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

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… 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).

Introduction Eemian periods (128000-115000 B.P.), large herbi vores were still present. These periods Naturally, the temperate forest zone of showed a very similar development of for- Central Europe would be a mostly uniform est reestablishment after glaciation in com- forest area (forest cover 95%), dominated parison with the early Holocene (10300 B.P.). by deciduous trees. Only dunes, marshes, Therefore, the stated signi cant impact of the bogs, rocky outcrops and few alpine regions large herbivores, which became extinct at the above the upper timberline were once un- start of the Holocene on the vegetation has to wooded (Ellenberg 1996). The vast progress be rejected (LWF 2000). made in palaeobotany and the amount of Conversely, Central Europe is also a zone data available prove conclusively that Europe of most intensive anthropogenic use. Since was natu rally forested, either with or without at least the second half of the sixth mille- mega-herbivores (Litt 2000). Pollen diagrams nium before Christ (B.C.), forests of natural from many localities across Central Europe origin were transformed progressively by show a clear pattern of post-glaciation refor- human in uence (Lang 1994, Küster 1996, estation, its tree species combinations and 1998a). In north-western Germany as well also the degree of openness of European as in some loess regions of central and forests, both during Eemian inter-glacial pe- southern Germany, the rst farm cultiva- riod and the beginning of the Holocene prior tions were established. The regions’ inhab- to human intervention. Pollen samples show itants farmed elds and bred cattle, pigs, exactly the same result: measured by the sheep and goats. Cattle pastured freely in ratio of non-tree pollen to tree-pollen, the the surrounding forests, a method still prac- European landscape was nearly covered by tised locally in extensively-managed areas woodland which formed forests in the true in the Balkan Peninsula and Carpathians. sense and without resembling a park land- In addition to the production of timber, re- scape. This is true for the entirety of Cen- wood, lumber and leaf litter, the early farm- tral Europe, since none of the known pollen ers also took fruit for their diet, to extract pro les deviate from this picture by showing oil or as a winter supply for their domestic signi cant amounts of non-tree pollen. In both animals. Further forest uses followed, such the Holstein (347000-362000 B.P.) and the as the production of charcoal or ash burning Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 63 and subsidiary uses such as bee keeping or Anthropogenic impact on Central Euro- tan-bark peeling. pean forests has differed both regionally and These forest uses effected a complex temporally. Thus, the original conditions of change of structural and ecological conditions these forests can only be estimated. Never- in the forests, alterations of species abun- theless, some quantitative changes can be dance and species combination as well as detected and qualitative effects assessed. a clear reduction of the forest area. Decisive These include: factors for the degree of anthropogenic trans- (1) diminishment of welfare and protective formation were duration, intensity and nature functions; of the impacts and the resultant ecological (2) alteration of tree species composition site conditions. In extensively-grazed areas, (including direct and indirect in uence non-natural open pastures with individual on the role of indigenous species for groups of bushes as well as park-like stages the vegetation and introduction of for- and groves, alternating with closed forest eign species); stands, were formed (Pott 1983, Ellenberg (3) modi cation of forest structure; 1996). Instead of natural forests, numerous (4) impact on forest dynamics/cyclic phas- substitute communities emerged, so that a di- es; verse mosaic of secondary forests, pastures, (4) modi cation of micro-climate; meadows, heaths and elds is found today. (5) change to substratum conditions, e.g. Fragmentation, soil degradation and isolation nutrient supply; of forests contributed to these effects. (6) alteration of ground cover species Present-day forests in Central Europe (abundance and species combination) have been modi ed completely by man. In and order to obtain an impression of the forests (7) shift of competition equilibrium; devel- before their anthropogenic transformation, opment of secondary vegetation forma- we must visit inaccessible and high-altitude tion and plant communities of varying forest areas. Some relics of virgin forests degrees of naturalness. worth mentioning are Rothwald (Northern In order to understand the radical altera- Limestone Alps, Austria) and Kubany (Krušné tion of the character of forest vegetation, an hory and Šumava, respectively; both Czech investigation of the ecological properties of Republic) (Zukrigl 1978, Leibundgut 1982, the tree species (e.g. geobotanical site fac- Zukrigl 1984). Due to previous and current tors, shade tolerance, coppice sprouting, dis- human impact, forest reserves and forest na- semination, reproduction) is crucial. Abrupt tional parks provide only a reduced picture of spatial changes in species composition of original forests. For numerous forest areas, Central European forests re ect small-scale the label virgin forest (e.g. Neuenburger Ur- patterns in topographical and geological site wald, Nitzschke 1932, see also Rüther and factors, as well as management in uences. Peppler-Lisbach 2007) is misleading. Most of Therefore, it is appropriate to de ne discrete these citations concern rare and irregularly- forest units, i.e. plant associations or vegeta- used (e.g. as forest pasture or for browsing) tion units generated by clustering, rather than or otherwise modi ed natural forests, but by arrange plots along underlying environmental no means original virgin forests. gradients. Well-de ned forest units are es- Available data on forest dynamics, struc- sential for mapping projects which are intend- ture, species composition, species propaga- ed for management strategies. For assess- tion and matter cycles and their interaction ment of the transformation caused by human with soil and vegetation are derived, almost inference to forests with reference to former exclusively, from investigations of secondary vegetation conditions, the post-glacial forest forests. However, these do not permit extra- history is outlined on the basis of palynologi- polation for forests of natural origin, such as cal data. those encountered by the rst Central Euro- pean farmers. 64 Rüther and Walentowski: Tree species composition and historic changes…

Outline of the area Site conditions

The boundaries of Central Europe vary Geology and morphology signi cantly depending on their historical, cultural, political, geological, climatic or geo- In terms of geology and morphology, Cen- botanical de nition. The simple geographical tral Europe may be divided into two principal term concentrates on the central part of the zones (Walter 1992): European continent and, from this, Ellenberg (1) Most of the Central European lowland, (1996) gave a combined climatic-geobotani- situated in the region’s north, is below 50m cal de nition with Central Europe lying be- a.s.l. and attains elevations of 200m a.s.l. in tween 47o and 53oN. Essentially, this covers few places only. Its tallest peak is Wieyca Germany, Poland, the Czech Republic, Slo- (329m a.s.l.), which is part of the Pomeranian vakia, Austria, Switzerland, Luxembourg and young moraine in Poland. Both the geologi- parts of the adjacent states. cal material and the shape of the surface are In our study we follow the boundaries of comparatively young. Vast quantities of sedi- Rubner and and Reinhold (1953) and Mayer ment were deposited during the Pleistocene (1984), which are de ned by natural forest re- ice ages and formed the surface texture. Dur- gions. The Central European oak/beech for- ing the inter-glacial and the Holocene periods, est region covers Germany (excluding East further relocating and modelling processes Frisia and the East Frisian islands), Denmark occurred. The base saturation of the soils dif- (excluding western Jutland), the southern fers depending on the age of the deposit: Old part of Norway as well as the southern point moraine landscapes are strongly decalci ed, of Sweden, the northern part of Poland (in- whereas young ones are rich in bases. The cluding the north-west and south-west re- pre-Quaternary underground is only revealed gions and the Silesian basin), the Czech in few places (e.g. shelly limestone occur- Republic (excluding the territory east of the rences at Rüdersdorf, Berlin). River Morava), as well as parts of non-Alpine (2) The low mountain range with heights Austria and Switzerland. Its western bound- between 200 and 1000m a.s.l. follows south- ary approximates the frontiers of Germany, wards. Individual summits [e.g. Feldberg France and the Benelux countries (Fig. 1). (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 Fig. 1: Natural forest regions of Europe (according sediments and with a surface texture with the to Rubner and Reinhold 1953, Mayer 1984). regional character of a hill country. Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 65

