Stefan G. Michalski

Contrasting changes in taxonomic, phylogenetic and functional diversity during a long-term succession: insights into assembly processes

Summary 1. Theory predicts that the processes generating biodiversity after disturbance will change during succession. Comparisons of phylogenetic and functional (alpha and beta) diversity with taxonomic diversity can provide insights into the extent to which community assembly is driven by deterministic or stochastic processes, but comparative approaches have yet to be applied to successional systems. 2. We characterized taxonomic, phylogenetic and functional plant (alpha and beta) diversity within and between four successional stages in a > 270-year-long arable-to-grassland chronosequence. Null models were used to test whether functional and phylogenetic turnover differed from random expectations, given the levels of species diversity. 3. The three facets of diversity showed different patterns of change during succession. Between early and early-mid succession, species richness increased but there was no increase in functional or phylogenetic diversity. Higher than predicted levels of functional similarity between species within the early and early-mid successional stages, indicate that abiotic filters have selected for sets of functionally similar species within sites. Between late-mid and late succession, there was no further increase in species richness, but a significant increase in functional alpha diversity, suggesting that functionally redundant species were replaced by functionally more dissimilar species. Functional turnover between stages was higher than predicted, and higher than within-stage turnover, indicating that different assembly processes act at different successional stages. 4. Synthesis. Analysis of spatial and temporal turnover in different facets of diversity suggests that deterministic processes generate biodiversity during post-disturbance ecosystem development and that the relative importance of assembly processes has changed over time. Trait-mediated abiotic filtering appears to play an important role in community assembly during the early and early-mid stages of arable-to-grassland succession, whereas the relative importance of competitive exclusion appears to have increased towards the later successional stages. Phylogenetic diversity provided a poor reflection of functional diversity and did not contribute to inferences about underlying assembly processes. Functionally deterministic assembly suggests that it may be possible to predict future post-disturbance changes in biodiversity, and associated ecosystem attributes, on the basis of species’ functional traits but not phylogeny.

Daphne: a dated phylogeny of a large European flora for phylogenetically informed ecological analyses

This data set represents a comprehensive, dated phylogeny of a large European flora comprising the vascular plants of the British Isles, Germany, The Netherlands, and Switzerland, totaling 4685 species. The phylogeny thus encompasses all species in the trait databases BIOLFLOR, PLANTATT, and BioBase 2003. The topology of the phylogentic tree is based on a backbone family phylogeny of the Angiosperm Phylogeny Group III. Subsequently, partial phylogenetic subtrees derived from a total of 518 recent molecular studies were manually pruned onto the backbone tree, using multi-gene consensus topologies if possible. Similarly, 1103 internal nodes and the root node were dated based on 261 recent studies. Finally, an ultrametric tree was calculated by placing undated nodes evenly between dated nodes. The phylogeny provides a reference data set for comparative analyses of trait correlations, trait evolution, trait based ecological processes, community assembly, or other phylogenetically informed analyses across a la...

Designing forest biodiversity experiments: general considerations illustrated by a new large experiment in subtropical China

Summary 1. Biodiversity–ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25� 8 9 25� 8m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial ‘ecoscape’ to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achiev

Mycorrhizas in the Central European flora: relationships with plant life history traits and ecology

Plant traits have been widely used to characterize different aspects of the ecology of plant species. Despite its wide distribution and its proven significance at the level of individuals, communities, and populations, the ability to form mycorrhizal associations has been largely neglected in these studies so far. Analyzing plant traits associated with the occurrence of mycorrhizas in plants can therefore enhance our understanding of plant strategies and distributions. Using a comparative approach, we tested for associations between mycorrhizal status and habitat characteristics, life history traits, and plant distribution patterns in 1752 species of the German flora (a major part of the Central European flora). Data were analyzed using log-linear models or generalized linear models, both accounting for phylogenetic relationships. Obligatorily mycorrhizal (OM) species tended to be positively associated with higher temperature, drier habitats, and higher pH; and negatively associated with moist, acidic, and fertile soils. Competitive species were more frequently OM, and stress tolerators were non-mycorrhizal (NM), while ruderal species did not show any preference. Facultatively mycorrhizal (FM) species showed the widest geographic and ecological amplitude. Indigenous species were more frequently FM and neophytes (recent aliens) more frequently OM than expected. FM species differed markedly from OM and NM species in almost all analyzed traits. Specifically, they showed a wider geographic distribution and ecological niche. Our study of the relationships between mycorrhizal status and other plant traits provides a comprehensive test of existing hypotheses and reveals novel patterns. The clear distinction between FM and OM + NM species in terms of their ecology opens up a new field of research in plant-mycorrhizal ecology.

