Community connectivity and local heterogeneity explain animal species co-occurrences within pond communities.

Metacommunity processes have the potential to determine most features of the community structure. However, species diversity has been the dominant focus of studies. Nestedness, modularity and checkerboard distribution of species occurrences are main components of biodiversity organisation. Within communities, these patterns emerge from the interaction between functional diversity, spatial heterogeneity and resource availability. Additionally, the connectivity determines the pool of species for community assembly and, eventually, the pattern of species co-occurrence within communities. Despite the recognised theoretical expectations, the change in occurrence patterns within communities along ecological gradients has seldom been considered. Here, we analyse the spatial occurrence of animal species along sampling units within 18 temporary ponds and its relationship with pond environments and geographic isolation. Isolated ponds presented a nested organisation of species with low spatial segregation-modularity and checkerboard-and the opposite was found for communities with high connectivity. A pattern putatively explained by high functional diversity in ponds with large connectivity and heterogeneity, which determines that species composition tracks changes in microhabitats. On the contrary, nestedness is promoted in dispersal-limited communities with low functional diversity, where microhabitat filters mainly affect richness without spatial replacement between functional groups. Vegetation biomass promotes nestedness, probably due to the observed increase in spatial variance in biomass with the mean biomass. Similarly, the richness of vegetation reduced the spatial segregation of animals within communities. This result may be due to the high plant diversity of the pond that is observed similarly along all sampling units, which promotes the spatial co-occurrence of species at this scale. In the study system, the spatial arrangement of species within communities is related to local drivers as heterogeneity and metacommunity processes by means of dispersal between communities. Patterns of species co-occurrence are interrelated with community biodiversity and species interactions, and consequently with most functional and structural properties of communities. These results indicate that understanding the interplay between metacommunity processes and co-occurrence patterns is probably more important than previously thought to understand biodiversity assembly and functioning.

Los procesos metacomunitarios tienen el potencial de determinar la mayoría de las características de la estructura de las comunidades. Sin embargo, los trabajos se han enfocado principalmente en los patrones de diversidad de especies. El anidamiento, la modularidad y la distribución en damero de la ocurrencia espacial de las especies son propiedades básicas de las comunidades. Estos patrones surgen de la interacción entre la diversidad funcional, la heterogeneidad espacial y la disponibilidad de recursos dentro de las comunidades. Además, el pool de especies disponibles para el ensamblaje está determinado por la conectividad de la comunidad, afectando así su patrón de co­ocurrencia de especies. A pesar de las reconocidas expectativas teóricas, el cambio en los patrones de ocurrencia dentro de las comunidades a lo largo de gradientes ecológicos ha sido poco considerado. Aquí, analizamos la ocurrencia espacial de especies animales dentro de 18 charcos temporales y su relación con las características ambientales y el aislamiento geográfico de los charcos. Los charcos aislados presentaron alto anidamiento espacial mientras que los charcos de alta conectividad una distribución de ocurrencias modular y en damero. Por un lado, la baja diversidad funcional en charcos aislados, determinaría que los filtros microambientales afecten la riqueza de especies sin reemplazo espacial entre grupos funcionales, promoviendo un arreglo anidado de ocurrencias. Por otro lado, la alta diversidad funcional en charcos con alta conectividad y heterogeneidad permitiría el reemplazo espacial de especies en gradientes microambientales, determinando los patrones de segregación observados. La biomasa vegetal promueve el anidamiento, probablemente debido al aumento observado en la variación espacial de la biomasa con la biomasa media. La riqueza vegetal también redujo la segregación espacial de los animales dentro de las comunidades. Este resultado puede deberse a que la alta diversidad de plantas de los charcos es también observada a nivel de unidades muestreales, favoreciendo esto la coexistencia espacial de especies. El arreglo espacial de especies dentro de las comunidades estudiadas estaría determinado tanto por factores locales como la heterogeneidad, como por procesos regionales operando a través de la dispersión de individuos entre comunidades. Los patrones de co­ocurrencia de especies están interrelacionados con la diversidad comunitaria y las interacciones bióticas, y consecuentemente con la mayoría de las propiedades estructurales y funcionales de las comunidades. Este estudio evidencia la importancia de la conexión entre procesos metacomunitarios y la co­ocurrencia espacial de especies para comprender el ensamblaje y funcionamiento de la biodiversidad.

