Contrasting processes drive alpha and beta taxonomic, functional and phylogenetic diversity of orthopteran communities in grasslands

Taxonomic, functional and phylogenetic diversities can respond differently to biotic and abiotic filters. However, biodiversity management tends to focus on a single index, generally taxonomic diversity, assuming a strong positive correlation among biodiversity components across scales. This can result in a mismanagement of functional and phylogenetic diversities with negative consequences for ecosystem functioning and long-term maintenance of services. Understanding the relationships among biodiversity components, how they change across scales and which their main drivers are can lead to more sustainable management of biodiversity and its associated ecosystem services and functions. We used an integrative approach of biodiversity where we investigated alpha and beta taxonomic (TD), functional (FD) and phylogenetic (PD) diversity of orthopteran communities as well as species and functional traits composition and their associated drivers at the local and landscape scales in permanent mesic grasslands of the French Jura Mountains. We assessed whether orthopteran TD, FD and PD were positively correlated. We also determined the drivers of TD, FD and PD and their changes across scales and among indices using data related to soil, agricultural practices, elevation, and biotic interactions with plants. Our results showed that (i) elevation was a strong driver of orthopteran community trait composition, (ii) orthopteran alpha TD, FD and PD were correlated among themselves and increased with plant species richness, and (iii) local beta diversity was not correlated with alpha diversity. Beta diversity had different drivers at local and landscape scale: it was influenced by soil chemistry and texture at the local scale and increased with the difference in elevation among plots at the landscape scale. This study evidences distinct processes driving alpha and beta diversity of orthopteran communities at both the local and landscape scales. It supports the hypothesis that less intensive agricultural practices enhance orthopteran diversity and highlights the importance to consider beta diversity at both local and landscape scale when designing and assessing the management regimes of grassland ecosystems. Prioritizing the importance of the different biodiversity components and spatial scales constitutes an important challenge for sustainable grassland management.

