Under natural conditions beech forest would be the most widespread habitat in Central Europe. Therefore the knowledge of this vegetation type, including the animal communities dwelling there, as well as the mechanisms forming these communities are of high interest for biodiversity research and conservation in Central Europe. However, historically, Central European beech forests were thought to harbor only relatively few species. Here, faunistic data of five beech forest reserves, which were generated as part of the Hessian strict forest reserves program, are analyzed to investigate diversity patterns of Hessian beech forests of low mountain ranges. We focus on species-level data from six organism groups: Aculeata, Araneae, Coleoptera, Heteroptera, Lumbricidae, and Macrolepidoptera. We show that with 2552 forest-dwelling species from these organism groups an unexpectedly high number of species was found in Hessian beech forests, and that a higher species richness can be expected in these groups. Observed species diversity ranges from 1245 to 1556 forest dwelling species in the five individual forest reserves. Overall, 36 % of the forest dwelling species of the considered species groups known from Germany were found with all applied methods in the forest reserves. Different forest reserves share 40-50 % of the species, representing 50-70 % of the species of the individual reserves, indicating high levels of regional and habitat-structure based differentiation. Only 21 % of the species are found in all five reserves. The low percentage of species found in all five reserves is, in addition to differing local conditions, probably a result of the communities being composed of few highly abundant and many rare species, leading to a high percentage of species only found by chance in our surveys. We also observe differences in community heterogeneity among the five reserves. Patterns differ between organism groups, clearly indicating that a focus on single taxa or a single indicator group falls short of revealing meaningful patterns. In spider communities, beta diversity is linked to the spatial distance between traps. In other organism groups community heterogeneity within reserves rather depends on structural heterogeneity. Species richness was associated with percentage of reserve area not covered with the most dominant habitat type, the deadwood amount, and with survey year. Being the potential natural vegetation of Germany and considering the unexpectedly high diversity of their associated fauna, beech forests bear a great conservation value. However, their widespread occurrence and dominance is likely to push them out of focus of conservation efforts. Yet protecting diverse and richly structured beech forests can contribute greatly to preserving the native arthropod fauna and should play a central role in biodiversity conservation efforts in Central Europe.
Since 2011 a list of the forest affinities of all forest species of vascular plants, bryophytes, and ferns known from Germany, is available. Recently, the first comparable lists of the forest affinities of animal species were published. These lists include a categorization of forest affinities for all species of Crassiclitellata: Lumbricidae (earth worms), Araneae (spiders), Opiliones (harvestmen), Pseudoscorpiones (pseudoscorpions), Hemiptera: Heteroptera (true bugs), Coleoptera (beetles), Hymenoptera: Aculeata (ants, bees, wasps), Lepidoptera: Macrolepidoptera (butterflies, moths partly) and Aves (birds) known from Germany. The lists are a valuable resource that have the potential to add new layers of discovery to analyses of forest animal communities. Here we present a summary of the lists. We also re-analyze existing data sets in light of forest affinities to highlight possible applications of these lists in ecological assessments and characterizations of animal communities in central European forests. We show that the forest affinity lists can add valuable new insights for analyzing community structure, temporal community dynamics or communities across ecological gradients, represent an important tool for developing conservation strategies, and can be used to clear a dataset of chance findings of allochtonous specimens prior to analysis.
