Can reed harvest be used as a management strategy for improving invertebrate biomass and diversity?

The succession-driven reed bed habitat hosts a unique flora and fauna including several endangered invertebrate species. Reed beds can be managed through commercial winter harvest, with implications for reed bed conservation. However, the effects of winter harvest on the invertebrate community are not well understood and vary across studies and taxonomic levels. The aim of this study was to investigate the effects of reed harvest on invertebrate communities. Ground-dwelling and aerial invertebrates were continuously sampled for 10 weeks in the largest coherent reed bed of Scandinavia in order to assess how time since last reed harvest (0, 3, and 25-years) influences invertebrate biomass, biodiversity and community structure across taxonomic levels. Biomass was measured and all specimens were sorted to order level, and Coleoptera was even sorted to species level. The invertebrate community showed distinct compositional differences across the three reed bed ages. Furthermore, biomass of both aerial and ground-dwelling invertebrates was highest in the age-0 reed bed and lowest in the age-25 reed bed. Generally, biodiversity showed an opposite trend with the highest richness and diversity in the age-25 reed bed. We conclude that it is possible to ensure high insect biomass and diversity by creating a mosaic of reed bed of different ages through small-scale harvest in the largest coherent reed bed in Scandinavia. The youngest red beds support a high invertebrate biomass whereas the oldest reed beds support a high biodiversity. Collectively, this elevate our understanding of reed harvest and the effects it has on the invertebrate communities, and might aid in future reed bed management and restoration.

eDNA metabarcoding for biodiversity assessment, generalist predators as sampling assistants.

With an accelerating negative impact of anthropogenic actions on natural ecosystems, non-invasive biodiversity assessments are becoming increasingly crucial. As a consequence, the interest in the application of environmental DNA (eDNA) survey techniques has increased. The use of eDNA extracted from faeces from generalist predators, have recently been described as "biodiversity capsules" and suggested as a complementary tool for improving current biodiversity assessments. In this study, using faecal samples from two generalist omnivore species, the Eurasian badger and the red fox, we evaluated the applicability of eDNA metabarcoding in determining dietary composition, compared to macroscopic diet identification techniques. Subsequently, we used the dietary information obtained to assess its contribution to biodiversity assessments. Compared to classic macroscopic techniques, we found that eDNA metabarcoding detected more taxa, at higher taxonomic resolution, and proved to be an important technique to verify the species identification of the predator from field collected faeces. Furthermore, we showed how dietary analyses complemented field observations in describing biodiversity by identifying consumed flora and fauna that went unnoticed during field observations. While diet analysis approaches could not substitute field observations entirely, we suggest that their integration with other methods might overcome intrinsic limitations of single techniques in future biodiversity surveys.

Native and alien flower visitors differ in partner fidelity and network integration.

Globalisation persistently fuels the establishment of non-native species outside their natural ranges. While alien plants have been intensively studied, little is known about alien flower visitors, and especially, how they integrate into natural communities. Here, we focus on mutualistic networks from five Galápagos islands to quantify whether alien and native flower visitors differ consistently in their pairwise interactions. We find that (1) alien flower visitors have more interaction partners and larger species strengths (i.e. plants are more connected to alien insects), (2) native insects tend to have higher partner fidelity as they deviate more from random partner utilisation, and iii) the difference between native and alien flower visitors in network integration intensifies with island degradation. Thus, native and alien flower visitors are not interchangeable, and alien establishment might have yet unforeseen consequences for the pairwise dynamics between plants and flower visitors on the Galápagos - especially on the heavily disturbed islands.

Disclosing the double mutualist role of birds on Galápagos.

Life on oceanic islands deviate in many ways from that on the mainland. Their biodiversity is relatively poor and some groups are well-represented, others not, especially not insects. A scarcity of insects forces birds to explore alternative food, such as nectar and fruit. In this way, island birds may pollinate and disperse seed to an extent unseen on any mainland; they may even first consume floral resources of a plant species and then later harvest the fruit of the same species. Through this biotic reuse, they may act as double mutualists. The latter have never been studied at the level of the network, because they are traditionally considered rare. We sampled pollination and seed-dispersal interactions on Galápagos and constructed a plant-bird mutualism network of 108 plant (12% being double mutualists) and 21 bird species (48% being double mutualists), and their 479 interactions, being either single (95%) or double mutualisms (5%). Double mutualists constitute the core in the pollination-dispersal network, coupling the two link types together. They may also initiate positive feedbacks (more pollination leading to more dispersal), which theoretically are known to be unstable. Thus, double mutualisms may be a necessary, but risky prerequisite to the survival of island biodiversity.

Geographical variation in mutualistic networks: similarity, turnover and partner fidelity.

Although species and their interactions in unison represent biodiversity and all the ecological and evolutionary processes associated with life, biotic interactions have, contrary to species, rarely been integrated into the concepts of spatial ß-diversity. Here, we examine ß-diversity of ecological networks by using pollination networks sampled across the Canary Islands. We show that adjacent and distant communities are more and less similar, respectively, in their composition of plants, pollinators and interactions than expected from random distributions. We further show that replacement of species is the major driver of interaction turnover and that this contribution increases with distance. Finally, we quantify that species-specific partner compositions (here called partner fidelity) deviate from random partner use, but vary as a result of ecological and geographical variables. In particular, breakdown of partner fidelity was facilitated by increasing geographical distance, changing abundances and changing linkage levels, but was not related to the geographical distribution of the species. This highlights the importance of space when comparing communities of interacting species and may stimulate a rethinking of the spatial interpretation of interaction networks. Moreover, geographical interaction dynamics and its causes are important in our efforts to anticipate effects of large-scale changes, such as anthropogenic disturbances.

Beta diversity of plant-pollinator networks and the spatial turnover of pairwise interactions.

