Contrasting Partners' Traits of Generalized and Specialized Species in Flower-Visitation Networks.

Much ecological research has focused on trying to understand why species are generalized or specialized in their interactions and how networks develop in a certain environment. It is now well known that traits such as phenology and abundance of a species are important determinants of its generalization level (i.e., number of different interactions or links to other species). Less information is available, however, on whether generalized and specialized species differ in particular traits of their interacting partners. Such partners might differ, for instance, in abundance and/or in the diversity of functional groups they belong to. Moreover, species might exhibit shifts through time (e.g., flowering season) in their partners' traits, though we know close to nothing on whether these changes do indeed occur. Assessing how such network links in both types of species are established is important for a better understanding of how different types of disturbance can affect community dynamics. Using data from four quantitative flower-visitation networks and independent measures of flower availability obtained when recording interactions, we test for such differences between species which have been previously categorized according to two specialization indexes: (1) number of partners (links), also named linkage level; this is a qualitative index and (2) complementary specialization d', named here selectiveness level; this is a quantitative index. We found that: (1) species with low linkage levels mainly interact with common species in the community whereas generalized species interact with a greater heterogeneity of partner's abundances and functional richness, (2) both selective and opportunistic species (with high and low d', respectively) interact with a similarly high functional richness (number of functional groups or families) of partners, and (3) generalized species are the only ones showing shifts along the season in their partners' traits, driven by changes in community species composition. The risk of extinction in front of a disturbance is generally expected to be highest for specialized species (with few partners) and selective species (which visit non-abundant or scarce partners). However, our findings show that by linking to abundant and/or to functionally diverse partners, respectively, these species may be maintained in the community and be less vulnerable to disturbances.

Linking plant specialization to dependence in interactions for seed set in pollination networks.

Studies on pollination networks have provided valuable information on the number, frequency, distribution and identity of interactions between plants and pollinators. However, little is still known on the functional effect of these interactions on plant reproductive success. Information on the extent to which plants depend on such interactions will help to make more realistic predictions of the potential impacts of disturbances on plant-pollinator networks. Plant functional dependence on pollinators (all interactions pooled) can be estimated by comparing seed set with and without pollinators (i.e. bagging flowers to exclude them). Our main goal in this study was thus to determine whether plant dependence on current insect interactions is related to plant specialization in a pollination network. We studied two networks from different communities, one in a coastal dune and one in a mountain. For ca. 30% of plant species in each community, we obtained the following specialization measures: (i) linkage level (number of interactions), (ii) diversity of interactions, and (iii) closeness centrality (a measure of how much a species is connected to other plants via shared pollinators). Phylogenetically controlled regression analyses revealed that, for the largest and most diverse coastal community, plants highly dependent on pollinators were the most generalists showing the highest number and diversity of interactions as well as occupying central positions in the network. The mountain community, by contrast, did not show such functional relationship, what might be attributable to their lower flower-resource heterogeneity and diversity of interactions. We conclude that plants with a wide array of pollinator interactions tend to be those that are more strongly dependent upon them for seed production and thus might be those more functionally vulnerable to the loss of network interaction, although these outcomes might be context-dependent.

Invaders of pollination networks in the Galapagos Islands: emergence of novel communities.

The unique biodiversity of most oceanic archipelagos is currently threatened by the introduction of alien species that can displace native biota, disrupt native ecological interactions, and profoundly affect community structure and stability. We investigated the threat of aliens on pollination networks in the species-rich lowlands of five Galápagos Islands. Twenty per cent of all species (60 plants and 220 pollinators) in the pooled network were aliens, being involved in 38 per cent of the interactions. Most aliens were insects, especially dipterans (36%), hymenopterans (30%) and lepidopterans (14%). These alien insects had more links than either endemic pollinators or non-endemic natives, some even acting as island hubs. Aliens linked mostly to generalized species, increasing nestedness and thus network stability. Moreover, they infiltrated all seven connected modules (determined by geographical and phylogenetic constraints) of the overall network, representing around 30 per cent of species in two of them. An astonishingly high proportion (38%) of connectors, which enhance network cohesiveness, was also alien. Results indicate that the structure of these emergent novel communities might become more resistant to certain type of disturbances (e.g. species loss), while being more vulnerable to others (e.g. spread of a disease). Such notable changes in network structure as invasions progress are expected to have important consequences for native biodiversity maintenance.

The dimensionality of ecological networks.

How many dimensions (trait-axes) are required to predict whether two species interact? This unanswered question originated with the idea of ecological niches, and yet bears relevance today for understanding what determines network structure. Here, we analyse a set of 200 ecological networks, including food webs, antagonistic and mutualistic networks, and find that the number of dimensions needed to completely explain all interactions is small ( < 10), with model selection favouring less than five. Using 18 high-quality webs including several species traits, we identify which traits contribute the most to explaining network structure. We show that accounting for a few traits dramatically improves our understanding of the structure of ecological networks. Matching traits for resources and consumers, for example, fruit size and bill gape, are the most successful combinations. These results link ecologically important species attributes to large-scale community structure.