Stable pollination service in a generalist high Arctic community despite the warming climate.

Insects provide key pollination services in most terrestrial biomes, but this service depends on a multistep interaction between insect and plant. An insect needs to visit a flower, receive pollen from the anthers, move to another conspecific flower, and finally deposit the pollen on a receptive stigma. Each of these steps may be affected by climate change, and focusing on only one of them (e.g., flower visitation) may miss important signals of change in service provision. In this study, we combine data on visitation, pollen transport, and single-visit pollen deposition to estimate functional outcomes in the high Arctic plant-pollinator network of Zackenberg, Northeast Greenland, a model system for global warming-associated impacts in pollination services. Over two decades of rapid climate warming, we sampled the network repeatedly: in 1996, 1997, 2010, 2011, and 2016. Although the flowering plant and insect communities and their interactions varied substantially between years, as expected based on highly variable Arctic weather, there was no detectable directional change in either the structure of flower-visitor networks or estimated pollen deposition. For flower-visitor networks compiled over a single week, species phenologies caused major within-year variation in network structure despite consistency across years. Weekly networks for the middle of the flowering season emerged as especially important because most pollination service can be expected to be provided by these large, highly nested networks. Our findings suggest that pollination ecosystem service in the high Arctic is remarkably resilient. This resilience may reflect the plasticity of Arctic biota as an adaptation to extreme and unpredictable weather. However, most pollination service was contributed by relatively few fly taxa (Diptera: Spilogona sanctipauli and Drymeia segnis [Muscidae] and species of Rhamphomyia [Empididae]). If these key pollinators are negatively affected by climate change, network structure and the pollination service that depends on it would be seriously compromised.

Time-calibrated phylogenies reveal mediterranean and pre-mediterranean origin of the thermophilous vegetation of the Canary Islands.

BACKGROUND AND AIMS: The Canary Islands have strong floristic affinities with the Mediterranean Basin. One of the most characteristic and diverse vegetation belts of the archipelago is the thermophilous woodland (between 200 and 900 m.a.s.l.). This thermophilous plant community consists of many non-endemic species shared with the Mediterranean Floristic Region together with Canarian endemic species. Consequently, phytogeographic studies have historically proposed the hypothesis of an origin of the Canarian thermophilous species following the establishment of the summer-dry mediterranean climate in the Mediterranean Basin around 2.8 million years ago. METHODS: Time-calibrated phylogenies for 39 plant groups including Canarian thermophilous species were primarily analysed to infer colonization times. In particular, we used 26 previously published phylogenies together with 13 new time-calibrated phylogenies (including newly generated plastid and nuclear DNA sequence data) to assess whether the time interval between stem and crown ages of Canarian thermophilous lineages postdates 2.8 Ma. For lineages postdating this time threshold, we additionally conducted ancestral area reconstructions to infer the potential source area for colonization. KEY RESULTS: A total of 43 Canarian thermophilous lineages were identified from 39 plant groups. Both mediterranean (16) and pre-mediterranean (9) plant lineages were found. However, we failed to determine the temporal origin for 18 lineages because a stem-crown time interval overlaps with the 2.8-Ma threshold. The spatial origin of thermophilous lineages was also heterogeneous, including ancestral areas from the Mediterranean Basin (nine) and other regions (six). CONCLUSIONS: Our findings reveal an unexpectedly heterogeneous origin of the Canarian thermophilous species in terms of colonization times and mainland source areas. A substantial proportion of the lineages arrived in the Canaries before the summer-dry climate was established in the Mediterranean Basin. The complex temporal and geographic origin of Canarian thermophilous species challenges the view of the Canary Islands (and Madeira) as a subregion within the Mediterranean Floristic Region.

Ego network analysis of the trophic structure of an island land bird through 300 years of climate change and invaders.