Interactions between species form complex networks that vary across space and time. Even without spatial or temporal constraints mutualistic pairwise interactions may vary, or rewire, across space but this variability is not well understood. Here, we quantify the beta diversity of species and interactions and test factors influencing the probability of turnover of pairwise interactions across space. We ask: 1) whether beta diversity of plants, pollinators, and interactions follow a similar trend across space, and 2) which interaction properties and site characteristics are related to the probability of turnover of pairwise interactions. Geographical distance was positively correlated with plant and interaction beta diversity. We find that locally frequent interactions are more consistent across space and that local flower abundance is important for the realization of pairwise interactions. While the identity of pairwise interactions is highly variable across space, some species-pairs form interactions that are locally frequent and spatially consistent. Such interactions represent cornerstones of interacting communities and deserve special attention from ecologists and conservation planners alike.

Establishing macroecological trait datasets: digitalization, extrapolation, and validation of diet preferences in terrestrial mammals worldwide.

Ecological trait data are essential for understanding the broad-scale distribution of biodiversity and its response to global change. For animals, diet represents a fundamental aspect of species' evolutionary adaptations, ecological and functional roles, and trophic interactions. However, the importance of diet for macroevolutionary and macroecological dynamics remains little explored, partly because of the lack of comprehensive trait datasets. We compiled and evaluated a comprehensive global dataset of diet preferences of mammals ("MammalDIET"). Diet information was digitized from two global and cladewide data sources and errors of data entry by multiple data recorders were assessed. We then developed a hierarchical extrapolation procedure to fill-in diet information for species with missing information. Missing data were extrapolated with information from other taxonomic levels (genus, other species within the same genus, or family) and this extrapolation was subsequently validated both internally (with a jack-knife approach applied to the compiled species-level diet data) and externally (using independent species-level diet information from a comprehensive continentwide data source). Finally, we grouped mammal species into trophic levels and dietary guilds, and their species richness as well as their proportion of total richness were mapped at a global scale for those diet categories with good validation results. The success rate of correctly digitizing data was 94%, indicating that the consistency in data entry among multiple recorders was high. Data sources provided species-level diet information for a total of 2033 species (38% of all 5364 terrestrial mammal species, based on the IUCN taxonomy). For the remaining 3331 species, diet information was mostly extrapolated from genus-level diet information (48% of all terrestrial mammal species), and only rarely from other species within the same genus (6%) or from family level (8%). Internal and external validation showed that: (1) extrapolations were most reliable for primary food items; (2) several diet categories ("Animal", "Mammal", "Invertebrate", "Plant", "Seed", "Fruit", and "Leaf") had high proportions of correctly predicted diet ranks; and (3) the potential of correctly extrapolating specific diet categories varied both within and among clades. Global maps of species richness and proportion showed congruence among trophic levels, but also substantial discrepancies between dietary guilds. MammalDIET provides a comprehensive, unique and freely available dataset on diet preferences for all terrestrial mammals worldwide. It enables broad-scale analyses for specific trophic levels and dietary guilds, and a first assessment of trait conservatism in mammalian diet preferences at a global scale. The digitalization, extrapolation and validation procedures could be transferable to other trait data and taxa.

Downscaling pollen-transport networks to the level of individuals.

Most plant-pollinator network studies are conducted at species level, whereas little is known about network patterns at the individual level. In fact, nodes in traditional species-based interaction networks are aggregates of individuals establishing the actual links observed in nature. Thus, emergent properties of interaction networks might be the result of mechanisms acting at the individual level. Pollen loads carried by insect flower visitors from two mountain communities were studied to construct pollen-transport networks. For the first time, these community-wide pollen-transport networks were downscaled from species-species (sp-sp) to individuals-species (i-sp) in order to explore specialization, network patterns and niche variation at both interacting levels. We used a null model approach to account for network size differences inherent to the downscaling process. Specifically, our objectives were (i) to investigate whether network structure changes with downscaling, (ii) to evaluate the incidence and magnitude of individual specialization in pollen use and (iii) to identify potential ecological factors influencing the observed degree of individual specialization. Network downscaling revealed a high specialization of pollinator individuals, which was masked and unexplored in sp-sp networks. The average number of interactions per node, connectance, interaction diversity and degree of nestedness decreased in i-sp networks, because generalized pollinator species were composed of specialized and idiosyncratic conspecific individuals. An analysis with 21 pollinator species representative of two communities showed that mean individual pollen resource niche was only c. 46% of the total species niche. The degree of individual specialization was associated with inter- and intraspecific overlap in pollen use, and it was higher for abundant than for rare species. Such niche heterogeneity depends on individual differences in foraging behaviour and likely has implications for community dynamics and species stability. Our findings highlight the importance of taking interindividual variation into account when studying higher-order structures such as interaction networks. We argue that exploring individual-based networks will improve our understanding of species-based networks and will enhance the link between network analysis, foraging theory and evolutionary biology.

Strong impact of temporal resolution on the structure of an ecological network.

Most ecological networks are analysed as static structures, where all observed species and links are present simultaneously. However, this is over-simplified, because networks are temporally dynamical. We resolved an arctic, entire-season plant-flower visitor network into a temporal series of 1-day networks and compared the properties with its static equivalent based on data pooled over the entire season. Several properties differed. The nested link pattern in the static network was blurred in the dynamical version, because the characteristic long nestedness tail of flower-visitor specialists got stunted in the dynamical networks. This tail comprised a small food web of pollinators, parasitoids and hyper-parasitoids. The dynamical network had strong time delays in the transmission of direct and indirect effects among species. Twenty percent of all indirect links were impossible in the dynamical network. Consequently, properties and thus also robustness of ecological networks cannot be deduced from the static topology alone.

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.