Dietary metabarcoding reveals the simplification of bird-pest interaction networks across a gradient of agricultural cover.

Agriculture is vital for supporting human populations, but its intensification often leads to landscape homogenization and a decline in non-provisioning ecosystem services. Ecological intensification and multifunctional landscapes are suggested as nature-based alternatives to intensive agriculture, using ecological processes like natural pest regulation to maximize food production. Birds are recognized for their role in increasing crop yields by consuming invertebrate pests in several agroecosystems. However, the understanding of how bird species, their traits and agricultural land cover influence the structure of bird-pest interactions remains limited. We sampled bird-pest interactions monthly for 1 year, at four sites within a multifunctional landscape, following a gradient of increasing agricultural land cover. We analysed 2583 droppings of 55 bird species with DNA metabarcoding and detected 225 pest species in 1139 samples of 42 bird species. As expected, bird-pest interactions were highly variable across bird species. Dietary pest richness was lower in the fully agricultural site, while predation frequency remained consistent across the agricultural land cover gradient. Network analysis revealed a reduction in the complexity of bird-pest interactions as agricultural coverage increased. Bird species abundance affected the bird's contribution to the network structure more than any of the bird traits analysed (weight, phenology, invertebrate frequency in diet and foraging strata), with more common birds being more important to network structure. Overall, our results show that increasing agricultural land cover increases the homogenization of bird-pest interactions. This shows the importance of maintaining natural patches within agricultural landscapes for biodiversity conservation and enhanced biocontrol.

A agricultura é essencial para suportar a população humana, mas a sua intensificação geralmente leva à homogeneização da paisagem e à redução dos serviços do ecossistema que não sejam de provisão. A intensificação ecológica e paisagens multifuncionais são sugeridas como alternativas naturais à agricultura intensiva, utilizando processos ecológicos como a regulação natural de pragas para maximizar a produção de alimentos. As aves são conhecidas pelo seu papel no aumento da produtividade das culturas por consumirem pragas em diversos agroecossistemas. Contudo, o conhecimento de como as espécies de aves, as suas características e a cobertura agrícola influenciam as interações entre aves e pragas são limitados. Nós amostrámos estas interações mensalmente durante um ano, em quatro locais, numa paisagem multifuncional, ao longo um gradiente de aumento da cobertura agrícola. Analisamos 2583 dejetos de 55 espécies de aves com DNA metabarcoding e detetamos 225 espécies praga em 1139 amostras de 42 espécies de aves. Como esperado, as interações entre aves e pragas foram muito distintas entre as várias espécies de aves. A riqueza de pragas na dieta foi menor no local completamente dominado por área agrícola, enquanto a frequência de predação de pragas foi constante ao longo do gradiente de cobertura agrícola. A análise de redes demonstrou uma redução na complexidade das interações entre aves e pragas à medida que a cobertura agrícola aumenta. A abundância das espécies de aves influenciou mais a contribuição das aves para a estrutura da rede do que qualquer uma das características analisadas (peso, fenologia, frequência de invertebrados na dieta e estrato de alimentação), sendo as aves mais comuns as mais importantes na estrutura da rede. De forma geral, os nossos resultados indicam que o aumento da cobertura agrícola aumenta a homogeneização das interações entre aves e pragas. Isto demonstra a importância de preservar áreas naturais em paisagem agrícolas para a conservação de biodiversidade e melhor controlo biológico.

EuDiS - A comprehensive database of the seed dispersal syndromes of the European flora.