Theoretical and Experimental Studies of Dendritic Metacommunities

Francesco Carrara

The present thesis deals with the understanding of the origins and the mechanisms of maintenance of biodiversity in natural landscapes, in particular by identifying key processes that define large-scale patterns of abundance and diversity. Biological communities often occur in spatially structured habitats where connectivity directly affects dispersal and metacommunity processes. Recent theoretical work suggests that dispersal constrained by the connectivity of specific habitat structures affects diversity patterns and species interactions. This is particularly relevant in dendritic networks epitomized by fluvial ecological corridors. This thesis addresses whether connectivity alone can explain observed features of biodiversity and selectively promote different components of community composition in river-like landscapes, such as local species richness or the among-community similarity. The relevance of this thesis lies in the major ecological challenges posed by the topic, and its fundamental importance to conservation biology. The studies pursued herein are also deemed relevant because of the influence of the spatial connectivity and dispersal on population dynamics and of the relevance of biodiversity to ecosystem functioning. Mechanisms of species coexistence were investigated with a blend of theoretical tools (broadly related to statistical mechanics and the theory of stochastic processes) and experimental work using laboratory microbial communities. The research tools ranged from aspects of modern coexistence theory in a local perspective to the recent concept of the metacommunity in spatial ecology, within a unified framework. The study of biodiversity in riverine ecosystems guided by observational data has been addressed by combining theoretical metacommunity models with laboratory experiments. The results are diverse. First, they show experimentally that connectivity per se shapes key components of biodiversity in metacommunities. Local dispersal in isotropic lattice landscapes homogenizes local species richness and leads to pronounced spatial persistence. By contrast, dispersal along dendritic landscapes leads to higher variability in local diversity and among-community composition. Although headwaters exhibit relatively lower species richness, they are crucial for the maintenance of regional biodiversity. By suitably arranging patch sizes within river-like networks the effect of local habitat capacity (i.e., the patch size) and dendritic connectivity on biodiversity can be experimentally disentangled in aquatic microcosm metacommunities. It is shown in this thesis that species coexistence and community assembly depend on intricate, non-trivial interactions of local community capacity and network positioning. Furthermore, an interaction of spatial arrangement of habitat capacity and dispersal along river-like networks also affects a key ecosystem descriptor, namely regional evenness. High regional evenness in community composition is found only in landscapes preserving geomorphological scaling properties of patch sizes. In riverine environments some of the rarer species sustained regionally more abundant populations and were better able to track their own niche requirements compared to landscapes with homogeneous patch size or landscapes with spatially uncorrelated patch size. All the experimental results were supported and extended by a theoretical analysis where the above mechanisms have been generalized. This thesis provides the first direct experimental evidence that spatially constrained dendritic connectivity is a key factor for community composition and population persistence in riverine landscapes. As such, this thesis assesses a longstanding issue in spatial community ecology. It offers unique insights into the ecological forces structuring natural communities in a key ecosytem, and demonstrates principles that can be further tested in theoretical metacommunity models possibly to be extended to real riverine ecosystems. Taken together, the analyses show how the structure of ecological networks interacts with the spatial environmental matrix in determining biodiversity patterns and the functioning of biological communities. The analyses also suggest that altering the natural linkage between dendritic connectivity and patch size strongly affects community properties at multiple scales. The first part of this thesis (chapters 2 and 3) addresses key aspects of biodiversity-ecosystem functioning research where the combination of theory-guided experiments and theoretical investigations shows how a stochastic implementation of population dynamics proves fundamental for key community properties such as species persistence and community stability. The diversity-productivity and diversity-stability relationships are explored. Both experimental findings and the results of a stochastic model fitted to the experimental interaction matrix, suggest the emergence of strong stabilizing forces when species from different functional groups interact in the same environment, increasing species coexistence and community biomass production. The last part (chapter 6) provides a synthesis of this thesis work, in that it aims at unifying aspects from niche-theory, usually adopted in spatially implicit models, with those characteristic of a spatially explicit context from a typical real-life mountainous regions. It is dedicated to the possible explanation for a macroecological pattern routinely observed from organisms in different domains of life, that is, the mid-elevational peak in local species richness. Guided by empirical observations on diversity of macroinvertebrates in Swiss river basins, a theoretical ansatz is provided which is deemed to capture the essential geomorphological drivers and controls relating species-fitness to altitude. A set of overarching conclusions and perspectives for future research are discussed in the concluding chapter.