Climate as well as the windward and leeward situation (Fig. 3). Increasing altitude above sea level is Climatically, Central Europe is affected by linked to a thermal and hygric gradient. For both atlantic and continental climates. In the the Bavarian Forest, Baumgartner (1970) in- west, the atlantic character predominates with dicated a decrease of 0.5°C per 100m differ- a balanced cool-humid climate, without tem- ence in altitude. This gradient is overlain by perature extremes and rainfall in all seasons a north-south decline. From north to south, (oceanic conditions). Eastwards, the conti- the altitude belts shift upwards because an- nental character increases with warmer sum- nual average temperatures on the same al- mers and colder winters; annual precipitation titude increase and the vegetation period is with brief and heavy rainfall concentrates on prolonged. Therefore, the upland parts of the the summer months (continental conditions). Black Forest with high annual temperatures From north to south, annual precipitation in- have milder winters in comparison with the creases notably towards the Alps. The rain Harz Mountains. However, with increasing catcher effect is pronounced in summer. It is continental conditions (i.e. eastwards) the caused by the humid Atlantic airstreams form vegetation levels are shifted vertically, due, the north-west meeting on the mountains primarily, to lower occurence of cloud cover (Fig. 2). and more intensive thermal radiation. In the low mountain ranges, the macro- With increasing altitude, precipitation in- climate is modi ed notably by the elevation creases and reaches a maximum on the

west middle east north middle south

Fig. 2: Climatic diagrams from the planar-colline to sub-montane level of Central Europe. Data base: www. klimadiagramme.de. 66 Rüther and Walentowski: Tree species composition and historic changes… mountain crests. Additionally, the amount of (e.g. the dry and hot summer 2003). Temper- rainfall is in uenced by the windward or lee- ate, cool-humid climate favours tree growth, ward position (Fig. 3). This contrast is particu- in particular summer-green hardwoods such larly pronounced when comparing the rain- as beech and oak. catching Harz Mountains and the adjacent leeward central German dry region. Altogether, the sub-oceanic to sub-conti- Late glacial and post-glacial immigration nental transition in climate in Central Europe of the tree species can be described as follows: the temperature extremes are not pronounced (summers are In the glacial stages of the Pleistocene rarely more than 30°C and winters rarely are epoch, an open peri-glacial dwarf shrub- and below -20°C). Expanded springs and au- grassland-tundra prevailed in Central Europe. tumns extend the vegetation period. Precipi- In addition to species of present-day Nordic tation can occur throughout the year. Longer tundra [e.g. Betula nana L. (Betulaceae) and hot or cold periods only arise exceptionally Dryas octopetala L. (Rosaceae)], species of

Harz Mountains Black Forest Bavarian Forest planar-colline luff sub-mont.-mont. high-montane planar-mont. lee

Fig. 3: Climatic diagrams from low mountain ranges in Central Europe. Data base: Schirmer (1969), Bay- FORKLIM (1996) and www.klimadiagramme.de. Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 67 the dry steppes such as Artemisia spp. (Aster- The post-glacial (Holocene) period began aceae) and Helianthemum spp. (Cistaceae), with a distinct warming. In the pre-boreal pe- as well as species of Chenopodiaceae, most riod (10300-8800 years ago) birch and pine of them common in eastern Europe (Firbas expanded their ranges again. In the follow- 1949/52, Lang 1994) are found. Only at its ing boreal climatic epoch (8800-7500 years northern margin the Central European area ago), there was a major expansion of Cory- was covered by nordic inland ice. The last lus L. (Betulaceae). In birch-dominated land- glacial period is therefore called the Weichse- scapes, pines spread out considerably for a lian glaciation. Further to the south, the Alps short time. At the same time, species of mixed were covered by a massive ice sheet, extend- oak forests [Ulmus L. (Ulmaceae), Quercus ing southwards to the northern Alpine foothills L. (Fagaceae), Acer L. (Aceraceae), Tilia L. (with its nal period being the Würm glacia- (Tiliaceae), Fraxinus excelsior L. (Oleaceae)] tion). Around some peaks of the low mountain expanded their ranges and rapidly replaced ranges (e.g. Black Forest, Bavarian Forest, the light-demanding pioneers. In the south- Krkonošé), there were local glaciations (Er- eastern low mountain ranges, Picea abies A. genzinger 1967, Liedtke 1975, Walter 1992). Dietr. (Pinaceae) appeared for the rst time. With the gradual warming in the last glacia- During the Atlantic period (7500-4500 years tion (up to 10000 years ago) the ice sheet ago), species of mixed oak forests became slowly melted. However, the post-glacial re- dominant. In the valleys of the northern low- forestation proceeding from south to north did lands and in other smaller areas, a mass not occur immediately. Theories about post- expansion of Alnus Miller (Betulaceae) oc- glacial migration differ signi cantly: trees and curred. Pinus was the only genus to persist as shrubs spread from their Pleistocene glacial a domi nant taxon in more continental areas refuges in south-eastern and south-western of the eastern lowlands. Picea covered the Europe via different routes and with different higher elevations of the eastern low mountain speed, so that they reached dominance suc- ranges to the Harz Mountains. In the second cessively or at the same time. The character- half of this period (from 6200 years ago) Fagus istic succession of the vegetation, depending L. [principally F. sylvatica L. (Fagaceae)] and on the climatic changes has been de ned as Abies alba Miller (Pinaceae) began to spread the so-called Mitteleuropäische Grundfolge out in the southern low mountain ranges of (Central European basic sequence) (Firbas the area. In loess regions, the rst Neolithic 1949/52). farmers interferred with the natural succes- The rst tree species, i.e. those resistant sion (as shown by palynological evidence). to frost, but also light demanding pioneer spe- The sub-boreal period (4500-2800 years ago) cies such as those of Betula L. and Pinus L. was characterised by a signi cant cooling and (Pinaceae), along with Salix L. and Populus increasing precipitation. The species of mixed tremula L. (both Salicaceae) colonised the oak forests were displaced gradually by the steppe areas. Forest succession started in the progressive propagation of beech and r as north-west and the west with high percentages well as of Carpinus betulus L. (Corylaceae) of Betula, whilst in the south-west, south-east and Picea in the north-east. The propagation and the east Pinus dominated. During the so- of beech may have been accelerated by the called Alleröd interstadial (11900-10800 years anthropogenic opening up of the mixed oak ago) the greatest forest density and cover of forests in the Bronze Age (e.g. Küster 1996, the last glaciation was reached. At the end of 1998a, Pott 1997a, 2000). this period a new relocation of the forests took In the older sub-Atlantic epoch (2800-1000 place. It was caused by a climatic setback. years ago) the competition balance shifted This nal glaciation (10800-10300 years ago) in favour of the beech (the so-called beech was characterised by an open landscape with period). This was caused by cool, humid, tundras poor in trees (birch and pine) and ar- sub-oceanic climatic conditions, similar to the eas devoid of forest (low mountain ranges). present-day climate, which permitted beech- es to colonise lowland and upland forests 68 Rüther and Walentowski: Tree species composition and historic changes…