Both local stability and dispersal contribute to metacommunity sensitivity to asynchronous habitat availability.

The stability of isolated communities depends on the complexity of their foodwebs. However, it remains unclear how local stability interacts with dispersal in multitrophic metacommunities to shape biodiversity patterns. This lack of understanding is deeper in the more realistic frame of landscapes that exhibit non-trivial and time-varying structures. Therefore, in this study, we aim to evaluate the influence of local stabilizing factors versus dispersal in determining the sensitivity of metacommunity biodiversity to increasing asynchrony of site availability. Additionally, we assess the role of foodweb complexity and landscape structure as modulating factors. To accomplish our goals we developed a model based on random matrices for local communities, which are linked by stochastic dispersal over explicit dynamic landscapes. We ran numerical simulations and computed the effect sizes of foodweb temperature, self-limitation, dispersal ability, and all pairwise combinations, on the sensitivity of biodiversity to landscape asynchrony. In our experiments we explored gradients of species richness, foodweb connectance, number of sites, and landscape modularity. Our results showed that asynchrony among site availability periods reduced α -diversity and increased ß -diversity. Asynchrony increased γ -diversity at high dispersal rates. Both local and regional stabilizing factors determined the sensitivity of metacommunities to landscape asynchrony. Local factors were more influential in landscapes with fewer sites and lower modularity, as well as in metacommunities composed of complex foodwebs. This research offers insights into the dynamics of metacommunities in dynamic landscapes, providing valuable knowledge about the interplay between local and regional factors in shaping ecological stability and species persistence.

Prey selection along a predators' body size gradient evidences the role of different trait-based mechanisms in food web organization.

An increase in prey richness, prey size and predator trophic position with predator body size has been consistently reported as prime features of food web organization. These trends have been explained by non-exclusive mechanisms. First, the increase in energy demand with body size determines that larger predators must reduce prey selectivity for achieving the required number of resources, being consumption relationships independent of prey traits. Second, when consumption is restricted by gape limitation, small predators are constrained to select among small prey. However, this selection weakens over large predators, which progressively consume more and larger prey. Finally, the optimal foraging mechanism predicts that larger predators optimize their diet by selecting only large prey with high energy reward. Each one of these mechanisms can individually explain the increase in prey richness, prey size and predator trophic position with predator body size but their relative importance or the direct evidence for their combined role was seldom considered. Here we use the community assembly by trait selection (CATS) theory for evaluating the support for each one of these mechanisms based on the prey selection patterns that they predict. We analyzed how prey body size and trophic guild determine prey selection by predators of increasing body size in a killifish guild from a temporary pond system. Results support the combination of the three mechanisms to explain the structural trends in our food web, although their strength is contingent on prey trophic group. Overall, high energy prey are preferred by larger predators, and small predators select small prey of all trophic status. However, large predators prefer large primary producers and avoid large carnivorous prey, probably because of the inherent risk of consuming other carnivorous. Our study provides a mechanistic understanding of how predator traits determine the selection of prey traits affecting food web assembly.

Scaling of biological rates with body size as a backbone in the assembly of metacommunity biodiversity.

The dispersal-body mass association has been highlighted as a main determinant of biodiversity patterns in metacommunities. However, less attention has been devoted to other well-recognized determinants of metacommunity diversity: the scaling in density and regional richness with body size. Among active dispersers, the increase in movement with body size may enhance local richness and decrease ß-diversity. Nevertheless, the reduction of population size and regional richness with body mass may determine a negative diversity-body size association. Consequently, metacommunity assembly probably emerges from a balance between the effect of these scalings. We formalize this hypothesis by relating the exponents of size-scaling rules with simulated trends in α-, ß- and γ-diversity with body size. Our results highlight that the diversity-body size relationship in metacommunities may be driven by the combined effect of different scaling rules. Given their ubiquity in most terrestrial and aquatic biotas, these scaling rules may represent the basic determinants-backbone-of biodiversity, over which other mechanisms operate determining metacommunity assembly. Further studies are needed, aimed at explaining biodiversity patterns from functional relationships between biological rates and body size, as well as their association with environmental conditions and species interactions.

Avoiding artifacts when varying the number of species in ecological models.