EPFL2013

Subalpine Pasture-Woodlands in a Changing Climate

Konstantin Svetlozarov Gavazov

Climate change in temperate mountain systems and associated increase in temperature and decrease in precipitation are expected to have strong implications for vegetation productivity, species diversity and carbon turnover in subalpine grasslands. Little is known, however, about the interaction between the effects of climate change and those of local land use management and possible changes in landscape structure. Pasture woodlands in the Swiss Jura Mountains are a traditional landscape, resulting from a long-lived sustainable use of grasslands and woodlands, and as such provide a suite of important ecosystem services to human society. These range from carbon sequestration and biodiversity preservation, to provision of timber and forage for livestock, and last but not least an aesthetic value, much appreciated by tourism. In this thesis various aspects of ecosystem functioning have been studied, investigating the combined effects of experimental climate change and land use on structurally different wooded pastures. An altitudinal gradient method has been used to simulate future climate change conditions, by imposing warmer and drier climate on subalpine turfs transplanted at lower elevation. The resulting gradient in mean annual temperature and precipitation – ranging from cold and wet in the subalpine zone, to warm and dry in the colline zone – has allowed for the detection of tipping points and altered states of ecosystem functioning in response to the treatments. The method employed provided also the possibility for a direct comparison of three land use types: unwooded pastures, sparsely wooded pastures, and densely wooded pastures (the result of pasture management intensity), in their response to climate perturbation. During the four years of experimental work, a series of observations have been made at the plot scale (square metre) in terms of plant performance and biogeochemical cycles, as well as at the landscape scale (hectare) in terms of forage production. A general threshold level for ecosystem resistance to experimental climate change was detected between the moderate IPCC scenario (+2 K mean annual temperature; -20 % annual precipitation) and the intensive IPCC scenario (+4 K mean annual temperature; -40 % annual precipitation). A concomitant gradient in ecosystem response to climate change was observed across the three land use types. The intensively managed unwooded pasture type was consistently more affected by the experimental treatment and rarely exhibited signs of resistance, especially under the intense climate change scenario. A drastic loss of plant species diversity, reduction of herbaceous biomass, impaired litter decomposition and soil microbial metabolic activity have all contributed to the altered state of ecosystem functioning. In contrast, the two extensively managed wooded pasture types showed considerable resistance to climate perturbation in terms of both above and belowground ecosystem processes. The reported inter-annual variation in herbaceous diversity and biomass production within these land use types demonstrated their resilience (recovery) potential too. Using a modelling approach for upscaling these results to the heterogeneous landscape of pasture woodlands in the Swiss Jura Mountains, has proven that extensively used wooded pastures could grant sustainable ecosystem services in terms of forage provision for cattle under climate change. Considering that the two experimental climate change intensities implemented this study are the projected ‘best’ and ‘worst’ case scenarios for the coming decades, the reported resistance of wooded pastures to climate change has to be embraced, and sustainable land use set as a goal in high altitude mountain pastures.

EPFL2013

Impact of management type and intensity on multiple facets of grassland biodiversity in the French Jura Mountains

François Gillet

Questions(1) Does grassland vegetation differ in its taxonomic, phylogenetic and functional alpha diversity between strictly grazed parcels (SGP) and mowed-and-grazed parcels (MGP)? (2) Do we observe differences in the responses of functional diversity (reflecting differences in the interspecific variation of a trait in the plant community) and community-weighted mean trait values (CWMs, reflecting the characteristics of the dominant species)? (3) How do diversity and CWM patterns respond to disturbance and fertilization gradients measured by agronomic descriptors in both parcel types? LocationFrench Jura Mountains. MethodsWe compared taxonomic, phylogenetic and functional diversities (based on Rao quadratic entropy) and CWMs between the two management types (SGP and MGP). We then used agronomic variables, describing both defoliation regime and fertilization intensity over the past 10yrs to explain diversity indices and CWMs in a partial redundancy analysis (pRDA). ResultsTaxonomic and phylogenetic diversities were higher in SGP than in MGP, as well as functional diversity computed for qualitative traits (clonal growth organs and leaf distribution along the stem). Contrary to expectations from literature, we found higher CWM for leaf dry matter content (LDMC) in intensively and regularly cut grasslands compared to only grazed grasslands, which can be explained by the higher dominance of grass species in hay meadows. In addition, constrained ordination revealed that taxonomic, phylogenetic and some functional diversity metrics were negatively affected by defoliation intensity (mowing frequency), while most CWMs were mainly influenced by fertilization intensity, either by the available N input or by the role of commercial inorganic fertilizers, depending on the trait considered. ConclusionsOur study revealed consistent patterns between the various facets of biodiversity in grasslands according to the main management regime. Taking agronomic gradients into account suggests the importance of considering both CWMs and functional diversity, which respond variously to agricultural practices. Conservation of high diversity in grasslands, whatever the facet considered, implies keeping some parcels exclusively grazed and limiting N inputs, especially those from inorganic commercial fertilizers.

Wiley-Blackwell2014

Theoretical and Experimental Studies of Dendritic Metacommunities

Francesco Carrara

EPFL2013

Subalpine Pasture-Woodlands in a Changing Climate

Konstantin Svetlozarov Gavazov

EPFL2013

Impact of management type and intensity on multiple facets of grassland biodiversity in the French Jura Mountains

François Gillet

Wiley-Blackwell2014