Manuel J. Steinbauer

Late Quaternary climate change shapes island biodiversity

Island biogeographical models consider islands either as geologically static with biodiversity resulting from ecologically neutral immigration–extinction dynamics, or as geologically dynamic with biodiversity resulting from immigration–speciation–extinction dynamics influenced by changes in island characteristics over millions of years. Present climate and spatial arrangement of islands, however, are rather exceptional compared to most of the Late Quaternary, which is characterized by recurrent cooler and drier glacial periods. These climatic oscillations over short geological timescales strongly affected sea levels and caused massive changes in island area, isolation and connectivity, orders of magnitude faster than the geological processes of island formation, subsidence and erosion considered in island theory. Consequences of these oscillations for present biodiversity remain unassessed. Here we analyse the effects of present and Last Glacial Maximum (LGM) island area, isolation, elevation and climate on key components of angiosperm diversity on islands worldwide. We find that post-LGM changes in island characteristics, especially in area, have left a strong imprint on present diversity of endemic species. Specifically, the number and proportion of endemic species today is significantly higher on islands that were larger during the LGM. Native species richness, in turn, is mostly determined by present island characteristics. We conclude that an appreciation of Late Quaternary environmental change is essential to understand patterns of island endemism and its underlying evolutionary dynamics.

Oceanic island biogeography through the lens of the general dynamic model: Assessment and prospect

The general dynamic model of oceanic island biogeography (GDM) has added a new dimension to theoretical island biogeography in recognizing that geological processes are key drivers of the evolutionary processes of diversification and extinction within remote islands. It provides a dynamic and essentially non‐equilibrium framework generating novel predictions for emergent diversity properties of oceanic islands and archipelagos. Its publication in 2008 coincided with, and spurred on, renewed attention to the dynamics of remote islands. We review progress, both in testing the GDMs predictions and in developing and enhancing ecological–evolutionary understanding of oceanic island systems through the lens of the GDM. In particular, we focus on four main themes: (i) macroecological tests using a space‐for‐time rationale; (ii) extensions of theory to islands following different patterns of ontogeny; (iii) the implications of GDM dynamics for lineage diversification and trait evolution; and (iv) the potential for downscaling GDM dynamics to local‐scale ecological patterns and processes within islands. We also consider the implications of the GDM for understanding patterns of non‐native species diversity. We demonstrate the vitality of the field of island biogeography by identifying a range of potentially productive lines for future research.

Scale- and taxon-dependent patterns of plant diversity in steppes of Khakassia, South Siberia (Russia)

The drivers of plant richness at fine spatial scales in steppe ecosystems are still not sufficiently understood. Our main research questions were: (i) How rich in plant species are the natural steppes of Southern Siberia compared to natural and semi-natural grasslands in other regions of the Palaearctic? (ii) What are the main environmental drivers of the diversity patterns in these steppes? (iii) What are the diversity–environment relationships and do they vary between spatial scales and among different taxonomic groups? We sampled the steppe vegetation (vascular plants, bryophytes and lichens) in Khakassia (Russia) with 39 nested-plot series (0.0001–100-m2 plot size) and 54 additional 10-m2 quadrats across the regional range of steppe types and measured various environmental variables. We measured β-diversity using z-values of power-law species–area relationships. GLM analyses were performed to assess the importance of environmental variables as predictors of species richness and z-value. Khakassian steppes showed both high α- and β-diversity. We found significant scale dependence for the z-values, which had their highest values at small spatial scales and then decreased exponentially. Total species richness was controlled predominantly by heat load index, mean annual precipitation, humus content and soil skeleton content. The positive role of soil pH was evident only for vascular plant species richness. Similar to other studies, we found that the importance of environmental factors strongly differed among taxonomic groups and across spatial scales, thus highlighting the need to study more than one taxon and more than one plot size to get a reliable picture.

Optimizing sampling approaches along ecological gradients

Summary Natural scientists and especially ecologists use manipulative experiments or field observations along gradients to differentiate patterns driven by processes from those caused by random noise. A well-conceived sampling design is essential for identifying, analysing and reporting underlying patterns in a statistically solid and reproducible manner, given the normal restrictions in labour, time and money. However, a technical guideline about an adequate sampling design to maximize prediction success under restricted resources is lacking. This study aims at developing such a solid and reproducible guideline for sampling along gradients in all fields of ecology and science in general. We conducted simulations with artificial data for five common response types known in ecology, each represented by a simple function (no response, linear, exponential, symmetric unimodal and asymmetric unimodal). In the simulations, we accounted for different levels of random and systematic error, the two sources of noise in ecological data. We quantified prediction success for varying total sample size, number of locations sampled along a spatial/temporal gradient and number of replicates per sampled location. The number of replicates becomes more important with increasing random error, whereas replicates become less relevant for a systematic error bigger than 20% of total variation. Thus, if high levels of systematic error are indicated or expected (e.g. in field studies with spatial autocorrelation, unaccountable additional environmental drivers or population clustering), continuous sampling with little to no replication is recommended. In contrast, sampling designs with replications are recommended in studies that can control for systematic errors. In a setting that is characteristic for ecological experiments and field studies strictly controlling for undeterminable systematic error (random error ≥10% and systematic error ≤10% of total variation), prediction success was best for an intermediate number of sampled locations along the gradient (10–15) and a low number of replicates per location (3). Our findings from reproducible, statistical simulations will help design appropriate and efficient sampling approaches and avoid erroneous conclusions based on studies with flawed sampling design, which is currently one of the main targets of public criticism against science.

