The role of pollination effectiveness on the attributes of interaction networks: from floral visitation to plant fitness (original) (raw)
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Interaction strength in plant-pollinator networks: Are we using the right measure?
PLOS ONE
Understanding how ecological networks are assembled is important because network structure reflects ecosystem functioning and stability. Quantitative network analysis incorporates measures of interaction strength as an estimate of the magnitude of the effect of interaction partners on one another. Most plant-pollinator network studies use frequency of interaction between individual pollinators and individual plants (encounter) as a surrogate of interaction strength. However, the number of flowers visited per encounter may strongly vary among pollinator and plant species, and therefore not all encounters are quantitatively equivalent. We sampled plant-pollinator interactions in a Mediterranean scrubland and tested whether using a measure of interaction strength based on the number of flowers visited resulted in changes in species (species strength, interaction species asymmetry, specialization) and network descriptors (nestedness, H2', interaction evenness, plant generality, pollinator generality) compared to the encounter-based measure. Several species (including some of the most abundant ones) showed important changes in species descriptors, notably in specialization. These changes were especially important in plant species with large floral displays, which became less specialized with the visit-based measure of interaction strength. At the network level we found significant changes in all properties analysed. With the encounterbased approach plant generality was much higher than pollinator generality (high specialization asymmetry between trophic levels). However, with the visit-based approach plant generality was greatly reduced so that plants and pollinators had similar levels of generalization. Interaction evenness also decreased strongly with the visit-based approach. We conclude that accounting for the number of flowers visited per encounter provides a more ecologically relevant measure of interaction strength. Our results have important implications for the stability of pollination networks and the evolution of plant-pollinator interactions. The use of a visit-based approach is especially important in studies relating interaction network structure and ecosystem function (pollination and/or exploitation of floral resources).
Facilitation between plants shapes pollination networks
Plants cluster together and interact among themselves and with other organisms, with fundamental consequences for biodiversity and ecological networks. However, linkages between interacting plants and plants interacting with mutualists are poorly understood in real-world ecosystems. Here, we report results of a field removal experiment with natural plant communities where we compared networks of pollinators interacting with dominant plant species, here referred to as foundation species, and their associated beneficiary plant species growing in clusters, with networks of pollinators interacting with the same foundation and beneficiary species growing alone. We tested the hypothesis that the plant--pollinator networks in multispecies clusters of foundation and beneficiary plant species are more nested and robust than the sum of networks in plant clusters containing foundation species or beneficiary species growing separately. We found that pollinator diversity and plant--pollinator ne...
Plant interactions shape pollination networks via nonadditive effects
Ecology
Plants grow in communities where they interact with other plants and with other living organisms such as pollinators. On the one hand, studies of plant-plant interactions rarely consider how plants interact with other trophic levels such as pollinators. On the other, studies of plant-animal interactions rarely deal with interactions within trophic levels such as plant-plant competition and facilitation.
Acta Oecologica, 2020
Most plant-pollinator networks are based on observations of contact between an insect and a flower in the field. Despite significant sampling efforts, some links are easier to report, while others remain unobserved. Therefore, visit-based networks represent a subsample of possible interactions in which the ignored part is variable. Pollen is a natural marker of insect visits to flowers. The identification of pollen found on insect bodies can be used as an alternative method to study plant-pollinator interactions, with a potentially lower risk of bias than the observation of visits, since it increases the number of interactions in the network. Here we compare plant-pollinator networks constructed (i) from direct observation of pollinator visits and (ii) from identification of pollen found on the same insects. We focused on three calcareous grasslands in France, with different plant and pollinator species diversities. Since pollen identification always yields richer, more connected networks, we focused our comparisons on sampling bias at equal network connectance. To do so, we first compared network structures with an analysis of latent blocks and motifs. We then compared species roles between both types of networks with an analysis of specialization and species positions within motifs. Our results suggest that the sampling from observations of insect visits does not lead to the construction of a network intrinsically different from the one obtained using pollen found on insect bodies, at least when field sampling strives to be exhaustive. Most of the significant differences are found at the species level, not at the network structure level, with singleton species accounting for a respectable fraction of these differences. Overall, this suggests that recording plant-pollinator interactions from pollinator visit observation does not provide a biased picture of the network structure, regardless of species richness; however, it provided less information on species roles than the pollen-based network.
