Consequences of plant invasions on compartmentalization and species' roles in plant-pollinator networks.

Compartmentalization-the organization of ecological interaction networks into subsets of species that do not interact with other subsets (true compartments) or interact more frequently among themselves than with other species (modules)-has been identified as a key property for the functioning, stability and evolution of ecological communities. Invasions by entomophilous invasive plants may profoundly alter the way interaction networks are compartmentalized. We analysed a comprehensive dataset of 40 paired plant-pollinator networks (invaded versus uninvaded) to test this hypothesis. We show that invasive plants have higher generalization levels with respect to their pollinators than natives. The consequences for network topology are that-rather than displacing native species from the network-plant invaders attracting pollinators into invaded modules tend to play new important topological roles (i.e. network hubs, module hubs and connectors) and cause role shifts in native species, creating larger modules that are more connected among each other. While the number of true compartments was lower in invaded compared with uninvaded networks, the effect of invasion on modularity was contingent on the study system. Interestingly, the generalization level of the invasive plants partially explains this pattern, with more generalized invaders contributing to a lower modularity. Our findings indicate that the altered interaction structure of invaded networks makes them more robust against simulated random secondary species extinctions, but more vulnerable when the typically highly connected invasive plants go extinct first. The consequences and pathways by which biological invasions alter the interaction structure of plant-pollinator communities highlighted in this study may have important dynamical and functional implications, for example, by influencing multi-species reciprocal selection regimes and coevolutionary processes.

Does the invasive Lupinus polyphyllus increase pollinator visitation to a native herb through effects on pollinator population sizes?

Invasive plants may compete with native species for abiotic factors as light, space and nutrients, and have also been shown to affect native pollination interactions. Studies have mainly focused on how invasive plants affect pollinator behaviour, i.e. attraction of pollinators to or away from native flowers. However, when an invasive plant provides resources utilized by native pollinators this could increase pollinator population sizes and thereby pollination success in natives. Effects mediated through changes in pollinator population sizes have been largely ignored in previous studies, and the dominance of negative interactions suggested by meta-analyses may therefore be biased. We investigated the impact of the invasive Lupinus polyphyllus on pollination in the native Lotus corniculatus using a study design comparing invaded and uninvaded sites before and after the flowering period of the invasive. We monitored wild bee abundance in transects, and visit rate and seed production of potted Lotus plants. Bumblebee abundance increased 3.9 times in invaded sites during the study period, whereas it was unaltered in uninvaded sites. Total visit rate per Lotus plant increased 2.1 times in invaded sites and decreased 4.4 times in uninvaded sites. No corresponding change in seed production of Lotus was found. The increase in visit rate to Lotus was driven by an increase in solitary bee visitation, whereas mainly bumblebees were observed to visit the invasive Lupinus. The mechanism by which the invasive increases pollinator visit rates to Lotus could be increased availability of other flower resources for solitary bees when bumblebees forage on Lupinus.

Evaluating sampling completeness in a desert plant-pollinator network.

1. The study of plant-pollinator interactions in a network context is receiving increasing attention. This approach has helped to identify several emerging network patterns such as nestedness and modularity. However, most studies are based only on qualitative information, and some ecosystems, such as deserts and tropical forests, are underrepresented in these data sets. 2. We present an exhaustive analysis of the structure of a 4-year plant-pollinator network from the Monte desert in Argentina using qualitative and quantitative tools. We describe the structure of this network and evaluate sampling completeness using asymptotic species richness estimators. Our goal is to assess the extent to which the realized sampling effort allows for an accurate description of species interactions and to estimate the minimum number of additional censuses required to detect 90% of the interactions. We evaluated completeness of detection of the community-wide pollinator fauna, of the pollinator fauna associated with each plant species and of the plant-pollinator interactions. We also evaluated whether sampling completeness was influenced by plant characteristics, such as flower abundance, flower life span, number of interspecific links (degree) and selectiveness in the identity of their flower visitors, as well as sampling effort. 3. We found that this desert plant-pollinator network has a nested structure and that it exhibits modularity and high network-level generalization. 4. In spite of our high sampling effort, and although we sampled 80% of the pollinator fauna, we recorded only 55% of the interactions. Furthermore, although a 64% increase in sampling effort would suffice to detect 90% of the pollinator species, a fivefold increase in sampling effort would be necessary to detect 90% of the interactions. 5. Detection of interactions was incomplete for most plant species, particularly specialists with a long flowering season and high flower abundance, or generalists with short flowering span and scant flowers. Our results suggest that sampling of a network with the same effort for all plant species is inadequate to sample interactions. 6. Sampling the diversity of interactions is labour intensive, and most plant-pollinator networks published to date are likely to be undersampled. Our analysis allowed estimating the completeness of our sampling, the additional effort needed to detect most interactions and the plant traits that influence the detection of their interactions.

Impact of alien plant invaders on pollination networks in two archipelagos.

Mutualistic interactions between plants and animals promote integration of invasive species into native communities. In turn, the integrated invaders may alter existing patterns of mutualistic interactions. Here we simultaneously map in detail effects of invaders on parameters describing the topology of both plant-pollinator (bi-modal) and plant-plant (uni-modal) networks. We focus on the invader Opuntia spp., a cosmopolitan alien cactus. We compare two island systems: Tenerife (Canary Islands) and Menorca (Balearic Islands). Opuntia was found to modify the number of links between plants and pollinators, and was integrated into the new communities via the most generalist pollinators, but did not affect the general network pattern. The plant uni-modal networks showed disassortative linkage, i.e. species with many links tended to connect to species with few links. Thus, by linking to generalist natives, Opuntia remained peripheral to network topology, and this is probably why native network properties were not affected at least in one of the islands. We conclude that the network analytical approach is indeed a valuable tool to evaluate the effect of invaders on native communities.