(demontane spreading), in part accompanied northward from one inter-glacial period to the by r and, in higher altitudes, by spruce. The next. north-western lowlands were still dominated (4) The west-east oriented Alps represent- by oak forests but, in the north-eastern low- ed migration barriers both for the retreat into lands, Carpinus was evident. The more re- the glacial refuges, and for the inter-glacial cent sub-Atlantic period (starting from 1000 expansion. The Mediterranean can be inter- years ago) was shaped by a growing anthro- preted as an additional barrier. pogenic impact. The cultural epoch of the Iron (5) In the glacial refuges, the climatic con- Age was characterised by increasing agricul- ditions were so extremely unfavourable that tural land-use together with decreasing forest none of the forest communities occurring in cover and an economical promotion of certain Central Europe could survive there (Beug timber species. Initially, pioneer species such 1977). These facts can be summarised as fol- as birch and pine and then species that could lows: be coppiced, such as hazel, elm, hornbeam (1) Since arboreal taxa became extinct and oak, pro ted from man-made clearings north of the Alps, e.g. during the Würm gla- and exploitation of forests. Finally, the signi - ciation, several thousand years elapsed until cance of broad-leaved trees decreased, whilst plant and animal species re-immigrated from spruce and pine gained in importance. Initially refuges in regions south of the Alps. This was this transition occured more or less unwitting- determined by climatic and biotic conditions ly by man. Later, this became active, caused but also by historical events. by the conversion of coppice (with standards) (2) Glacial refuges are thought to have systems to high forest systems, which were contained a large share of the intra-speci c harvested by clear-cutting. biodiversity of the temperate biota (e.g. hap- lotype diversity in Fagus has a strong latitudi- nal gradient). Potentially natural forest vegetation of (3) Modern forest communities started to today develop in the Holocene during re-immigra- tion of tree species. Beech forests were as- Tree species and forest communities sessed to be rather young in comparison with other Central European forest communities, The Central European ora of the Quater- since Fagus re-immigrated as one of the last nary was characterised by a massive spe- tree species. In the north of Central Europe, cies depletion, in particular with respect to they are not older than approximately 3000 to the arboreal taxa (the Pleistocene extinc- 4000 years, which is not more than 30 to 60 tion). The pre-Pleistocene dendro ora, once tree generations (Pott 1997a, 2000). rich in species, decreased to approximately (4) Less than 40 tree species (30 decidu- 50 genera (Walter and Straka 1970, Ellen- ous and 7 coniferous) occur in extant forest berg 1996). Several factors may have con- communities of Central Europe. 90% of the tributed to the extent of these extinctions forest communities investigated were domi- (Lang 1992). nated by one or two principal tree species (1) At least ve prolonged cold glacial peri- only (Fig. 4). ods alternated with shorter warm inter-glacial (5) Some relict communities which re- periods. semble tree species combinations of earlier (2) Climate changes between glacial and Holocene periods inter-glacial periods occurred very rapidly. - The Betula pubescens Ehrh./Sorbus au- (3) The glacial periods became progres- cuparia L. (Rosaceae) community, colonising sively colder. None of today’s dominating nutrient-poor boulder beds with considerable broad-leaved trees could have survived accumulation of humus, represents an exam- the nal Würm glaciation north of the Alps. ple from the early Holocene, similar to some Gradually, re-immigration distances became relict pine forests (Ellenberg 1996, Rüther greater, so that the species migrated less far 2003). Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 69

- Tilio-Acerion communities are examples - oxygen on waterlogged soils; for vegetation pictures during the oak/mixed - nutrients on acid podsols; woodland time (Atlantic period) (Müller in - iron and phosphorus on calcareous sites Oberdorfer 1992) on unstable and nutrient- and rich skeletal soils. - stability on unconsolidated landslides. Many Central European tree species were Tree species composition related to site fac- able to occupy similar habitats, i.e. their phys- tors and competition iological amplitude overlapped within a wide range (Ellenberg 1996). The ecological ampli- The rate of plant growth is limited by the tude refers only to those habitats where a spe- environmental resource which is in least sup- cies could grow and reproduce successfully ply. On a regional or greater scale, the range in competition with others. In Central Europe, of a tree species is mainly determined by the Fagus is a particularly competitive and shade- climate. For example, temperature may be a tolerant tree species, driving other species limiting factor in locations of higher altitude. out. Its ecological amplitude is nearly identical According the Map of the Natural Vegeta- to its physiological behaviour (Fig. 5). tion of Europe (Bohn 2000/2003), beech is The competitiveness of beech had a great common from planar-colline up to sub-alpine effect on the number of tree species in Cen- levels, with an emphasis on sub-montane tral European forests. Fig. 6 shows that beech to montane regions. In the montane to high dominated upland forests both on acidic and montane levels, beech is accompanied by r on calcareous sites. Other tree species occur- and spruce. However, a climatically-caused ring on nutrient-rich beech forest habitats are drought limit could not be identi ed for beech classi ed as associated trees (see Hordely- in the area. Even in the central German dry mo-Fagetum in Fig. 4), which dominated only region east of the Harz Mountains, beech temporarily in certain dynamic phases or at forests prosper at an annual precipitation of the ecological margins of a forest community. 500mm or less (i.e. large areas in the forest The in uence of beech in reducing species of Ziegelroda between Querfurt and Artern; diversity is illustrated in Fig. 6. A high diversity Leuschner 1998). In sub-continental lowlands of tree species can only be expected in nu- of eastern Central Europe with warm sum- tritious habitats outside natural beech domi- mers, the vegetation consists of oak/horn- nance. In particular, these characteristics ap- beam forests. On over-exploited, poor and plied to: dry sandy soils in this region, pine prevails. (1) lowland riparian forests (Ulmenion mi- In the lowlands, particularly in the west, aci- noris; Fig. 4, no. 35) and ravine forests (Tilio dophilous oak/mixed forests are widespread, Acerion: Fig. 4, nos. 16 and 22) on sites with but their degree of naturalness is question- high dynamics which provided a large pool of able due to their former utilisation in coppice temporary niches and spatial micro-habitats and incomplete post-glacial re-immigration of and beech. In higher low mountain ranges (e.g. (2) warm-dry, isolated oak/shrub forests Harz Mountains, Black Forest, Bavarian For- (Quercion pubescentis-petraeae; Fig. 4, est, Krušné hory, Sudetic and Carpathian no. 25) and oak/hornbeam forest habitats Mountain ranges) coniferous forests with (Carpinion; Fig. 4, no. 23). 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; 70 Rüther and Walentowski: Tree species composition and historic changes…

high-altitudes from lowlands up to mountains Alnus viridis Salix appendiculata Pinus mugo Pinus cembra Larix decidua Sorbus aucuparia Acer pseudoplatanus Picea abies Abies alba Fagus sylvatica Ulmus glabra Fraxinus excelsior Alnus glutinosa baccata Taxus 1 2 3 4 5 6 7 8 9 11 12 13 14 10 1 Alnetum viridis 2 Rhododendro-Pinetum mughi a subalpine 3 Vaccinio-Pinetum cembrae 4 Adenostylo glabrae- / Homogyne-Piceetum l 5 Calamagrostio villosae-Piceetum t 6 Aceri-Fagetum 7 Ulmo glabrae-Aceretum pseudoplatani i 8 Sorbo ariae-Aceretum 9 Luzulo luzuloidis-Fagetum, montane t montane 10 Galio-Fagetum, montan u 11 Galio rotundifolii-Abietetum 12 Aposerido-Fagetum d 13 Seslerio-Fagetum i 14 Calamagrostio variae-Pinetum 15 Fraxino-Aceretum n 16 Adoxo-Aceretum 17 Vaccinio vitis-idaeae-Abietetum a submontane 18 Hordelymo-Fagetum l 19 Luzulo luzuloidis-Fagetum, colline-submontane 20 Galio-Fagetum, colline-submontane 21 Carici-Fagetum 22 Aceri platanoidis-Tilietum platyphylli 23 Galio sylvatici-Carpinetum z 24 Stellario holosteae-Carpinetum colline 25 Potentillo albae- und Cytiso-Quercetum o 26 Calamagrostio arundinaceae-Quercetum 27 Luzulo luzuloidis-Quercetum n 28 Leucobryo-Pinetum e planar 29 baltic Melica uni ora-beech forest 30 Deschampsia exuosa-beech forest 31 Betulo-Quercetum riparian 32 Carici remotae-Fraxinetum forests 33 Pruno padis-Fraxinetum 34 Alnetum incanae 35 Querco roboris-Ulmetum minoris 36 Salicetum albae alder carr 37 Carici elongatae-Alnetum glutinosae 38 Vaccinio uliginosi-Betuletum pubescentis peatland 39 Vaccinio uliginosi-Pinetum sylvestris forests 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. Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 71

lowlands and uplands pioneer character riparian forests div. spec. div. Tilia cordata Tilia Carpinus betulus Quercus robur Quercus petraea Acer campestre Prunus avium platyphyllos Tilia Acer platanoides Sorbus torminalis Betula pendula Populus tremula Sorbus aria Pinus sylvestris Malus, Pyrus Betula pubescens agg. Prunus padus Ulmus carpinifolia Ulmus laevis Alnus incana Populus nigra Salix alba Salix

