Seed-dispersal Networks Research Articles

Loss of Generalist Plant Species and Functional Diversity Decreases the Robustness of a Seed Dispersal Network

SummaryUnderstanding cascading effects of species loss is a major challenge for ecologists. Traditionally, the robustness of ecological networks has been evaluated based on simulation studies where primary extinctions occur at random or as a function of species specialization, ignoring other important biological factors. Here, we estimate the robustness of a seed dispersal network from a grassland–forest mosaic in southern Brazil, simulating distinct scenarios of woody plant species extinction, including scenarios where species are eliminated based on their evolutionary and functional distinctiveness. Our results suggest that the network is more robust when species are eliminated based on their evolutionary uniqueness, followed by random extinctions, the extinction of the most specialist species, functional distinctiveness and, at last, when the most generalist species are sequentially eliminated. Our results provide important information for grassland–forest mosaic management, as they indicate that loss of generalist species and functional diversity makes the system more likely to collapse.

Network resilience of mutualistic ecosystems and environmental changes: an empirical study.

It is theorized that a mutualistic ecosystem's resilience against perturbations (e.g. species extinction) is determined by a single macroscopic parameter (network resilience), calculable from the network. Given that such perturbations occur owing to environmental changes (e.g. climate change and human impact), it has been predicted that mutualistic ecosystems that exist despite extensive environmental changes exhibit higher network resilience; however, such a prediction has not been confirmed using real-world data. Thus, in this study, the effects of climate change velocity and human activities on mutualistic network resilience were investigated. A global dataset of plant–animal mutualistic networks was used, and spatial analysis was performed to examine the effects. Moreover, the potential confounding effects of network size, current climate and altitude were statistically controlled. It was demonstrated that mutualistic network resilience was globally influenced by warming velocity and human impact, in addition to current climate. Specifically, pollination network resilience increased in response to human impact, and seed-dispersal network resilience increased with warming velocity. The effect of environmental changes on network resilience for plants was remarkable. The results confirmed the prediction obtained based on the theory and imply that real-world mutualistic networks have a structure that increases ecosystem resilience against environmental changes. These findings will enhance the understanding of ecosystem resilience.

How intraspecific variation in seed-dispersing animals matters for plants.

Seed dispersal by animals is a complex phenomenon, characterized by multiple mechanisms and variable outcomes. Most researchers approach this complexity by analysing context-dependency in seed dispersal and investigating extrinsic factors that might influence interactions between plants and seed dispersers. Intrinsic traits of seed dispersers provide an alternative way of making sense of the enormous variation in seed fates. I review causes of intraspecific variability in frugivorous and granivorous animals, discuss their effects on seed dispersal, and outline likely consequences for plant populations and communities. Sources of individual variation in seed-dispersing animals include sexual dimorphism, changes associated with growth and ageing, individual specialization, and animal personalities. Sexual dimorphism of seed-dispersing animals influences seed fate through diverse mechanisms that range from effects caused by sex-specific differences in body size, to influences of male versus female cognitive functions. These differences affect the type of seed treatment (e.g. dispersal versus predation), the number of dispersed seeds, distance of seed dispersal, and likelihood that seeds are left in favourable sites for seeds or seedlings. The best-documented consequences of individual differences associated with growth and ageing involve quantity of dispersed seeds and the quality of seed treatment in the mouth and gut. Individual specialization on different resources affects the number of dispersed plant species, and therefore the connectivity and architecture of seed-dispersal networks. Animal personalities might play an important role in shaping interactions between plants and dispersers of their seeds, yet their potential in this regard remains overlooked. In general, intraspecific variation in seed-dispersing animals often influences plants through effects of these individual differences on the movement ecology of the dispersers. Two conditions are necessary for individual variation to exert a strong influence on seed dispersal. First, the individual differences in traits should translate into differences in crucial characteristics of seed dispersal. Second, individual variation is more likely to be important when the proportions of particular types of individuals fluctuate strongly in a population or vary across space; when proportions are static, it is less likely that intraspecific differences will be responsible for changes in the dynamics and outcomes of plant-animal interactions. In conclusion, focusing on variation among foraging animals rather than on species averages might bring new, mechanistic insights to the phenomenon of seed dispersal. While this shift in perspective is unlikely to replace the traditional approach (based on the assumption that all important variation occurs among species), it provides a complementary alternative to decipher the enormous variation observed in animal-mediated seed dispersal.

