeneration and regeneration complexes. However, successful regenera-tion occurred after pasturing had ended.

In some low mountain ranges an alternat-ing forestry and � eld crops system was devel-oped including agriculture, forest pasture and coppicing (e.g. Black Forest: Schwabe-Braun 1980, Bavarian Forest: Lippert 1984, Reif and Oberdorfer 1990, Siegerland: Pott and Speier 1993). Thus, on the same sites, forest de-generation stages continually took place (Fig. 11), whether induced by man or animal. Re-generation was possible during fallow periods only. Anthropogenic in� uence on forest devel-opment is particularly evident in the coppice system: timber was cut in a regular cycle of 15 to 25 years. Comparable natural disturbances and interactions, with frequent environmental changes and similar mechanical demands on and damages to trees, are strictly limited to habitats along untamed rivers and avalanche lanes in high mountains.

Even in modern high forest systems with site-adapted near-natural tree species com-bination, a high degree of naturalness may be inferred only if the factor time is neglected. Within a conventional rotation period of 100 to 140 years (Mayer 1992), the characteris-tic complexity, heterogeneity and biodiversity are reduced drastically (chapters “tree spe-cies composition related to time” and “forest structures”, see above).

Herbaceous layer, brushwood

Due to the different forest utilisation and management types, the composition of the


within the loess-covered uplands are explic-itly favourable for these species. In southern Germany, thus far, only a few authors have made (relatively small-scale) investigations of forest � oor species, comparing ancient and recent forests (e.g. Schneider and Poschlod 1999). Such studies are badly needed, partic-ularly with regard to forest management and forest conservation.

In addition to qualitative changes in spe-cies, quantitative changes have been ob-served. Oxalis acetosella L. (Oxalidaceae) and numerous mosses of the ground layer, such as Eurhynchium angustirete (Broth.) T.J.Kop. (Brachytheciaceae), Thuidium tama-riscinum (Hedw.) Bruch et al. (Thuidiaceae), Hypnum cupressiforme Hedw. (Hypnaceae), Polytrichum formosum Hedw. (Polytrichace-ae) and Hylocomium splendens (Hedw.) Bruch et al. (Hylocomiaceae), have spread out in the man-made Fagetalia-spruce forest-communities. Originally, these species at-tained their ecological optimum in montane, herb-rich mixed coniferous forests on accu-mulated humus. Recently disturbed clear-cut areas may be colonised rapidly by tillering and creeping dwarf shrubs [e.g. Rubus spp.

ground layer, the habitat tradition played a de-cisive role. Many investigations have shown that the species composition of ancient forests (i.e. forests with a long tradition) differs from that of recent forests (see literature surveys in Wulf 1997, Hermy et al. 1999). A summary of numerous local lists of indicator plants of ancient forests revealed that only � ve species [Carex sylvatica Huds. (Cyperaceae), Chrys-osplenium alternifolium L. (Saxifragaceae), Lamiastrum galeobdolon (L.) Ehrend. and Polatschek (Lamiaceae), Melica uni� ora Retz. (Poaceae) and Paris quadrifolia L. (Lil-iaceae)] appear widespread in Europe (Wulf 1997). Most of the indicator plants are char-acterised by a very limited temporal and spa-tial dispersal,thus having a poor ability to col-onise new forest stands. The main causes of this are short-distance dispersal strategy (e.g. autochory, myrmecochory), low production of satellite populations and low competitive abil-ity. Another reason is the modern reduction of dispersal processes (primarily of forest pas-ture) in comparison with the historical man-made landscape (Bonn and Poschlod 1998). In addition, the isolation and fragmentation of current forest stands affects the aggregate ability to colonise (Rackham 1980, Peterken and Game 1984). Furthermore, most of the indicator plants have no persistent seed bank record (Bossuyt and Hermy 2001). Finally, fa-vourable stand climate and humus condition are necessary for regeneration and/or resto-ration.