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 n 122-5 115-7 212-5 1-6310 114-6 21328 13318 214-7 12227 124-7 216110 12418 117-9 1-7-8 415-10 2111216 215210 1 -14217 115310 1111215 1-11113 3111-15 336315 227314 211114 23319 125311 1-618 117211 113510 11417 246416 228416 129-12 3 310117 115-7 12328 11125 124-7 131-5 1-618

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

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 Fig. 6: Approximate tree species numbers in the oak/mixed woodlands and sub-continental Quer- tree layer of important alliances and sub-alliances cus/Carpinus forests (range a), dominance of of Central European forest communities (Leusch ner Fagus is adapted to closed stands (a favourable 1998). The numbers apply to well-developed (i.e. micro-climate) and intact humus cover. In nutrient- comparatively species-rich) stands and are based, poor and strongly acidic habitats (range b) Fagus mainly, on relevés in Oberdorfer (1992). In these can predominate if intact humus cover guarantees tables, only those tree species which achieved an a comparatively favourable nutrient supply. Moist abundance of at least 25% constancy in the rele- soil locations (range c) can favour Fagus, partic- vant associations were considered. ularly on sandy substrates and under an oceanic climate (lower drought endangerment of the upper cies typical for the region and adapted to the soil), contrasting Quercus and Carpinus. site (such as Quercus, Fraxinus or Acer) may react rapidly and become established as ser- al species. In mixed montane forests, beech Tree species composition related to time has a shorter average life expectancy (200- 300 years) in comparison with spruce (300- During a development cycle of up to 600 400 years) and r (400-500 years). Hence years, a virgin forest passes through different the r becomes dominant particularly in late phases (cyclic changes, Fig. 7). These include development phases, i.e. late over-mature growing, ageing and dying of tree individuals phase and regeneration phase. and show completely different structures. The The opening up in the over-mature phase succession of a complex virgin forest system is crucial (e.g. in terms of gap- and patch- is undetermined and subject to random uc- dynamics). Leibundgut (1982) proved that tuation (factors from outside the forest which in beech-dominated mixed montane forests, affect growth, e.g. insect attacks, wind and most collapse and rejuvenation took place snow damage and others). on small-scale by loss of individual old trees. Most different micro-climates, light regimes Usually, on these sites the next beech gen- and niches provide a temporally-graduated eration followed directly, either evenly spread and species-rich rejuvenation. In particular, or in clusters. It may be concluded that only temporal heterogeneity in virgin forests caus- in larger gaps or patches, rapidly-growing es large structural variety and creates spatial species, such as Fraxinus excelsior L. and patterns of different dimensions and ecologi- Acer pseudoplatanus L., reached a tempo- cal niches. Due to random uctuation, some rary dominance, which contributed to an in- light-demanding and semi-tolerant tree spe- creased species diversity. Comparable tree Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 73

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- Fig. 7: Cyclic changes in mixed mountain forests, il- ducing annual crops, effected pronounced lustrated by the example of virgin forest relics of the lower Austrian limestone Alps (according to Zukrigl dynamics (secondary successions). Farm- et al. 1963). ing practices depended on traditional know- ledge. Natural forest vegetation was cleared by burning without removing the roots. Land species successions were also observed in use was rarely permanent, since it was aban- the former woodland pastures of north-wes- doned for recuperation when yields declined tern Germany (Koop 1982). markedly (e.g. Kossack 1982, 1995, Lüning et al. 1997). Under this in uence of shifting cultivation, the proportion of the forest area Anthropogenic changes: transition of the remained relatively high. From the Neolithic forest vegetation period, human settlement behaviour changed. Instead of a nomadic existence, peoples be- Duration and intensity as well as effects came increasingly settled and established of anthropogenic in uence vary within differ- permanent dwellings. These profound altera- ent growth ranges of Central European for- tions instigated by the beginning of agricul- ests. The comparison of palyonogical data ture are called the Neolithic Revolution (e.g. indicates temporal and regional differences Küster 1998a, 1998b). between coastal, lowland, upland, and low In the Bronze and Iron Ages (2200-750 mountain regions (Pott 1997b). The degree of B.C. and 750-15 B.C., respectively) and early landscape changes were determined by not Antiquity, human impact on soil and vegeta- only (i) the varying intensity of human activi- tion became greater with increasing popula- ties on a regional scale but also (ii) by the lo- tion, with need for fodder (due to the unfa- cal site conditions or (iii) the regenerative abil- vourable winter temperatures) and with new ity of the habitats (see the overview in Pott cultivation techniques on arable land (Lang 1988). As a result, an intensely structured 1994). Initially, human disturbance concen- landscape with a great number of vegetation trated, almost exclusively, on lowlands and types developed during the period of exten- uplands that were easy to cultivate. sive management (Burrichter 1977). From the early Middle Ages, unaffected forest regions of the low mountain ranges, Forest area such as the Bavarian Forest, became more heavily settled and cultivated (Stalling 1987, Under anthropogenic in uence, the ex- Nelle 2002, Rüther and Nelle 2006). Due to pansion of the forest area was subjected to adverse climatic conditions unfavourable for permanent change. Most spatial alterations sustained agriculture in many places, the set- occurred in times of upheaval or major crises. tlements were later abandoned (Fehn 1963) Palynological data show that, for example, and forest cover remained comparatively during the Migration of Nations (its culmination high. 74 Rüther and Walentowski: Tree species composition and historic changes…