Connecting Resource Tracking by Frugivores to Temporal Variation in Seed Dispersal Networks

Mutualistic interactions between species are crucial for the maintenance and functioning of the community. Current research has shown the importance of not only understanding the structure of these mutualistic interaction networks but also their temporal dynamics. Temporal changes in species abundance, such as those caused by vertebrate resource tracking, may create temporal variability in a network though neighborhood effects and influence the directedness of dispersal. While much research has been done on resource tracking, neighborhood effects, and directed dispersal individually, little research has been done into their interrelationships and, therefore, theory of how resource tracking can influence frugivory and seed dispersal networks remains poorly developed. We use the available literature to show the prevalence of resource tracking by vertebrate frugivores and hypothesize how, through neighborhood effects, resource tracking may influence short-term variation in network properties. We then discuss how resource tracking can influence long-term network properties by altering the dispersal of plant species. Lastly, we use this information to hypothesize how the introductions of new species into a community may alter the influence of resource tracking on frugivory and seed dispersal. While trait matching and links that are not possible between species in a community (i.e., forbidden links) play a large role in determining the structure of a network, temporal change in the abundance of species due to resource tracking may also affect the properties of networks. By increasing our understanding of the role resource tracking has in the temporal variability of frugivory and seed dispersal networks, we can better determine the full extent of species interactions and provide valuable information for the conservation of ecological communities.

Defaunation effects on plant recruitment depend on size matching and size trade-offs in seed-dispersal networks.

Defaunation by humans causes a loss of large animals in many ecosystems globally. Recent work has emphasized the consequences of downsizing in animal communities for ecosystem functioning. However, no study so far has integrated network theory and life-history trade-offs to mechanistically evaluate the functional consequences of defaunation in plant-animal networks. Here, we simulated an avian seed-dispersal network and its derived ecosystem function seedling recruitment to assess the relative importance of different size-related mechanisms. Specifically, we considered size matching (between bird size and seed size) and size trade-offs, which are driven by differences in plant or animal species abundance (negative size-quantity relationship) as well as in recruitment probability and disperser quality (positive size-quality relationship). Defaunation led to impoverished seedling communities in terms of diversity and seed size, but only if models accounted for size matching. In addition, size trade-off in plants, in concert with size matching, provoked rapid decays in seedling abundance in response to defaunation. These results underscore a disproportional importance of large animals for ecosystem functions. Downsizing in ecological networks will have severe consequences for ecosystem functioning, especially in interaction networks that are structured by size matching between plants and animals.

Drivers of species' role in avian seed-dispersal mutualistic networks.

The mutualistic interaction between frugivore birds and the fruiting plants they disperse presents an asymmetric interaction pattern, with some species having a more important role (i.e. being essential) for maintaining the structure and functioning of the interaction network. The identification of the biological characteristics of these species is of major importance for the understanding and conservation of seed-dispersal interactions. In this study, I use a network approach and avian seed-dispersal networks from 23 different geographical areas to test five hypotheses about species characteristics determining the structure of the assemblage. I expected (i) large birds to forage on a large number of fruits and (ii) large fruits to be dispersed by few bird species (because of morphological constraints), and (iii) highly energetic fruits to be dispersed by more bird species (in accordance with optimal foraging theory). Besides the number of interacting partners, I also expected (iv) large birds and (v) small energetic fruits to be important for the maintenance of the structure of the interactions in seed-dispersal networks. As species that are closely related are more likely to be similar to each other, I performed phylogenetically corrected analyses to account for this data dependence. Although bird size was not associated to species important in the maintenance of the structure of the seed-dispersal community, I identified that bird species whose diet was strongly dependent on fruits were important for the structure of the network. Regarding the plants, I found that large fruits were dispersed by fewer species, but the most important attribute to predict the role of a fruit was its energy content (higher energy, more bird species dispersing the plant, but low-energy fruits being of conservation concern because they are dispersed by specific species). The results of this study suggest that the role of the species in seed-dispersal assemblages seems to be determined by the role of the species as consumers (frugivory degree for animals) or by their nutritional inputs (energy content for fruits) rather than by morphological constrains.

