W. Daniel Kissling

The role of biotic interactions in shaping distributions and realised assemblages of species: Implications for species distribution modelling

Predicting which species will occur together in the future, and where, remains one of the greatest challenges in ecology, and requires a sound understanding of how the abiotic and biotic environments interact with dispersal processes and history across scales. Biotic interactions and their dynamics influence species relationships to climate, and this also has important implications for predicting future distributions of species. It is already well accepted that biotic interactions shape species spatial distributions at local spatial extents, but the role of these interactions beyond local extents (e.g. 10 km2 to global extents) are usually dismissed as unimportant. In this review we consolidate evidence for how biotic interactions shape species distributions beyond local extents and review methods for integrating biotic interactions into species distribution modelling tools. Drawing upon evidence from contemporary and palaeoecological studies of individual species ranges, functional groups, and species richness patterns, we show that biotic interactions have clearly left their mark on species distributions and realised assemblages of species across all spatial extents. We demonstrate this with examples from within and across trophic groups. A range of species distribution modelling tools is available to quantify species environmental relationships and predict species occurrence, such as: (i) integrating pairwise dependencies, (ii) using integrative predictors, and (iii) hybridising species distribution models (SDMs) with dynamic models. These methods have typically only been applied to interacting pairs of species at a single time, require a priori ecological knowledge about which species interact, and due to data paucity must assume that biotic interactions are constant in space and time. To better inform the future development of these models across spatial scales, we call for accelerated collection of spatially and temporally explicit species data. Ideally, these data should be sampled to reflect variation in the underlying environment across large spatial extents, and at fine spatial resolution. Simplified ecosystems where there are relatively few interacting species and sometimes a wealth of existing ecosystem monitoring data (e.g. arctic, alpine or island habitats) offer settings where the development of modelling tools that account for biotic interactions may be less difficult than elsewhere.

Biodiversity, Species Interactions and Ecological Networks in a Fragmented World

Biodiversity is organised into complex ecological networks of interacting species in local ecosystems, but our knowledge about the effects of habitat fragmentation on such systems remains limited. We consider the effects of this key driver of both local and global change on both mutualistic and antagonistic systems at different levels of biological organisation and spatiotemporal scales. n nThere is a complex interplay of patterns and processes related to the variation and influence of spatial, temporal and biotic drivers in ecological networks. Species traits (e.g. body size, dispersal ability) play an important role in determining how networks respond to fragment size and isolation, edge shape and permeability, and the quality of the surrounding landscape matrix. Furthermore, the perception of spatial scale (e.g. environmental grain) and temporal effects (time lags, extinction debts) can differ markedly among species, network modules and trophic levels, highlighting the need to develop a more integrated perspective that considers not just nodes, but the structural role and strength of species interactions (e.g. as hubs, spatial couplers and determinants of connectance, nestedness and modularity) in response to habitat fragmentation. n nMany challenges remain for improving our understanding: the likely importance of specialisation, functional redundancy and trait matching has been largely overlooked. The potentially critical effects of apex consumers, abundant species and super-generalists on network changes and evolutionary dynamics also need to be addressed in future research. Ultimately, spatial and ecological networks need to be combined to explore the effects of dispersal, colonisation, extinction and habitat fragmentation on network structure and coevolutionary dynamics. Finally, we need to embed network approaches more explicitly within applied ecology in general, because they offer great potential for improving on the current species-based or habitat-centric approaches to our management and conservation of biodiversity in the face of environmental change.

Food plant diversity as broad-scale determinant of avian frugivore richness

The causes of variation in animal species richness at large spatial scales are intensively debated. Here, we examine whether the diversity of food plants, contemporary climate and energy, or habitat heterogeneity determine species richness patterns of avian frugivores across sub-Saharan Africa. Path models indicate that species richness of Ficus (their fruits being one of the major food resources for frugivores in the tropics) has the strongest direct effect on richness of avian frugivores, whereas the influences of variables related to water–energy and habitat heterogeneity are mainly indirect. The importance of Ficus richness for richness of avian frugivores diminishes with decreasing specialization of birds on fruit eating, but is retained when accounting for spatial autocorrelation. We suggest that a positive relationship between food plant and frugivore species richness could result from niche assembly mechanisms (e.g. coevolutionary adaptations to fruit size, fruit colour or vertical stratification of fruit presentation) or, alternatively, from stochastic speciation–extinction processes. In any case, the close relationship between species richness of Ficus and avian frugivores suggests that figs are keystone resources for animal consumers, even at continental scales.