Ego net analysis is a well-known practice in social sciences, where an ego net (EN) consists of a focal node, the ego, and its links to other nodes, called alters, and alter-alter links may also be included. An EN describes how a focal node is embedded in its interaction context. Here, I introduce EN analysis to ecology in a study of the trophic network of a sub-Antarctic land bird, Lesser Sheathbill (Chionis minor). Data originate from the sheathbill population on Marion Island in the Southern Ocean. The bird is ego and its enemies and food are alters. The EN is organized along three dimensions: habitat, interaction type, and time (from before human arrival in 1803 and until a future year 2100). Ten EN descriptors are defined, estimated, and used to track the 300 years of change in sheathbill EN structure. Since 1803, the EN has passed two major, but reversible shifts-seal exploitation in the 19th century and presence of cats from 1949 to 1991. These shifts can be read as structural changes in the sheathbill EN. In the future, a third, perhaps irreversible change is predicted, driven by climate change and a surprising, recent shift to seabird predation by House Mouse, the most detrimental of all extant invaders on Marion. In a warmer and drier future, the mouse will proliferate, and if this forces seabirds to abandon the island, their accumulation of detritus runs dry, starving a rich invertebrate detritivore fauna, which also is a key food source to sheathbills. These detritivores together with plants have also constituted the main food sources of mice. The EN descriptors quantify that story. In the future, these events may lead to a collapse of the island ecosystem, including extinction of the sheathbill-unless plans for mouse eradication are implemented.

Evaluating competition for forage plants between honey bees and wild bees in Denmark.

A recurrent concern in nature conservation is the potential competition for forage plants between wild bees and managed honey bees. Specifically, that the highly sophisticated system of recruitment and large perennial colonies of honey bees quickly exhaust forage resources leading to the local extirpation of wild bees. However, different species of bees show different preferences for forage plants. We here summarize known forage plants for honey bees and wild bee species at national scale in Denmark. Our focus is on floral resources shared by honey bees and wild bees, with an emphasis on both threatened wild bee species and foraging specialist species. Across all 292 known bee species from Denmark, a total of 410 plant genera were recorded as forage plants. These included 294 plant genera visited by honey bees and 292 plant genera visited by different species of wild bees. Honey bees and wild bees share 176 plant genera in Denmark. Comparing the pairwise niche overlap for individual bee species, no significant relationship was found between their overlap and forage specialization or conservation status. Network analysis of the bee-plant interactions placed honey bees aside from most other bee species, specifically the module containing the honey bee had fewer links to any other modules, while the remaining modules were more highly inter-connected. Despite the lack of predictive relationship from the pairwise niche overlap, data for individual species could be summarized. Consequently, we have identified a set of operational parameters that, based on a high foraging overlap (>70%) and unfavorable conservation status (Vulnerable+Endangered+Critically Endangered), can guide both conservation actions and land management decisions in proximity to known or suspected populations of these species.

Diet composition of the lizard Podarcis lilfordi (Lacertidae) on 2 small islands: an individual-resource network approach.

Despite it is widely accepted that intrapopulation variation is fundamental to ecological and evolutionary processes, this level of information has only recently been included into network analysis of species/population interactions. When done, it has revealed non-random patterns in the distribution of trophic resources. Nestedness in resource use among individuals is the most recurrent observed pattern, often accompanied by an absence of modularity, but no previous studies examine bipartite modularity. We use network analysis to describe the diet composition of the Balearic endemic lizard Podarcis lilfordi in 2 islets at population and individual levels, based on the occurrence of food items in fecal samples. Our objectives are to 1) compare niche structure at both levels, 2) characterize niche partition using nestedness and modularity, and 3) assess how size, sex, season, and spatial location influence niche structure. At population-level niche width was wide, but narrow at the level of the individual. Both islet networks were nested, indicating similar ranking of the food preferences among individuals, but also modular, which was partially explained by seasonality. Sex and body size did not notably affect diet composition. Large niche overlap and therefore possibly relaxed competition were observed among females in one of the islets and during spring on both islets. Likewise, higher modularity in autumn suggests that higher competition could lead to specialization in both populations, because resources are usually scarce in this season. The absence of spatial location influence on niche might respond to fine-grained spatio-temporally distribution of food resources. Behavioral traits, not included in this study, could also influence resource partitioning.