Untangling the Tangled Bank: A Novel Method for Partitioning the Effects of Phylogenies and Traits on Ecological Networks

Understanding how evolutionary and ecological processes shape species interaction networks remains as one of the main challenges in eco-evolutionary studies. Here, we present an integrative analytical framework to partition the effects of phylogenies and functional traits on the structure of ecological networks. The method combines fuzzy set theory and matrix correlation, implemented under a Monte Carlo framework. We designed a simulation study in order to estimate the accuracy of the methods proposed here, measuring Type I Error rates. The simulation study shows that the method is accurate, i.e., incorrectly rejecting a true null hypothesis in ~5% of the cases and falling within the confidence interval. We illustrate our framework using data from a seed dispersal network from southern Brazil. Our analyses suggest that birds must have specific traits in order to consume their plant resources, and that phylogenetic resemblance has no explanatory power for species traits and species interactions in this seed-dispersal network.

Quantifying species contributions to ecosystem processes: a global assessment of functional trait and phylogenetic metrics across avian seed-dispersal networks.

Quantifying the role of biodiversity in ecosystems not only requires understanding the links between species and the ecological functions and services they provide, but also how these factors relate to measurable indices, such as functional traits and phylogenetic diversity. However, these relationships remain poorly understood, especially for heterotrophic organisms within complex ecological networks. Here, we assemble data on avian traits across a global sample of mutualistic plant-frugivore networks to critically assess how the functional roles of frugivores are associated with their intrinsic traits, as well as their evolutionary and functional distinctiveness. We find strong evidence for niche complementarity, with phenotypically and phylogenetically distinct birds interacting with more unique sets of plants. However, interaction strengths-the number of plant species dependent on a frugivore-were unrelated to evolutionary or functional distinctiveness, largely because distinct frugivores tend to be locally rare, and thus have fewer connections across the network. Instead, interaction strengths were better predicted by intrinsic traits, including body size, gape width and dietary specialization. Our analysis provides general support for the use of traits in quantifying species ecological functions, but also highlights the need to go beyond simple metrics of functional or phylogenetic diversity to consider the multiple pathways through which traits may determine ecological processes.

Ecological networks are more sensitive to plant than to animal extinction under climate change.

Impacts of climate change on individual species are increasingly well documented, but we lack understanding of how these effects propagate through ecological communities. Here we combine species distribution models with ecological network analyses to test potential impacts of climate change on >700 plant and animal species in pollination and seed-dispersal networks from central Europe. We discover that animal species that interact with a low diversity of plant species have narrow climatic niches and are most vulnerable to climate change. In contrast, biotic specialization of plants is not related to climatic niche breadth and vulnerability. A simulation model incorporating different scenarios of species coextinction and capacities for partner switches shows that projected plant extinctions under climate change are more likely to trigger animal coextinctions than vice versa. This result demonstrates that impacts of climate change on biodiversity can be amplified via extinction cascades from plants to animals in ecological networks.

Refaunation and the reinstatement of the seed-dispersal function in Gorongosa National Park.

Large animals are important seed dispersers; however, they tend to be under a high extinction risk worldwide. There is compelling evidence that the global biodiversity crisis is leading to the deterioration of several ecosystem functions, but there is virtually no information on how large-scale refaunation efforts can reinstate seed dispersal. We evaluated the effectiveness of a 62-km2 wildlife sanctuary, which was established to recover populations of large mammals in Gorongosa National Park (Mozambique), in restoring seed dispersal. We collected animal scats during the dry season of 2014 (June-August) along 5 transects inside and 5 transects outside the sanctuary fence (50 km total) with the same type of plant community, identified animal and plant species in the transects, and quantified the number of seeds in each scat. Based on these data, we built bipartite networks and calculated network and species-level descriptor values, and we compared data collected inside and outside the sanctuary. There were more scats (268 vs. 207) and more scats containing seeds (132 vs. 94) inside than outside the sanctuary. The number of mammal dispersers was also higher inside (17) than outside the sanctuary (11). Similarly, more seeds (2413 vs. 2124) and plant species (33 vs. 26) were dispersed inside than outside the sanctuary. Overall, the seed-dispersal network was less specialized (0.38 vs. 0.44) and there was a greater overlap (0.16 vs. 0.07) inside than outside the sanctuary. Both networks were significantly modular and antinested. The high number and richness of seeds dispersed inside the sanctuary was explained mostly by a higher abundance of dispersers rather than by disperser identity. Our results suggest conservation efforts aimed at recovering populations of large mammals are helping to reestablish not only target mammal species but also their functional roles as seed dispersers in the ecosystem.