Spiders (Araneae) of the Strict Forest Reserve “Goldbachs- und Ziebachsrück” (Hesse, Germany). Investigation period 1994-1996 A total of 166 spider species (24 % of the species known from Hesse) was recorded in the investigation area, with 148 in the Strict Forest Reserve and 143 in the managed sites. The recorded spiders are considered to document the fauna of the area representatively and nearly completely. One species, Improphantes nitidus THORELL, 1875 (Linyphiidae, 1 ♀), was found for the first time in Hesse. Two species which are very rarely recorded in Germany have been found: Gongylidiellum edentatum MILLER, 1951 (Linyphiidae, 3 ♂♂, 44 ♀♀) and Pseudomaro aenigmaticus DENIS, 1966 (Linyphiidae, 1 ♀). Gongylidiellum edentatum is regarded as a potential indicator for historic old forests. The composition of the spider fauna according to the species’ frequency, habitat requirements, stratum preference, size classes, geographical distribution, altitudinal distribution, phenology and degree of endangerment (6 species are listed in the German Red Data Book) is analysed. The similarity of species assemblages obtained by comparable methods is analysed and visualised using an ordination technique of non-metric multidimensional scaling (NMS). The results are compared mainly with those from other Strict Forest Reserves in Hesse, as only these have been investigated using analogous methodology. The Strict Forest Reserve Goldbachs- und Ziebachsrück supports a typical spider coenosis and, compared with the other Hessian Strict Forest Reserves, a moderately high number of spider species. The spider fauna is dominated by typical forest species (66 % of the species, 84 % of the individuals); species of open habitats have only minor importance. Due to the presence of spring and brook areas, species of wet and humid habitats (21 %, 18 %) are slightly better represented than species of dry habitats (22 %, 10 %), which occur on the warm edges and borders. Predominantly tree-living (arboricolous) spiders constitute 24 % of the species and 28 % of the individuals, respectively. 8 % of the species have a restricted distribution within Europe (covering clearly less than half of the area of Europe). Altogether 48 % of the species occur only in Europe and adjacent areas, 52 % have at least a Palaearctic distribution. Only 8 % of the species (5 % of the individuals) are restricted to climax habitats, 67 % (80 %) also live in semi-natural habitats, the remaining 25 % (15 %) can also be found in disturbed and artificial synanthropic habitats. The spider fauna of the Strict Forest Reserve and the managed sites do not differ significantly in any of these aspects. The highest numbers of species were recorded with the eclectors on living and dead standing beeches (118 species) and with the pitfall traps (100 species). The dominant species among the 18,081 adult spiders collected were Amaurobius fenestralis (STRÖM, 1768) (Amaurobiidae, 17 %), Walckenaeria cuspidata BLACKWALL, 1833 (Linyphiidae, 13 %) and Diplocephalus cristatus (BLACKWALL, 1833) (Linyphiidae, 6 %), which were predominantly caught in the eclectors, and Coelotes terrestris (WIDER, 1834) (Amaurobiidae, 7 %) and Tapinocyba pallens (O. P.-CAMBRIDGE, 1872) (Linyphiidae, 6 %), both predominantly caught in the pitfall traps. Altogether 35 spider species are discussed in detail, phenology of the 28 most abundant species (≥ 100 animals) is illustrated by graphs, and distribution maps for Germany are presented for 8 species. This investigation yielded additional knowledge on the ecology and phenology of numerous spider species, e. g. in revealing that epigeic species such as T. pallens also occurred in large numbers on the bark of tree trunks. The results highlight the conspicuously high biodiversity of indigenous forests. Research was conducted in cooperation with and financially supported by “Landesbetrieb Hessen-Forst”.
The spider fauna of four Strict Forest Reserves in Hesse (Germany) is analysed in respect to characterisations of the recorded species: frequency in Germany, total distribution range, summarised types of habitats. The data are compared with seven (mostly unpublished) projects in SE- and NE-Germany with a comparable species number. The number of species per project ranges from 159 to 205. In the four Strict Forest Reserves in Hesse a total of 278 has been recorded. Higher numbers of species occur on the trunk than on the ground. Rare spiders are more frequent in forests than in open land. Spider species with the most restricted distributions are best represented in forests of low mountain ranges in Hesse and Bavaria. Even a rough classification of preferred habitats yields valuable information. The biodiversity of spiders in commonly occurring types of forests is much higher than expected.