Potential role of lava lizards as pollinators across the Galápagos Islands.

Lizards have been reported as important pollinators on several oceanic islands. Here we evaluate the potential role of Galápagos lava lizards (Microlophus spp.) as pollinators across their radiation. Over 3 years, we sampled pollen transport by 9 lava lizard species on the 10 islands where they are present, including 7 single-island endemics. Overall, only 25 of 296 individuals sampled (8.4%) transported pollen of 10 plant species, the most common being Prosopis juliflora, Exodeconus miersii, Sesuvium sp. and Cordia leucophlyctis. At least 8 of these plant species were native, and none were confirmed as introduced to the archipelago. Despite the low overall proportion of individuals carrying pollen, this was observed in 7 of the nine lizard species, and on 8 of the ten main islands (Española, Fernandina, Floreana, Isabela, Marchena, Pinta, Santa Cruz and Santiago), suggesting that this is a widespread interaction. The results reported here support the potential role of lava lizards as pollinators across their radiation, although they may represent a relatively modest contribution when compared with birds and insects. However, we cannot discard that lizards may be ecologically significant for particular plant species and ecosystems given the specific climatic condition and functional diversity of each island.

Small size does not restrain frugivory and seed dispersal across the evolutionary radiation of Galápagos lava lizards.

Frugivory in lizards is often assumed to be constrained by body size; only large individuals are considered capable of consuming fruits, with the potential of acting as seed dispersers. However, only one previous study has tested the correlation of frugivory with body and head size at an archipelago scale across closely related species. All nine lava lizards (Microlophus spp.) were studied on the eleven largest Galápagos islands from 2010 to 2016 to investigate whether frugivory is related to body and head size. We also tested whether fruit abundance influences fruit consumption and explored the effect of seed ingestion on seedling emergence time and percentage. Our results showed that across islands, lava lizards varied considerably in size (64-102 mm in mean snout-vent length) and level of frugivory (1-23%, i.e., percentage of droppings with seeds). However, level of frugivory was only weakly affected by size as fruit consumption was also common among small lizards. Lava lizards consumed fruits throughout the year and factors other than fruit abundance may be more important drivers of fruit selection (e.g., fruit size, energy content of pulp). From 2,530 droppings, 1,714 seeds of at least 61 plant species were identified, 76% of the species being native to the Galápagos. Most seeds (91%) showed no external structural damage. Seedling emergence time (44 versus 118 days) and percentage (20% versus 12%) were enhanced for lizard-ingested seeds compared to control (uningested) fruits. De-pulping by lizards (i.e., removal of pulp with potential germination inhibitors) might increase the chances that at least some seeds find suitable recruitment conditions. We concluded that lizards are important seed dispersers throughout the year and across the whole archipelago, regardless of body size.

Divergence in floral trait preferences between nonflower-specialized birds and insects on the Galápagos.