In the late Middle Ages and in the begin- Wood pasture is an ancient type of multi- ning of modernity, reduction of the forest area functional use. The resulting structure was reached its greatest extent. Decisive factors a mixture of woodland and grassland that were industry and mining, in particular the iron developed through a long history of grazing and glass industries, as well as salt mines, by domestic herbivores under gnarled trees which required much wood or wood products (often pollarded, see below). Wood pastures (i.e. charcoal and potash). Major substantial were widespread in lowland and upland land- disturbances were found in partial regions of scapes through the Middle Ages and until the the Harz Mountains (Hillebrecht 1982, Bartels 19th century. The resulting timber was small 1996), the Siegerland (Pott and Speier 1993), in diameter and was harvested for fuel, con- the Black Forest (Brückner 1981, Golden- struction of wooden items and possibly char- berg 1996), the Upper Palatinate (Lutz 1941, coal. The effects of wood pasture were: Vangerow 1987) as well as the Bavarian Al- (1) opening up of the forest landscape; pine Uplands (Bülow 1962, Knott 1988, Zi- (2) encouragement of the thicket stage of erhut 2003). At the same time, reforestation trees and ground layer species; measures took place in many deserted areas; (3) promotion of tree species with vigorous for example, these have been documented sprouting and regeneration capacity for the Nuremberg Reichswald in 1368 (Man- (e.g. Carpinus, Fraxinus, Corylus and tel 1968). In the course of modern forestry, Ulmus). at the end of 18th century, reforestation had (4) selection of woody species which bore become mandatory. Since this period, the for- spines or thorns [e.g. Juniperus L. (Cu- est area in different regions of Central Europe pressaceae), Rosa L., Prunus L. and has increased again. Crategus L. (all Rosaceae)] or toxic for Today, approximately 30% of Central Eu- the cattle (e.g. Alnus). rope are covered by forest (Mayer 1984). The (5) suppression of shade tolerant, but less largest connected forest areas can be found in regenerative climax trees like beech. the central mountain regions (e.g. the Pfälzer Similar species selections were caused by Wald, the Odenwald-Spessart-Rhön hills and pollarding techniques. Pruning was performed the Bavarian Forest). However, the forest ar- for winter feeding and bedding (e.g. Pott 1983, eas are reduced greatly and have become Haas and Rasmussen 1993). Many of the pol- isolated and fragmentated, particularly in the larded trees are now ancient. Some of these loess-covered lowland and upland regions relicts of both wood-pasture and pollarding (e.g. Franconian plateau, northern Alpine remain in parts of Central Europe (e.g. Neu- foothills), where cultivation was easy. This enburger Urwald, Nitzschke 1932, see also also applied to offshore areas [e.g. Schles- the overview for north-western Germany in wig-Holstein (northern Germany), Denmark Pott and Hüppe 1991). The complex structure and Skåne (southern Sweden)]. of the gnarled old trees provided a number of micro-habitats, such as bark crevices and Tree species composition tree holes, which are not found in high for- est and this made them suitable for species Due to human activity and usage, the tree of fungi, invertebrates and bats. The effects species composition of the original forests of the wood-pasture on the growth habit and was changed markedly. Until the Middle Ages the phenotype of beech were shown for the these changes occurred more or less unwit- Bavarian Forest and the Black Forest (Sey- tingly by man. Commencing in the late Middle fert 1975, Schwabe and Kratochwil 1987, Ages and becoming more widespread in the respectively).Today, the ground vegetation of modern era, organised forestry was under- such in uenced forest stands is dominated by taken in western Europe (Köstler 1956) with Agrostis L. and/or Holcus lanatus L., with Fes- the intention of promoting certain tree species tuca rubra L., Deschampsia exuosa (L.) Trin. (e.g. 1529: Waldpuech für das Reichenhalle- (all Poaceae) and herbs, such as Potentilla sche Waldwesen). erecta (L.) Raeusch. (Rosaceae) and Galium Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 75 saxatile L. (Rubiaceae). It contained some rst plantings of Pinus were documented in woodland edge species, such as Teucrium the14th century for the Nuremberg Reichswald scorodonia L. (Lamiaceae), but few typical (Mantel 1968). This introduced a gradual woodland species. change from broad-leaved to conifer-domi- Coppicing is the oldest form of forestry; nated forests (Fig. 8). As a pioneer with few woven hazel screens used for shing have demands, Pinus established itself spontane- been dated back to 5000 B.C. (in the Neolith- ously on strongly degraded soils (e.g. in the ic period). Much of the ancient semi-natural Upper Palatinate, Lutz 1941, Diepold 1945). woodland in Central Europe developed un- Pinus bene tted from anthropogenic exploi- der the coppice management system. Many tation and deforestation of the landscape, as broad-leaved trees, including Corylus, can did Picea in the low mountain regions. Both be cut down to the stumps, which re-grow genera expanded outside of their native ar- producing multiple stems called poles. Usu- eas (e.g. Black Forest, Harz Mountains, lower ally, the poles were harvested every 15 to 25 Bavarian Forest; Brückner 1970, Schubart years as fuel, the poles of hazel were har- 1978, Rüther 2003, respectively). vested approximately every eight years and Finally, from the middle of the 18th and until converted into a wide range of products. Tra- the beginning of 19th century, organised for- ditionally, willow [Salix L. (Salicaceae)] poles estry promoted high forest systems (Hartig were used in hurdles. In addition to Corylus 1791, Cotta 1817). Silviculture (as a theory and Salix, Carpinus, Tilia, Acer, Fraxinus and and practice of controlling forest establish- Alnus were promoted. Quercus, Ulmus and ment, composition and growth) was devel- Populus (Salicaceae) are also adapted to oped as discipline for the manipulation of for- coppicing, but have less regeneration poten- ests in order to enhance certain forest values tial. Like wood pasturing, pollarding coppicing or products. Due to previous forest devasta- suppressed not only Fagus, but also conifers tion and exploitation of the natural resources, such as Abies, Picea and Pinus. Requiring a lack of timber, perhaps inspired further by longer rotation periods, beech-dominated the idea of a rapid economic pro t (theory of coppices were limited to some regions with nancial rotation), resulted in the planting of milder winters, such as the margins of the low age-class forests, dominated by Picea, Pinus mountain ranges in north-western Germany and Pseudotsuga menziesii (Mirb.) Franco (e.g. Pott 1981) . (Pinaceae) (introduced from Paci c North Coppicing with standards, a variation of America). Table 1 shows that the relationship traditional coppicing, was established in the between broad-leaved trees and conifers Middle Ages. Individual trees were excluded has become reversed in comparison with the from short-term activities. Quercus, in particu- original situation. An additional and signi cant lar, was promoted, since it served in pig-keep- factor was that considerable areas of native ing (e.g. Hesmer and Schröder 1963, Schöller broad-leaved forests were cleared for farm- 2001). This form of management was particu- ing. The proportion of Picea increased notably, larly common in densely-settled loess regions whilst the percentage of Abies declined over of Central Europe. This sometimes led to the last two hundred years. A well-known ex- shortages of wood (Ellenberg 1996). ample of the decline of Abies is from the Fran- This preservation of standards and the conian Forest, where, in the Middle Ages, a oak mast for pig-keeping promoted a con- r-promoting selection system was operated. scious tree species for the rst time. Also in It was based on the export of raft-wood along low mountain regions, human activities in- the rivers Main and Rhine to the Netherlands uenced the tree species composition of the (Fig. 9). Selective timber production resulted forests to a greater extent [e.g. plenter forests in a change of tree species combination: prior (i.e. individual tree selection, group selection to the 16th Century, we estimate a ratio of 40% system) promoting Abies directly, Wirth 1956, of broad-leaved trees and 60% of Abies. An Rüther 2005)]. At the same time, reforesta- initial consequence was the elimination, by tions of many deserted sites took place. The creation of forest pasture, of rare high-quality 76 Rüther and Walentowski: Tree species composition and historic changes…

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.. Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 77

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 conifers beech oak others pine spruce r trees original 66 34 38 20 8 20 7 7 current 29 71 14 10 5 44 25 2 hardwood. This was followed by disorganised withered before the roots had reached min- use, including selective removal of timber eral soil. In 1830, productive forests were (e.g. material for joinery). Fagus, unwieldy in formed exclusively by Abies and Picea, but, growth habit and incapable of otation, was after 1850 the former was nally displaced used less. In 1665 the share of broad-leaved by the latter. Reforestations often succeeded trees (exclusively Fagus) was not more than with Pinus and Picea only, and the decline of 10%, whilst Abies and Picea covered 80% Abies was accelerated by large-scale open- and 10%, respectively (Wirth 1956). ings caused by undifferentiated management Exploitation and the breaking up of the methods. Another important reason for the canopy led to an increasing cover of Picea. dramatic decline of Abies was intensive man- Within the raw humus cover and dwarf shrub agement of game. Comparable simultaneous blankets with intensively-matted roots, Fagus decreases and increases of Abies and Picea, could either not germinate or its seedlings 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 Fig. 9: Changes of tree species combination in the timber trade and rapid staff reduction within in Franconian forest due to human impact. forest administration. Nevertheless, sustaina- 78 Rüther and Walentowski: Tree species composition and historic changes…

