Michael Kleyer

Predictors of species sensitivity to fragmentation

We reviewed empirical data and hypotheses derived from demographic, optimal foraging, life-history, community, and biogeographic theory for predicting the sensitivity of species to habitat fragmentation. We found 12 traits or trait groups that have been suggested as predictors of species sensitivity: population size; population fluctuation and storage effect; dispersal power; reproductive potential; annual survival; sociality; body size; trophic position; ecological specialisation, microhabitat and matrix use; disturbance and competition sensitive traits; rarity; and biogeographic position. For each trait we discuss the theoretical justification for its sensitivity to fragmentation and empirical evidence for and against the suitability of the trait as a predictor of fragmentation sensitivity. Where relevant, we also discuss experimental design problems for testing the underlying hypotheses. There is good empirical support for 6 of the 12 traits as sensitivity predictors: population size; population fluctuation and storage effects; traits associated with competitive ability and disturbance sensitivity in plants; microhabitat specialisation and matrix use; rarity in the form of low abundance within a habitat; and relative biogeographic position. Few clear patterns emerge for the remaining traits from empirical studies if examined in isolation. Consequently, interactions of species traits and environmental conditions must be considered if we want to be able to predict species sensitivity to fragmentation. We develop a classification of fragmentation sensitivity based on specific trait combinations and discuss the implications of the results for ecological theory.

The global spectrum of plant form and function

Earth is home to a remarkable diversity of plant forms and life histories, yet comparatively few essential trait combinations have proved evolutionarily viable in today’s terrestrial biosphere. By analysing worldwide variation in six major traits critical to growth, survival and reproduction within the largest sample of vascular plant species ever compiled, we found that occupancy of six-dimensional trait space is strongly concentrated, indicating coordination and trade-offs. Three-quarters of trait variation is captured in a two-dimensional global spectrum of plant form and function. One major dimension within this plane reflects the size of whole plants and their parts; the other represents the leaf economics spectrum, which balances leaf construction costs against growth potential. The global plant trait spectrum provides a backdrop for elucidating constraints on evolution, for functionally qualifying species and ecosystems, and for improving models that predict future vegetation based on continuous variation in plant form and function.

Leaf traits capture the effects of land use changes and climate on litter decomposability of grasslands across Europe

Land use and climate changes induce shifts in plant functional diversity and community structure, thereby modifying ecosystem processes. This is particularly true for litter decomposition, an essential process in the biogeochemical cycles of carbon and nutrients. In this study, we asked whether changes in functional traits of living leaves in response to changes in land use and climate were related to rates of litter potential decomposition, hereafter denoted litter decomposability, across a range of 10 contrasting sites. To disentangle the different control factors on litter decomposition, we conducted a microcosm experiment to determine the decomposability under standard conditions of litters collected in herbaceous communities from Europe and Israel. We tested how environmental factors (disturbance and climate) affected functional traits of living leaves and how these traits then modified litter quality and subsequent litter decomposability. Litter decomposability appeared proximately linked to initial litter quality, with particularly clear negative correlations with lignin-dependent indices (litter lignin concentr tion, lignin:nitrogen ratio, and fiber component). Litter quality was directly related to community-weighted mean traits. Lignin-dependent indices of litter quality were positively correlated with community-weighted mean leaf dry matter content (LDMC), and negatively correlated with community-weighted mean leaf nitrogen concentration (LNC). Consequently, litter decomposability was correlated negatively with community-weighted mean LDMC, and positively with community-weighted mean LNC. Environmental factors (disturbance and climate) influenced community-weighted mean traits. Plant communities experiencing less frequent or less intense disturbance exhibited higher community-weighted mean LDMC, and therefore higher litter lignin content and slower litter decomposability. LDMC therefore appears as a powerful marker of both changes in land use and of the pace of nutrient cycling across 10 contrasting sites.

