Modularity, pollination systems, and interaction turnover in plant-pollinator networks across space.

Mutualistic interaction networks have been shown to be structurally conserved over space and time while pairwise interactions show high variability. In such networks, modularity is the division of species into compartments, or modules, where species within modules share more interactions with each other than they do with species from other modules. Such a modular structure is common in mutualistic networks and several evolutionary and ecological mechanisms have been proposed as underlying drivers. One prominent explanation is the existence of pollination syndromes where flowers tend to attract certain pollinators as determined by a set of traits. We investigate the modularity of seven community level plant-pollinator networks sampled in rupestrian grasslands, or campos rupestres, in SE Brazil. Defining pollination systems as corresponding groups of flower syndromes and pollinator functional groups, we test the two hypotheses that (1) interacting species from the same pollination system are more often assigned to the same module than interacting species from different pollination systems and; that (2) interactions between species from the same pollination system are more consistent across space than interactions between species from different pollination systems. Specifically we ask (1) whether networks are consistently modular across space; (2) whether interactions among species of the same pollination system occur more often inside modules, compared to interactions among species of different pollination systems, and finally; (3) whether the spatial variation in interaction identity, i.e., spatial interaction rewiring, is affected by trait complementarity among species as indicated by pollination systems. We confirm that networks are consistently modular across space and that interactions within pollination systems principally occur inside modules. Despite a strong tendency, we did not find a significant effect of pollination systems on the spatial consistency of pairwise interactions. These results indicate that the spatial rewiring of interactions could be constrained by pollination systems, resulting in conserved network structures in spite of high variation in pairwise interactions. Our findings suggest a relevant role of pollination systems in structuring plant-pollinator networks and we argue that structural patterns at the sub-network level can help us to fully understand how and why interactions vary across space and time.

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.

Determinants of bird species richness, endemism, and island network roles in Wallacea and the West Indies: is geography sufficient or does current and historical climate matter?

Island biogeography has greatly contributed to our understanding of the processes determining species' distributions. Previous research has focused on the effects of island geography (i.e., island area, elevation, and isolation) and current climate as drivers of island species richness and endemism. Here, we evaluate the potential additional effects of historical climate on breeding land bird richness and endemism in Wallacea and the West Indies. Furthermore, on the basis of species distributions, we identify island biogeographical network roles and examine their association with geography, current and historical climate, and bird richness/endemism. We found that island geography, especially island area but also isolation and elevation, largely explained the variation in island species richness and endemism. Current and historical climate only added marginally to our understanding of the distribution of species on islands, and this was idiosyncratic to each archipelago. In the West Indies, endemic richness was slightly reduced on islands with historically unstable climates; weak support for the opposite was found in Wallacea. In both archipelagos, large islands with many endemics and situated far from other large islands had high importance for the linkage within modules, indicating that these islands potentially act as speciation pumps and source islands for surrounding smaller islands within the module and, thus, define the biogeographical modules. Large islands situated far from the mainland and/or with a high number of nonendemics acted as links between modules. Additionally, in Wallacea, but not in the West Indies, climatically unstable islands tended to interlink biogeographical modules. The weak and idiosyncratic effect of historical climate on island richness, endemism, and network roles indicates that historical climate had little effects on extinction-immigration dynamics. This is in contrast to the strong effect of historical climate observed on the mainland, possibly because surrounding oceans buffer against strong climate oscillations and because geography is a strong determinant of island richness, endemism and network roles.

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.