Human Impacts and Climate Change Influence Nestedness and Modularity in Food-Web and Mutualistic Networks

Theoretical studies have indicated that nestedness and modularity—non-random structural patterns of ecological networks—influence the stability of ecosystems against perturbations; as such, climate change and human activity, as well as other sources of environmental perturbations, affect the nestedness and modularity of ecological networks. However, the effects of climate change and human activities on ecological networks are poorly understood. Here, we used a spatial analysis approach to examine the effects of climate change and human activities on the structural patterns of food webs and mutualistic networks, and found that ecological network structure is globally affected by climate change and human impacts, in addition to current climate. In pollination networks, for instance, nestedness increased and modularity decreased in response to increased human impacts. Modularity in seed-dispersal networks decreased with temperature change (i.e., warming), whereas food web nestedness increased and modularity declined in response to global warming. Although our findings are preliminary owing to data-analysis limitations, they enhance our understanding of the effects of environmental change on ecological communities.

Geographical variation in the heterogeneity of mutualistic networks.

Plant–animal mutualistic networks are characterized by highly heterogeneous degree distributions. The majority of species interact with few partner species, while a small number are highly connected to form network hubs that are proposed to play an important role in community stability. It has not been investigated, however, if or how the degree distributions vary among types of mutualisms or communities, or between plants and animals in the same network. Here, we evaluate the degree distributions of pollination and seed-dispersal networks, which are two major types of mutualistic networks that have often been discussed in parallel, using an index based on Pielou's evenness. Among 56 pollination networks we found strong negative correlation of the heterogeneity between plants and animals, and geographical shifts of network hubs from plants in temperate regions to animals in the tropics. For 28 seed-dispersal networks, by contrast, the correlation was positive, and there is no comparable geographical pattern. These results may be explained by evolution towards specialization in the presence of context-dependent costs that occur if plants share the animal species as interaction partner. How the identity of network hubs affects the stability and resilience of the community is an important question for future studies.

Linking structure and functionality in mutualistic networks: do core frugivores disperse more seeds than peripheral species?

Seed dispersal networks are often organized in nested structures in a way that a few core species can disproportionally affect the remaining species in a network, in both the ecological and evolutionary sense. Yet, the relative importance of core and peripheral species has not been properly tested in seed dispersal network studies. We determined core species from 10 local seed dispersal networks composed by fleshy‐fruited plants and frugivorous birds. Each of those local quantitative networks was characterized with a core quality value, a core score for each species and a threshold value between core and peripheral species. From a total of 52 bird and 69 plant species that interacted in the study area, only 8 and 15, respectively, were identified as core. Each local network had a core that comprised a small number of birds and plants, always lower than 30% of the interacting species. There was no difference in the quantitative component of seed dispersal effectiveness (QC) provided by the frugivorous bird assemblage to plant functional groups clustered according to their growth form and fruit characteristics. Core birds dispersed seeds from each of these plant functional groups with a higher QC than peripheral species. Thus, we empirically demonstrate for the first time that seed dispersal networks at a regional scale have a small core set of fruit‐eating birds, upon which heavily rely most fleshy‐fruited plants for their seed removal. Hence, the activity of just a few core frugivores could deeply impact the demography of an entire assemblage of fleshy‐fruited plants.

Quantity and spatial distribution of seeds dispersed by a western lowland gorilla population in south-east Cameroon

