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Many vertebrate species act as both plant pollinators and seed-dispersers, thus interconnecting these processes, particularly on islands. Ecological multilayer networks are a powerful tool to explore interdependencies between processes; however, quantifying the links between species engaging in different types of interactions (i.e. inter-layer edges) remains a great challenge. Here, we empirically measured inter-layer edge weights by quantifying the role of individually marked birds as both pollinators and seed-dispersers of Galápagos plant species over an entire year. Although most species (80%) engaged in both functions, we show that only a small proportion of individuals actually linked the two processes, highlighting the need to further consider intra-specific variability in individuals' functional roles. Furthermore, we found a high variation among species in linking both processes, i.e. some species contribute more than others to the modular organization of the multilayer network. Small and abundant species are particularly important for the cohesion of pollinator seed-dispersal networks, demonstrating the interplay between species traits and neutral processes structuring natural communities.

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., (© 2019 Botanical Society of America.)

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.

Owing to food scarcity and to the high densities that vertebrates often reach on islands, typical insect- and seed-eaters widen their feeding niche and interact with a greater fraction of species than their mainland counterparts. This phenomenon, coined here 'interaction release', has been previously reported for single species but never for an entire community. During 4 years, we gathered data on bird-flower visitation on 12 Galápagos islands. We show that all sampled land birds exploit floral resources and act as potential pollinators across the entire archipelago, in all major habitats and all year round. Although species and link composition varies among islands, strong interaction release takes place on all islands, making their bird-flower network highly generalized. Interaction release is crucial to the survival of native birds but simultaneously threatens the unique biodiversity of this archipelago, as the birds also visit invading plants, likely facilitating their integration into pristine native communities.

Seed dispersal effectiveness (SDE) is the contribution of dispersers to plant recruitment and is estimated as the product of the number of seeds dispersed (quantity) and the probability of recruitment of each dispersed seed (quality). Although SDE is a key concept in seed dispersal ecology, few studies estimate SDE and none has a community approach. Oceanic islands, with simple communities, are ideal for this purpose. In this study, we compared the SDE of the main types of dispersers (lizards and passerine birds) at the community level in a given habitat. We estimated SDE using a stochastic simulation model parameterized with empirical data on quantity and quality components measured throughout the recruitment process. Although lizards are highly frugivorous and their density was approximately 20 times higher than that of birds, lizards and birds dispersed a similar quantity of seeds. This may be due to lower intake of seeds by lizards due to their slower metabolism (approximately 20 times lower than birds). This low metabolic rate limits the importance of lizards as seed dispersers, but it is compensated by extraordinarily high lizard densities in the study area (approximately 9600 individuals/km2). High densities of lizards are typical of islands, and this helps to explain why dispersal by lizards seems mainly an island phenomenon. Birds and lizards showed functional complementarity, especially regarding seed dispersal distribution patterns. In fact, lizards dispersed more seeds in shrublands and open sites, and birds in woodlands and beneath canopies, with their joint contribution helping to maximize recruitment. Lizards provided higher SDE than birds for 7 out of 11 plant species. The disperser with a higher quantity for a given plant generally had the higher quality, and plants could be classified as bird- or lizard-dependent for dispersal. This dependence increased when considering SDE instead of dispersal quantity only. Moreover, quality was a better predictor of SDE than quantity, which should be considered when parameterizing interaction networks, as this might affect inferences about their architecture.

This study assessed the impact of introduced black rats (Rattus rattus) on Cory's shearwater (Calonectris diomedea borealis) in a multi-invaded insular ecosystem where rats are mesopredators. We hypothesized that black rats should have little impact on Cory's shearwaters in the presence of cats as superpredators. Stomach contents and stable isotope analysis (SIA) in tissues of black rats were analyzed to assess the trophic ecology and the importance of Cory's shearwater in their diet. We also studied the isotopic signature in tissues of house mouse (Mus domesticus) to confirm previous data showing no predation of this species on Cory's shearwaters. For both rodent species, temporal variation in diet composition in response to the availability of seabird prey was evaluated, and short- and long-term consistency in diet was tested using different tissues from the same individual. For black rats a Bayesian isotope mixing model (SIAR) was applied to determine the relative contribution of each prey to the individual diet. SIA of mouse tissues varied between the Cory's shearwater breeding and non-breeding periods. However, no significant differences were found in diet and SIA for black rats. In contrast, individuals of both species showed a strong consistency in diet which apparently benefited their body condition index. Although black rats supplement their diet with Cory's shearwater eggs and chicks (8.3% in stomach contents and 10.6% in the SIAR model), their current impact on the Cory's shearwater population appears to be small, probably due to several factors including the small size of the rat population and a high level of rat predation by cats., (Copyright © 2014 Elsevier GmbH. All rights reserved.)

