Höhengradienten in der Biodiversitäts-Forschung: eine Übersicht unter Berücksichtigung des Klimawandels

Mountains, with their isolated position and altitudinal belts, are hotspots of biodiversity. Their flora and fauna have been observed worldwide since the days of Alexander von Humboldt, which has led to basic knowledge and understanding of species composition and the most important driving forces of ecosystem differentiation in such altitudinal gradients. Systematically designed analyses of changes in species composition with increasing elevation have been increasingly implemented since the 1990s. Since global climate change is one of the most important problems facing the world this century, a focus on such ecosystem studies is urgently needed. To identify the main future needs of such research we analyze the studies dealing with species changes of diverse taxonomical groups along altitudinal gradients (0 to 6,400 m a.s. l.) on all continents, published during the past one to two decades. From our study we can conclude that although mountains are powerful for climate change research most studies have to face the challenge of separating confounding effects driving species assemblages along altitudinal gradients. Our study therefore supports the view of the need of a global altitudinal concept including that (1) not only one or a few taxonomical groups should be analyzed, but rather different taxonomical groups covering all ecosystem functions simultaneously; (2) relevant site conditions should be registered to reveal direct environmental variables responsible for species distribution patterns and to resolve inconsistent effects along the altitudinal gradients; (3) transect design is appropriate for analyzing ecosystem changes in site gradients and over time; (4) both the study design and the individual methods should be standardized to compare the data collected worldwide; and (5) a long-term perspective is important to quantify the degree and direction of species changes and to validate species distribution models. (6) Finally we suggest to develop experimental altitudinal approaches to overcome the addressed problems of biodiversity surveys.Gebirge mit ihrer mehr oder weniger isolierten Lage und ihren vielfältigen Höhenstufen stellen „Hotspots der Biodiversität“ dar. Ihre Flora und Fauna wird seit den ersten Beschreibungen durch Alexander von Humboldt immer wieder untersucht. Systematisch angelegte Studien zur räumlichen Verteilung von Arten bzw. Artengruppen in Höhengradienten erlangen eine zunehmend größere Bedeutung seit den 1990iger Jahren, insbesondere im Rahmen der Klimafolgenforschung. Um für die zukünftige Biodiversitätsforschung die heute als wesentlich angesehenen Forschungsansätze zu identifizieren, wurden in dieser Studie Literaturangaben der letzten zwei Jahrzehnte über die sich verändernden Artenzusammensetzungen in Höhengradienten aus allen Kontinenten, verschiedenen Artengruppen und einem Höhenbereich von 0 bis 6.400 m ü. NN ausgewertet. Diese Zusammenstellung zeigt, dass Untersuchungen der Muster der Biodiversität von Höhengradienten in Gebirgen einen wichtigen Beitrag für das Verständnis der Herausbildung und Änderung von Biodiversitätsmustern gerade im Rahmen des zu erwartenden Klimawandels leisten können. Die Musterbildung im Höhengradienten ist für verschiedene Artengruppen durchaus unterschiedlich. Diese Literaturauswertung macht aber auch die Notwendigkeit eines weltweit einheitlichen Konzeptes für die Forschung in Höhengradienten deutlich. Ein solches Konzept sollte folgende Gesichtspunkte beinhalten: (1) Nicht nur ausgewählte Artengruppen, sondern mehrere verschiedene Artengruppen gemeinsam analysieren; nur so lassen sich die vielfältigen Ökosystemfunktionen und Prozesse berücksichtigen. (2) Begleitend möglichst viele Umweltfaktoren erfassen, um herausarbeiten zu können, welche der Umweltfaktoren für die Musterbildung wirklich maßgeblich sind. (3) Die Transekte so anlegen, dass sie nicht nur eine einmalige räumliche Analyse ermöglichen, sondern auch zukünftig im Sinne von Zeitreihen zur Verfügung stehen. (4) Sowohl das Erhebungsflächendesign als auch die Aufnahmemethoden für die einzelnen Artengruppen standardisieren, um die Daten weltweit vergleichen zu können. (5) Langzeitstudien starten; denn nur sie bieten die Möglichkeit, erwartete oder aus Modellen abgeleitete Veränderungen der Artenzusammensetzungen bzw. von Verschiebungen von einzelnen Arten im Höhengradienten zu validieren. (6) Gezielt konzipierte Experimente einsetzen, um offene Fragen zur Verteilung der Arten in Höhengradienten untersuchen zu können