Background: Seed dispersal is a critical process in plant colonisation and demography. Fruits and seeds can be transported by several vectors (typically animals, wind and water), which may have exerted strong selective pressures on plant's morphological traits. The set of traits that favour dispersal by a specific vector have been historically considered as seed dispersal syndromes. As seed dispersal syndromes have a great potential to predict how seeds move (i.e. the relative importance of the standard mechanisms of seed dispersal), they have attracted the attention of naturalists and researchers for centuries. However, given that observations of actual dispersal events and colonisation are seldom reported, there is still much confusion in current studies failing to properly discriminate between seed dispersal syndromes (i.e. sets of traits that favour a particular mechanism) and actual seed dispersal (i.e. the vector that moves a given seed in one dispersal event). This distinction is important because the presence of any seed dispersal syndrome does not preclude the seed being occasionally dispersed by other non-standard mechanisms (i.e. different from the one predicted). Similarly, the absence of seed dispersal syndromes does not prevent seeds from being dispersed. The correct coding of seed dispersal syndromes thus requires a systematic and evolutive, rather than a phenomenological approach. Unfortunately, such approach has rarely been implemented at a community-level and no comprehensive datasets of seed dispersal syndromes are yet available for any entire flora. New information: This database contains categorisation of the native European flora into eight seed dispersal syndromes. Information for a total of 9,874 species retrieved from the volumes of Flora Europaea were analysed. Earlier versions of this database, which only coded for the presence of four long-distance dispersal syndromes (endozoochorous, epizoochorous, thalassochorous and anemochorous diaspores), were used in four previous studies. Here, we present a fully revised and expanded database, including the presence of four additional short-distance dispersal syndromes (myrmecochorous, vertebrate hoarding, freshwater hydrochorous and ballochorous diaspores), a nomenclatural update for all species and the codification of 416 additional species.Roughly half (51.3%) of the native European flora produce diaspores without traits clearly associated with facilitating seed dispersal. The other half (48.7%) of the European plant species produces diaspores with some specialised traits associated with seed dispersal, most of which (79.9%) with a potential to facilitate long-distance dispersal events. The most common diaspores are those with anemochorous (23.5%), epizoochorous (8.0%), endozoochorous (7.8%), myrmecochorous (7.2%), thalassochorous (2.3%), freshwater dispersal (2.1%), ballochorous (4.6%) and vertebrate hoarding associated traits (0.2%). Two-thirds (66.3%) of the European shrub and tree species have diaspores with some specialisation for biotic seed dispersal.

Changing with the times: Seasonal environmental gradients unveil dynamic bat assemblages and vulnerability.

Uncovering the temporal and spatial dynamics of biological communities in response to biotic and abiotic drivers is essential to predict the effects of environmental change on biodiversity. Similarly, estimating species vulnerability in the face of such dynamics is crucial for implementing effective conservation actions. We explored how bat diversity changes over the year across an altitudinal gradient and identified the environmental drivers that shape bat communities. By analysing species' marginality within the biophysical niche space, we evaluated bats' vulnerability to foreseeable environmental changes. Our results suggest that altitude, the proportion of forest cover and shrub cover are the main drivers shaping bat communities year-round. Additionally, while some bat species are restricted to a single ecological assemblage (or ecological preferences group), others show greater plasticity throughout the year. Importantly, we found that although bats associated with highland habitats and forests could be particularly vulnerable to environmental changes (in particular Myotis mystacinus), this vulnerability correlates poorly with their national conservation status. We suggest that species' ecological plasticity is critical for the resilience of biological communities exposed to environmental changes and should be considered when planning tailored conservation strategies.

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.

Global and regional ecological boundaries explain abrupt spatial discontinuities in avian frugivory interactions.

Species interactions can propagate disturbances across space via direct and indirect effects, potentially connecting species at a global scale. However, ecological and biogeographic boundaries may mitigate this spread by demarcating the limits of ecological networks. We tested whether large-scale ecological boundaries (ecoregions and biomes) and human disturbance gradients increase dissimilarity among plant-frugivore networks, while accounting for background spatial and elevational gradients and differences in network sampling. We assessed network dissimilarity patterns over a broad spatial scale, using 196 quantitative avian frugivory networks (encompassing 1496 plant and 1004 bird species) distributed across 67 ecoregions, 11 biomes, and 6 continents. We show that dissimilarities in species and interaction composition, but not network structure, are greater across ecoregion and biome boundaries and along different levels of human disturbance. Our findings indicate that biogeographic boundaries delineate the world's biodiversity of interactions and likely contribute to mitigating the propagation of disturbances at large spatial scales.