Most of the low- and upland beech for-est habitats have a tradition of disturbance or perturbation. They regenerated from cop-pice forests or were afforested (Dierschke and Bohn 2004), so that most of them may be termed recent forests. In Bavaria, some plant species typical of beech forests and also indi-cator plants for ancient forests are absent in speci� c landscapes. The large-area overview for Galium odoratum (L.) Scop. (Rubiaceae) shows a patchy occurence in the loess re-gions of Franconia and Upper Bavaria (Fig. 12). This absence can be interpreted only by the aforementioned reasons (principally short-distance dispersal strategy, isolation of forest stands and reduction of dispersal processes), since both climatic and edaphic conditions

Fig. 12: Spatial distribution of Galium odoratum in Bavaria (Schönfelder and Bresinsky 1990). Rea-sons for absence - a: anthropogenic, e: edaphic, c: climatic.


centrated on the humus layer and the upper humus/mineral soil layer (Ellenberg 1996, Gulder 1998).

With the abandonment of forest litter use at the beginning of the 20th century, the regen-eration of the humus layer commenced. This development has been accelerated greatly by air pollution and raw humus amelioration over the last 50 years, but has led to eutrophica-tion, which is extreme in some regions. In ad-dition to large-scale nitrogen deposition, there are many small-scale impacts on the soils. Soils affected by man (e.g. soil compaction, lime substrate from forest path construction, small nitrogenous patches) may be found along forests roads. Additional eutrophication from adjacent farmland occurs along forest borders.

Conclusions and suggested management responses

The present tree species composition of the Central European oak/beech region has been determined largely by historic in� u-ences. Formative processes were the Pleis-tocene extinction and the post-glacial migra-tion of trees in close connection with human impacts. Managing biodiversity of our forest habitats should aim not for the maximum but for locally speci� c species diversity (Granke et al. 2004). To strive for a maximum species di-versity would primarily mean to support a pool of non-speci� c and ubiquitous species, intro-duced by anthropogenic disturbance. Howev-er, due to the dramatic loss of wilderness, the assessment should relate to the original and speci� c composition, structure and dynamics of natural forests. In other words, biodiversity is only an important operational standard of value for nature protection in connection with the degree of naturalness. To measure this degree of naturalness, three different compo-nents have to be considered:

(1) Tradition of old-growth trees (at least 200 years) with biocoenoses of decaying trees (e.g. Gastero cercus depressirostris. Fabricius (Coleoptera: Curculionidae) and Bolitophagus reticu latus. L. (Coleoptera: Ten-ebrionidae) as indicators.

(Rosaceae)] such as blackberry and rasp-berry and grasses [e.g. Calamagrostis spp. (Poaceae)]. Further soil compaction encour-ages the clonal spread of Carex brizoides L. (Cyperaceae), once harvested as an eel grass [Zostera L. (Zosteraceae)] substitute in mattress � llings.

Soils and matter cycles

The numerous forest utilisation and man-agement types have not only affected the vegetation but also the nutrient content of the soils. Due to the enduring nutrient depletion, the forest litter utilisation and the removal of sods led to a lasting soil degeneration, par-ticularly on deep, badly buffered sites with low levels of lime and bases. Since the end of the Middle Ages, in southern Germany, initially in the Upper Palatinate but also in Middle and Lower Franconia, a massive forest litter utilisation was conducted (Rebel 1920, e.g. Nuremberg Reichswald), Ott-Eschke 1946, 1951, Sperber 1968, large woodlands in the Upper Palatinate hill and basin landscapes, Lutz 1941, 1942). In many cases, the ground litter and upper humus-rich soil layers were removed. Forest litter use also damaged or destroyed the thicket stage of the trees, limit-ing the regeneration of the forests. As a con-sequence, both the yield and the vitality de-creased (Rebel 1920).

In terms of sustainability, the use of forest litter did more damage than any other type of forest exploitaiton. Generally, silviculture was able to compensate its effects on the forest vegetation within a reasonable period of time, for example a few decades. However, chang-es in soil composition have a lasting effect over several centuries (Hasel and Schwartz 2002). The acidi� cation and subsequent nutri-ent depletion of soils is a natural and gradual process that was accelerated unwittingly by man, and to a far greater extent than in prim-eval forests (Walter 1984). When evaluating forest vegetation, changes in site conditions in many regions, including degradation of the soil, must be considered. Most of the moss- or lichen-dominated coniferous forests (i.e. of Picea and Pinus) are degraded types of beech forest, where the nutrient cycle con-


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(2) Natural tree species composition (rela-ted to site factors, competition and time) with abundance and dominance of principal, se-condary, companion and pioneer trees of the natural vegetation unit as indicators.