Pollinators visit related plant species across 29 plant-pollinator networks
Ecology and Evolution, 2014
Understanding the evolution of specialization in host plant use by pollinators is often complicated by variability in the ecological context of specialization. Flowering communities offer their pollinators varying numbers and proportions of floral resources, and the uniformity observed in these floral resources is, to some degree, due to shared ancestry. Here, we find that pollinators visit related plant species more so than expected by chance throughout 29 plant-pollinator networks of varying sizes, with "clade specialization" increasing with community size. As predicted, less versatile pollinators showed more clade specialization overall. We then asked whether this clade specialization varied with the ratio of pollinator species to plant species such that pollinators were changing their behavior when there was increased competition (and presumably a forced narrowing of the realized niche) by examining pollinators that were present in at least three of the networks. Surprisingly, we found little evidence that variation in clade specialization is caused by pollinator species changing their behavior in different community contexts, suggesting that clade specialization is observed when pollinators are either restricted in their floral choices due to morphological constraints or innate preferences. The resulting pollinator sharing between closely related plant species could result in selection for greater pollinator specialization.
Linkage Rules for Plant–Pollinator Networks: Trait Complementarity or Exploitation Barriers?
PLoS Biology, 2007
Recent attempts to examine the biological processes responsible for the general characteristics of mutualistic networks focus on two types of explanations: nonmatching biological attributes of species that prevent the occurrence of certain interactions (''forbidden links''), arising from trait complementarity in mutualist networks (as compared to barriers to exploitation in antagonistic ones), and random interactions among individuals that are proportional to their abundances in the observed community (''neutrality hypothesis''). We explored the consequences that simple linkage rules based on the first two hypotheses (complementarity of traits versus barriers to exploitation) had on the topology of plant-pollination networks. Independent of the linkage rules used, the inclusion of a small set of traits (two to four) sufficed to account for the complex topological patterns observed in real-world networks. Optimal performance was achieved by a ''mixed model'' that combined rules that link plants and pollinators whose trait ranges overlap (''complementarity models'') and rules that link pollinators to flowers whose traits are below a pollinator-specific barrier value (''barrier models''). Deterrence of floral parasites (barrier model) is therefore at least as important as increasing pollination efficiency (complementarity model) in the evolutionary shaping of plant-pollinator networks.
Plant-pollinator networks: adding the pollinator’s perspective
Ecology Letters, 2009
Pollination network studies are based on pollinator surveys conducted on focal plants. This plant-centred approach provides insufficient information on flower visitation habits of rare pollinator species, which are the majority in pollinator communities. As a result, pollination networks contain very high proportions of pollinator species linked to a single plant species (extreme specialists), a pattern that contrasts with the widely accepted view that plant-pollinator interactions are mostly generalized. In this study of a Mediterranean scrubland community in NE Spain we supplement data from an intensive field survey with the analysis of pollen loads carried by pollinators. We observed 4265 contacts corresponding to 19 plant and 122 pollinator species. The addition of pollen data unveiled a very significant number of interactions, resulting in important network structural changes. Connectance increased 1.43-fold, mean plant connectivity went from 18.5 to 26.4, and mean pollinator connectivity from 2.9 to 4.1. Extreme specialist pollinator species decreased 0.6-fold, suggesting that ecological specialization is often overestimated in plant-pollinator networks. We expected a greater connectivity increase in rare species, and consequently a decrease in the level of asymmetric specialization. However, new links preferentially attached to already highly connected nodes and, as a result, both nestedness and centralization increased. The addition of pollen data revealed the existence of four clearly defined modules that were not apparent when only field survey data were used. Three of these modules had a strong phenological component. In comparison to other pollination webs, our network had a high proportion of connector links and species. That is, although significant, the four modules were far from isolated.