The spider fauna active on the bark of trees in forests on eight sites in different regions in Germany was investigated. Trunk eclectors at about 2-4 meter height on living trees were used in different regions of Germany (SW Bavaria, Hesse, Brandenburg) in the last two decades. In Hesse eclectors were also used on dead beech trees (standing and lying). Data, mainly from beech (Fagus sylvatica) and spruce (Picea abies), from May to October are compared – only data for Hesse are available for the whole year (including winter). A total of 334 spider species were been recorded with these bark traps, i.e. about one third of the spider species known from Germany. On average each of the eight regions yielded 140.5 (± 26.2) species, each single tree 40.5 (± 12.2) species and 502 (± 452) adult spiders per season (i.e. May to Oct.). The 20 most abundant species are listed and characterised (totals, exclusivity/regional focus, tendency to conifer/broadleaf, notes). Six of the 20 species were not known as abundant bark species, 3 prefer conifers and 3 beech/broadleaf. Even in winter (December-March) there is remarkably high activity on the trunks. But only a few species occur exclusively or mainly in the winter. Finally, the rarity of some bark spider species is discussed and details (all known records in Germany, phenology) of four of them are presented (Clubiona leucaspis, Gongylidiellum edentatum, Kratochviliella bicapitata, Oreonetides quadridentatus). The diversity and importance of the spider fauna on bark in Central Europe is still underestimated.
Introduction and Methodology For all species of Lumbricidae (earth worms), Araneae (spiders), Opiliones (daddy long-legs), Pseudoscorpiones (pseudoscorpions), Heteroptera (true bugs), Coleoptera (beetles), Aculeata (ants, bees and wasps), Macrolepidoptera (moths and butterflies) and Aves (birds) present in Germany we present lists summarizing their affinities to forest habitats. w: strong affinity to forest habitats, without known preference for light or closed forests wg: mainly found in forests, with strong affinity to closed forest habitats wl: mainly found in forests, with strong affinity to open forests, forest edges or glades m: occurring in both open landscapes and forest habitats, but without a preference for for-est habitats (assignment to mm or mo not possible) mm: occurring equally in open landscapes and forest habitats mo: strong affinity to open landscapes, but also regularly occurring in forests, at for-est edges or in glades o: only occurring in open landscapes or other habitats without forest cover like caves or buildings u: unknown The selected animal groups cover the most important taxa of forest biocoenoes according to Winter et al. (1999) and in total 25 % of the 46.500 species known from Germany. Summary of the forest affinity of animal species of Germany About one fourth of Germany’s animal species are classified into eight categories based on their affinity to forest habitats. These species comprise the most important groups for biodi-versity monitoring in forests. The classification is centered on the association with forest habitats, i. e., species categorized as w, wg, wl mainly occur in forests, and species catego-rized as m, mm, mo occur in forests and open landscapes but without a preference for for-est habitats. Species categorized as ‘o’ exclusively live in open habitats. It is important to note that both ‘w*’ and ‘m*’ species are typical inhabitants of forests. Only taxa categorized as ‘o’ should be considered as nonresident species in forest habitats. A prolonged pres-ence of large numbers of ‘o’-species point to disturbance of the forest habitat. Therefore, authors should consider excluding ‘o’-species from analyses concentrated on forest biodi-versity. The presented classification is a valuable tool for distinction between members of the forest biocoenoses and accidental finds of open-land-species and to compare results from differ-ent forests concerning the degree of specialization of the species. Based on the presented list, more than one third of the species in Germany live exclusively in open habitats, another 15 % live mainly in open habitats, but are also regularly found in forest habitats. 15 % are more or less equally distributed in open and forested habitats. Species of open light forest and forest margins represent 10 % of the fauna, while very few species are restricted to closed dense forests (less than 1 %). The large number of mo-species (16.9 %) and wl-species (9.8 %) is conspicuous. These are animals, which in forested areas prefer light forests, glades and/or forest margins. The high proportion of mo and wl species points to the well known fact, that many arthropods prefer sunny, dry and warm habitats. Because forestry today creates relatively dense and uniformly structured stands (concerning tree species and tree age), the conservation of species of light forests, glades and/or forest margins presents an important aim opportunity for sustainable forest management. Generally our results demonstrate, that management of open landscapes (especially agri-cultural areas, urban and industrial wastelands) is the most important task in nature conser-vation. The large number of species living as well in open as in forested areas points to the collective responsibility of forestry and agriculture. But because of the intensification of agri-culture in the last decades, this part is transmitted mainly to forestry. For 26.1 % of the spe-cies only forestry is responsible.
