The number of interactions with flower visitor species differs considerably among insect pollinated plants. Knowing the causes for this variation is central to the conservation of single species as well as whole plant Á/flower visitor communities. Species specific constraints on flower visitor numbers are seldom investigated at the community level. In this study we tested whether flower size parameters set constraints on the morphology of the potential nectar feeding visitors and thus determine the number of visitor species. We studied three possible constraints: the depth and width of tubular structures hiding the nectar (nectar holder depth and width) and the size of flower parts that visitors can land on (size of the alighting place). In addition we assess the role of flower abundance on this relationship. We hypothesized that the stronger size constraints and the smaller flower abundance, the smaller the number of visitor species will be. Our study of a Mediterranean plant Á/flower visitor community revealed that nectar holder depth, nectar holder width and number of flowers explained 71% of the variation in the number of visitor species. The size of the alighting place did not restrict the body length of the visitors and was not related to visitor species number. In a second step of the analyses we calculated for each plant species the potential number of visitors by determining for each insect species of the local visitor pool whether it passed the morphological limits set by the plant. These potential numbers were highly correlated with the observed numbers (r 2 0/0.5, p B/0.001).For each plant species we tested whether the observed visitors were a random selection out of these potential visitors by comparing the mean of the observed and expected proboscis length distributions. For most plant species the observed mean was not significantly different from the random means. Our findings shed light on the way plant Á/flower visitor networks are structured. Knowing the constraints on interaction patterns will be an important prerequisite to formulate realistic null models and understand patterns of resource partitioning as well as coevolutionary processes.M. Stang, P. G. L. Klinkhamer and E. van der Meijden, Institute of Biology Leiden, Leiden Univ., PO Box 9516, NL-2300 RA Leiden, the Netherlands (stang@ rulsfb.leidenuniv.nl).Plants pollinated by animals differ greatly in the number of interactions with visitor species, varying from one to more than hundred animal species (Moldenke 1975, Jordano 1987, Ellis and Ellis-Adam 1993, Waser et al. 1996. The mechanisms leading to this variation are still poorly understood (Johnson and Steiner 2000). Especially the importance of species specific constraints on this variation has seldom been investigated at the community level (Waser et al. 1996, Vazquez 2005. In order to illustrate the role of constraints, we will use traits that are thought to have an important impact on flower visitors even if they are rarely tested as a factor determining the number of visitor spe...

The results suggest that in addition to size thresholds and species abundances, size distributions are important for understanding interaction patterns in plant-pollinator webs. It is likely that the understanding will be improved further by characterizing for entire communities how nectar production of flowers and energetic requirements of pollinators covary with size, and how sampling methods influence the observed interaction patterns.

Summary1. Biotic interactions do not occur in isolation but are imbedded in a network of species interactions. Network analysis facilitates the compilation and understanding of the complexity found in natural ecosystems and is a powerful tool to reveal information on the degree of specialization of the interacting partners and their niches. The indices measuring these properties are based on qualitative or quantitative observations of interactions between partners from different trophic levels, which informs about the structure of network patterns, but not about the underlying mechanisms. Functional traits may control the interaction strength between partners and also the (micro-) structure of networks. Here, we ask whether flower visitors specialize on certain plant traits and how this trait specialization contributes to niche partitioning and interaction partner diversity. 2. We introduce two novel statistical approaches suited to evaluate the dimension of the realized niche and to analyse which traits determine niches. As basis for our analysis, we measured 10 quantitative flower traits and evaluated whether 31 arthropod taxa i visited flowers displaying only subsets of the available trait characteristics, indicating a specialization on these traits by narrow trait-widths 〈S i 〉. The product of 10 trait-and species-specific trait-widths 〈S i 〉 was defined as trait-volume V i (expansion of a n-dimensional hypervolume) occupied by each taxon i. These indices are applicable beyond flower-visitor interactions to quantify realized niches based on various biotic and abiotic factors. 3. Each flower visitor species showed some degree of specialization to a unique set of flower traits (realized niche). Overall, our data suggested a hierarchical sequence of flower traits influencing the flower visitors' behaviour and thus network structure: flowering phenology was found to have the strongest effect, followed by flower height, nectar-tube depth and floral reflectance. Less important were pollen-mass/flower, sugar/flower, anther position, phylogeny, display size and abundance. 4. The species-specific specialization on traits suggests that plant communities with more diverse floral niches may sustain a larger number of flower visitors with non-redundant fundamental niches. Our study and statistical approach provide a basis for a better understanding of how plant traits shape interactions between flowers and their visitors and thus network structure.

