Anke Stein

Environmental heterogeneity as a universal driver of species richness across taxa, biomes and spatial scales

Environmental heterogeneity is regarded as one of the most important factors governing species richness gradients. An increase in available niche space, provision of refuges and opportunities for isolation and divergent adaptation are thought to enhance species coexistence, persistence and diversification. However, the extent and generality of positive heterogeneity-richness relationships are still debated. Apart from widespread evidence supporting positive relationships, negative and hump-shaped relationships have also been reported. In a meta-analysis of 1148 data points from 192 studies worldwide, we examine the strength and direction of the relationship between spatial environmental heterogeneity and species richness of terrestrial plants and animals. We find that separate effects of heterogeneity in land cover, vegetation, climate, soil and topography are significantly positive, with vegetation and topographic heterogeneity showing particularly strong associations with species richness. The use of equal-area study units, spatial grain and spatial extent emerge as key factors influencing the strength of heterogeneity-richness relationships, highlighting the pervasive influence of spatial scale in heterogeneity-richness studies. We provide the first quantitative support for the generality of positive heterogeneity-richness relationships across heterogeneity components, habitat types, taxa and spatial scales from landscape to global extents, and identify specific needs for future comparative heterogeneity-richness research.

Terminology and quantification of environmental heterogeneity in species-richness research

Spatial environmental heterogeneity (EH) is an important driver of species diversity, and its influence on species richness has been analysed for numerous taxa, in diverse ecological settings, and over a large range of spatial scales. The variety and ambiguity of concepts and terminology, however, have hampered comparisons among studies. Based on a systematic literature survey of 192 studies including 1148 data points, we provide an overview of terms and measures related to EH, and the mechanisms that relate EH to species richness of plants and animals in terrestrial systems. We identify 165 different measures used to quantify EH, referred to by more than 350 measure names. We classify these measures according to their calculation method and subject area, finding that most studies have analysed heterogeneity in land cover, topography, and vegetation, whereas comparatively few studies have focused on climatic or soil EH. Overall, elevation range emerged as the most frequent measure in our dataset. We find that there is no consensus in the literature about terms (such as ‘habitat diversity’ or ‘habitat complexity’), their meanings and associated quantification methods. More than 100 different terms have been used to denote EH, with largely imprecise delimitations. We reveal trends in use of terms and quantification with respect to spatial scales, study taxa, and locations. Finally, we discuss mechanisms involved in EH–richness relationships, differentiating between effects on species coexistence, persistence, and diversification. This review aims at guiding researchers in their selection of heterogeneity measures. At the same time, it shows the need for precise terminology and avoidance of ambiguous synonyms to enhance understanding and foster among‐study comparisons and synthesis.

EDITOR'S CHOICE: REVIEW: Effects of land use on plant diversity – A global meta‐analysis

Summary Plant diversity is globally threatened by anthropogenic land use including management and modification of the natural environment. At regional and local scales, numerous studies world-wide have examined land use and its effects on plant diversity, but evidence for declining species diversity is mixed. This is because, first, land use comes in many variations, hampering comparisons of studies. Second, land use directly affects the environment, but indirect effects extend beyond the boundaries of the land in use. Third, land-use effects greatly depend on the environmental, historical and socio-economic context. To evaluate the generality and variation of studies’ findings about land-use effects, we undertook a quantitative synthesis using meta-analytic techniques. Using 572 effect sizes from 375 studies distributed globally relating to 11 classes of land use, we found that direct and indirect effects of land use on plant diversity (measured as species richness) are variable and can lead to both local decreases and increases. Further, we found evidence (best AIC model) that land-use-specific covariables mostly determine effect-size variation and that in general land-use effects differ between biomes. Synthesis and applications. This extensive synthesis provides the most comprehensive and quantitative overview to date about the effects of the most widespread and relevant land-use options on plant diversity and their covariables. We found important covariables of specific land-use classes but little evidence that land-use effects can be generally explained by their environmental and socio-economic context. We also found a strong regional bias in the number of studies (i.e. more studies from Europe and North America) and highlight the need for an overarching and consistent land-use classification scheme. Thereby, our study provides a new vantage point for future research directions.

The evolution and function of vessel and pit characters with respect to cavitation resistance across 10 Prunus species

Various structure-function relationships regarding drought-induced cavitation resistance of secondary xylem have been postulated. These hypotheses were tested on wood of 10 Prunus species showing a range in P50 (i.e., the pressure corresponding to 50% loss of hydraulic conductivity) from -3.54 to -6.27 MPa. Hydraulically relevant wood characters were quantified using light and electron microscopy. A phylogenetic tree was constructed to investigate evolutionary correlations using a phylogenetically independent contrast (PIC) analysis. Vessel-grouping characters were found to be most informative in explaining interspecific variation in P50, with cavitation-resistant species showing more solitary vessels than less resistant species. Co-evolution between vessel-grouping indices and P50 was reported. P50 was weakly correlated with the shape of the intervessel pit aperture, but not with the total intervessel pit membrane area per vessel. A negative correlation was found between P50 and intervessel pit membrane thickness, but this relationship was not supported by the PIC analysis. Cavitation resistance has co-evolved with vessel grouping within Prunus and was mainly influenced by the spatial distribution of the vessel network.

How drought and deciduousness shape xylem plasticity in three Costa Rican woody plant species

This paper explores the phenotypic plasticity of xylem in tropical trees in order to test the hypothesis that different leaf phenological patterns influence levels of xylem plasticity. Wood anatomy was studied in the wet-deciduous species Cordia alliodora (Ruiz & Pav.) Oken, the dry-deciduous species Tabebuia rosea (Bertol.) DC., and the evergreen species Ocotea veraguensis (Meisn.) Mez., collected from seasonally dry forest and tropical cloud forest in Costa Rica. Xylem plasticity and trait conservatism were examined by analysing the coefficient of variation (CV) and the relative distance plasticity index (RDPI) of xylem anatomical traits. The two deciduous species exhibited wider vessels, lower wood density, and higher Huber values than the evergreen species. Furthermore, intervessel connectivity was highest for the two deciduous species in seasonally dry forest compared to cloud forest, whereas the opposite was found for the evergreen species. Overall highest trait variability was found for all plants at the seasonally dry site. The evergreen species O. veraguensis had the highest plasticity values compared to the deciduous species. Highest plasticity was found for vessel composition index (vessel area/vessel number), whilestrongest trait conservatism was found for vessel diameter. In conclusion, our data indicate that evergreen O. veraguensis does not show less xylem plasticity than the two deciduous tree species studied.