In hessischen Naturwaldreservaten werden im Zuge der zoologischen Begleitforschung seit 2002 Fledermäuse untersucht. Neben einem vereinfachten Erfassungsprogramm aus akustischen Erhebungen und Netzfängen wurden in 2015 drei Naturwaldreservate intensiver hinsichtlich der Artenvielfalt und Aktivitätsdichte in den Totalreservaten und den bewirtschafteten Vergleichsflächen analysiert. In den 14 untersuchten hessischen Naturwaldreservaten konnten auf einer Gesamtfläche von 878 ha (0,02 % der Landesfläche Hessens) 17 Fledermausarten gefunden, was > 90% aller in Hessen vorkommenden Arten entspricht. Dabei zeigte sich eine Abhängigkeit der Artenzahl und Anzahl reproduktiver Arten von der Temperatur und Höhenlage des NWR. So steigt die Artenvielfalt als auch die Anzahl reproduzierender Arten mit Zunahme der Jahresdurchschnittstemperatur und mit Abnahme der Höhenlage. Klima und Höhenlage sind dabei korrespondierende Faktoren. Bei gleicher Waldstruktur beeinflussen mildere klimatische Bedingungen das verfügbare Nahrungsangebot, die Phänologie der Beutetiere sowie deren nächtliche Aktivität günstig. Innerhalb der NWR konnte eine signifikant höhere Artendiversität und –aktivität im Totalreservat als in der bewirtschafteten Vergleichsflächen nachgewiesen werden. Die eingestellte Bewirtschaftung auf den Flächen des Totalreservats hat einen günstigen Effekt auf das Vorkommen von Fledermäusen.
The fauna of the Strict Forest Reserve ”Kinzigaue“ (Hesse, Germany) was investigated over a period of two years using diverse techniques (pitfall traps; different types of eclectors on tree trunks, stumps, dead branches; blue, white and yellow pans; window traps; hand sampling, observations, light traps and red wine bait for moths, traps for bark beetles and glue rings, population density mapping for birds). All material was sorted to order and is preserved in a permanent sample bank. Annelida, Araneae, Heteroptera, Coleoptera (volume 2), Aculeata (volume 2), Macrolepidoptera and Aves were analysed completely at species level. Their ecology is discussed in detail. For further groups determination lists are compiled. A complete list of all determined species is presented at the end of the second volume of this monograph. The reserve is situated in the lowland area of southern Hesse, in the lowlands of the Upper Rhine and Rhine-Main area about 12 km east of the town Hanau at 105–110 m a.s.l. (coordinates 8° 59’ E, 50° 9’ N). The mean annual temperature is 9.6° C, mean annual precipitation 712 mm. The investigated area has a size of 18.1 ha, was declared as Natural Forest Reserve in 1993 (without an unmanaged site for comparison) and has also the status as a nature reserve. Soil: silty loam on a Holocene floodplain ground. The potential natural plant cover is an alder-elm floodplain forest. The forest is dominated by the common oak (Quercus robur), other frequent trees are lime (Tilia sp.), hornbeam (Carpinus betulus), ash (Fraxinus excelsior), and maple (Acer sp.). During the investigations the dominating old oak stands had an age of 161 to 206 years. Research was conducted in cooperation with and financially supported by “Landesbetrieb Hessen-Forst”.
Senckenberg Gesellschaft für Naturforschung, Germany Since 1970, strict forest reserves have been established all over Germany to gain a spectrum of total reserves, which allow the development of "primeval forests of tomorrow" and are available for research and as reference areas for silviculture. The first Hessian reserves were established in 1988; today there exist 31 which cover 1200ha. 22 have adjacent managed sites for comparison. The reserves represent over all altitude zones and geological landscapes the spectrum of forest types in Hesse: mainly beechforests, but also oak, pine and spruce forests. The forestal and botanical research is conducted by the "Northwest-German Forestry Research Station, Göttingen", the zoological investigations by Senckenberg. Research is conducted in cooperation with and financially supported by "Landesbetrieb Hessen-Forst". A broad set of traps is used: pitfall traps, eclectors at living or dead (standing or lying) trunks, stubs and dead branches, window traps and blue, yellow and white pan traps. The traps are in use continuously over two years. Additionally, hand sampling, light trapping, and bait trapping for Macrolepidoptera and population density mapping for birds is conducted. All material is sorted to order. Seven standard groups are studied: Lumbricidae, Araneae, Heteroptera, Coleoptera, Aculeata, Macrolepidoptera, Aves. Additionally, other animal groups – as many as possible – are studied by honorary collaborators (ATBI-approach). For each reserve a monograph is published with extensive reports on the standard groups and species lists for all determined species.The fauna in a Central European beech forest, which had been managed lately, is with 5000–6000 species 2–3 times more speciouse than expected. Especially open structures provide habitats for many species. Specialists of old trees and deadwood are underrepresented. Biodiversity assessments have to be conducted on species level and with detailed analyses of the species' ecological requirements. Numerous new findings in the fields of ecology and faunistics were acquired.