PREMISE OF THE STUDY: The characteristic scarcity of insects on remote oceanic islands has driven nonflower-specialized vertebrates to broaden their trophic niches and explore floral resources. From our previous studies in the Galápagos, we know that native insectivorous and frugivorous birds visit a wide range of entomophilous flowers and can also act as effective pollinators. Here, we tested whether opportunistic Galápagos birds show any preference for specific floral traits, and if so, this preference differs from that of insects. METHODS: Sixteen floral morphology and nectar traits of 26 native species were studied, as well as the frequency with which they are visited by birds and insects. Nonmetric multidimensional scaling (NMDS) was used to evaluate the distribution of flower traits values along two main dimensions and measure the similarity between the plants visited mostly by birds versus those by insects. KEY RESULTS: NMDS of floral traits resulted in two species groups: (1) bell-shaped, white flowers with wider corollas at nectary level and higher nectar volume, associated with high bird visitation rates; and (2) bowl and tubular-shaped flowers with narrower corollas at nectary level and lower nectar volume, associated with high insect visitation rates. CONCLUSIONS: Despite the divergence in floral trait preferences between opportunistic Galápagos birds and insects, bird-visited flowers display mixed traits not fitting the classical ornithophilous syndrome. This finding is compatible with the existence of a transitional or bet-hedging phenotype between insect and bird visitors and underscores the importance of coevolution and floral diversification in nonspecialized plant-visitor interactions.

Analysing ecological networks of species interactions.

Network approaches to ecological questions have been increasingly used, particularly in recent decades. The abstraction of ecological systems - such as communities - through networks of interactions between their components indeed provides a way to summarize this information with single objects. The methodological framework derived from graph theory also provides numerous approaches and measures to analyze these objects and can offer new perspectives on established ecological theories as well as tools to address new challenges. However, prior to using these methods to test ecological hypotheses, it is necessary that we understand, adapt, and use them in ways that both allow us to deliver their full potential and account for their limitations. Here, we attempt to increase the accessibility of network approaches by providing a review of the tools that have been developed so far, with - what we believe to be - their appropriate uses and potential limitations. This is not an exhaustive review of all methods and metrics, but rather, an overview of tools that are robust, informative, and ecologically sound. After providing a brief presentation of species interaction networks and how to build them in order to summarize ecological information of different types, we then classify methods and metrics by the types of ecological questions that they can be used to answer from global to local scales, including methods for hypothesis testing and future perspectives. Specifically, we show how the organization of species interactions in a community yields different network structures (e.g., more or less dense, modular or nested), how different measures can be used to describe and quantify these emerging structures, and how to compare communities based on these differences in structures. Within networks, we illustrate metrics that can be used to describe and compare the functional and dynamic roles of species based on their position in the network and the organization of their interactions as well as associated new methods to test the significance of these results. Lastly, we describe potential fruitful avenues for new methodological developments to address novel ecological questions.

Disentangling the role of floral sensory stimuli in pollination networks.

Despite progress in understanding pollination network structure, the functional roles of floral sensory stimuli (visual, olfactory) have never been addressed comprehensively in a community context, even though such traits are known to mediate plant-pollinator interactions. Here, we use a comprehensive dataset of floral traits and a novel dynamic data-pooling methodology to explore the impacts of floral sensory diversity on the structure of a pollination network in a Mediterranean scrubland. Our approach tracks transitions in the network behaviour of each plant species throughout its flowering period and, despite dynamism in visitor composition, reveals significant links to floral scent, and/or colour as perceived by pollinators. Having accounted for floral phenology, abundance and phylogeny, the persistent association between floral sensory traits and visitor guilds supports a deeper role for sensory bias and diffuse coevolution in structuring plant-pollinator networks. This knowledge of floral sensory diversity, by identifying the most influential phenotypes, could help prioritize efforts for plant-pollinator community restoration.

Disclosing the double mutualist role of birds on Galápagos.

Life on oceanic islands deviate in many ways from that on the mainland. Their biodiversity is relatively poor and some groups are well-represented, others not, especially not insects. A scarcity of insects forces birds to explore alternative food, such as nectar and fruit. In this way, island birds may pollinate and disperse seed to an extent unseen on any mainland; they may even first consume floral resources of a plant species and then later harvest the fruit of the same species. Through this biotic reuse, they may act as double mutualists. The latter have never been studied at the level of the network, because they are traditionally considered rare. We sampled pollination and seed-dispersal interactions on Galápagos and constructed a plant-bird mutualism network of 108 plant (12% being double mutualists) and 21 bird species (48% being double mutualists), and their 479 interactions, being either single (95%) or double mutualisms (5%). Double mutualists constitute the core in the pollination-dispersal network, coupling the two link types together. They may also initiate positive feedbacks (more pollination leading to more dispersal), which theoretically are known to be unstable. Thus, double mutualisms may be a necessary, but risky prerequisite to the survival of island biodiversity.