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 Fig. 10: Natural and current stocking related to the to atmospheric nitrogen immissions up total forest area of Bavaria. The natural stocking to 30 kg/ha/a and was projected approximately over the area of the (3) global warming, as it supports decidu- natural forest communities. The current stocking ous trees of temperate zones, with data is based on the national forest inventory. conifers dominating boreal, sub-boreal or montane zones. However, in some landscapes of Central ble forest management is increasingly orient- Europe, Fagus or Quercus disappeared en- ed towards the natural biological resources of tirely or became so rare that rejuvenation on a landscape (by taking advantage of primary suitable habitats did not occur spontaneously. production, thus minimising the need for in- The lack of nursery trees, selective game tervention). damage, unfavourable micro-climate, frag- The following factors require special con- mentation and isolation of forests or dense/ sideration: impenetrable ground layer vegetation compli- (1) ecology (soil conditions, indicator spe- cated resettlement. cies groups and vegetation units/natu- ral forest communities with their princi- Forest structures pal, secondary and pioneer trees) and (2) forest genetics (sources, genetic vari- The structure of the stand, in particular the ability of tree populations). complexity and dynamics of the forest cano- Individual stand characteristics (stand his- py, is of fundamental importance to the for- tory, genetics, current and natural vegetation, est dynamics, e.g. spatial heterogeneity and succession stage, ecological site character- temporal change in understorey vegetation, istics) are analysed for the suitability of tree patterns in regeneration mosaics, and micro- species. Recommendations are given for dif- climatic variation (Norman and Campbell ferentiated, economically rational and ecologi- 1989, Song et al. 1997). The structure of the cally sound sylvicultural options. Site-adapted forests was modi ed by different forms and biodiversity and rich structural heterogeneity intensities of utilisation. The wide range of cli- (chapter “forest structures”, see below) of the matic, geological and geobotanical conditions stands are considered as indicators for a sus- in Central Europe produced a differentiated tainable forest management under the risks settlement history and speci c management of climatic change/global warming (e.g. Ferris techniques, resulting in regionally different and Humphrey 1999). The maintenance or re- forest structure types. Fig. 11 shows the pos- establishment of forests in as natural a state sible modi cations of the natural forest in the as possible is a strategy for sustainability [e.g. hill and mountain country depending on the as shown by LÖWE (=langfristige ökologische use. The dynamics between utilisation and Waldentwicklung, Otto 1989), a forestry pro- abandonment produced numerous transition Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 79 stages between the individual structure types, can affect substantially the adaptive abilities leading to a considerable structural diversity of tree stands and genetic resources of tree even on small sites. species (Müller-Starck 1996), and which may There are very little data on the structure be occupied by the entire range of regionally of original virgin forests in Central Europe typical admixed/secondary tree species, in- (chapter “tree species composition related to cluding rare, sensitive and endemic species time”, see above). Relicts of virgin forests re- (e.g. Otto 1989, Schölch et al. 2003). mained small-scale in climatically rough and inaccessible locations within montane and Stages of forest development sub-alpine zones. Observations within these locations cannot be applied by inference to Each kind of forest utilisation disrupts the lowland forests. The principal differences be- development cycle of woodlands, in particu- tween commercial forest and virgin forest are lar with regard to old-growth and over-mature the high degree of order (regular structure), stages (chapter “tree species composition the limited heterogeneity, breaks in continu- related to time”, see above). In prehistoric ity of old-growth-habitats and the quantity and times, the human and animal impacts oc- quality of deadwood (Walter 1986). curred temporarily only and were limited spa- With reference to stand structure, (i) the tially (chapter “forest area”, see above). After size and shape of gaps and patches, (ii) the utilisation, the sites were able to re-grow, so uneven cutting of individual trees (resulting that, apart from site changes, these impacts in small and large groups) and (iii) selective could be compared with some natural events working resemble natural forest conditions. like forest re or wind throw. In addition, nature-oriented forestry practice From the Neolithic period onwards human tries to create niches of different size, which impact became more intense, due to new

Forest structures lowand and upland regions Forest structures mountainous regions

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). 80 Rüther and Walentowski: Tree species composition and historic changes… cultivation systems. Forests and elds were ground layer was modi ed profoundly. Forest cultivated permanently and systematically, so litter utilisation and removal of sods displaced that the natural development of forests was the original forest oor species and supported affected profoundly. Forest pasture caused the invasion of indicators of soil degradation different vegetation types, with degeneration such as Calluna vulgaris (L.) Hull (Ericace- and regeneration complexes occurring simul- ae). Particularly large areas of heathland de- taneously (Pott 1988). With the decrease of veloped on the sandy soils (on old moraine pasturing, regressive reforestation processes landscapes) in north-western Germany and occurred. Due to enduring extensive pasture, in Denmark (Ellenberg 1996). Heather also thorn bushes played a decisive role in forest spread in the forests of low mountain ranges regeneration, due to their protective function such as the upper Bavarian Forest (Rüther for the natural thicket stage. The intensity of 2003). the pasture instigated many dynamic, spa- The forests opened up due to woodland tially and temporally heterogenous systems, pasturing, so that the stand climate changed. consisting of degeneration and regeneration Speci c shade-tolerant species on the forest complexes. However, successful regenera- oor disappeared and were replaced by light- tion occurred after pasturing had ended. demanding herbs and shrubs. Additionally, In some low mountain ranges an alternat- some grass species, adapted to grazing but ing forestry and eld crops system was devel- unknown from forests, were able to invade. In oped including agriculture, forest pasture and addition to tree species, grazing animals also coppicing (e.g. Black Forest: Schwabe-Braun enabled the spread of numerous herbs and 1980, Bavarian Forest: Lippert 1984, Reif and shrubs, which possessed toxic ingredients or Oberdorfer 1990, Siegerland: Pott and Speier thorns (this is dependent on the natural zone, 1993). Thus, on the same sites, forest de- e.g. Pteridium aquilinum (L.) Kuhn (Denns- generation stages continually took place (Fig. taedtiaceae), J. communis L., Ilex aquifolium 11), whether induced by man or animal. Re- L. (Aquifoliaceae), Prunus spinosa L., Cra- generation was possible during fallow periods taegus spp., Rosa spp., (all three Rosaceae), only. Anthropogenic in uence on forest devel- Genista spp. (Fabaceae), Thymus spp. (La- opment is particularly evident in the coppice miaceae). system: timber was cut in a regular cycle of 15 All of (i) the method, frequency and inten- to 25 years. Comparable natural disturbances sity of utilisation, (ii) vertical and horizontal and interactions, with frequent environmental structure of the stand, (iii) canopy closure, (iv) changes and similar mechanical demands on mixture of tree species, (v) isolation, fragmen- and damages to trees, are strictly limited to tation and partition of the forest area, (vi) the habitats along untamed rivers and avalanche history of the forest stand and (vii) the stage lanes in high mountains. of forest development are potentially deter- Even in modern high forest systems with minant factors of the species composition of site-adapted near-natural tree species com- the ground layer in today’s commercial for- bination, a high degree of naturalness may ests. Man-made forest edges or openings are be inferred only if the factor time is neglected. habitats for some light-demanding species Within a conventional rotation period of 100 which are absent in natural forests or occur to 140 years (Mayer 1992), the characteris- in natural open stages of forest development tic complexity, heterogeneity and biodiversity only. These species immigrate from either ad- are reduced drastically (chapters “tree spe- jacent agricultural areas or from natural open- cies composition related to time” and “forest ings such as rocks, stone elds, debris cones structures”, see above). and riparian areas. In forests with a young habitat tradition, some relicts of former land Herbaceous layer, brushwood use (e.g. dry grasslands, alpine pastures, lit- ter meadows, vineyards and orchards) may Due to the different forest utilisation and be present. management types, the composition of the With reference to the composition of the Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 81 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 Fig. 12: Spatial distribution of Galium odoratum in Bavaria (Schönfelder and Bresinsky 1990). Rea- satellite populations and low competitive abil- sons for absence - a: anthropogenic, e: edaphic, ity. Another reason is the modern reduction of c: climatic. dispersal processes (primarily of forest pas- ture) in comparison with the historical man- made landscape (Bonn and Poschlod 1998). within the loess-covered uplands are explic- In addition, the isolation and fragmentation of itly favourable for these species. In southern current forest stands affects the aggregate Germany, thus far, only a few authors have ability to colonise (Rackham 1980, Peterken made (relatively small-scale) investigations of and Game 1984). Furthermore, most of the forest oor species, comparing ancient and indicator plants have no persistent seed bank recent forests (e.g. Schneider and Poschlod record (Bossuyt and Hermy 2001). Finally, fa- 1999). Such studies are badly needed, partic- vourable stand climate and humus condition ularly with regard to forest management and are necessary for regeneration and/or resto- forest conservation. ration. In addition to qualitative changes in spe- Most of the low- and upland beech for- cies, quantitative changes have been ob- est habitats have a tradition of disturbance served. Oxalis acetosella L. (Oxalidaceae) or perturbation. They regenerated from cop- and numerous mosses of the ground layer, pice forests or were afforested (Dierschke such as Eurhynchium angustirete (Broth.) and Bohn 2004), so that most of them may be T.J.Kop. (Brachytheciaceae), Thuidium tama- termed recent forests. In Bavaria, some plant riscinum (Hedw.) Bruch et al. (Thuidiaceae), species typical of beech forests and also indi- Hypnum cupressiforme Hedw. (Hypnaceae), cator plants for ancient forests are absent in Polytrichum formosum Hedw. (Polytrichace- speci c landscapes. The large-area overview ae) and Hylocomium splendens (Hedw.) for Galium odoratum (L.) Scop. (Rubiaceae) Bruch et al. (Hylocomiaceae), have spread shows a patchy occurence in the loess re- out in the man-made Fagetalia-spruce forest- gions of Franconia and Upper Bavaria (Fig. communities. Originally, these species at- 12). This absence can be interpreted only by tained their ecological optimum in montane, the aforementioned reasons (principally short- herb-rich mixed coniferous forests on accu- distance dispersal strategy, isolation of forest mulated humus. Recently disturbed clear-cut stands and reduction of dispersal processes), areas may be colonised rapidly by tillering since both climatic and edaphic conditions and creeping dwarf shrubs [e.g. Rubus spp. 82 Rüther and Walentowski: Tree species composition and historic changes…