Abstract:Understanding the ecological function of large frugivores in tropical forests is important considering their central position in seed-dispersal networks. This study illustrates the seed-dispersal effectiveness of a western lowland gorilla (Gorilla gorilla gorilla) population in Cameroon by looking at the interaction of seed quantity, species regeneration traits and the environmental characteristics of the deposition sites (sleeping sites vs. feeding trails) in order to discuss the role of gorillas within the ecosystem. The analysis of 1030 faecal samples collected over a 3-y period (September 2009–August 2012) showed that gorillas dispersed a diverse array of species (58 spp. identified), with an average of 289 intact seeds (>5 mm) d−1 per individual. Seed damage during mouth and gut processing occurred for 12 spp., generally accounting for <5% of specific seed load. Germination trials suggested that germination success of passed seeds remained unchanged (N = 6 spp.) or improved (N = 5 spp.) after gut passage compared with hand-cleaned seeds, with one exception. Indicator species identification (IndVal method) and General Discriminant Analysis suggested that sleeping sites (N = 36), as a main type of seed deposition site, correspond to regenerating microhabitats and correspondingly degree of canopy openness is one variable discriminating significantly this type of deposition site with random locations in the forest (N = 36) used as a proxy for non-directed seed deposition on feeding trails. With a mean canopy openness of 19–27%, sleeping sites offer optimal light conditions for the seedling recruitment of non-pioneer light-demanding species, the seeds of which encompass 47.4% of the total seed load dispersed over the 3-y period. From this study, it is hypothesized that the gorilla performed directed-dispersal of some light-demanding plant species at sleeping sites, and hence, by being actively implicated in gap-phase dynamics, fulfils a valuable ecological function within its ecosystem.

Functional importance of avian seed dispersers changes in response to human-induced forest edges in tropical seed-dispersal networks.

Although seed-dispersal networks are increasingly used to infer the functioning of ecosystems, few studies have investigated the link between the properties of these networks and the ecosystem function of seed dispersal by animals. We investigate how frugivore communities and seed dispersal change with habitat disturbance and test whether relationships between morphological traits and functional roles of seed dispersers change in response to human-induced forest edges. We recorded interaction frequencies between fleshy fruited plants and frugivorous bird species in tropical montane forests in the Bolivian Andes and recorded functional bird traits (body mass, gape width and wing tip length) associated with quantitative (seed-removal rate) and qualitative (seed-deposition pattern) components of seed-dispersal effectiveness. We found that the abundance and richness of frugivorous birds were higher at forest edges. More fruits were removed and dispersed seeds were less clustered at edges than in the interior. Additionally, functional and interaction diversity were higher at edges than in the interior, but functional and interaction evenness did not differ. Interaction strength of bird species increased with body mass, gape width and wing tip length in the forest interior, but was not related to bird morphologies at forest edges. Our study suggests that increases in functional and interaction diversity and an even distribution of interaction strength across bird morphologies lead to enhanced quantity and tentatively enhanced quality of seed dispersal. It also suggests that the effects of species traits on ecosystem functions can vary along small-scale gradients of human disturbance.

Partitioning of seed dispersal services between birds and bats in a fragment of the Brazilian Atlantic Forest

Community-level network studies suggest that seed dispersal networks may share some universal properties with other complex systems. However, most of the datasets used so far in those studies have been strongly biased towards temperate birds, including not only dispersers, but also seed predators. Recent evidence from multi-taxon networks suggests that seed dispersal networks are not all alike and may be more complex than previously thought. Here, we used network theory to evaluate seed dispersal in a strongly impacted Atlantic Forest fragment in northeastern Brazil, where bats and birds are the only extant dispersers. We hypothesized that the seed dispersal network should be more modular then nested, and that the dispersers should segregate their services according to dispersal syndromes. Furthermore, we predicted that bat and bird species that are more specialized in frugivory would be more important for maintaining the network structure. The mixed network contained 56 plant species, 12 bat species, and eight bird species, and its structure was more modular (M = 0.58) then nested (NODF = 0.21) compared with another multi-taxon network and 21 single-taxon networks (with either bats or birds). All dispersed fruits had seeds smaller than 9 mm. Bats dispersed mainly green fruits, whereas birds dispersed fruits of various colors. The network contained eight modules: five with birds only, two with bats only, and one mixed. Most dispersers were peripheral, and only specialized frugivores acted as hubs or connectors. Our results strongly support recent studies, suggesting that seed dispersal networks are complex mosaics, where different taxa form separate modules with different properties, which in turn play complementary roles in the maintenance of the associated ecosystem functions and services.

Reconstructing past ecological networks: the reconfiguration of seed-dispersal interactions after megafaunal extinction.