Ecological and evolutionary legacy of megafauna extinctions.

For hundreds of millions of years, large vertebrates (megafauna) have inhabited most of the ecosystems on our planet. During the late Quaternary, notably during the Late Pleistocene and the early Holocene, Earth experienced a rapid extinction of large, terrestrial vertebrates. While much attention has been paid to understanding the causes of this massive megafauna extinction, less attention has been given to understanding the impacts of loss of megafauna on other organisms with whom they interacted. In this review, we discuss how the loss of megafauna disrupted and reshaped ecological interactions, and explore the ecological consequences of the ongoing decline of large vertebrates. Numerous late Quaternary extinct species of predators, parasites, commensals and mutualistic partners were associated with megafauna and were probably lost due to their strict dependence upon them (co-extinctions). Moreover, many extant species have megafauna-adapted traits that provided evolutionary benefits under past megafauna-rich conditions, but are now of no or limited use (anachronisms). Morphological evolution and behavioural changes allowed some of these species partially to overcome the absence of megafauna. Although the extinction of megafauna led to a number of co-extinction events, several species that likely co-evolved with megafauna established new interactions with humans and their domestic animals. Species that were highly specialized in interactions with megafauna, such as large predators, specialized parasites, and large commensalists (e.g. scavengers, dung beetles), and could not adapt to new hosts or prey were more likely to die out. Partners that were less megafauna dependent persisted because of behavioural plasticity or by shifting their dependency to humans via domestication, facilitation or pathogen spill-over, or through interactions with domestic megafauna. We argue that the ongoing extinction of the extant megafauna in the Anthropocene will catalyse another wave of co-extinctions due to the enormous diversity of key ecological interactions and functional roles provided by the megafauna.

Community-wide integration of floral colour and scent in a Mediterranean scrubland.

Angiosperm flowers have evolved a dazzling palette of colours and a rich bouquet of scents, principally serving to attract pollinators. Despite recent progress in the ecology of pollination, the sensory floral traits that are important for communication with pollinators (for example, colour and scent) have not been assessed in an unbiased, integrative sense within a community context. Nonetheless, floral sensory stimuli are known key factors that mediate flower visitation, thus affecting community dynamics. Here we show that flowers of the phrygana, a natural Mediterranean scrubland, display integrated patterns of scent composition and colour (as perceived by pollinators). Surprisingly, the data reveal predictive relationships between patterns of volatile composition and flower reflectance spectra. The presence of nectar is related to visual cues and the qualitative composition of floral aromas. Our results reveal a coordinated phenotypic integration consistent with the sensory abilities and perceptual biases of bees, suggesting potential facilitative effects for pollination and highlighting the fundamental importance of bees in Mediterranean-type ecosystems. We offer our unbiased approach as a starting point for more extensive, global investigations of the diversity of floral sensory phenotypes and its role in the community ecology of plant-pollinator interactions.

Species-rich networks and eco-evolutionary synthesis at the metacommunity level.

Understanding how ecological and evolutionary processes interdependently structure biosphere dynamics is a major challenge in the era of worldwide ecosystem degradation. However, our knowledge of 'eco-evolutionary feedbacks' depends largely on findings from simple systems representing limited spatial scales and involving few species. Here we review recent conceptual developments for the understanding of multispecies coevolutionary processes and then discuss how new lines of concepts and methods will accelerate the integration of ecology and evolutionary biology. To build a research workflow for integrating insights into spatiotemporal dynamics of species-rich systems, we focus on the roles of 'metacommunity hub' species, whose population size and/or genetic dynamics potentially control landscape- or regional-scale phenomena. As large amounts of network data are becoming available with high-throughput sequencing of various host-symbiont, prey-predator, and symbiont-symbiont interactions, we suggest it is now possible to develop bases for the integrated understanding and management of species-rich ecosystems.

