Article

The list of forest affinities for animals in Central Europe – a valuable resource for ecological analysis and monitoring in forest animal communities?

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Abstract

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.

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... Only species categorized as forest dwellers by Dorow et al. (2019) (categories: f, fc, fl, m, mm, mo (see Schneider et al. (2021) for an assignment of English and German abbreviations)) are included in the analyses. Species dwelling exclusively in open habitats (category: o) are considered as chance findings of transient specimens and therefore excluded from the analyses (Dorow et. ...
... Species dwelling exclusively in open habitats (category: o) are considered as chance findings of transient specimens and therefore excluded from the analyses (Dorow et. al. 2019, Schneider et al. 2021). ...
Article
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.
... Pararge aegeria is usually considered a forest (edge) species with a widespread distribution across Europe (van Swaay et al. 2006;Schneider et al. 2021), but during the last decades, it has-at least in North-West Europe-successfully expanded its ecological niche and habitat use into more open, agricultural environments (e.g., Vandewoestijne and Van Dyck 2010). Butterflies of agricultural and forest ecotypes (i.e., landscapes with fragmented vs. connected resource distributions, respectively) have been shown to differ in the way they interact with their environment. ...
Article
Local adaptation, early-developmental and behavioral plasticity have all been suggested to be underlying mechanisms of behav-ioral variation. They allow organisms to cope with resource heterogeneity in time and space. However, the relative contribution of each of these drivers on niche-expansion success is still unclear. We addressed this issue by studying oviposition behavior in a butterfly (Pararge aegeria) that used to be confined to forest, but recently colonized anthropogenic areas too. We caught females of either forest or agricultural ecotype and allowed their larval offspring to experience early-habitat cues in either open field or woodland conditions. For 110 females of this transplant experiment, we monitored all oviposition-related behaviors (i.e., site selection, searching, motivation, learning) using repeated trials in outdoor flight cages with naturally occurring host grasses. We observed changes in oviposition site preference, with increasing preference for semi-shaded host grass (by vegetation or vertical structures) over consecutive egg-lays. Motivation to oviposit was most affected by larval experience (in interaction with cage environment). We also show evidence for ecotypic differentiation with the agricultural ecotype showing reduced search effort during the first trial compared to the forest ecotype. We argue that females of agricultural landscapes with fragmented resource distributions adopt a more risk-aversive oviposition strategy. Our detailed behavioral tracking under ecologically relevant conditions, stresses the importance of considering different types of behavioral plasticity to unravel a species' success during niche expansion.
... The list was subsequently improved and supplemented with lists of forest-dwelling bryophyte and lichen species by Schmidt et al. (2011). More recently, a list of selected groups of forest animals has also been published for Germany, using a slightly modified approach (Dorow et al., 2019, see also Schneider et al., 2021). ...
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Question When evaluating forests in terms of their biodiversity, distinctiveness and naturalness, the affinity of the constituent species to forests is a crucial parameter. Here we ask to what extent are vascular plant species associated with forests, and does species’ affinity to forests vary between European regions? Location Temperate and boreal forest biome of Northwestern and Central Europe. Methods We compiled EuForPlant, a new extensive list of forest vascular plant species in 24 regions spread across 13 European countries using vegetation databases and expert knowledge. Species were region‐specifically classified into four categories reflecting the degree of their affinity to forest habitats: 1.1, species of forest interiors; 1.2, species of forest edges and forest openings; 2.1, species that can be found in forest as well as open vegetation; and 2.2, species that can be found partly in forest, but mainly in open vegetation. An additional “O” category was distinguished, covering species typical for non‐forest vegetation. Results EuForPlant comprises 1,726 species, including 1,437 herb‐layer species, 159 shrubs, 107 trees, 19 lianas and 4 epiphytic parasites. Across regions, generalist forest species (with 450 and 777 species classified as 2.1 and 2.2, respectively) significantly outnumbered specialist forest species (with 250 and 137 species classified as 1.1 and 1.2, respectively). Even though the degree of shifting between the categories of forest affinity among regions was relatively low (on average, 17.5%), about one‐third of the forest species (especially 1.2 and 2.2) swapped categories in at least one of the study regions. Conclusions The proposed list can be used widely in vegetation science and global change ecology related to forest biodiversity and community dynamics. Shifting of forest affinity among regions emphasizes the importance of a continental‐scale forest plant species list with regional specificity.