Interactions for pollinator visitation and their consequences for reproduction in a plant community
Acta Oecologica, 2012
Competition and facilitation in species interactions attract much attention in ecology, but their relative importance has seldom been evaluated in a community context. We assessed competitive and facilitative interactions for pollinator visitation among co-flowering species in a plant community, investigated the subsequent consequences for plant reproduction, and investigated whether effects could be trait-based. We removed the flowers of two species attractive to pollinators, in two separate experiments and assessed the effects on pollinator visitation rates and components of reproductive success in 11 coflowering focal herb species. Overall, most focal species appear not to interact with the removal species with respect to pollinator visitation and subsequent reproduction (neutral interactions). Three focal species in the community had significantly higher reproductive responses (fruit production and seed weight) in the presence of the attractive removal species (facilitative interactions), but species interaction effects were less pronounced in species' flower visitation rates. A community-wide metaanalysis demonstrated that the two experiments did not have a significant effect on either facilitation or competition, and that there was no overall correlation between effect sizes for visitation and reproduction. Based on species-specific responses, it seems likely that floral traits such as similar flower colors contribute to interspecific facilitation of pollinator visitation and, in particular, that high pollinator dependence for plant reproduction, and associated pollen limitation, may contribute to subsequent interaction effects on reproduction in the focal species.
Structure and dynamics of pollination networks: the role of alien plants
Oikos, 2009
Research on ecological communities, and plant–pollinator mutualistic networks in particular, has increasingly benefited from the theory and tools of complexity science. Nevertheless, up to now there have been few attempts to investigate the interplay between the structure of real pollination networks and their dynamics. This study is one of the first contributions to explore this issue. Biological invasions, of major concern for conservation, are also poorly understood from the perspective of complex ecological networks. In this paper we assess the role that established alien species play within a host community by analyzing the temporal changes in structural network properties driven by the removal of non-native plants. Three topological measures have been used to represent the most relevant structural properties for the stability of ecological networks: degree distribution, nestedness, and modularity. Therefore, we investigate for a detailed pollination network, 1) how its dynamics, represented as changes in species abundances, affect the evolution of its structure, 2) how topology relates to dynamics focusing on long-term species persistence; and 3) how both structure and dynamics are affected by the removal of alien plant species. Network dynamics were simulated by means of a stochastic metacommunity model. Our results showed that established alien plants are important for the persistence of the pollination network and for the maintenance of its structure. Removal of alien plants decreased the likelihood of species persistence. On the other hand, both the full network and the subset native network tended to lose their structure through time. Nevertheless, the structure of the full network was better preserved than the structure of the network without alien plants. Temporal topological shifts were evident in terms of degree distribution, nestedness, and modularity. However the effects of removing alien plants were more pronounced for degree distribution and modularity of the network. Therefore, elimination of alien plants affected the evolution of the architecture of the interaction web, which was closely related to the higher species loss found in the network where alien plants were removed.
Does the sociality of pollinators shape the organisation of pollination networks?
Oikos
A striking structural pattern of pollination networks is the presence of a few highly connected species which has implications for ecological and evolutionary processes that create and maintain diversity. To understand the structure and dynamics of pollination networks we need to know which mechanisms allow the emergence of highly connected species. We investigate whether social pollinator species are highly connected in pollination networks, and whether network structure is affected by the presence of high proportions of social pollinator species. Social insects are abundant, with long activity periods and, at the highest level of social organisation, specialised foraging castes. These three attributes are likely to increase the number of interactions of social species and, consequently, their role in pollination networks. We find that social species have, on average, more prominent network roles than solitary species, a possible mechanism being the individual-rich colonies of social insects. However, when accounting for the shared evolutionary history of pollinators, sociality is only associated with highly interactive roles in Apidae. For apid bees, our structural equation analysis shows that the effect of sociality on species network roles is an indirect result of their high levels of interaction frequency. Despite the relative importance of sociality at a species-level, an increasing proportion of social species in pollination networks did not affect overall network structure. Our results suggest that behavioural traits may shape patterns of interaction of individual species but not the network-level organisation of species interactions. Instead, network structure appears to be determined by more general aspects of ecological systems such as interaction intimacy, patterns of niche overlap, and species abundance distributions.