The BIOKLIM project: biodiversity research between climate change and wilding in a temperate montane forest : the conceptual framework

To understand the rapid rate of change in global biodiversity, it is necessary to analyse the present condition of ecosystems and to elucidate relationships of species to their environment. The BIOKLIM Project (Biodiversity and Climate Change Project) is intended to close this gap in our knowledge of montane and high montane forests of Central European low mountain ranges, one of the most threatened mixed montane systems worldwide. The Bavarian Forest National Park is characterised by its altitude range of ca. 800 m and a strongly developed gradient of forest structure. Relicts of old growth forests (areas of former local nature reserves) and dead stands, mostly killed by bark beetles, are accompanied by widely varying levels of woody debris and light. The gradients comprise a wide range of abiotic and forest structure factors, making the study area well suited for a multidisciplinary investigation of biodiversity. Unconstrained ordination (CA) of six taxa (vascular plants, wood inhabiting fungi, birds, carabids, spiders and molluscs) indicate the altitudinal gradient to be the main driver for distribution patterns of species assemblages. Objectives, structure, study design and data sampling of the BIOKLIM Project are described in detail. We set up 293 sampling plots along four main straight transects following the altitudinal gradient. All abiotic and stand structure data regarded as relevant are available for each plot. Vascular plants, wood inhabiting fungi and birds were sampled or mapped on all 293 plots. For the other 22 investigated taxa we used subsamples pre-stratified according to the sampling methods. The necessity of dealing with spatial autocorrelation, arising from sampling along linear transects, is described. Finally, study approach of our biodiversity project is compared with others involving altitudinal gradients. Worldwide, only a few multidisciplinary biodiversity studies have been previously conducted on long altitudinal gradients. However, in most cases sampling techniques were similar to ours, which allows comparison of results between continents. Keywords: Climate Change, Biodiversity, species-environment relationshipsUm die rasante Veränderung globaler Biodiversität zu verstehen, ist es erforderlich, den gegenwärtigen Zustand von Ökosystemen zu analysieren und die Zusammenhänge zwischen Arten und deren Umwelt aufzulösen. Das BIOKLIMProjekt (Biodiversität und Klima Projekt) hat zum Ziel, diese Wissenslücken für Wälder montaner und hochmontaner Mittelgebirge zu schließen. Der Nationalpark Bayerischer Wald ist neben dem Höhengradient (ca. 800 m) durch einen starken Strukturgradient geprägt. Dieser resultiert aus Restvorkommen sehr alter Bestände (ehem. Naturschutzgebiete) sowie dem Wirken des Borkenkäfers seit ca. zwei Jahrzehnten und einem dadurch verbundenen z. T. sehr hohen Totholzvorrat. Die Gradienten umfassen eine breite Spanne von abiotischen Faktoren und Bestandesstrukturen und machen den Nationalpark zu einem gut geeigneten Untersuchungsgebiet für interdisziplinäre Biodiversitätsforschung. Korrespondenzanalysen (CA) für 6 taxonomische Gruppen (Gefäßpflanzen, Holzpilze, Vögel, Laufkäfer, Spinnen und Mollusken) machen die starke Abhängigkeit der Artengruppen vom Höhengradienten deutlich. Es werden detailliert die Zielsetzungen, Projektaufbau, das Untersuchungsdesign sowie die Erfassungsmethoden des BIOKLIM-Projektes beschrieben. 293 Probepunkte wurden entlang von 4 Transekten, welche dem Höhengradienten folgen, eingerichtet. Zu jedem Probekreis stehen alle als relevant erachteten Daten zur Abiotik und Bestandesstruktur zur Verfügung. Gefäßpflanzen, Holzpilze und Vögel wurden auf allen 293 Probepunkten erfasst. Für die anderen 22 untersuchten Artengruppen wurde in Abhängigkeit von der Methode ein stratifiziertes Design gewählt. Lösungsansätze zum Umgang mit Autokorrelation, die durch die Anordnung von Probekreisen entlang von Linien (Transekte) bedingt ist, werden dargestellt. Schließlich wird das BIOKLIM-Projekt mit den wenigen weltweiten Biodiversitätsprojekten verglichen und diskutiert. In den meisten Fällen sind die Erhebungsmethoden ähnlich, sodass Vergleiche der Ergebnisse zwischen verschiedenen Kontinenten möglich werden. Schlüsselwörter: Klimawandel, Biodiversität, Arten-Umwelt-Beziehun