Canopy arthropod declines along a gradient of olive farming intensification.

Arthropod declines have been linked to agricultural intensification. However, information about the impacts of intensification is still limited for many crops, as is our understanding of the responses of different arthropod taxa and trophic groups, thus hindering the development of effective mitigation measures. We investigated the impacts of olive farming intensification on canopy-dwelling arthropods in the Mediterranean region. Intensification involves the increased use of agrochemicals, mechanisation and irrigation, but also structural changes from traditional orchards with low densities of large and old trees, to intensive and superintensive orchards with high to very high densities of smaller and younger trees, respectively. Canopy arthropods were vacuum-sampled at 53 sites representing the three orchard intensification levels, in spring, summer and autumn 2017. We evaluated how the arthropod community varied across intensification levels, and in response to orchard structure, management and landscape context. We found no changes in the diversity of arthropod taxa across intensification levels after correcting for sample coverage, but arthropod abundance declined markedly along the intensification gradient. Decreased abundance was associated with changes in orchard structure, lower herbaceous cover, and higher herbicide and insecticide use. The abundance of a specialized olive pest was lower in landscapes with higher woodland cover. The negative effects of intensification were stronger in spring and summer than in autumn, and parasitoids and predators were particularly affected. Overall, results suggest that retaining herbaceous cover, reducing agrochemical inputs and preserving natural woody elements in the landscape, may contribute to mitigate impacts of olive farming intensification on canopy arthropods, particularly on beneficial species.

Limited potential for bird migration to disperse plants to cooler latitudes.

Climate change is forcing the redistribution of life on Earth at an unprecedented velocity1,2. Migratory birds are thought to help plants to track climate change through long-distance seed dispersal3,4. However, seeds may be consistently dispersed towards cooler or warmer latitudes depending on whether the fruiting period of a plant species coincides with northward or southward migrations. Here we assess the potential of plant communities to keep pace with climate change through long-distance seed dispersal by migratory birds. To do so, we combine phenological and migration information with data on 949 seed-dispersal interactions between 46 bird and 81 plant species from 13 woodland communities across Europe. Most of the plant species (86%) in these communities are dispersed by birds migrating south, whereas only 35% are dispersed by birds migrating north; the latter subset is phylogenetically clustered in lineages that have fruiting periods that overlap with the spring migration. Moreover, the majority of this critical dispersal service northwards is provided by only a few Palaearctic migrant species. The potential of migratory birds to assist a small, non-random sample of plants to track climate change latitudinally is expected to strongly influence the formation of novel plant communities, and thus affect their ecosystem functions and community assembly at higher trophic levels.

Dimensions of invasiveness: Links between local abundance, geographic range size, and habitat breadth in Europe's alien and native floras.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders-abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions-for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions.

Establishment, spread and early impacts of the first biocontrol agent against an invasive plant in continental Europe.

Classical biocontrol is key for the successful management of invasive alien plants; yet, it is still relatively new in Europe. Although post-release monitoring is essential to evaluate the effectiveness of a biocontrol agent, it is often neglected. This study reports the detailed post-release monitoring of the first biocontrol agent intentionally introduced against an invasive plant in continental Europe. The Australian bud-galling wasp Trichilogaster acaciaelongifoliae (Frogatt) is used to control the invasive Acacia longifolia (Andr.) Willd., with a long history of success in South Africa. This biocontrol agent was first released in Europe in 2015 at several sites along the Portuguese coast. We monitored the establishment, spread and early impacts of T. acaciaelongifoliae on target-plants in Portugal, across 61 sites, from 2015 to 2020. Initial release of adults emerging from galls imported from South Africa and the subsequent releases from galls established in Portugal (2018 onwards) was compared, assessing the implications of the hemisphere shift. The impacts on the reproductive output and vegetative growth of A. longifolia were evaluated in more detail at three sites. From 2015 to 2019, 3567 T. acaciaelongifoliae were released at 61 sites, with establishment confirmed at 36 sites by 2020. The transfer of the wasp from the southern hemisphere limited its initial establishment, but increased rates of establishment followed with synchronization of its life cycle with northern hemisphere conditions. Therefore, after an initial moderate establishment, T. acaciaelongifoliae adapted to the northern hemisphere conditions and experienced an exponential growth (from 66 galls by 2016, to 24000 galls by 2018). Galled A. longifolia branches produced significantly fewer pods (-84.1%), seeds (-95.2%) and secondary branches (-33.3%) and had fewer phyllodes but increased growth of the main branch compared to ungalled branches. Trichilogaster acaciaelongifoliae successfully established in the northern hemisphere, despite the initial phenological mismatch and adverse weather conditions. To achieve this, it had to establish and synchronize its life cycle with the phenology of its host-plant, after which it developed exponentially and began to show significant impacts on the reproductive output of A. longifolia.