Ecosystem restoration strengthens pollination network resilience and function.

Land degradation results in declining biodiversity and the disruption of ecosystem functioning worldwide, particularly in the tropics. Vegetation restoration is a common tool used to mitigate these impacts and increasingly aims to restore ecosystem functions rather than species diversity. However, evidence from community experiments on the effect of restoration practices on ecosystem functions is scarce. Pollination is an important ecosystem function and the global decline in pollinators attenuates the resistance of natural areas and agro-environments to disturbances. Thus, the ability of pollination functions to resist or recover from disturbance (that is, the functional resilience) may be critical for ensuring a successful restoration process. Here we report the use of a community field experiment to investigate the effects of vegetation restoration, specifically the removal of exotic shrubs, on pollination. We analyse 64 plant-pollinator networks and the reproductive performance of the ten most abundant plant species across four restored and four unrestored, disturbed mountaintop communities. Ecosystem restoration resulted in a marked increase in pollinator species, visits to flowers and interaction diversity. Interactions in restored networks were more generalized than in unrestored networks, indicating a higher functional redundancy in restored communities. Shifts in interaction patterns had direct and positive effects on pollination, especially on the relative and total fruit production of native plants. Pollinator limitation was prevalent at unrestored sites only, where the proportion of flowers producing fruit increased with pollinator visitation, approaching the higher levels seen in restored plant communities. Our results show that vegetation restoration can improve pollination, suggesting that the degradation of ecosystem functions is at least partially reversible. The degree of recovery may depend on the state of degradation before restoration intervention and the proximity to pollinator source populations in the surrounding landscape. We demonstrate that network structure is a suitable indicator for pollination quality, highlighting the usefulness of interaction networks in environmental management.

Oceanic island biogeography through the lens of the general dynamic model: assessment and prospect.

The general dynamic model of oceanic island biogeography (GDM) has added a new dimension to theoretical island biogeography in recognizing that geological processes are key drivers of the evolutionary processes of diversification and extinction within remote islands. It provides a dynamic and essentially non-equilibrium framework generating novel predictions for emergent diversity properties of oceanic islands and archipelagos. Its publication in 2008 coincided with, and spurred on, renewed attention to the dynamics of remote islands. We review progress, both in testing the GDM's predictions and in developing and enhancing ecological-evolutionary understanding of oceanic island systems through the lens of the GDM. In particular, we focus on four main themes: (i) macroecological tests using a space-for-time rationale; (ii) extensions of theory to islands following different patterns of ontogeny; (iii) the implications of GDM dynamics for lineage diversification and trait evolution; and (iv) the potential for downscaling GDM dynamics to local-scale ecological patterns and processes within islands. We also consider the implications of the GDM for understanding patterns of non-native species diversity. We demonstrate the vitality of the field of island biogeography by identifying a range of potentially productive lines for future research.

Nested species interactions promote feasibility over stability during the assembly of a pollinator community.

The foundational concepts behind the persistence of ecological communities have been based on two ecological properties: dynamical stability and feasibility. The former is typically regarded as the capacity of a community to return to an original equilibrium state after a perturbation in species abundances and is usually linked to the strength of interspecific interactions. The latter is the capacity to sustain positive abundances on all its constituent species and is linked to both interspecific interactions and species demographic characteristics. Over the last 40 years, theoretical research in ecology has emphasized the search for conditions leading to the dynamical stability of ecological communities, while the conditions leading to feasibility have been overlooked. However, thus far, we have no evidence of whether species interactions are more conditioned by the community's need to be stable or feasible. Here, we introduce novel quantitative methods and use empirical data to investigate the consequences of species interactions on the dynamical stability and feasibility of mutualistic communities. First, we demonstrate that the more nested the species interactions in a community are, the lower the mutualistic strength that the community can tolerate without losing dynamical stability. Second, we show that high feasibility in a community can be reached either with high mutualistic strength or with highly nested species interactions. Third, we find that during the assembly process of a seasonal pollinator community located at The Zackenberg Research Station (northeastern Greenland), a high feasibility is reached through the nested species interactions established between newcomer and resident species. Our findings imply that nested mutualistic communities promote feasibility over stability, which may suggest that the former can be key for community persistence.