Epiphytic lichens indicating process protection and ecological continuity in the Bavarian Forest National Park

Lichens are one of the prominent taxonomical indicator groups for changes in forests. Within the course of the research project BIOKLIM, we sampled 125 epiphytic lichens, 5 lichenicolous fungi, and 1 non-lichenized ascomycete in wilding areas, managed areas, and relicts of old-growth forests along four transects in the Bavarian Forest National Park. Our analyses showed that old-growth relicts and wilding areas have a significantly higher diversity and a higher number of red-listed species than managed forests, demonstrating the importance of these areas for the protection and dispersal of endangered lichens and lichenicolous fungi. The rare and threatened lichenicolous fungi Chaenothecopsis viridialba, Sphinctrina anglica, and Sphinctrina turbinata and the critically threatened lichen Mycobilimbia sphaeroides were new records for the Bavarian Forest. The lichen-fungus Sphinctrina tubiformis, presumed extinct in Germany, was recovered in the old-growth relict Rachelseeurwald. The following species were classified as indicators of old, nearnatural forests of the Bavarian Forest: Lecanactis abietina, Pyrenula nitida, Micarea cinerea, Microcalicium disseminatum, Loxospora cismonica, and Thelotrema lepadinum. We propose management implications for the managed areas of montane and high montane forests in Central Europe.Flechten gehören zu den herausragenden Indikatorgruppen für Veränderungen in Wäldern. Im Rahmen des Forschungsprojektes „Biodiversität und Klima“ (BIOKLIM) wurden im Jahr 2007 im Nationalpark Bayerischer Wald entlang von 4 Forschungslinien 125 baumbewohnende Flechten, 5 lichenicole Pilze sowie 1 nicht lichenisierter Ascomycet festgestellt. Alte Wälder und Prozessschutzwälder wiesen dabei signifikant die größte Artendiversität und die höchste Anzahl an bedrohten Arten im Vergleich mit den noch gemanagten Wäldern auf. Die herausragende Bedeutung von alten Wäldern und Prozessschutzflächen für den Erhalt und die Ausbreitung gefährdeter Flechten und lichenicoler Pilze für den ostbayerischen Mittelgebirgsraum wird dargestellt. Die sehr seltenen und hochgradig gefährdeten lichenicolen Pilze Chaenothecopsis viridialba, Sphinctrina anglica und Sphinctrina turbinata und die Flechte Mycobilimbia sphaeroides werden als Neufunde für den Bayerischen Wald genannt. Der in der Bundesrepublik Deutschland als verschollen geltende Flechtenpilz Sphinctrina tubiformis wurde im „Rachelseeurwald“ wiederentdeckt. Für alte naturnahe Wälder des Bayerischen und Böhmischen Waldes werden folgende Zeigerarten beschrieben: Lecanactis abietina, Pyrenula nitida, Micarea cinerea, Microcalicium disseminatum, Loxospora cismonica und Thelotrema lepadinum. Abschließend werden für das Management montaner und hochmontaner Wälder Mitteleuropas Empfehlungen abgeleitet