Co-flowering plant species commonly share flower visitors, and thus have the potential to influence each other's pollination. In this study we analysed 750 quantitative plant-pollinator networks from 28 studies representing diverse biomes worldwide. We show that the potential for one plant species to influence another indirectly via shared pollinators was greater for plants whose resources were more abundant (higher floral unit number and nectar sugar content) and more accessible. The potential indirect influence was also stronger between phylogenetically closer plant species and was independent of plant geographic origin (native vs. non-native). The positive effect of nectar sugar content and phylogenetic proximity was much more accentuated for bees than for other groups. Consequently, the impact of these factors depends on the pollination mode of plants, e.g. bee or fly pollinated. Our findings may help predict which plant species have the greatest importance in the functioning of plant-pollination networks.

A recently discovered feature of plant-flower visitor webs is the asymmetric specialization of the interaction partners: specialized plants interact mainly with generalized flower visitors and specialized flower visitors mainly with generalized plants. Little is known about the factors leading to this asymmetry and their consequences for the extinction risk of species. Previous studies have proposed random interactions proportional to species abundance as an explanation. However, the simulation models used in these studies did not include potential biological constraints. In the present study, we tested the potential role of both morphological constraints and species abundance in promoting asymmetric specialization. We compared actual field data of a Mediterranean plant-flower visitor web with predictions of Monte Carlo simulations including different combinations of the potential factors structuring the web. Our simulations showed that both nectar-holder depth and abundance were able to produce asymmetry; but that the expected degree of asymmetry was stronger if based on both. Both factors can predict the number of interaction partners, but only nectar-holder depth was able to predict the degree of asymmetry of a certain species. What is more, without the size threshold the influence of abundance would disappear over time. Thus, asymmetric specialization seems to be the result of a size threshold and, only among the allowed interactions above this size threshold, a result of random interactions proportional to abundance. The simulations also showed that asymmetric specialization could not be the reason that the extinction risk of specialists and generalists is equalized, as suggested in the literature. In asymmetric webs specialists clearly had higher short-term extinction risks. In fact, primarily generalist visitors seem to profit from asymmetric specialization. In our web, specialists were less abundant than generalists. Therefore, including abundance in the simulation models increased the difference between specialists and generalists even more.

A long‐standing question in ecology is how species interactions are structured within communities. Although evolutionary theory predicts close size matching between floral nectar tube depth and pollinator proboscis length of interacting species, such size matching has seldom been shown and explained in multispecies assemblages. Here, we investigated the degree of size matching among Asteraceae and their pollinators and its relationship with foraging efficiency. The majority of pollinators, especially Hymenoptera, choose plant species on which they had high foraging efficiencies. When proboscides were shorter than nectar tubes, foraging efficiency rapidly decreased because of increased handling time. When proboscides were longer than nectar tubes, a decreased nectar reward rather than an increased handling time made shallow flowers more inefficient to visit. Altogether, this led to close size matching. Overall, our results show the importance of nectar reward and handling time as drivers of plant–pollinator network structure.

Identifying the determinants of biological interactions in mutualistic networks is key to understanding the rules that govern the organization of biodiversity. We used structural equation modeling and dissimilarities in nine ecological variables to investigate community processes underlying the turnover of species and their interaction frequencies (interaction pattern) among highly resolved plant-pollinator networks. Floral and pollinator community composition, i.e., species identities and their abundances, were strong determinants of the microstructure of pairwise interactions among the networks, explaining almost 69% of their variation. Flower and pollinator traits were directly related to interaction patterns, but were partly masked in the model by shared variance with community composition. Time of year and geographic location, floral and pollinator abundances independent of species identity, and relative abundance of exotic flowers had indirect and relatively weak effects on interaction patterns. Our analyses lead to precise predictions about the processes behind the interaction patterns in mutualistic networks. Future understanding of these processes will be aided by studies that evaluate these predictions experimentally at the network level.

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