Homogenizing and diversifying effects of intensive agricultural land-use on plant species beta diversity in Central Europe — A call to adapt our conservation measures

The prevention of biodiversity loss in agricultural landscapes to protect ecosystem stability and functions is of major importance to stabilize overall diversity. Intense agriculture leads to a loss in species richness and homogenization of species pools, but the processes behind are poorly understood due to a lack of systematic case studies: The specific impacts by agriculture in contrast to other land-use creating open habitat are not studied as such landscapes hardly exist in temperate regions. Applying systematic grids, we compared the plant species distribution at the landscape scale between an active military training areas in Europe and an adjacent rather intensively used agricultural landscape. As the study areas differ mainly in the type of disturbance regime (agricultural vs. non-agricultural), differences in species pattern can be traced back more or less directly to the management. Species trait analyses and multiple measures of beta diversity were applied to differentiate between species similarities between plots, distance-decay, or nestedness. Contrary to our expectation, overall beta diversity in the agricultural area was not reduced but increased under agricultural. This was probably the result of species nestedness due to fragmentation. The natural process of increasing dissimilarity with distance (distance-decay) was suppressed by intense agricultural land-use, generalist and long-distance dispersers gained importance, while rare species lost continuity. There are two independent processes that need to be addressed separately to halt biodiversity loss in agricultural land. There is a need to conserve semi-natural open habitat patches of diverse size to favor poor dispersers and specialist species. At the same time, we stress the importance of mediating biotic homogenization caused by the decrease of distance-decay: The spread of long-distance dispersers in agricultural fields may be acceptable, however, optimized fertilizer input and erosion control are needed to stop the homogenization of environmental gradients due to nitrogen input into semi-natural habitat.

Influence of tree hollow characteristics on saproxylic beetle diversity in a managed forest

Tree hollows are key structures in forest ecosystems constituting long-lasting habitats and nutritional resources for many saproxylic arthropod species. Due to diverse microhabitat structures and conditions in tree hollows, they can support a broad range of species. However, in the past intensive management practices in parts of Europe reduced the abundance of tree hollows, resulting in a decrease and endangerment of species specialised in this tree habitat. We investigated 40 beech trees with hollows in 2014 and a subset of 23 of these trees in 2015 in a managed forest landscape in Germany. Using emergence traps we collected 89 beetle species of which 33% were on the Bavarian Red List. We described the tree characteristics, physical hollow characteristics, and their surrounding environment investigating their influence on α-diversity of non-Red List and Red List species. Furthermore, we investigated spatial (between tree hollows) and temporal (same tree hollow but different years) β-diversity, considering the importance of turnover and nestedness components on β-diversity. α-Diversity decreased with increasing decomposition of wood mould and increased with increasing area of hollow entrance in both years. Additional characteristics differed between years and between non-Red List and Red List species. β-Diversity was related to diameter at breast height, number of surrounding tree hollows, area of hollow entrance and a temperature gradient. We found a higher species turnover than nestedness between tree hollows and between years, indicating highly dynamic beetle communities spatially as well as temporally. To support and maintain the diversity of saproxylic beetles inhabiting tree hollows, the heterogeneity of microhabitats is important and should be supported by maintaining the diversity of differently structured and sized tree hollows.

Plant invasion and speciation along elevational gradients on the oceanic island La Palma, Canary Islands

Abstract Ecosystems that provide environmental opportunities but are poor in species and functional richness generally support speciation as well as invasion processes. These processes are expected not to be equally effective along elevational gradients due to specific ecological, spatial, and anthropogenic filters, thus controlling the dispersal and establishment of species. Here, we investigate speciation and invasion processes along elevational gradients. We assess the vascular plant species richness as well as the number and percentage of endemic species and non‐native species systematically along three elevational gradients covering large parts of the climatic range of La Palma, Canary Islands. Species richness was negatively correlated with elevation, while the percentage of Canary endemic species showed a positive relationship. However, the percentage of Canary–Madeira endemics did not show a relationship with elevation. Non‐native species richness (indicating invasion) peaked at 500 m elevation and showed a consistent decline until about 1,200 m elevation. Above that limit, no non‐native species were present in the studied elevational gradients. Ecological, anthropogenic, and spatial filters control richness, diversification, and invasion with elevation. With increase in elevation, richness decreases due to species–area relationships. Ecological limitations of native ruderal species related to anthropogenic pressure are in line with the absence of non‐native species from high elevations indicating directional ecological filtering. Increase in ecological isolation with elevation drives diversification and thus increased percentages of Canary endemics. The best preserved eastern transect, including mature laurel forests, is an exception. The high percentage of Canary–Madeira endemics indicates the cloud forests environmental uniqueness—and thus ecological isolation—beyond the Macaronesian islands.