Distribution of plant functional types along gradients of disturbance intensity and resource supply in an agricultural landscape

Abstract. In this study, plant functional types are understood as groups of plants with similar biological traits displaying significant optima or maxima on a gradient plane of resource supply and disturbance intensity. The biological traits refer to expansion, vegetative regeneration, generative reproduction, dispersal and seed bank longevity. 129 vegetation samples were taken in an agricultural landscape in southwestern Germany, covering a wide range of terrestrial vegetation types – but with the exception of forests and wetlands. For each site, also soil data were recorded. Mean daily soil moisture was estimated with a simple model. Soil moisture, balanced nitrogen supply and available phosphorus were combined into a factor ‘resource supply’. In addition, disturbance intensity was estimated for each site. This factor was based on (1) frequency of disturbance, (2) disturbance depth below or above the soil surface, and (3) proportion of the area affected by a discrete disturbance event. 30 plant groups with similar biological characteristics resulted from a cluster analysis, based on a compilation of 19 biological traits for a regional species pool. Logistic regression on a gradient plane of disturbance intensity and resource supply yielded response curves for 28 groups. The dependent variable was defined as the probability of encountering all members of a group in a sample. 17 groups display a significant response curve on the gradient plane. Plants with a potential for long- range dispersal are concentrated on sites with low or high disturbance intensities (e.g. fallow land, fields, lawns). On sites with medium disturbance intensity (e.g. meadows) and low to medium resource supply, small-range dispersal predominates. There are no distinct trends concerning seed bank longevity. The potential for vertical and lateral expansion increases with decreasing disturbance intensity. Only at medium disturbance intensities does vertical expansion correlate positively with resource supply. Rapid detachment of daughter individuals occurs more often on productive sites than on less productive sites. Diversity of groups with similar biological traits is highest on sites with medium disturbance intensities.

BIOPOP - A DATABASE OF PLANT TRAITS AND INTERNET APPLICATION FOR NATURE CONSERVATION

In scientific literature there is a wealth of information about the ecological and life-history traits of plant species. This information would be very valuable for a functional analysis of the response of species and communities to environmental changes and subsequent vulnerability/viability analysis or predictive modelling. However, the information is scattered in a multiple of different sources. Researchers often are discouraged from searching all available information.We are compiling a database of plant traits of the flora of Germany. Our aim is to create a comprehensive, well documented database to be made freely accessible within FLORAWEB, an Internet portal of the German Federal Agency for Nature Conservation (BfN). An interactive user-interface will enable the data to be readily used for extracting biological risk factors and for decision-making in nature conservation.A further aim of the project is to develop an application to enable simple predictions of plant population responses to habitat conservation or restoration management.

Life-history traits of the Northwest European flora: The LEDA database

Abstract An international group of scientists is building a ‘trait base’, an open internet database of life-history traits of the Northwest European flora (LEDA) that can be used as a tool in planning, in nature conservation and restoration, and in other applied research. The species-trait matrix will comprise referenced information under control of an editorial board, for species of the Northwest European flora, combining existing information and additional measurements. The focus will be on 26 plant traits that describe three key features of plant dynamics: persistence, regeneration, and dispersability. Currently 35% of the species-trait matrix has been filled; however, as the LEDA trait base consortium aims to deliver a database as complete as possible, all input from the scientific community is welcome.

A parsimonious combination of functional traits predicting plant response to disturbance and soil fertility

Abstract Questions: 1. How many traits associated with persistence and regeneration are necessary to predict the response of plants to soil fertility and disturbance? 2. Are correlated changes in trait expressions linked to the response of functional groups to fertility and disturbance? Location: Lower Frankonia, Germany. Methods: On 120 plots located in managed and abandoned grasslands, fields, and vineyards, we recorded species composition, disturbance intensity, soil water and nutrients, and ten candidate traits for 75 species. We used a novel method which is based on three steps: (1) logistic regression to separate responsive from non-responsive species; (2) iterative clustering of all possible combinations of the candidate traits including regression of each cluster in response to the environmental variables; (3) selection of the trait combination that performed best with respect to goodness of fit of all clusters from this combination. Bivariate trait relationships across functional groups were analysed with reduced major axis regression (RMA). Results: The parsimonious trait combination consisted of life span, specific leaf area (SLA), canopy height, and seed number. The ‘acquisitive’ functional groups in terms of SLA and height were linked to higher fertility and earlier disturbance, while the ‘retentive’ groups related to lower fertility and later disturbance. Investment in reproduction, however, displayed a reverse relation. SLA and canopy height showed correlated shifts in two pairs of co-occurring functional groups. Conclusions: A small number of traits is sufficient to predict the response of species. Plants need a higher increment in SLA to reach the same height, if start of disturbance is earlier. Linkages between traits shift from generative to vegetative with increasing fertility and earlier start of disturbance. Functional groups enable shifts in scaling relationships between traits to be analysed, in contrast to the analysis of single traits. Nomenclature: Rothmaler (1994).