the strict forest reserve Kinzigaue more than 950,000 animals were trapped in a period of two years using a broad set of trap types and specific hand sampling. A wide spectrum of animal groups was determined to species level, representing 27.1 % of the German fauna. In total 1,923 species were determined, which by extrapolation indicates that more than 6,500 species occur in the area. A hitherto undescribed species of Bethylidae is recorded, one fly species (Diptera) is new to Germany and 22 species new to Hesse: 14 beetles (Coleoptera), 4 bees and wasps (Hymenoptera), 2 true bugs (Heteroptera), 1 lacewing (Neuroptera), 1 barklouse (Psocoptera). 238 species are included in the Red Data Lists of Germany or Hesse. Most of the endangered species are Coleoptera and Lepidoptera. The reserve Kinzigaue is the first oak dominated, first inundated and first lowland forest investigated in the Strict Forest Reserve program in Hesse. For several groups of animals (spiders, true bugs, beetles, butterflies and moths, pseudoscorpions, woodlice) Kinzigaue until now is the most species rich forest reserve, partly at a clear distance to the beech forests in the low mountain range, which had been studied previously. Research was conducted in cooperation with and financially supported by “Landesbetrieb Hessen-Forst”.
In a long-term survey of the Strict Forest Reserves in Hesse (Central Germany) a faunistic inventory was conducted using a wide range of traps and hand sampling over a period of two years. Five of the six sites are beech forest types (dominating: Fagus sylvatica), and one is a floodplain oak forest (dominating: Quercus robur). Although no special research program was run for the earthworms, pitfall traps and eclectors at tree trunks and logs provided an unexpectedly high number (9 to 13) of species of earthworms. A comparison of data from pitfall traps and trunk eclectors shows that highly different sets of lumbricids do occur in both types. In the pitfall traps Lumbricus rubellus (22–43 % of adults) and Dendrobaena octaedra (10–16 % of adults) were dominant. In the trunk eclectors the community consisted mainly (60–83 %) of Allolobophoridella eiseni. In addition, Dendrodrilus rubidus (3–25 %) and Dendrobaena octaedra (6–16 %) were regularly found in these eclectors. Based on our findings Allolobophoridella eiseni is a regular element of the invertebrate fauna of forests, occurring both in decaying logs and predominantly at the bark of living and dead trees. Therefore, this species can be classified as predominantly corticolous. For the first time it could be proven that Allolobophoridella eiseni regularly climbs on trees, which it does surprisingly mainly in late-autumn and in the winter.
The Heteroptera fauna of the Strict Forest Reserve ” Kinzigaue “ (Hesse, Germany) was investigated with a broad spectrum of methods (pitfall traps, blue, white and yellow pans, window traps, different types of eclectors at tree trunks, stumps, dead branches, hand sampling). In total 7.880 specimens of 124 species of Heteroptera of 23 families were collected. This equals 13.9 % of Germany's and 17.6 % of the Hessian true bug fauna in an area of only 18.1 ha. This is the largest amount of species so far collected in all five Hessian strict forest reserves and four managed areas for comparison. Statistics (Jackknife 1 and 2) calculate 162 to 190 as the total number of species. Xyloecocoris ovatulus is reported for the first time for Hesse and the lost species Brachynotocoris puncticornis is reported again for this state.