Identifying Conservation and Restoration Priorities for Saproxylic and Old-Growth Forest Species: A Case Study in Switzerland

Saproxylic (dead-wood-associated) and old-growth species are among the most threatened species in European forest ecosystems, as they are susceptible to intensive forest management. Identifying areas with particular relevant features of biodiversity is of prime concern when developing species conservation and habitat restoration strategies and in optimizing resource investments. We present an approach to identify regional conservation and restoration priorities even if knowledge on species distribution is weak, such as for saproxylic and old-growth species in Switzerland. Habitat suitability maps were modeled for an expert-based selection of 55 focal species, using an ecological niche factor analyses (ENFA). All the maps were then overlaid, in order to identify potential species' hotspots for different species groups of the 55 focal species (e.g., birds, fungi, red-listed species). We found that hotspots for various species groups did not correspond. Our results indicate that an approach based on "richness hotspots" may fail to conserve specific species groups. We hence recommend defining a biodiversity conservation strategy prior to implementing conservation/restoration efforts in specific regions. The conservation priority setting of the five biogeographical regions in Switzerland, however, did not differ when different hotspot definitions were applied. This observation emphasizes that the chosen method is robust. Since the ENFA needs only presence data, this species prediction method seems to be useful for any situation where the species distribution is poorly known and/or absence data are lacking. In order to identify priorities for either conservation or restoration efforts, we recommend a method based on presence data only, because absence data may reflect factors unrelated to species presence.

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

Dead wood in managed forests: how much and how much is enough?

Rita Bütler Sauvain

The aims of this research were twofold: to develop an efficient method for the quantification of large spruce snags (standing dying and dead trees), and to establish snag target values for sustainable forest management. We answer the two basic questions: how much dead wood is currently available in managed forests? And how much dead wood is enough for biodiversity conservation? It is widely accepted that modern forest management has to be sustainable. One generally recognised criterion of sustainability is the maintenance of biodiversity. Since this concept encompasses a large range of scales and features (landscapes, ecosystems, species and genes; components, processes, functions, etc.), biodiversity indicators have been identified for measurement and monitoring purposes. "Dead wood" has been recognised as a key indicator for biodiversity in forest ecosystems. Verifying and assessing progress towards biodiversity maintenance or restoration hence requires the measurement of different kinds of dead wood. Yet cost-efficient and rapid methods are still lacking. That is why we developed in this thesis an efficient method for the quantification of large spruce snags. Being based on infrared Aerial Photos and a Geographic Information System (GIS), it is called the AP-GIS-method. It enables mapping of snags and calculation of the spatial snag-density, i.e. the number of snags per hectare, and can be used to answer the question: How much dead wood is in managed forests? Beside techniques to assess dead-wood quantities, forest managers need quantitative target values, i.e. guidelines in order to know how much dead wood should be maintained in managed forests for biodiversity conservation. Natural forests may be used as reference systems to define such targets. However, since dead-wood amounts in natural forests may be extremely high, up to 30% of dead trunks, the retention of such amounts in managed forests would hardly be compatible with economic objectives. Another possibility for defining guidelines is the translation of the habitat requirements of deadwood- dependent species into management targets. The Three-toed woodpecker Picoides tridactylus has previously been recognised as a potential indicator species of features characteristic for forests with natural dynamics (especially old-growth). Although ecological studies had demonstrated its need of dying and dead trees for foraging, nesting and drumming, the required density of such trees has never been quantified. In this thesis, we analysed the dependence on dead wood for both European subspecies, the Alpine Picoides tridactylus alpinus and the northern P. tr. tridactylus. The study was conducted in sub-Alpine spruce forests in Switzerland and boreal forests in central Sweden. In both countries, we found a significant non-linear response of the probability of woodpecker presence to different amounts of dead trees, and identified critical ecological thresholds for the local presence of this species. Clearly, the Three-toed woodpecker depends on relatively high amounts of dying and dead trees. In Switzerland, the road network density negatively affected the presence of this woodpecker species, since a high road density facilitates forest management intensification and the removal of diseased and dead trees. Based on our results, and since several other links with biodiversity have previously been demonstrated, we suggest that Three-toed woodpeckers be considered indicators of dead wood and habitat quality. This species has therefore been used in this thesis to find an answer to the question: How much dead wood is enough in managed forests? In order to ascertain dead-tree targets, we developed a bioenergetic model for Threetoed woodpeckers, enabling estimation of snag amounts required by this species to satisfy its energy needs. By comparing modelling results with the previously identified critical dead-wood thresholds, we were able to derive reliable targets, since both approaches resulted in similar critical values. We recommend, for both boreal and sub-Alpine spruce forests, aiming for 5% of the standing tree basal area, or volume to be dead. Such snag-retention levels, to be applied over an area of about 100 ha, correspond in sub-Alpine forests to a basal area of ≥ 1.6 m2 ha-1, or a volume of ≥ 18 m3 ha-1, or a minimum of 14 snags with a diameter ≥ 21 cm per hectare. Considered as a pragmatic way to stimulate forest managers' interest in dead wood maintenance or restoration, we analysed the potential usefulness of Three-toed woodpeckers as natural agents against bark beetles. By defining three scenarios for different levels of woodpecker effectiveness, we compared the numbers of bark beetles consumed with those caught in pheromone traps used in forestry. We demonstrate that woodpeckers catch 2-19 times more bark beetles than traps do. This result is true for both cases, when one woodpecker is compared with one trap, and when the whole Swiss woodpecker population is compared to all traps installed in Swiss forests.