... We addressed the issue of oviposition site selection, microhabitat exploration and related experience-induced changes in the speckled wood butterfly, Pararge aegeria. This is usually considered a forest (edge) species (Schneider et al. 2021), but in northwestern Europe its ecological niche has expanded from forest to anthropogenic environments over the last few decades (Vandewoestijne & Van Dyck, 2010). This expansion is assumed to explain its regional increase in distribution and abundance under rapid human-induced environmental change (Van Dyck et al. 2009). ...
Article
Organisms that colonize new habitats may gain fitness advantages from changes in habitat selection including oviposition behaviour. Variation in oviposition site selection may contribute to ecological differentiation in response to novel ecological conditions. Both inherited and environmentally induced or learned differences may contribute to variation in preoviposition search behaviour and oviposition site selection. We addressed this issue with the speckled wood butterfly, Pararge aegeria. Over recent decades, this species has expanded its habitat use by colonizing anthropogenic environments with populations that show differences in several phenotypic traits (i.e. ecotypes). Here we tested whether preoviposition search behaviour and oviposition site selection differ between field-derived females from forested and agricultural ecotypes, and whether prior experience would change their behaviour differently. We used F1-offspring reared under common garden laboratory conditions and observed individuals of both ecotypes repeatedly in an indoor experimental arena. The arena offered compartments with host plants under simulated open and woody landscape conditions, as well as control compartments. In contrast with predictions, ecotypic differences in preference were found not for a particular microhabitat, but rather for oviposition site versus control compartment. Nevertheless, exploration and microhabitat use differed between females of agricultural and forest ecotypes. Furthermore, we provide evidence of short-term (i.e. a few hours after initial exposure) learning in the context of preoviposition search behaviour, pointing at the often ignored role of habitat familiarization. Our study sheds new light on behavioural changes that relate to habitat use in novel environments for a species that thrives successfully in landscapes under rapid human-induced environmental change.
... Das Insektensterben macht auch vor dem Wald nicht halt: In Deutschland nahmen in Forstbeständen in drei untersuchten Regionen zwischen 2008 und 2017 die Artenzahl und die Biomasse an Arthropoden ab(Seibold et al. 2019). Wenn auch der größere Teil bedrohter Arten verschiedener Arthropodengruppen im Offenland zu finden ist, so gibt es unter den Rote-Liste-Arten auch an Wald gebundene Arten, für die die Forstwirtschaft eine besondere Verantwortung hat(Schneider et al. 2021). ...
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The massive outbreaks of the Gypsy Moth in South Hesse in 1993/94 revisited (Lymantria dispar, Lepidoptera: Erebidae, Lymantriinae) Abstract: In 1993 and 1994, massive outbreaks of the Gypsy Moth in South Hesse forests were treated with Dimilin and Bacillus thuringiensis kurstaki on large areas by airplane or helicopter. We surveyed the nocturnal Lepidoptera fauna with light traps in oak stands (mainly Quercus robur) over two years. There was a clear decrease of the counts of species as well as individuals over the two years (year of poisoning and following year) especially in the Dimilin area, and especially for species living as caterpillars in canopies or bushes. No such decrease was observed in small untreated comparative stands within the treatmet areas in spite of full defoliation. Similar results were found in surveys in Bavaria and Baden-Wurttemberg by other observers. We describe the present situation on basis of acute climate change, strong general decrease of insect populations and oak forest (the latter mainly caused by drought) and demand that any poison treatment should only be done by respecting the actual administrative regulations and never without a regular long-term monitoring of the effects on non-target organisms and efficiency of the measures.