Dispersal traits determine plant response to habitat connectivity in an urban landscape

Identification of trait syndromes that make species vulnerable to habitat fragmentation is essential in predicting biodiversity change. Plants are considered particularly vulnerable if their capacities for persistence in and for dispersal among local habitats are low. Here we investigated the influence of easily measured functional traits on the presence of 45 plant species in an urban landscape in north-west Germany where patches were separated by distances consistent with regular plant dispersal range. To describe the spatial configuration of patches we calculated species-specific patch connectivities. Then we assessed plant connectivity responses in distribution models calculated from connectivities and environmental predictors. Twenty (45%) of the analysed species showed a positive connectivity response after accounting for species-specific habitat requirements. These species differed from non-responsive species by functional traits associated with dispersal, including reduced seed numbers and higher terminal velocities relative to non-responsive species. Persistence traits played however no role which we attribute to the environmental conditions of urban habitats and their spatiotemporal characteristics. Our study underlines that even ruderal plants experience dispersal limitation and demonstrates that easily measured functional traits may be used as indicators of fragmentation vulnerability in urban systems allowing generalizations to larger species sets.

Plant species richness, vegetation structure and soil resources of urban brownfield sites linked to successional age

Brownfield sites contribute significantly to urban biodiversity due to their high spatio-temporal dynamics and their transient character. Plant species richness is, among other factors, contingent on vegetation structure. In this study, we examined plant species richness, vegetation height, vegetation density and soil parameters of a chronosequence of urban brownfield sites in Bremen and Berlin, Germany. These parameters were linked to successional age using single and multiple linear regression. Most biotic and abiotic variables differed significantly between sites with and without brick rubble in the soil, indicating a strong effect of site history on vegetation development. Soil parameters of the sites were not clearly linked to site age. Vegetation height and density increased significantly over time. Additionally, height and density increased with soil phosphorus content and water permeability of the soil, whilst plant available water only contributed to the model of vegetation density. Species richness increased with vegetation height but decreased with vegetation density. This indicates that species richness is maximised when a community comprises a mixture of early and mid-successional species. The results suggest that high plant species richness on sandy brownfield sites can be achieved by strong disturbances at an interval of 5 (±2) years. However, limiting soil resources can prolong this interval considerably. Management aiming to maximise plant species richness in urban brownfield sites should therefore take into account the interplay between soil resources and site age.

Validation of plant functional types across two contrasting landscapes

Abstract The validation of plant functional type models across contrasting landscapes is seen as a step towards the claim that plant functional types should recur regionally or even globally. I sampled the vegetation of an urban landscape on a range of sites representing gradients of resource supply and disturbance intensity. A group of plants with similar attributes was considered a ‘functional type’, if the species significantly co-occurred in a certain segment of the gradient plane of resource supply and disturbance intensity. Vegetative and regeneration traits were considered. A similar study was performed in a nearby agricultural landscape (Kleyer 1999). The logistic regression models from the urban landscape were applied to the data set of the agricultural landscape and vice versa. Although the overall environment of the two landscapes was very different, recurrent patterns of several functional types were found. At high fertility and high disturbance levels, annual species predominated with a persistent seed bank, high seed output, and short vertical expansion. When disturbances changed from below-ground to above-ground, the sexual regeneration mode was replaced by the vegetative mode, while vertical expansion remained low. At medium disturbance intensities, the vertical expansion and vegetative regeneration increased with fertility, while the seed bank remained mostly transient to short-term persistent and lateral expansion and sexual regeneration was intermediate. At low disturbances and low resource supplies, seed bank longevity, and vertical and lateral expansion tended to be long. Diversity of groups of plants with similar attributes was highest at intermediate disturbance levels and low fertility. These results correspond with Grime’s humped-back model and Connell’s intermediate disturbance hypothesis. Nomenclature: Ellenberg et al. (1991) for plants; Anon. (1988; modified) for soil types.