High selfing and high inbreeding depression in peripheral populations of Juncus atratus

The mating system of a plant is the prime determinant of its population genetic structure. However, mating system effects may be modified by postzygotic mechanisms like inbreeding depression. Furthermore, historical as well as contemporary ecological factors and population characteristics, like the location within the species range can contribute to genetic variability. Using microsatellite markers we assessed the population genetic structure of the wind‐pollinated Juncus atratus in 16 populations from peripheral and nearly central areas of the distribution range and studied the mating system of the species. In three peripheral populations, outcrossing rates at seeds stage were low (mean tm = 5.6%), suggesting a highly autogamous mating system. Despite this fact, on adult stage both individual heterozygosity (mean HO = 0.48) and gene diversity (mean HE = 0.58) were high even in small populations. Inbreeding coefficients were consistently low among all populations (mean FIS = 0.15). Within the three peripheral populations indirect estimates of lifetime inbreeding depression were surprisingly high (δeq = 0.96) and inbreeding depression could be shown to act mostly on early seedling establishment. Similar conditions of autogamy combined with high inbreeding depression are typical for plants with a large lifetime genomic mutation rate that cannot avoid selfing by geitonogamy. However, the results presented here are unexpected for small‐statured, herbaceous plants. Substantial genetic differentiation among all populations was found (mean FST = 0.24). An isolation‐by‐distance pattern was apparent on large scale but not on local scale suggesting that the overall pattern was largely influenced by historical factors, e.g. colonization, whereas locally genetic drift was of greater importance than gene flow. Peripheral populations exhibited lower genetic diversity and higher inbreeding coefficients when compared with subcentral populations.

Biodiversity Promotes Tree Growth during Succession in Subtropical Forest

Losses of plant species diversity can affect ecosystem functioning, with decreased primary productivity being the most frequently reported effect in experimental plant assemblages, including tree plantations. Less is known about the role of biodiversity in natural ecosystems, including forests, despite their importance for global biogeochemical cycling and climate. In general, experimental manipulations of tree diversity will take decades to yield final results. To date, biodiversity effects in natural forests therefore have only been reported from sample surveys or meta-analyses with plots not initially selected for diversity. We studied biomass and growth of subtropical forests stands in southeastern China. Taking advantage of variation in species recruitment during secondary succession, we adopted a comparative study design selecting forest plots to span a gradient in species richness. We repeatedly censored the stem diameter of two tree size cohorts, comprising 93 species belonging to 57 genera and 33 families. Tree size and growth were analyzed in dependence of species richness, the functional diversity of growth-related traits, and phylogenetic diversity, using both general linear and structural equation modeling. Successional age covaried with diversity, but differently so in the two size cohorts. Plot-level stem basal area and growth were positively related with species richness, while growth was negatively related to successional age. The productivity increase in species-rich, functionally and phylogenetically diverse plots was driven by both larger mean sizes and larger numbers of trees. The biodiversity effects we report exceed those from experimental studies, sample surveys and meta-analyses, suggesting that subtropical tree diversity is an important driver of forest productivity and re-growth after disturbance that supports the provision of ecological services by these ecosystems.

Pollination mode and life form strongly affect the relation between mating system and pollen to ovule ratios.

Pollen to ovule (P : O) ratios have been hypothesized to correlate with the degree of outcrossing and thus with the mating system of a plant. Also, P : O ratios are likely to vary with respect to pollination mode (i.e. wind pollination or animal pollination). Furthermore, constraints on the evolution of mating systems depending on life form may affect P : O ratios. We compiled P : O ratios and outcrossing rates for 107 angiosperm species and analyzed the relation between these traits considering pollination mode, life form and phylogenetic relatedness among species. In general, P : O ratios correlated significantly with outcrossing rates. However, when taking additional factors into account, the relation became ambiguous. The correlation was significantly positive in wind-pollinated species, but only marginally so in animal-pollinated species. Wind-pollinated species had higher P : O ratios than animal-pollinated taxa. In woody perennials, outcrossing was the predominant mating system and outcrossing rates did not correlate with P : O ratios. The results were not altered by accounting for phylogenetic relatedness among species. The results indicate that P : O ratios vary more strongly with pollination mode and life form than with the mating system.