... The Speckled wood (Pararge aegeria L.) is a frequently used species for the study of ecology and evolution in insects (Watt & Boggs, 2003). This woodland and woodland edge butterfly is widespread throughout Europe (Kudrna et al., 2011;Schneider et al., 2021) and in the majority of its range, it flies in two, or even more, generations a year depending on its flexible life cycle regulation (e.g. Wiklund & Friberg, 2011). ...
Article
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The main objective of the European Union (EU) funded project AQEM1was to develop a framework of an assessment system for streams in Europe based on benthic macroinvertebrates that fulfils the requirements of the EU Water Framework Directive. Initial assessment methods for 28 European stream types and more generally applicable tools for stream biomonitoring in Europe were generated. The development of the system was based on a newly collected data set covering stream types in Austria, the Czech Republic, Germany, Greece, Italy, The Netherlands, Portugal and Sweden. Altogether, 901 benthic invertebrate samples were taken using a standardised multi-habitat sampling procedure and a large number of parameters describing the streams and their catchments was recorded for all sampling sites. From the stream and catchment characteristics measures of stress were derived. A large number of metrics was tested independently for each of the stream types, to identify the response of each metric to degradation of a site. This process resulted in up to 18 core metrics for the individual stream types, which were combined into a different multimetric index in each country. The multimetric AQEM assessment system is used to classify a stream stretch into an Ecological Quality Class ranging from 5 (high quality) to 1 (bad quality) and often provides information on the possible causes of degradation. AQEM provides a taxa list of 9557 European macroinvertebrate taxa with associated autecological information, a software package for performing all the calculations necessary for applying the multimetric AQEM assessment system and a manual describing all aspects of the application of the system from site selection to data interpretation.
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The Rothamsted Insect Survey has operated a Great Britain-wide network of light-traps since 1968. From these data we estimated the first ever national abundance indices and 35-year population trends for 338 species of common macro-moths. Although the number of trap sites which run each year is not constant, there is a representative, well-distributed core of traps that have run for 15years. The proportion of operating sites catching a species and the annual geometric mean catch of successful traps were used to provide estimates of species range and absolute abundance. T, an index of long-term population trends, was used to compare trends among species. T was not biased by trap site turnover. The percentage of species displaying significant decreases (54%) was more than double that displaying increases (22%). Species found throughout Great Britain are decreasing most rapidly in the south and especially the southeast but species with a southerly distribution are increasing. Results of a preliminary overview suggest habitat and climate change may both play a role in changing species dynamics. The existence of estimates of abundances and trends for such a large species pool opens the way for much further research, linking trends with land-use changes, climate change and inter-specific dynamics.
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During the last 50 years, agricultural intensification has caused many wild plant and animal species to go extinct regionally or nationally and has profoundly changed the functioning of agro-ecosystems. Agricultural intensification has many components, such as loss of landscape elements, enlarged farm and field sizes and larger inputs of fertilizer and pesticides. However, very little is known about the relative contribution of these variables to the large-scale negative effects on biodiversity. In this study, we disentangled the impacts of various components of agricultural intensification on species diversity of wild plants, carabids and ground-nesting farmland birds and on the biological control of aphids.
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The changes in population size of carabid beetles inhabiting an ancient woodland were analysed in a long-term investigation covering a period of nine years (1994–2002). The amplitude of fluctuations of the relative density of these species was estimated and compared with those of carabid beetles inhabiting recent woodlands and more unstable habitats in open landscapes. To analyse this, 8 pitfall traps were set in an ancient beech and oak woodland in the nature reserve “Lüneburger Heide” in northern Germany and were emptied every fortnight the whole year round.Catching rates for some species (e.g. Carabus problematicus and Abax parallelepipedus) fluctuated only slightly, whereas those of other species (e.g. C. violaceus and C. auronitens) varied as much as ten-fold. Comparison of these results with those of other long-term investigations of recent woodlands and also of more open habitats showed that in each of the three habitat types some species varied very little in abundance and others, very widely. So it seems that the amplitude of fluctuations in abundance is a feature of each single species rather than a special attribute of their habitats.