Insects Overshoot the Expected Upslope Shift Caused by Climate Warming

Along elevational gradients, climate warming may lead to an upslope shift of the lower and upper range margin of organisms. A recent meta-analysis concluded that these shifts are species specific and considerably differ among taxonomic lineages. We used the opportunity to compare upper range margins of five lineages (plants, beetles, flies, hymenoptera, and birds) between 1902-1904 and 2006-2007 within one region (Bavarian Forest, Central Europe). Based on the increase in the regional mean annual temperature during this period and the regional lapse rate, the upslope shift is expected to be between 51 and 201 m. Averaged across species within lineages, the range margin of all animal lineages shifted upslope, but that of plants did not. For animals, the observed shifts were probably due to shifts in temperature and not to changes in habitat conditions. The range margin of plants is therefore apparently not constrained by temperature, a result contrasting recent findings. The mean shift of birds (165 m) was within the predicted range and consistent with a recent global meta-analysis. However, the upslope shift of the three insect lineages (>260 m) exceeded the expected shift even after considering several sources of uncertainty, which indicated a non-linear response to temperature. Our analysis demonstrated broad differences among lineages in their response to climate change even within one region. Furthermore, on the considered scale, the response of ectothermic animals was not consistent with expectations based on shifts in the mean annual temperature. Irrespective of the reasons for the overshooting of the response of the insects, these shifts lead to reorganizations in the composition of assemblages with consequences for ecosystem processes

Traits and phylogenies modulate the environmental responses of wood-inhabiting fungal communities across spatial scales

Identifying the spatial scales at which community assembly processes operate is fundamental for gaining a mechanistic understanding of the drivers shaping ecological communities. In this study, we examined whether and how traits and phylogenetic relationships structure fungal community assembly across spatial scales. We applied joint species distribution modelling to a European-scale dataset on 215 wood-inhabiting fungal species, which includes data on traits, phylogeny and environmental variables measured at the local (log-level) and regional (site-level) scales. At the local scale, wood-inhabiting fungal communities were mostly structured by deadwood decay stage, and the trait and phylogenetic patterns along this environmental gradient suggested the lack of diversifying selection. At regional scales, fungal communities and their trait distributions were influenced by climatic and connectivity-related variables. The fungal climatic niches were not phylogenetically structured, suggesting that diversifying selection or stabilizing selection for climatic niches has played a strong role in wood-inhabiting communities. In contrast, we found a strong phylogenetic signal in the responses to connectivity-related variables, revealing phylogenetic homogenization in small and isolated forests. Synthesis. Altogether, our results show that species-level traits and phylogenies modulate the responses of wood-inhabiting fungi to environmental processes acting at different scales. This result suggests that the evolutionary histories of fungal traits diverge along different environmental axes.Peer reviewe

Seven Micarea (Pilocarpaceae) species new to Germany and notes on deficiently known species in the Bavarian Forest

We report new records of 19, predominantly rare, Micarea species, mostly from dead wood in mixed montane forests characterized mainly by Norway spruce, European beech and silver fir in the Bavarian Forest National Park on the German-Czech border. Their ecology and key morphological features are discussed. Micarea contexta, M. fallax, M. melanobola, M. pseudomicrococca, M. pusilla, M. soralifera and M. tomentosa are reported for the first time from Germany. Micarea anterior, M. byssacea, M. elachista, M. laeta, M. micrococca and M. nowakii, in addition to the aforementioned, are reported as new for the Bavarian Forest National Park.We report new records of 19, predominantly rare, Micarea species, mostly from dead wood in mixed montane forests characterized mainly by Norway spruce, European beech and silver fir in the Bavarian Forest National Park on the German-Czech border. Their ecology and key morphological features are discussed. Micarea contexta, M. fallax. M. melanobola, M. pseudomicrococca, M. pusilla, M. soralifera and M. tomentosa are reported for the first time from Germany. Micarea anterior, M. byssacea, M. elachista, M. laeta, M. rnicrococca and M. nowakii, in addition to the aforementioned, are reported as new for the Bavarian Forest National Park.Peer reviewe