(Rosaceae)] such as blackberry and rasp- centrated on the humus layer and the upper berry and grasses [e.g. Calamagrostis spp. humus/mineral soil layer (Ellenberg 1996, (Poaceae)]. Further soil compaction encour- Gulder 1998). ages the clonal spread of Carex brizoides With the abandonment of forest litter use L. (Cyperaceae), once harvested as an eel at the beginning of the 20th century, the regen- grass [Zostera L. (Zosteraceae)] substitute in eration of the humus layer commenced. This mattress llings. development has been accelerated greatly by air pollution and raw humus amelioration over Soils and matter cycles the last 50 years, but has led to eutrophica- tion, which is extreme in some regions. In ad- The numerous forest utilisation and man- dition to large-scale nitrogen deposition, there agement types have not only affected the are many small-scale impacts on the soils. vegetation but also the nutrient content of the Soils affected by man (e.g. soil compaction, soils. Due to the enduring nutrient depletion, lime substrate from forest path construction, the forest litter utilisation and the removal of small nitrogenous patches) may be found sods led to a lasting soil degeneration, par- along forests roads. Additional eutrophication ticularly on deep, badly buffered sites with low from adjacent farmland occurs along forest levels of lime and bases. Since the end of the borders. Middle Ages, in southern Germany, initially in the Upper Palatinate but also in Middle and Lower Franconia, a massive forest litter Conclusions and suggested management utilisation was conducted (Rebel 1920, e.g. responses Nuremberg Reichswald), Ott-Eschke 1946, 1951, Sperber 1968, large woodlands in the The present tree species composition of Upper Palatinate hill and basin landscapes, the Central European oak/beech region has Lutz 1941, 1942). In many cases, the ground been determined largely by historic in u- litter and upper humus-rich soil layers were ences. Formative processes were the Pleis- removed. Forest litter use also damaged or tocene extinction and the post-glacial migra- destroyed the thicket stage of the trees, limit- tion of trees in close connection with human ing the regeneration of the forests. As a con- impacts. Managing biodiversity of our forest sequence, both the yield and the vitality de- habitats should aim not for the maximum but creased (Rebel 1920). for locally speci c species diversity (Granke et In terms of sustainability, the use of forest al. 2004). To strive for a maximum species di- litter did more damage than any other type of versity would primarily mean to support a pool forest exploitaiton. Generally, silviculture was of non-speci c and ubiquitous species, intro- able to compensate its effects on the forest duced by anthropogenic disturbance. Howev- vegetation within a reasonable period of time, er, due to the dramatic loss of wilderness, the for example a few decades. However, chang- assessment should relate to the original and es in soil composition have a lasting effect speci c composition, structure and dynamics over several centuries (Hasel and Schwartz of natural forests. In other words, biodiversity 2002). The acidi cation and subsequent nutri- is only an important operational standard of ent depletion of soils is a natural and gradual value for nature protection in connection with process that was accelerated unwittingly by the degree of naturalness. To measure this man, and to a far greater extent than in prim- degree of naturalness, three different compo- eval forests (Walter 1984). When evaluating nents have to be considered: forest vegetation, changes in site conditions (1) Tradition of old-growth trees (at least in many regions, including degradation of the 200 years) with biocoenoses of decaying soil, must be considered. Most of the moss- trees (e.g. Gastero cercus depressirostris. or lichen-dominated coniferous forests (i.e. Fabricius (Coleoptera: Curculionidae) and of Picea and Pinus) are degraded types of Bolitophagus reticu latus. L. (Coleoptera: Ten- beech forest, where the nutrient cycle con- ebrionidae) as indicators. Floren and Schmidl (eds.) 2008: Canopy arthropod research in Europe 83