Ecological theory recognizes the importance of the variety of species for maintaining the functioning of ecosystems and their derived services. We assert that when studying the effects of shifts in biodiversity levels using mathematical models, their dynamics must be sensitive to the variety of species traits but not to raw species numbers, a property that we call order-invariance. We present a testing procedure for verifying order-invariance of ecological network models -with or without trait adaptation- expressed as ODEs. Furthermore, we applied our test to several influential models used for evaluating biodiversity effects on ecosystem functioning. In most of the surveyed studies the equations failed our test. This raises doubts about the validity of previous results and calls for revisiting the theory derived from these studies. Our results foster the creation of artifact-free models, a necessary step towards building a more robust theory of biodiversity-driven ecosystem functioning.

Core-periphery dynamics in a plant-pollinator network.

Mutualistic networks are highly dynamic, characterized by high temporal turnover of species and interactions. Yet, we have a limited understanding of how the internal structure of these networks and the roles species play in them vary through time. We used 6 years of observation data and a novel statistical method (dynamic stochastic block models) to assess how network structure and species' structural position within the network change throughout subseasons of the flowering season and across years in a quantitative plant-pollinator network from a dryland ecosystem in Argentina. Our analyses revealed a core-periphery structure persistent through subseasons and years. Yet, species structural position as core or peripheral was highly dynamic: virtually all species that were at the core in some subseasons were also peripheral in other subseasons, while many other species always remained peripheral. Our results illuminate our understanding of the dynamics of mutualistic networks and have important implications for ecosystem management and conservation.

Geographical origin determines responses to salinity of Mediterranean caddisflies.

Many freshwater ecosystems worldwide, and particularly Mediterranean ones, show increasing levels of salinity. These changes in water conditions could affect abundance and distribution of inhabiting species as well as the provision of ecosystem services. In this study we conduct laboratory experiments using the macroinvertebrate Smicridea annulicornis as a model organism. Our factorial experiments were designed to evaluate the effects of geographical origin of organisms and salinity levels on survival and behavioral responses of caddisflies. The experimental organisms were captured from rivers belonging to three hydrological basins along a 450 Km latitudinal gradient in the Mediterranean region of Chile. Animals were exposed to three conductivity levels, from 180 to 1400 µS/cm, close to the historical averages of the source rivers. We measured the behavioral responses to experimental stimuli and the survival time. Our results showed that geographical origin shaped the behavioral and survival responses to salinity. In particular, survival and activity decreased more strongly with increasing salinity in organisms coming from more dilute waters. This suggests local adaptation to be determinant for salinity responses in this benthic invertebrate species. In the current scenario of fast temporal and spatial changes in water levels and salt concentration, the conservation of geographic intra-specific variation of aquatic species is crucial for lowering the risk of salinity-driven biodiversity loss.

Non-trophic interactions strengthen the diversity-functioning relationship in an ecological bioenergetic network model.

Ecological communities are undeniably diverse, both in terms of the species that compose them as well as the type of interactions that link species to each other. Despite this long recognition of the coexistence of multiple interaction types in nature, little is known about the consequences of this diversity for community functioning. In the ongoing context of global change and increasing species extinction rates, it seems crucial to improve our understanding of the drivers of the relationship between species diversity and ecosystem functioning. Here, using a multispecies dynamical model of ecological communities including various interaction types (e.g. competition for space, predator interference, recruitment facilitation in addition to feeding), we studied the role of the presence and the intensity of these interactions for species diversity, community functioning (biomass and production) and the relationship between diversity and functioning.Taken jointly, the diverse interactions have significant effects on species diversity, whose amplitude and sign depend on the type of interactions involved and their relative abundance. They however consistently increase the slope of the relationship between diversity and functioning, suggesting that species losses might have stronger effects on community functioning than expected when ignoring the diversity of interaction types and focusing on feeding interactions only.

Phenology determines the robustness of plant-pollinator networks.