(3) Natural ground layer vegetation (re-lated to typical forest character species) with presence or absence of species of the natural vegetation unit (e.g. G. odoratum), weakness to propagate (forest speci� c short-distance dispersal strategy), shade tolerance, low pro-duction of satellites, temporary seed banks as indicators.

Modern management methods need to consider biodiversity and the degree of natu-ralness by paying attention to heterogeneity, complexity and scale (Walter 1986, Ott et al. 1997, McCoy and Bell 1991): ”Heterogeneity encompasses the variation due to the relative abundance of different structural components, whether both vertically or horizontally. Com-plexity refers to the variation resulting from absolute abundance of individual structural components, and the scale takes account of the variation due to the size of the area or volume used to measure heterogeneity and complexity.” (Ferris and Humphrey 1999).

Acknowledgements

We would like to thank Neil Springate for immensely helpful comments, criticisms and suggestions.

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Canopy Arthropod Research in Europe Basic and applied studies from the high frontier

Edited by Andreas Floren (Univ. Würzburg) & Jürgen Schmidl (Univ. Erlangen-Nuremberg)

Foreword by K.E. Linsenmair, University of Würzburg, Dep. of Animal Ecology and Tropical Biology 2008, softcover, 576 pp., ISBN 978-3-935654-01-2, price 49,90 € Contributors: U. Ammer, R. Asshoff, J. Bail, R. Bolz, H. Bussler, M. Dolek, K. vdDunk, A. Floren, M. Gossner, I. Gierde, A. Gruppe, W. Güthler, H. Hacker, D.V. Hagan, A. Häusler, K. Horstmann, P.J. Horchler, C. Kampichler, S. Keel, C. Körner, A. Liegl, K.E. Linsenmair, R. Market, A. Mitchell, W. Morawetz, J. Müller, H. Nickel, S. Otto, C. Rüther, J. Schmidl, U. Simon, O. Schmidt, B. Seifert, R. Siegwolf, S. Sobek, P. Sprick, A. Stark, H. Stark, R. Szadziewski, H. Walentowski and G. Weigmann

Aims & Scope: In contrast to tropical ecosystems, in temperate zones the importance of canopy ecology is underestimated and underrepresented in science projects. Recent surveys and studies show that also in temperate forest canopies a diverse arthropod fauna exists, containing specialized and endangered species and even species new to science. Species and guild compositions of canopy arthropods in European forests are not yet described sufficiently, and many functional aspects of temperate forests still are not understood or studied. The present volume tries to reduce this gap by summari-zing studies and papers dealing with canopy arthropods in Europe. Aspects of diversity, function, structure and dynamics of canopy arthropod as well as aspects of nature conservation and transmission of scientific results into forestry and management practice are central aims of this book. Contents & Chapters: Foreword � Introduction � General forest ecological aspects � Arthropod diversity, guilds and structure related communities � Stratification and distribution of arthropods in tree habitats � Anthropogenic and natural disturbance structuring arthropod communities � Canopy research and its impact on forestry and nature protection practice. The volume is fully refereed

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Canopy Arthropod Research in Europe Basic and applied studies from the high frontier.

Edited by Andreas Floren and Jürgen Schmidl

“As the global community comes to realise that our climatic future is intimately tied up with the health of our forests so canopy studies take their rightful place in the forefront of forest science. This book will ensure that studies of temperate forest canopies no longer remain the 'poor cousins' of tropical canopy studies. The research described will stimulate new and exciting activities in temperate canopy studies as well as giving the newcomer to the field an invaluable insight into what has gone before.” Roger Kitching, Professor of Ecology, Griffith University, Brisbane