Ecological Indicator Values of Europe (EIVE) 1.0: a powerful open-access tool for vegetation scientists

Background: Ecological indicator values (EIVs) have a long tradition in vegetation ecological research in Europe. EIVs characterise the ecological optimum of species along major environmental gradients using ordinal scales. Calculating mean indicator values per plot is an effective way of bioindication. Following first systems in Russia and Central Europe, about two dozen EIV systems have been published for various parts of Europe. Aims: As there was no EIV system available at European scale that could be used for broad- scale analyses, e.g. in the context of the European Vegetation Archive (EVA), we develop such a system for the first time for the vascular plants of Europe. Location: Europe. Methods: We compiled all national and major regional EIV systems and harmonized their plant nomenclature with a newly developed contemporary European taxonomic backbone (EuroSL 1.0). Using regression, we rescaled the individual EIV systems for the main parameters to continent-wide quasi-metric scales, ranging from 1 to 99. The data from each individual system were then translated into a probability curve approximated with a normal distribution, weighed with the logarithm of the area represented and summed up across the systems. From the European density curve we extracted then a mean and a variance, which characterise the distribution of this species along this particular ecological gradient. Results and conclusions: Our consensus approach of integrating the expert knowledge of all existing EIV systems allowed deriving the first consistent description of the ecological behaviour for a significant part of the European vascular flora. The resulting Ecological Indicator Values of Europe (EIVE) 1.0 will be published open access to allow bioindication beyond country borders. Future releases of EIVE might contain more parameters, non- vascular plants and regionalisation or could be re-adjusted and extended to hitherto non- covered species through co-occurrence data from EVA.

Biogeography, Patterns in

Biogeographic patterns result from environmental influences interacting with historic legacies and biotic characteristics. The emergence of biogeographic patterns is often scale dependent and the identification of causal processes is difficult due to complex cross-scale interactions. Prominent biogeographic patterns emerge particularly along strong environmental gradients such as latitude and elevation (species richness, range size, body size, coloration) or under isolated conditions like on islands (island gigantism/dwarfism, island woodiness, and dispersal loss). Historic legacies (such as colonization progression rules) or repeated evolutionary patterns (taxon cycle) may influence current distribution patterns. Yet, global patterns in species traits or growth forms can be clearly associated with specific environmental conditions (e.g., giant rosette plants, trait variability in Solanum ).

A second horizon scan of biogeography: Golden Ages, Midas touches, and the Red Queen

Are we entering a new ‘Golden Age’ of biogeography, with continued development of infrastructure and ideas? We highlight recent developments, and the challenges and opportunities they bring, in light of the snapshot provided by the 7th biennial meeting of the International Biogeography Society (IBS 2015). We summarize themes in and across 15 symposia using narrative analysis and word clouds, which we complement with recent publication trends and ‘research fronts’. We find that biogeography is still strongly defined by core sub-disciplines that reflect its origins in botanical, zoological (particularly bird and mammal), and geographic (e.g., island, montane) studies of the 1800s. That core is being enriched by large datasets (e.g. of environmental variables, ‘omics’, species’ occurrences, traits) and new techniques (e.g., advances in genetics, remote sensing, modeling) that promote studies with increasing detail and at increasing scales; disciplinary breadth is being diversified (e.g., by developments in paleobiogeography and microbiology) and integrated through the transfer of approaches and sharing of theory (e.g., spatial modeling and phylogenetics in evolutionary–ecological contexts). Yet some subdisciplines remain on the fringe (e.g., marine biogeography, deep-time paleobiogeography), new horizons and new theory may be overshadowed by popular techniques (e.g., species distribution modelling), and hypotheses, data, and analyses may each be wanting. Trends in publication suggest a shift away from traditional biogeography journals to multidisciplinary or open access journals. Thus, there are currently many opportunities and challenges as biogeography increasingly addresses human impacts on, and stewardship of, the planet (e.g., Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services). As in the past, biogeographers doubtless will continue to be engaged by new data and methods in exploring the nexus between biology and geography for decades into the future. But golden ages come and go, and they need not touch every domain in a discipline nor affect subdisciplines at the same time; moreover, what appears to be a Golden Age may sometimes have an undesirable ‘Midas touch’. Contexts within and outwith biogeography—e.g., methods, knowledge, climate, biodiversity, politics—are continually changing, and at times it can be challenging to establish or maintain relevance. In so many races with the Red Queen, we suggest that biogeography will enjoy greatest success if we also increasingly engage with the epistemology of our discipline.