Plant-pollinator systems are essential for ecosystem functioning, which calls for an understanding of the determinants of their robustness to environmental threats. Previous studies considering such robustness have focused mostly on species' connectivity properties, particularly their degree. We hypothesized that species' phenological attributes are at least as important as degree as determinants of network robustness. To test this, we combined dynamic modeling, computer simulation and analysis of data from 12 plant-pollinator networks with detailed information of topology of interactions as well as species' phenology of plant flowering and pollinator emergence. We found that phenological attributes are strong determinants of network robustness, a result consistent across the networks studied. Plant species persistence was most sensitive to increased larval mortality of pollinators that start earlier or finish later in the season. Pollinator persistence was especially sensitive to decreased visitation rates and increased larval mortality of specialists. Our findings suggest that seasonality of climatic events and anthropic impacts such as the release of pollutants is critical for the future integrity of terrestrial biodiversity.

Species traits and network structure predict the success and impacts of pollinator invasions.

Species invasions constitute a major and poorly understood threat to plant-pollinator systems. General theory predicting which factors drive species invasion success and subsequent effects on native ecosystems is particularly lacking. We address this problem using a consumer-resource model of adaptive behavior and population dynamics to evaluate the invasion success of alien pollinators into plant-pollinator networks and their impact on native species. We introduce pollinator species with different foraging traits into network models with different levels of species richness, connectance, and nestedness. Among 31 factors tested, including network and alien properties, we find that aliens with high foraging efficiency are the most successful invaders. Networks exhibiting high alien-native diet overlap, fraction of alien-visited plant species, most-generalist plant connectivity, and number of specialist pollinator species are the most impacted by invaders. Our results mimic several disparate observations conducted in the field and potentially elucidate the mechanisms responsible for their variability.

Niche partitioning due to adaptive foraging reverses effects of nestedness and connectance on pollination network stability.

Much research debates whether properties of ecological networks such as nestedness and connectance stabilise biological communities while ignoring key behavioural aspects of organisms within these networks. Here, we computationally assess how adaptive foraging (AF) behaviour interacts with network architecture to determine the stability of plant-pollinator networks. We find that AF reverses negative effects of nestedness and positive effects of connectance on the stability of the networks by partitioning the niches among species within guilds. This behaviour enables generalist pollinators to preferentially forage on the most specialised of their plant partners which increases the pollination services to specialist plants and cedes the resources of generalist plants to specialist pollinators. We corroborate these behavioural preferences with intensive field observations of bee foraging. Our results show that incorporating key organismal behaviours with well-known biological mechanisms such as consumer-resource interactions into the analysis of ecological networks may greatly improve our understanding of complex ecosystems.

A conceptual framework for studying the strength of plant-animal mutualistic interactions.

The strength of species interactions influences strongly the structure and dynamics of ecological systems. Thus, quantifying such strength is crucial to understand how species interactions shape communities and ecosystems. Although the concepts and measurement of interaction strength in food webs have received much attention, there has been comparatively little progress in the context of mutualism. We propose a conceptual scheme for studying the strength of plant-animal mutualistic interactions. We first review the interaction strength concepts developed for food webs, and explore how these concepts have been applied to mutualistic interactions. We then outline and explain a conceptual framework for defining ecological effects in plant-animal mutualisms. We give recommendations for measuring interaction strength from data collected in field studies based on a proposed approach for the assessment of interaction strength in plant-animal mutualisms. This approach is conceptually integrative and methodologically feasible, as it focuses on two key variables usually measured in field studies: the frequency of interactions and the fitness components influenced by the interactions.

Zooplankton competition promotes trade-offs affecting diapause in rotifers.

Facultative diapause should be favoured by natural selection in temporary variable habitats. Diapause patterns are evolutionary constrained because producing diapause is resource demanding, which might have implications for competitive dynamics and competitor coexistence through mechanisms such as the storage effect. Besides these implications, competition intensity might affect the quality of the diapausing stages and the reproductive success of the offspring emerging from them. We experimentally analysed traits involved in diapause in the cyclically parthenogenetic rotifer Brachionus calyciflorus, in relation to the presence of its competitor, the cladoceran Moina micrura. Under competition with Moina, Brachionus produced fewer diapausing eggs, most of which possessed visual attributes indicating a higher quality. These diapausing eggs produced under a competitive environment had a higher hatching success; however, the hatchlings exhibited a lower population growth rate. We propose the existence of trade-offs between traits related to diapause: the number of produced eggs, quality of these eggs and hatchling performance. Our results show that interspecific competition may cause fine-tuned changes in the life cycle patterns of the populations. Furthermore, these changes could affect that abundance and performance of competitors and thereby drive back effects on the competitive output. These diapause-driven feedback mechanisms may have strong implications for the dynamics of the natural communities.