Galápagos land iguana (Conolophus subcristatus) as a seed disperser.

The role of the most common land iguana (Conolophus subcristatus) in the Galápagos Islands as an effective seed disperser is explored in this study. A total of 5705 seeds of 32 plant species were identified from 160 scats, 4545 of which (80%) appeared visually undamaged. Germination trials of 849 seeds from 29 species revealed that at least 10 species remained viable after passing through the iguana's gut, although only a small proportion of those seeds (4%) germinated. In any case, we argue that C. subcristatus exerts an important role on the 7 Galapagos islands where it occurs because of its abundance and capacity to ingest and disperse seeds at long distances. Our results strongly suggest that the Galápagos C. subcristatus plays an important role as a seed disperser of not only of native species but also some introduced plants in the Galápagos Islands.

Below-ground plant-fungus network topology is not congruent with above-ground plant-animal network topology.

In nature, plants and their pollinating and/or seed-dispersing animals form complex interaction networks. The commonly observed pattern of links between specialists and generalists in these networks has been predicted to promote species coexistence. Plants also build highly species-rich mutualistic networks below ground with root-associated fungi, and the structure of these plant-fungus networks may also affect terrestrial community processes. By compiling high-throughput DNA sequencing data sets of the symbiosis of plants and their root-associated fungi from three localities along a latitudinal gradient, we uncovered the entire network architecture of these interactions under contrasting environmental conditions. Each network included more than 30 plant species and hundreds of mycorrhizal and endophytic fungi belonging to diverse phylogenetic groups. The results were consistent with the notion that processes shaping host-plant specialization of fungal species generate a unique linkage pattern that strongly contrasts with the pattern of above-ground plant-partner networks. Specifically, plant-fungus networks lacked a "nested" architecture, which has been considered to promote species coexistence in plant-partner networks. Rather, the below-ground networks had a conspicuous "antinested" topology. Our findings lead to the working hypothesis that terrestrial plant community dynamics are likely determined by the balance between above-ground and below-ground webs of interspecific interactions.

Space, time and aliens: charting the dynamic structure of Galápagos pollination networks.

Oceanic archipelagos are threatened by the introduction of alien species which can severely disrupt the structure, function and stability of native communities. Here we investigated the pollination interactions in the two most disturbed Galápagos Islands, comparing the three main habitats and the two seasons, and assessing the impacts of alien plant invasions on network structure. We found that the pollination network structure was rather consistent between the two islands, but differed across habitats and seasons. Overall, the arid zone had the largest networks and highest species generalization levels whereas either the transition between habitats or the humid habitat showed lower values. Our data suggest that alien plants integrate easily into the communities, but with low impact on overall network structure, except for an increase in network selectiveness. The humid zone showed the highest nestedness and the lowest modularity, which might be explained by the low species diversity and the higher incidence of alien plants in this habitat. Both pollinators and plants were also more generalized in the hot season, when networks showed to be more nested. Alien species (both plants and pollinators) represented a high fraction (∼56 %) of the total number of interactions in the networks. It is thus likely that, in spite of the overall weak effect we found of alien plant invasion on pollination network structure, these introduced species influence the reproductive success of native ones, and by doing so, they affect the functioning of the community. This certainly deserves further investigation.