Geographical ecology of the palms (Arecaceae): determinants of diversity and distributions across spatial scales

BACKGROUNDnThe palm family occurs in all tropical and sub-tropical regions of the world. Palms are of high ecological and economical importance, and display complex spatial patterns of species distributions and diversity.nnnSCOPEnThis review summarizes empirical evidence for factors that determine palm species distributions, community composition and species richness such as the abiotic environment (climate, soil chemistry, hydrology and topography), the biotic environment (vegetation structure and species interactions) and dispersal. The importance of contemporary vs. historical impacts of these factors and the scale at which they function is discussed. Finally a hierarchical scale framework is developed to guide predictor selection for future studies.nnnCONCLUSIONSnDeterminants of palm distributions, composition and richness vary with spatial scale. For species distributions, climate appears to be important at landscape and broader scales, soil, topography and vegetation at landscape and local scales, hydrology at local scales, and dispersal at all scales. For community composition, soil appears important at regional and finer scales, hydrology, topography and vegetation at landscape and local scales, and dispersal again at all scales. For species richness, climate and dispersal appear to be important at continental to global scales, soil at landscape and broader scales, and topography at landscape and finer scales. Some scale-predictor combinations have not been studied or deserve further attention, e.g. climate on regional to finer scales, and hydrology and topography on landscape and broader scales. The importance of biotic interactions - apart from general vegetation structure effects - for the geographic ecology of palms is generally underexplored. Future studies should target scale-predictor combinations and geographic domains not studied yet. To avoid biased inference, one should ideally include at least all predictors previously found important at the spatial scale of investigation.

Cenozoic imprints on the phylogenetic structure of palm species assemblages worldwide

Despite long-standing interest in the origin and maintenance of species diversity, little is known about historical drivers of species assemblage structure at large spatiotemporal scales. Here, we use global species distribution data, a dated genus-level phylogeny, and paleo-reconstructions of biomes and climate to examine Cenozoic imprints on the phylogenetic structure of regional species assemblages of palms (Arecaceae), a species-rich plant family characteristic of tropical ecosystems. We find a strong imprint on phylogenetic clustering due to geographic isolation and in situ diversification, especially in the Neotropics and on islands with spectacular palm radiations (e.g., Madagascar, Hawaii, and Cuba). Phylogenetic overdispersion on mainlands and islands corresponds to biotic interchange areas. Differences in the degree of phylogenetic clustering among biogeographic realms are related to differential losses of tropical rainforests during the Cenozoic, but not to the cumulative area of tropical rainforest over geological time. A largely random phylogenetic assemblage structure in Africa coincides with severe losses of rainforest area, especially after the Miocene. More recent events also appear to be influential: phylogenetic clustering increases with increasing intensity of Quaternary glacial-interglacial climatic oscillations in South America and, to a lesser extent, Africa, indicating that specific clades perform better in climatically unstable regions. Our results suggest that continental isolation (in combination with limited long-distance dispersal) and changing climate and habitat loss throughout the Cenozoic have had strong impacts on the phylogenetic structure of regional species assemblages in the tropics.

Space Use of Bumblebees (Bombus spp.) Revealed by Radio-Tracking

Background Accurate estimates of movement behavior and distances travelled by animals are difficult to obtain, especially for small-bodied insects where transmitter weights have prevented the use of radio-tracking. Methodology/Principal Findings Here, we report the first successful use of micro radio telemetry to track flight distances and space use of bumblebees. Using ground surveys and Cessna overflights in a Central European rural landscape mosaic we obtained maximum flight distances of 2.5 km, 1.9 km and 1.3 km for Bombus terrestris (workers), Bombus ruderatus (worker), and Bombus hortorum (young queens), respectively. Bumblebee individuals used large areas (0.25–43.53 ha) within one or a few days. Habitat analyses of one B. hortorum queen at the landscape scale indicated that gardens within villages were used more often than expected from habitat availability. Detailed movement trajectories of this individual revealed that prominent landscape structures (e.g. trees) and flower patches were repeatedly visited. However, we also observed long (i.e. >45 min) resting periods between flights (B. hortorum) and differences in flower-handling between bumblebees with and without transmitters (B. terrestris) suggesting that the current weight of transmitters (200 mg) may still impose significant energetic costs on the insects. Conclusions/Significance Spatio-temporal movements of bumblebees can now be tracked with telemetry methods. Our measured flight distances exceed many previous estimates of bumblebee foraging ranges and suggest that travelling long distances to food resources may be common. However, even the smallest currently available transmitters still appear to compromise flower handling performance and cause an increase in resting behavior of bees. Future reductions of transmitter mass and size could open up new avenues for quantifying landscape-scale space use of insect pollinators and could provide novel insights into the behavior and requirements of bumblebees during critical life stages, e.g. when searching for mates, nest locations or hibernation sites.