Decadal trends in the pollinator assemblage of Eucryphia cordifolia in Chilean rainforests.

Long-term studies of plant-pollinator interactions are almost nonexistent in the scientific literature. The objective of the present study was to determine changes and trends in the pollinator assemblage of ulmo (Eucryphia cordifolia; Cunoniaceae), a canopy-emergent tree found in Chilean temperate rainforests. We assessed the temporal variability of the pollinator assemblage and identified possible modulators of the observed temporal shifts. We sampled insect visitors to the flowers of 16 individual trees of E. cordifolia during 10 consecutive flowering seasons (2000-2009), recording a total of 137 pollinator species with a mean number of species per year of 44. Only three pollinator species (2.2%) were recorded every year. Two bee species accounted for 50% of all insect visits to flowers. One bee species, Bombus dahlbomii (native), was dominant in one season, whereas Apis mellifera (exotic) dominated during the next season. These interannual shifts in population abundances presented first-order dynamics that were characterized by oscillations with a period of 2 years. Changes in the abundances of the dominant pollinators, as well as differences in temperature and precipitation during insect emergence and flowering, led to a nested temporal structure of pollinator composition. Furthermore, the abundances of less common pollinators were sensitive to the abundance of the dominant bee species and to monthly maximum temperatures and the average precipitation during spring and summer. Based on our results and those from other studies, we predict a decline in the numbers of Bombus dahlbomii and nondominant native pollinators in response to new exotic arrivals.

Topological plasticity increases robustness of mutualistic networks.

1. Earlier studies used static models to evaluate the responses of mutualistic networks to external perturbations. Two classes of dynamics can be distinguished in ecological networks; population dynamics, represented mainly by changes in species abundances, and topological dynamics, represented by changes in the architecture of the web. 2. In this study, we model the temporal evolution of three empirical plant-pollination networks incorporating both population and topological dynamics. We test the hypothesis that topological plasticity, realized through the ability of animals to rewire their connections after depletion of host abundances, enhances tolerance of mutualistic networks to species loss. We also compared the performance of various rewiring rules in affecting robustness. 3. The results show that topological plasticity markedly increased the robustness of mutualistic networks. Our analyses also revealed that network robustness reached maximum levels when animals with less host plant availability were more likely to rewire. Also, preferential attachment to richer host plants, that is, to plants exhibiting higher abundance and few exploiters, enhances robustness more than other rewiring alternatives. 4. Our results highlight the potential role of topological plasticity in the robustness of mutualistic networks to species extinctions and suggest some plausible mechanisms by which the decisions of foragers may shape the collective dynamics of plant-pollinator systems.

Retrospective qualitative analysis of ecological networks under environmental perturbation: a copper-polluted intertidal community as a case study.

The coast of Chañaral Bay in northern Chile has been affected by copper mine wastes for decades. This sustained perturbation has disrupted the intertidal community in several ways, but the mechanisms behind the observed shifts in local biodiversity remain poorly understood. Our main goal was to identify the species (lumped into trophic groups) belonging to the Chañaral intertidal community that, being directly affected by copper pollution, contributed primarily to the generation of the observed changes in community structure. These groups of species were called initiators. We applied a qualitative modelling approach based only on the sign and direction of effects among species, and present a formula for predicting changes in equilibrium abundances considering stress on multiple variables simultaneously. We then applied this technique retrospectively to identify the most likely set of initiators. Our analyses allowed identification of a unique set of four initiators in the studied intertidal system (a group of algae, sessile invertebrates, a group of herbivores and starfish), which were hypothesized to be the primary drivers of the observed changes in community structure. In addition, a hypothesis was derived about how the perturbation affected these initiators. The hypothesis is that pollution affected negatively the population growth rate of both algae and sessile invertebrates and suppressed the interaction between herbivores and starfish. Our analytic approach, focused on identifying initiators, constitutes an advance towards understanding the mechanisms underlying human-driven ecosystem disruption and permits identifying species that may serve as a focal point for community management and restoration.

Consequences of adaptive behaviour for the structure and dynamics of food webs.