50 Years of Cumulative Open-Source Data Confirm Stable and Robust Biodiversity Distribution Patterns for Macrofungi

Fungi are a hyper-diverse kingdom that contributes significantly to the regulation of the global carbon and nutrient cycle. However, our understanding of the distribution of fungal diversity is often hindered by a lack of data, especially on a large spatial scale. Open biodiversity data may provide a solution, but concerns about the potential spatial and temporal bias in species occurrence data arising from different observers and sampling protocols challenge their utility. The theory of species accumulation curves predicts that the cumulative number of species reaches an asymptote when the sampling effort is sufficiently large. Thus, we hypothesize that open biodiversity data could be used to reveal large-scale macrofungal diversity patterns if these datasets are accumulated long enough. Here, we tested our hypothesis with 50 years of macrofungal occurrence records in Norway and Sweden that were downloaded from the Global Biodiversity Information Facility (GBIF). We first grouped the data into five temporal subsamples with different cumulative sampling efforts (i.e., accumulation of data for 10, 20, 30, 40 and 50 years). We then predicted the macrofungal diversity and distribution at each subsample using the maximum entropy (MaxEnt) species distribution model. The results revealed that the cumulative number of macrofungal species stabilized into distinct distribution patterns with localized hotspots of predicted macrofungal diversity with sampling efforts greater than approximately 30 years. Our research demonstrates the utility and importance of the long-term accumulated open biodiversity data in studying macrofungal diversity and distribution at the national level.publishedVersio

Increasing disturbance demands new policies to conserve intact forest

Ongoing controversy over logging the ancient Białowieża Forest in Poland symbolizes a global problem for policies and management of the increasing proportion of the earth's intact forest that is subject to postdisturbance logging. We review the extent of, and motivations for, postdisturbance logging in protected and unprotected forests globally. An unprecedented level of logging in protected areas and other places where green‐tree harvest would not normally occur is driven by economic interests and a desire for pest control. To avoid failure of global initiatives dedicated to reducing the loss of species, five key policy reforms are necessary: (1) salvage logging must be banned from protected areas; (2) forest planning should address altered disturbance regimes for all intact forests to ensure that significant areas remain undisturbed by logging; (3) new kinds of integrated analyses are needed to assess the potential economic benefits of salvage logging against its ecological, economic, and social costs; (4) global and regional maps of natural disturbance regimes should be created to guide better spatiotemporal planning of protected areas and undisturbed forests outside reserves; and (5) improved education and communication programs are needed to correct widely held misconceptions about natural disturbances.We thank the Bavarian Forest National Park administration for funding the travel costs of RN and DL in support of this manuscript. ABL acknowledges support from grants P12-RNM-2705 and FJCI- 2015–23687

Identification of factors influencing the Puumala virus seroprevalence within its reservoir in aMontane Forest Environment.

Puumala virus (PUUV) is a major cause of mild to moderate haemorrhagic fever with renal syndrome and is transmitted by the bank vole (Myodes glareolus). There has been a high cumulative incidence of recorded human cases in South-eastern Germany since 2004 when the region was first recognized as being endemic for PUUV. As the area is well known for outdoor recreation and the Bavarian Forest National Park (BFNP) is located in the region, the increasing numbers of recorded cases are of concern. To understand the population and environmental effects on the seroprevalence of PUUV in bank voles we trapped small mammals at 23 sites along an elevation gradient from 317 to 1420m above sea level. Generalized linear mixed effects models(GLMEM) were used to explore associations between the seroprevalence of PUUV in bank voles and climate and biotic factors. We found that the seroprevalence of PUUV was low (6%-7%) in 2008 and 2009, and reached 29% in 2010. PUUV seroprevalence was positively associated with the local species diversity and deadwood layer, and negatively associated with mean annual temperature, mean annual solar radiation, and herb layer. Based on these findings, an illustrative risk map for PUUV seroprevalence prediction in bank voles was created for an area of the national park. The map will help when planning infrastructure in the national park (e.g., huts, shelters, and trails)