EPFL2003

Functional responses of multitaxa communities to disturbance and stress gradients in a restored floodplain

Bertrand Fournier, François Gillet, Renée-Claire Le Bayon, Edward Mitchell

Trait-based approaches can reveal the mechanisms through which disturbances or stress impact communities, allowing comparisons of the role of different mechanisms in shaping communities among taxonomic groups. Such information can lead to higher comparability, transferability and predictability of the outcome of restoration projects. However, multitaxa trait-based approaches were rarely used in the context of ecosystem restoration. We investigated the responses to environmental gradients of seven taxa (vascular plants, staphylinid and carabid beetles, spiders, isopods, diplopods and earthworms) in a restored floodplain using a species traits approach. We assessed the impact of flood disturbances and soil hydric stress on the functional diversity (FD) and community-weighted mean (CWM) response of traits for each taxon. Ordination of hydrological variables revealed two main gradients. The first was related to the spatiotemporal dynamics of flood disturbances and the second to the average changes in soil hydric conditions. The analysis of CWM revealed that larger, poorly mobile species with narrow ecological tolerances were filtered by regular floods and/or changes in soil hydric conditions. Functional diversity patterns differed between the two gradients: decreasing with increasing flood disturbance, but increasing along the soil hydric stress gradient. This suggests that the mechanisms shaping community composition differ between the two gradients with environmental filtering being dominant with increasing flood disturbances and competition decreasing with more soil hydric stress.Synthesis and applications. Our study shows that the impact of restored flood disturbances and soil hydric stress on plant and invertebrate functional diversity and community-weighted mean can be positive, negative or more complex depending on the taxonomic group and environmental gradient considered. The patterns can to some extent be explained by the specific characteristics of each group. Larger, poorly mobile species with narrow ecological tolerances were particularly vulnerable to changes in disturbance and stress regime following floodplain restoration. These species may therefore be lost in the initial phases of restoration projects, but other more characteristic species of dynamic floodplains will be favoured. Understanding the consequences of these contrasted responses for biodiversity conservation and ecosystem functioning constitutes the next challenge for ecosystem restoration. Our study shows that the impact of restored flood disturbances and soil hydric stress on plant and invertebrate functional diversity and community-weighted mean can be positive, negative or more complex depending on the taxonomic group and environmental gradient considered. The patterns can to some extent be explained by the specific characteristics of each group. Larger, poorly mobile species with narrow ecological tolerances were particularly vulnerable to changes in disturbance and stress regime following floodplain restoration. These species may therefore be lost in the initial phases of restoration projects, but other more characteristic species of dynamic floodplains will be favoured. Understanding the consequences of these contrasted responses for biodiversity conservation and ecosystem functioning constitutes the next challenge for ecosystem restoration.

Wiley-Blackwell2015