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.

Native and alien flower visitors differ in partner fidelity and network integration.

Globalisation persistently fuels the establishment of non-native species outside their natural ranges. While alien plants have been intensively studied, little is known about alien flower visitors, and especially, how they integrate into natural communities. Here, we focus on mutualistic networks from five Galápagos islands to quantify whether alien and native flower visitors differ consistently in their pairwise interactions. We find that (1) alien flower visitors have more interaction partners and larger species strengths (i.e. plants are more connected to alien insects), (2) native insects tend to have higher partner fidelity as they deviate more from random partner utilisation, and iii) the difference between native and alien flower visitors in network integration intensifies with island degradation. Thus, native and alien flower visitors are not interchangeable, and alien establishment might have yet unforeseen consequences for the pairwise dynamics between plants and flower visitors on the Galápagos - especially on the heavily disturbed islands.

Integrating plant species contribution to mycorrhizal and seed dispersal mutualistic networks.

Mutualistic interactions like those established between plants and mycorrhizal fungi or seed dispersers are key drivers of plant population dynamics and ecosystem functioning; however, these interactions have rarely been explored together. We assembled a tripartite fungi-plant-disperser network in the Gorongosa National Park-Mozambique, to test (1) if diversity and importance of plant mutualists above- and belowground are correlated, and (2) whether biotically and abiotically dispersed plants are associated with distinct arbuscular mycorrhizal fungi (AMF). We quantified seed dispersal by animals for 1 year and characterized the AMF of 26 common plant species. Sixteen plant species were dispersed by 15 animals and colonized by 48 AMF virtual taxa (VT), while the remaining 10 plant species were not dispersed by animals and associated with 34 AMF VT. We found no evidence for a correlation between the number of plant partners above- and belowground or on plant specialization on both types of partners. We also found no evidence for differentiation of AMF communities between biotically and abiotically dispersed plants. Our results suggest that the establishment of plant interactions with seed dispersers and mycorrhizal fungi is largely independent and that both biotically and abiotically dispersed plants seem to associate with similar communities of AMF.

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.

First evidence for the joint dispersal of mycorrhizal fungi and plant diaspores by birds.

Seed dispersal allows plants to colonise new sites and escape from pathogens and intraspecific competition, maintaining plant genetic diversity and regulating plant distribution. Conversely, most plant species form mutualistic associations with arbuscular mycorrhizal (AM) fungi in a symbiosis established immediately after seed germination. Because AM fungi are obligate symbionts, using the same dispersal vector as their host should be highly advantageous for their survival, but the co-dispersal of seeds and AM fungal spores has never been confirmed. We aim to clarify the potential role of European birds, essential dispersers for many plant species, as co-dispersers of seeds and AM fungal spores. In total, 63 bird droppings with intact seeds were placed in sterilised soil and maintained for 4 months in a protected environment to avoid contamination. Additionally, 173 bird droppings and 729 gauze swabs used to clean birds' feet were inspected for AM fungal spores. Although no spores were detected by direct observation of these samples, seven Rubus ulmifolius seedlings obtained from four independent droppings of Erithacus rubecula and Sylvia melanocephala were colonised by AM fungi. Our results show that birds can effectively co-disperse viable seeds and AM fungal spores, potentially over long distances, providing a pivotal mechanism to understand the cosmopolitan distribution of AM fungi.