The late Quaternary megafaunal extinction impacted ecological communities worldwide, and affected key ecological processes such as seed dispersal. The traits of several species of large-seeded plants are thought to have evolved in response to interactions with extinct megafauna, but how these extinctions affected the organization of interactions in seed-dispersal systems is poorly understood. Here, we combined ecological and paleontological data and network analyses to investigate how the structure of a species-rich seed-dispersal network could have changed from the Pleistocene to the present and examine the possible consequences of such changes. Our results indicate that the seed-dispersal network was organized into modules across the different time periods but has been reconfigured in different ways over time. The episode of megafaunal extinction and the arrival of humans changed how seed dispersers were distributed among network modules. However, the recent introduction of livestock into the seed-dispersal system partially restored the original network organization by strengthening the modular configuration. Moreover, after megafaunal extinctions, introduced species and some smaller native mammals became key components for the structure of the seed-dispersal network. We hypothesize that such changes in network structure affected both animal and plant assemblages, potentially contributing to the shaping of modern ecological communities. The ongoing extinction of key large vertebrates will lead to a variety of context-dependent rearranged ecological networks, most certainly affecting ecological and evolutionary processes.

Ecological, historical and evolutionary determinants of modularity in weighted seed‐dispersal networks

Modularity is a recurrent and important property of bipartite ecological networks. Although well-resolved ecological networks describe interaction frequencies between species pairs, modularity of bipartite networks has been analysed only on the basis of binary presence-absence data. We employ a new algorithm to detect modularity in weighted bipartite networks in a global analysis of avian seed-dispersal networks. We define roles of species, such as connector values, for weighted and binary networks and associate them with avian species traits and phylogeny. The weighted, but not binary, analysis identified a positive relationship between climatic seasonality and modularity, whereas past climate stability and phylogenetic signal were only weakly related to modularity. Connector values were associated with foraging behaviour and were phylogenetically conserved. The weighted modularity analysis demonstrates the dominating impact of ecological factors on the structure of seed-dispersal networks, but also underscores the relevance of evolutionary history in shaping species roles in ecological communities.

Large vertebrates as the missing components of seed-dispersal networks

Large-bodied frugivores may play a key role in the networks of plants and their seed dispersers. These species, however, are often threatened by human impacts that lead to defaunation. In this paper, we discuss the potential implications of the loss of large frugivores for seed-dispersal networks. First, we review the role of large vertebrates as seed dispersers in different tropical ecosystems to show that these species are likely to be important components of seed-dispersal networks. Second, we showed that, despite their importance, large vertebrates are absent from most of the seed-dispersal networks described in the available literature. We identified three main reasons for this absence: (1) large vertebrates have already died out in the studied areas; (2) studies focus on particular taxonomic groups that consist mostly of small birds; and (3) it is inherently difficult to describe the interactions of naturally rare and secretive species such as large vertebrates. We argue that a proper evaluation of the importance of large frugivores within seed-dispersal networks would benefit from considering aspects other than the frequency of interactions. We suggest weighting pairwise interactions by their ecological consequences to quantify the contribution of large frugivores to outcomes of seed dispersal, such as landscape connectivity.

Seed dispersal interactions in the Mediterranean Region: contrasting patterns between islands and mainland

AbstractAim We studied, for the first time, the effect of insularity on plant–seed disperser networks at both the community and species level. We focused on the Mediterranean shrubland, comparing different biogeographical scenarios (insular and continental) that share the same basic species composition.Location The study was conducted within one mainland (southern Spain) and four island localities: two in the Canary Islands (oceanic origin) and two in the Balearic Islands (continental origin).Methods We built qualitative (presence/absence of interaction) and quantitative seed dispersal networks (number of consumed fruits) and used different descriptor parameters, such as connectance, nestedness, interaction asymmetry and various interaction diversity indices that describe their topology. To assemble the interaction networks, we used data on the presence of different species of fruits in systematically collected droppings of the different seed dispersers; seeds in droppings were identified by means of a binocular lens whereas a microscope was used to identify pericarp tissue remains.Results Island networks were smaller and less complex (a lower number of observed links than expected for their size) than the mainland network. As expected, connectance was higher within islands than on the mainland. By contrast, nestedness was consistently high at all sites, although relative nestedness (which accounts for network size) was lower within islands, whether continental or oceanic. At both community and species level (especially for animals) interactions tended to be more specialized and symmetric within the islands.Main conclusions The lower species number and greater specialization in insular seed dispersal systems appears to lead to the prevalence of more symmetric interactions than those found on the mainland. This indicates that insular mutualistic interactions and interacting species are more vulnerable than mainland ones to disturbances, as previous work has suggested that asymmetrical interactions facilitate the maintenance of biodiversity and act as a resilience mechanism against species extinction.