(2) Natural tree species composition (rela- Mittelalter. Auswirkungen auf Mensch und ted to site factors, competition and time) with Umwelt. - Vierteljahrschrift für Sozial- und abundance and dominance of principal, se- Wirtschaftsgesch., Beiheft 121:112-127. condary, companion and pioneer trees of the Baumgartner, A. 1970. Klima und Erholung im natural vegetation unit as indicators. Bayerischen Wald - Verhandlungen Deut- (3) Natural ground layer vegetation (re- scher Beauftragter für Naturschutz und lated to typical forest character species) with Landschaftsp ege 17:39-53. presence or absence of species of the natural Bayerische Landesanstalt für Wald und Forst- vegetation unit (e.g. G. odoratum), weakness wirtschaft (LWF, ed) 2000. Großtiere als to propagate (forest speci c short-distance Landschaftsgestalter - Wunsch oder Wirk- dispersal strategy), shade tolerance, low pro- lichkeit? - Berichte LWF 27, Freising. duction of satellites, temporary seed banks as BayFORKLIM 1996. Klimaatlas von Bayern. indicators. - Bayerischer Klimaforschungsverbund, Modern management methods need to München. consider biodiversity and the degree of natu- Beug, H.-J. 1977. Waldgrenzen und Waldbe- ralness by paying attention to heterogeneity, stand in Europa während des Eiszeitalters. complexity and scale (Walter 1986, Ott et al. - Göttinger Rektoratsreden 61:1-23. 1997, McCoy and Bell 1991): ”Heterogeneity Bohn, U., Neuhäusl, R., Gollub, G., Hettwer, encompasses the variation due to the relative C., Neuhäuslova, Z., Schlüter, H., Weber abundance of different structural components, H. 2000/2003. Karte der natürlichen Vege- whether both vertically or horizontally. Com- tation Europas/Map of the Natural Vegeta- plexity refers to the variation resulting from tion of Europe. Maßstab/Scale 1:2500000 absolute abundance of individual structural - Teil 1: Erläuterungstext mit CD-ROM components, and the scale takes account (2003): 655 pp., Teil 2: Legende (2000), of the variation due to the size of the area or Teil 3: Karten. Münster. volume used to measure heterogeneity and Bonn, S. and Poschlod, P. 1998. Bedeutung complexity.” (Ferris and Humphrey 1999). dynamischer Prozesse für die Ausbreitung von P anzenarten seit dem Postglazial. - Schriftenreihe für Landschaftsp ege und Acknowledgements Naturschutz 56:147-171. Bossuyt, B. and Hermy, M. 2001. In uence of We would like to thank Neil Springate for land use history on seed banks in Europe- immensely helpful comments, criticisms and an temperate forest ecosystems: a review. suggestions. - Ecography 24:225-238. Brückner, H. 1970. Der Wald im Feldbergge- biet. Eine wald- und forstgeschichtliche References Untersuchung des Südschwarzwaldes. - Veröffentlichungen des Alemannischen Andersen, S. T. and Berglund, E. 1994. Institutes Freiburg 28:1-128. Maps of terrestrial non-tree-pollen (NAP) Brückner, H. 1981. Die Entwicklung der Wäl- percentages in north and Central Europe der des Schwarzwaldes durch die Nutzung 1800 and 1450 years B.P. - In: Frenzel, B. vergangener Jahrhunderte und ihre heuti- (ed.), Evaluation of land surfaces cleared ge Bedeutung. - In: Liehl, E. and Sick, W. from forest. Special issue: ESE Project. - D. (eds), Der Schwarzwald. Beiträge zur European Palaeoclimate and Man 7:119- Landeskunde. - Veröffentlichungen des 134. Alemannischen Institutes Freiburg 47:155- Bartels, C. 1996. Montani und Silvani im 180. Harz. Mittelalterlicher und frühneuzeit- Burrichter, E. 1977. Vegetationsbereichung licher Bergbau und seine Ein üsse auf und Vegetationsverarmung unter dem die Umwelt. - In: Jockenhövel, A. (ed.), Ein uss des prähistorischen und histo- Bergbau, Verhüttung und Waldnutzung im rischen Menschen. - Natur und Heimat 84 Rüther and Walentowski: Tree species composition and historic changes…

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tersuchungen in Urwaldresten der nieder- österreichen Kalkalpen. - Mitteilungen der Forstlichen Bundesversuchsanstalt Wien 62:1-244. submitted 02.02.2005, accepted 19.10.2005 new book information from bioform entomology & equipment 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

4 Stratification and distribution of arthropods in Table of Content tree habitats Tracing arthropod movement in a deciduous forest canopy Foreword, by the editors using stable isotopes, by Roman Asshoff, Sonja G. Keel, Foreword, by Karl Eduard Linsenmair Rolf T. W. Siegwolf and Christian Körner 1 Introduction Oribatid mites in the canopy of a Central European mixed forest: species richness and species similarity between tree Canopy arthropod research in Europe, by Andreas Floren species and habitat types, by Stephanie Sobek, Christian and Jürgen Schmidl Kampichler and Gerd Weigmann Canopy research on a worldwide scale: biodiversity, climate Stratification of 'macro-Lepidoptera' in northern Bavarian change and forest canopies, by Andrew Mitchell, Director forest stands dominated by different tree species, by Global Canopy Programme Hermann Hacker and Jörg Müller 2 General forest ecological aspects Vertical and horizontal distribution of arthropods in Canopy structure and its effect on canopy organisms: a temperate forests, by Axel Gruppe, Martin Goßner, Kerstin general introduction and some first findings of the Leipzig Engel and Ulrich Simon Canopy Crane Project with special reference to vertical 5 Anthropogenic and natural disturbance stratification, by Peter J. Horchler and Wilfried Morawetz structuring arthropod communities Microclimatic variability in the canopy of a temperate forest, Introduced tree species as an anthropogenic disturbance of by Ophir Tal, Martin Freiberg and Wilfried Morawetz arthropod communities in tree crowns of managed forests - Tree species composition and historic changes of the a case study of native Heteroptera communities on Central European oak/beech region, by Carsten Rüther and introduced red oak, by Martin Goßner Helge Walentowski The diversity of moths communities in different structured Tree crowns and forest systems from a forestry point of oak-hornbeam forests - a comparison of different states of view, by Hans Stark and Olaf Schmidt succession in coppice with standard and forests with high 3 Arthropod diversity, guilds, and resource standard trees, by Ralf Bolz related communities The impact of flooding and forestry on the species Species of the genus Oedalea Meigen, 1820 (Diptera: composition of xylobiontic and phytophagous beetles on Hybotidae): An element of the canopy fauna in European oak canopies of the Bavarian Danube floodplain, by forests?, by Andreas Stark Johannes Bail and Jürgen Schmidl Heteroptera communities in tree crowns of beech, oak and Ichneumonidae from the canopies of primary and managed spruce in managed forests: diversity, seasonality, guild oak forests in eastern Poland and southern Germany, by structure, and tree specificity, by Martin Goßner Klaus Horstmann and Andreas Floren Diversity of Neuropterida in mixed forest stands in Do spider communities in primary forests differ from those Germany, by Axel Gruppe in forest-plantations? A canopy study in the Biaowiea- The ants of Central European tree canopies (Formicidae) - Forest (Poland), by Andreas Floren, Stefan Otto and Karl Eduard Linsenmair an underestimated population?, by Bernhard Seifert Diptera (Brachycera) in oak forest canopies - management Tracking the elusive: leafhoppers and planthoppers (Hemiptera) in tree canopies of European deciduous and stand openness gradient determine diversity and community structure, by Klaus von der Dunk and Jürgen forests, by Herbert Nickel Schmidl Search in the canopies and you will find new species records of insects, by Karl H. Thunes, Ivar Gjerde, Daniel V. Xylobiontic beetle guild composition and diversity driven by forest canopy structure and management, by Jürgen Hagan and Ryszard Szadziewski Schmidl and Heinz Bussler Species richness and historical relations of arboreal phytophagous beetles - a study based on fogging samples 6 Canopy research and its impact on forestry and nature conservation from primeval forests of Poland, Romania and Slovenia (Chrysomeloidea, Curculionoidea), by Peter Sprick and Integrating tree crown science with the development of Andreas Floren ‘Near-to-Nature’ forest management practices: examples from Bavaria, by Ulrich Ammer, Martin Goßner, Axel Species list and feeding guilds of arboreal phytophagous Gruppe and Ulrich Simon beetles (Chrysomelidae, Curculionoidea) in Germany, by Peter Sprick Conservation of coppice with standards for canopy arthropods: the Bavarian Conservation Programme for Abundance and ordinal composition of arboreal arthropod Forests, by Alois Liegl and Matthias Dolek communities of various trees in old primary and managed forests, by Andreas Floren Conservation efforts and strategies for forest canopies in Germany: a review of conservation programmes, by Andreas Häusler, Matthias Dolek, Wolfram Güthler and Renate Market