Genetic differentiation and regional adaptation among seed origins used for grassland restoration: lessons from a multispecies transplant experiment

Summary One of the key questions in ecosystem restoration is the choice of seed material for restoring plant communities. More and more scientists and practitioners are currently advocating the use of regional seed sources, based on the argument that plants are often adapted to local or regional environmental conditions, and thus, regional seed sources should provide the best restoration success. However, there is still substantial debate about this approach, partly because of a lack of solid empirical data. We conducted a multispecies transplant experiment in which we compared the performance of eight seed origins of seven plant species frequently used in grassland restoration in four common gardens across Germany. We found that, on average, plants of regional origins produced 10% more inflorescences and 7% more biomass than those of foreign origins. There were substantial differences among species in the strength of these effects, but in the majority of the study species fitness decreased with increasing geographical distance of seed origins or with increasing climatic differences between plant origins and experimental sites. In addition to these effects on plant fitness, increasing geographical or climatic distances of origin were often also correlated with increasing differences in plant phenology. Since phenology is important for biotic interactions, especially with pollinators and seed predators, using foreign seed sources may have cascading effects on local ecosystems. Synthesis and applications. Genetic differentiation is widespread in grassland species and often shows the patterns of regional adaptation. Our study thus supports the use of regional seed sources in restoration. Moreover, using non-regional seed sources in grassland restoration may not only decrease the performance of plants, but it will likely also affect their biotic interactions.

Isolation by Elevation: Genetic Structure at Neutral and Putatively Non-Neutral Loci in a Dominant Tree of Subtropical Forests, Castanopsis eyrei

Background The distribution of genetic diversity among plant populations growing along elevational gradients can be affected by neutral as well as selective processes. Molecular markers used to study these patterns usually target neutral processes only, but may also be affected by selection. In this study, the effects of elevation and successional stage on genetic diversity of a dominant tree species were investigated controlling for neutrality of the microsatellite loci used. Methodology/Principal Findings Diversity and differentiation among 24 populations of Castanopsis eyrei from different elevations (251–920 m) and successional stages were analysed by eight microsatellite loci. We found that one of the loci (Ccu97H18) strongly deviated from a neutral model of differentiation among populations due to either divergent selection or hitchhiking with an unknown selected locus. The analysis showed that C. eyrei populations had a high level of genetic diversity within populations (AR = 7.6, HE = 0.82). Genetic variation increased with elevation for both the putatively selected locus Ccu97H18 and the neutral loci. At locus Ccu97H18 one allele was dominant at low elevations, which was replaced at higher elevations by an increasing number of other alleles. The level of genetic differentiation at neutral loci was similar to that of other Fagaceae species (FST = 0.032,  = 0.15). Population differentiation followed a model of isolation by distance but additionally, strongly significant isolation by elevation was found, both for neutral loci and the putatively selected locus. Conclusions/Significance The results indicate higher gene flow among similar elevational levels than across different elevational levels and suggest a selective influence of elevation on the distribution of genetic diversity in C. eyrei. The study underlines the importance to check the selective neutrality of marker loci in analyses of population structure.

Wolbachia Infections Mimic Cryptic Speciation in Two Parasitic Butterfly Species, Phengaris teleius and P. nausithous (Lepidoptera: Lycaenidae)

Deep mitochondrial divergence within species may result from cryptic speciation, from phylogeographic isolation or from endosymbiotic bacteria like Wolbachia that manipulate host reproduction. Phengaris butterflies are social parasites that spend most of their life in close relationship with ants. Previously, cryptic speciation has been hypothesised for two Phengaris species based on divergent mtDNA sequences. Since Phengaris species are highly endangered, the existence of cryptic species would have drastic consequences for conservation and management. We tested for cryptic speciation and alternative scenarios in P. teleius and P. nausithous based on a comprehensive sample across their Palaearctic ranges using COI gene sequences, nuclear microsatellites and tests for Wolbachia. In both species a deep mitochondrial split occurring 0.65–1.97 myrs ago was observed that did not correspond with microsatellite data but was concordant with Wolbachia infection. Haplotypes previously attributed to cryptic species were part of the Wolbachia-infected clades. In both species remaining phylogeographic structure was largely consistent between mitochondrial and nuclear genomes. In P. teleius several mitochondrial and nuclear groups were observed in East Asia while a single haplogroup and nuclear cluster prevailed across continental Eurasia. Neutrality tests suggested rapid demographic expansion into that area. In contrast, P. nausithous had several mitochondrial and nuclear groups in Europe, suggesting a complex phylogeographic history in the western part of the species range. We conclude that deep intraspecific divergences found in DNA barcode studies do not necessarily need to represent cryptic speciation but instead can be due to both infection by Wolbachia and phylogeographic structure.