Long-term impacts of fuelwood extraction on a tropical montane cloud forest.

Fuelwood extracted from natural forests serves as a principal energy source in rural regions of many tropical countries. Although fuelwood extraction (even low intensities) might strongly impact the structure and species composition of natural forests, long-term studies remain scarce. Here, we estimate the potential long-term impacts (over several hundred years) of such repeated harvesting of single trees on tropical montane cloud forest in central Veracruz, Mexico, by applying a process-based forest growth model. We simulate a wide range of possible harvesting scenarios differing in wood volume harvested and preferred tree species and sizes, and use a set of indicators to compare their impacts on forest size structure and community composition. Results showed that the overall impact on forest structure and community composition increased linearly with the amount of harvested wood volume. Even at low levels of harvesting, forest size structure became more homogeneous in the long term because large old trees disappeared from the forest, but these changes might take decades or even centuries. Although recruitment of harvested species benefited from harvesting, species composition shifted to tree species that are not used for fuelwood. Our results demonstrate that fuelwood extraction can have marked long-term impacts on tropical montane cloud forests. The results also offer the possibility to support the design of management strategies for the natural species-rich forests that achieve a balance between economic needs and ecological goals of the stakeholders.

Mammal predator and prey species richness are strongly linked at macroscales

Predator-prey interactions play an important role for species composition and community dynamics at local scales, but their importance in shaping large-scale gradients of species richness remains unexplored. Here, we use global range maps, structural equation models (SEM), and comprehensive databases of dietary preferences and body masses of all terrestrial, non-volant mammals worldwide, to test whether (1) prey bottom-up or predator top-down relationships are important drivers of broad-scale species richness gradients once the environment and human influence have been accounted for, (2) predator-prey richness associations vary among biogeographic regions, and (3) body size influences large-scale covariation between predators and prey. SEMs including only productivity, climate, and human factors explained a high proportion of variance in prey richness (R2=0.56) but considerably less in predator richness (R2=0.13). Adding predator-to-prey or prey-to-predator paths strongly increased the explained variance in both cases (prey R2=0.79, predator R2=0.57), suggesting that predator-prey interactions play an important role in driving global diversity gradients. Prey bottom-up effects prevailed over productivity, climate, and human influence to explain predator richness, whereas productivity and climate were more important than predator top-down effects for explaining prey richness, although predator top-down effects were still significant. Global predator-prey associations were not reproduced in all regions, indicating that distinct paleoclimate and evolutionary histories (Africa and Australia) may alter species interactions across trophic levels. Stronger cross-trophic-level associations were recorded within categories of similar body size (e.g., large prey to large predators) than between them (e.g., large prey to small predators), suggesting that mass-related energetic and physiological constraints influence broad-scale richness links, especially for large-bodied mammals. Overall, our results support the idea that trophic interactions can be important drivers of large-scale species richness gradients in combination with environmental effects.

Invasion ecology of the alien tussock grass Nardus stricta (Poaceae) at Lake Pukaki, Canterbury, New Zealand

Abstract The European matgrass Nardus stricta has naturalised in New Zealand, often on damp soils within wetlands and grasslands. In this paper, we present for the first time field data on the ecology of this alien invader in New Zealand, from eight kettle‐hole wetlands on lateral moraine along the western side of Lake Pukaki, South Canterbury. The invaded wetland sites were all acidic but varied in other soil characteristics. Nardus stricta was the most dominant species within these wetland communities with 40% of all plots showing more than 50% coverage, and 21% having more than 90% cover. Species richness (including vascular plants and mosses) at some sites was relatively high (c. 40 species), but species richness and abundance were significantly reduced in quadrats with high Nardus stricta density. Seedling densities of Nardus stricta were high but variable (overall mean of 38.6 ± 116 seedlings per m2), and establishment mainly occurred on the cushion‐forming sedge Oreobolus pectinatus. Most reproductive tussocks (90%) produced up to 1000 florets, but single tussocks were able to produce up to 10000 and more florets. Our results suggest that Nardus stricta is still expanding its local range and, therefore, is likely to increase in abundance and dominance, resulting in further negative effects on native species diversity.