Table of Content Foreword, by the editors Foreword, by Karl Eduard Linsenmair 1 Introduction Canopy arthropod research in Europe, by Andreas Floren and Jürgen Schmidl Canopy research on a worldwide scale: biodiversity, climate change and forest canopies, by Andrew Mitchell, Director Global Canopy Programme 2 General forest ecological aspects Canopy structure and its effect on canopy organisms: a general introduction and some first findings of the Leipzig Canopy Crane Project with special reference to vertical stratification, by Peter J. Horchler and Wilfried Morawetz Microclimatic variability in the canopy of a temperate forest, by Ophir Tal, Martin Freiberg and Wilfried Morawetz Tree species composition and historic changes of the Central European oak/beech region, by Carsten Rüther and Helge Walentowski Tree crowns and forest systems from a forestry point of view, by Hans Stark and Olaf Schmidt 3 Arthropod diversity, guilds, and resource related communities Species of the genus Oedalea Meigen, 1820 (Diptera: Hybotidae): An element of the canopy fauna in European forests?, by Andreas Stark Heteroptera communities in tree crowns of beech, oak and spruce in managed forests: diversity, seasonality, guild structure, and tree specificity, by Martin Goßner Diversity of Neuropterida in mixed forest stands in Germany, by Axel Gruppe The ants of Central European tree canopies (Formicidae) - an underestimated population?, by Bernhard Seifert Tracking the elusive: leafhoppers and planthoppers (Hemiptera) in tree canopies of European deciduous forests, by Herbert Nickel Search in the canopies and you will find new species records of insects, by Karl H. Thunes, Ivar Gjerde, Daniel V. Hagan and Ryszard Szadziewski Species richness and historical relations of arboreal phytophagous beetles - a study based on fogging samples from primeval forests of Poland, Romania and Slovenia (Chrysomeloidea, Curculionoidea), by Peter Sprick and Andreas Floren Species list and feeding guilds of arboreal phytophagous beetles (Chrysomelidae, Curculionoidea) in Germany, by Peter Sprick Abundance and ordinal composition of arboreal arthropod communities of various trees in old primary and managed forests, by Andreas Floren

4 Stratification and distribution of arthropods in tree habitats Tracing arthropod movement in a deciduous forest canopy using stable isotopes, by Roman Asshoff, Sonja G. Keel, Rolf T. W. Siegwolf and Christian Körner Oribatid mites in the canopy of a Central European mixed forest: species richness and species similarity between tree species and habitat types, by Stephanie Sobek, Christian Kampichler and Gerd Weigmann Stratification of 'macro-Lepidoptera' in northern Bavarian forest stands dominated by different tree species, by Hermann Hacker and Jörg Müller Vertical and horizontal distribution of arthropods in temperate forests, by Axel Gruppe, Martin Goßner, Kerstin Engel and Ulrich Simon 5 Anthropogenic and natural disturbance structuring arthropod communities Introduced tree species as an anthropogenic disturbance of arthropod communities in tree crowns of managed forests - a case study of native Heteroptera communities on introduced red oak, by Martin Goßner The diversity of moths communities in different structured oak-hornbeam forests - a comparison of different states of succession in coppice with standard and forests with high standard trees, by Ralf Bolz The impact of flooding and forestry on the species composition of xylobiontic and phytophagous beetles on oak canopies of the Bavarian Danube floodplain, by Johannes Bail and Jürgen Schmidl Ichneumonidae from the canopies of primary and managed oak forests in eastern Poland and southern Germany, by Klaus Horstmann and Andreas Floren Do spider communities in primary forests differ from those in forest-plantations? A canopy study in the Bia�owie�a-Forest (Poland), by Andreas Floren, Stefan Otto and Karl Eduard Linsenmair Diptera (Brachycera) in oak forest canopies - management and stand openness gradient determine diversity and community structure, by Klaus von der Dunk and Jürgen Schmidl Xylobiontic beetle guild composition and diversity driven by forest canopy structure and management, by Jürgen Schmidl and Heinz Bussler 6 Canopy research and its impact on forestry and nature conservation Integrating tree crown science with the development of ‘Near-to-Nature’ forest management practices: examples from Bavaria, by Ulrich Ammer, Martin Goßner, Axel Gruppe and Ulrich Simon Conservation of coppice with standards for canopy arthropods: the Bavarian Conservation Programme for Forests, by Alois Liegl and Matthias Dolek Conservation efforts and strategies for forest canopies in Germany: a review of conservation programmes, by Andreas Häusler, Matthias Dolek, Wolfram Güthler and Renate Market

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