Pollination networks from natural and anthropogenic-novel communities show high structural similarity.

The Anthropocene is marked by an unprecedented homogenisation of the world's biota, confronting species that never co-occurred during their evolutionary histories. Interactions established in these novel communities may affect ecosystem functioning; however, most research has focused on the impacts of a minority of aggressive invasive species, while changes inflicted by a less conspicuous majority of non-invasive alien species on community structure are still poorly understood. This information is critical to guide conservation strategies, and instrumental to advance ecological theory, particularly to understand how non-native species integrate in recipient communities and affect the interactions of native species. We evaluated how the structure of 50 published pollination networks changes with the proportion of alien plant species and found that network structure is largely unaffected. Although some communities were heavily invaded, the proportion of alien plant species was relatively low (mean = 10%; max. = 38%). We further characterized the pollination network in a botanic garden with a plant community dominated by non-invasive alien species (85%). We show that the structure of this novel community is also not markedly different from native-dominated communities. Plant-pollinator interactions revealed no obvious differences regarding plant origin (native vs. alien) or the native bioregion of the introduced plants. This overall similarity between native and alien plants is likely driven by the contrasting patterns of invasive plants (promoting generalism), and non-invasive aliens, suggested here to promote specialization.

Author Correction: Multilayer networks reveal the spatial structure of seed-dispersal interactions across the Great Rift landscapes.

The original version of this Article contained Figshare links in the Code availability statement that were not functional. The correct Figshare links to MATLAB scripts and R code used in this study are https://doi.org/10.6084/m9.figshare.4955651 and https://doi.org/10.6084/m9.figshare.4836383, respectively. These errors have now been corrected in both the PDF and HTML versions of the Article.

Should I stay or should I go? Mycorrhizal plants are more likely to invest in long-distance seed dispersal than non-mycorrhizal plants.

Seed dispersal and mycorrhizal associations are key mutualisms for the functioning and regeneration of plant communities; however, these processes have seldom been explored together. We hypothesised that obligatory mycorrhizal plants will be less likely to have long-distance dispersal (LDD) syndromes since the probability of finding suitable mycorrhizal partners is likely to decrease with distance to the mother plant. We contrasted the mycorrhizal status and LDD syndromes for 1960 European plant species, using phylogenetically corrected log-linear models. Contrary to our expectation, having specialised structures for LDD is more frequent in obligate mycorrhizal plants than in non-mycorrhizal plants, revealing that lack of compatible mutualists does not constrain investment in LDD structures in the European Flora. Ectomycorrhizal plants associated with wind-dispersing fungi are also more likely to have specialised structures for wind dispersal. Habitat specificity and narrower niche of non-mycorrhizal plants might explain the smaller investment in specialised structures for seed dispersal.

Multilayer networks reveal the spatial structure of seed-dispersal interactions across the Great Rift landscapes.

Species interaction networks are traditionally explored as discrete entities with well-defined spatial borders, an oversimplification likely impairing their applicability. Using a multilayer network approach, explicitly accounting for inter-habitat connectivity, we investigate the spatial structure of seed-dispersal networks across the Gorongosa National Park, Mozambique. We show that the overall seed-dispersal network is composed by spatially explicit communities of dispersers spanning across habitats, functionally linking the landscape mosaic. Inter-habitat connectivity determines spatial structure, which cannot be accurately described with standard monolayer approaches either splitting or merging habitats. Multilayer modularity cannot be predicted by null models randomizing either interactions within each habitat or those linking habitats; however, as habitat connectivity increases, random processes become more important for overall structure. The importance of dispersers for the overall network structure is captured by multilayer versatility but not by standard metrics. Highly versatile species disperse many plant species across multiple habitats, being critical to landscape functional cohesion.