Species coexistence within ecosystems and the stability of patterns of temporal changes in population sizes are central topics in ecological theory. In the last decade, adaptive behaviour has been proposed as a mechanism of population stabilization. In particular, widely distributed adaptive trophic behaviour (ATB), the fitness-enhancing changes in individuals' feeding-related traits due to variation in their trophic environment, may play a key role in modulating the dynamics of feeding relationships within natural communities. In this article, we review and synthesize models and results from theoretical research dealing with the consequences of ATB on the structure and dynamics of complex food webs. We discuss current approaches, point out limitations, and consider questions ripe for future research. In spite of some differences in the modelling and analytic approaches, there are points of convergence: (1) ATB promotes the complex structure of ecological networks, (2) ATB increases the stability of their dynamics, (3) ATB reverses May's negative complexity-stability relationship, and (4) ATB provides resilience and resistance of networks against perturbations. Current knowledge supports ATB as an essential ingredient for models of community dynamics, and future research that incorporates ATB will be well positioned to address questions important for basic ecological research and its applications.

A network analysis of plant-pollinator interactions in temperate rain forests of Chiloé Island, Chile.

This study characterizes the structure of a plant-pollinator network in a temperate rain forest of Chiloé Island, southern Chile, where woody species are strongly dependent on biotic pollinators, and analyzes its robustness to the loss of participating species. Degree distribution, nestedness, and expected species persistence were evaluated. In addition, we assessed the roles of predefined subsets of plants (classified by life forms) and pollinators (grouped by taxonomic orders) in the network's structure and dynamics. For this, we simulated the complete removal of each plant and pollinator subset and analyzed the resultant connectivity patterns, as well as the expected long-term species losses by running a stochastic model. Finally, we evaluated the sensitivity of the network structure to the loss of single species in order to identify potential targets for conservation. Our results show that the plant-pollinator network of this Chilean temperate rain forest exhibits a nested structure of interactions, with a degree distribution best described by a power law model. Model simulations revealed the importance of trees and hymenopterans as pivotal groups that maintain the core structure of the pollination network and guarantee overall species persistence. The hymenopterans Bombus dahlbomii and Diphaglossa gayi, the shrubs Tepualia stipularis and Ugni molinae, the vines Mitraria coccinea and Asteranthera ovata, and the entire set of tree species exerted a disproportionately large influence on the preservation of network structure and should be considered as focal species for conservation programs given current threats from selective logging and habitat loss.

Understanding and predicting effects of modified interactions through a qualitative analysis of community structure.

Models of ecological communities are traditionally based on relationships between pairs of species, where the strengths of per capita interactions are fixed and independent of population abundance. A growing body of literature, however; describes interactions whose strength is modified by the density of either a third species or by one of the species involved in a pairwise interaction. These modified interactions have been treated as indirect effects, and the terminology addressing them is diverse and overlapping. In this paper we develop a general analytical framework based on a qualitative analysis of community structure to account for the consequence of modified interactions in complex ecological communities. Modified interactions are found to create both direct and indirect effects between species. The sign of a direct effect can change in some instances depending on the magnitude of a key variable or parameter, which leads to a threshold change in system structure and dynamics. By considering alternative structures of a community, we extend our ability to model perturbations that move the system far from a previous equilibrium. Using specific examples, we reinterpret existing results, develop hypotheses to guide experiments or management interventions, and explore the role of modified interactions and positive feedback in creating and maintaining alternative stable states. Through a qualitative analysis of community structure, system feedback is demonstrated as being key in understanding and predicting the dynamics of complex ecological communities.

Population dynamics of prey exhibiting inducible defenses: the role of associated costs and density-dependence.

The effect of antipredator behavior on the dynamics of a resource-consumer model was analyzed in relation to the magnitude of associated costs, and the strength of density-dependence. For this purpose, I present a deterministic continuous resource-consumer model that exhibits biomass conversion, structural homogeneity, and competition for renewable and fixed resources as separate processes. Antipredator behavior is incorporated as an inducible response to consumer density, and has metabolic and feeding costs. By means of numerical methods, I show: (1) that antipredator behavior is stabilizing for certain parameter ranges, where other stabilizing forces do not dominate the dynamics; (2) intraspecific competition for both fixed and renewable resources have a stabilizing role; (3) metabolic cost is always stabilizing, and feeding cost can be stabilizing or destabilizing, depending on the relative strength of the two competition forces.