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Agricultural dynamics and associated changes in the structure of habitat patches affect species composition and distribution in the landscape. Land-use, landscape changes and vegetation changes of weeds were analysed in a 4 km2 area in Central Germany (Saxony-Anhalt) from 1953 to 2000. This period includes the collectivisation (1952–1968), the agricultural industrialisation (1969–1989) and the privatisation of agricultural land following the political changes in East Germany in 1990. For the analyses, historic and current aerial photographs and vegetation data were used. Landscape indices and the average amount of mineral fertilizers were used as indicators for landscape structure and land-use intensity. Intensification of agriculture and the collectivisation in East Germany in the fifties and sixties led to a decline of the spatial heterogeneity of the landscape matrix (arable fields). The average number and cover of weed species, especially archaeophytes, decreased significantly since 1957. However, the total number of weed species increased. There was a remarkably high number of species with an average cover below 0.05%, called “chance” species in 2000. Out of 17 tested landscape indices only mean patch size and mean patch fractal dimension were significantly correlated with the average number of weed species. The average amount of the mineral fertilizer potash used as land-use intensity indicator was significantly negatively correlated with the total number of weed species. However, there was an increase in the number of farms after 1990 without changes in landscape structure and arable weed vegetation. The results suggest that structural variability of the landscape and habitat quality are the principal correlates of plant species diversity.
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Development of farming practices has caused drastic changes in European agricultural landscapes during the past 50 years. As a consequence of these changes, insect diversity is widely expected to decline. We performed a comparative analysis with long-term data of three insect groups: Auchenorrhyncha, Heteroptera and Orthoptera. In 2009, we revisited nine grassland sites in northern Germany that were originally sampled in 1951 using the same techniques and during a similar time frame. We found that the insect community exhibited no consistent trends between years. Species richness of Auchenorrhyncha and Heteroptera increased on plot level as well as on landscape level but remained unchanged for Orthoptera. Abundance of Auchenorrhyncha and Orthoptera significantly decreased, while Heteroptera increased. There is a strong trend towards homogeneity in community composition for Heteroptera and a weak one for Auchenorrhyncha. The frequency and abundance of species preferring disturbed and/or eutrophic habitats increased, whereas the number of species preferring low-productive habitats declined. This trend is especially pronounced in Auchenorrhyncha. Generalistic species were more abundant in relative proportions as well as in absolute numbers. We hypothesize that these trends arise from alterations of Central European landscapes because of agricultural intensification over the last several decades.
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Aim To determine whether arthropod richness and abundance for combined taxa, feeding guilds and broad taxonomic groups respond in a globally consistent manner to a range of agricultural land-use and management intensification scenarios. Location Mixed land-use agricultural landscapes, globally. Methods We performed a series of meta-analyses using arthropod richness and abundance data derived from the published literature. Richness and abundance were compared among land uses that commonly occur in agricultural landscapes and that represent a gradient of increasing intensification. These included land-use comparisons, such as wooded native vegetation compared with improved pasture, and a management comparison, reduced-input cropping compared with conventional cropping. Data were analysed using three different meta-analytical techniques, including a simple vote counting method and a formal fixed-effects/random-effects meta-analysis. Results Arthropod richness was significantly higher in areas of less intensive land use. The decline in arthropod richness was greater between native vegetation and agricultural land uses than among different agricultural land uses. These patterns were evident for all taxa combined, predators and decomposers, but not herbivorous taxa. Overall, arthropod abundance was greater in native vegetation than in agricultural lands and under reduced-input cropping compared with conventional cropping. Again, this trend was largely mirrored by predators and decomposers, but not herbivores. Main conclusions The greater arthropod richness found in native vegetation relative to agricultural land types indicates that in production landscapes still containing considerable native vegetation, retention of that vegetation may well be the most effective method of conserving arthropod biodiversity. Conversely, in highly intensified agricultural landscapes with little remaining native vegetation, the employment of reduced-input crop management and the provision of relatively low-intensity agricultural land uses, such as pasture, may prove effective in maintaining arthropod diversity, and potentially in promoting functionally important groups such as predators and decomposers.
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