Predator activity around the nests induces stress in breeding birds, which may have weaker immunity and are therefore more susceptible to parasite infections. The influence of predators on parasites has only been observed in land birds, and most studies are experimental. As seabird colonies offer excellent conditions for parasites, here we assess the influence of mouse, rat, and cat activity on parasites in Cory's shearwater (Calonectris diomedea borealis). Adults were examined for blood parasites and one adult and the juvenile from 53 nests were inspected for ectoparasites over two consecutive years (2010 and 2011). Nests differed in their physical characteristics and indices of mammal predator activity and were checked weekly to assess breeding success. Our results showed absence of blood parasites. Among the environmental factors, predator pressure received the most support (89 %) from the data as influencing nest ectoparasite intensity. Birds most infected had worse body condition, and breeding success was negatively influenced by predator activity and ectoparasite intensity. To our knowledge, this is the first analysis of the interaction between mammal predators and ectoparasites in seabird species and supports greater protection through eradication efforts. In addition, we provide the first data on the endoparasite fauna of Cory's shearwater.

Many fleshy-fruited plants from the Mediterranean and Macaronesian islands are dispersed through endozoochory. In mainland Mediterranean areas, reciprocal adaptations have been found between plants and animals, although evidence is scarce. On small isolated oceanic islands, such reciprocal adaptations might well be more prevalent due to intrinsic island traits. Here we evaluate the existence of selective pressures exerted by two different disperser guilds (lizards and birds) on two seed traits (seed coat thickness and seed germination pattern) of two congeneric species present on Mediterranean and Macaronesian islands. In the continental Balearic Islands, Rubia peregrina has evolved mostly with birds, although frugivorous lizards are present in some of these islands and are known to eventually consume its fruits. By contrast, R. fruticosa, endemic to the Macaronesian archipelago, has evolved mostly interacting with lizards and only recently with birds. We hypothesized that R. fruticosa would be especially adapted to saurochory, with thicker seed coats and higher germination proportion, whereas R. peregrina would be more adapted to ornithocory, with thinner seed coats and showing a lower germination percentage after being ingested by lizards. Captivity experiments of seed ingestions by natural and non-natural dispersers (i.e., frugivores that have not evolved with those plants) were conducted. Results suggest that dispersers did not exert any strong enough selective pressure to induce changes in germination patterns. We attribute this to the fact that the Rubiaceae is an ancestral family in the Mediterranean (both on continent and islands) and thus probably interacted with lizards in the past. Lastly, although we hold that the seed coat structure of R. fruticosa is probably associated with its evolutionary success after a long interaction with insular lizards, our findings support the idea that the relationship between endozoochorous plants and the guild of dispersers with whom they evolved is rather unspecific.

Alien plants are a growing threat to the Galápagos unique biota. We evaluated the impact of alien plants on eight seed dispersal networks from two islands of the archipelago. Nearly 10 000 intact seeds from 58 species were recovered from the droppings of 18 bird and reptile dispersers. The most dispersed invaders were Lantana camara, Rubus niveus and Psidium guajava, the latter two likely benefiting from an asynchronous fruit production with most native plants, which facilitate their consumption and spread. Lava lizards dispersed the seeds of 27 species, being the most important dispersers, followed by small ground finch, two mockingbirds, the giant tortoise and two insectivorous birds. Most animals dispersed alien seeds, but these formed a relatively small proportion of the interactions. Nevertheless, the integration of aliens was higher in the island that has been invaded for longest, suggesting a time-lag between alien plant introductions and their impacts on seed dispersal networks. Alien plants become more specialized with advancing invasion, favouring more simplified plant and disperser communities. However, only habitat type significantly affected the overall network structure. Alien plants were dispersed via two pathways: dry-fruited plants were preferentially dispersed by finches, while fleshy fruited species were mostly dispersed by other birds and reptiles.