Species are linked to each other by a myriad of positive and negative interactions. This complex spectrum of interactions constitutes a network of links that mediates ecological communities' response ...to perturbations, such as exploitation and climate change. In the last decades, there have been great advances in the study of intricate ecological networks. We have, nonetheless, lacked both the data and the tools to more rigorously understand the patterning of multiple interaction types between species (i.e., "multiplex networks"), as well as their consequences for community dynamics. Using network statistical modeling applied to a comprehensive ecological network, which includes trophic and diverse non-trophic links, we provide a first glimpse at what the full "entangled bank" of species looks like. The community exhibits clear multidimensional structure, which is taxonomically coherent and broadly predictable from species traits. Moreover, dynamic simulations suggest that this non-random patterning of how diverse non-trophic interactions map onto the food web could allow for higher species persistence and higher total biomass than expected by chance and tends to promote a higher robustness to extinctions.
Species co-occurrence networks Freilich, Mara A.; Wieters, Evie; Broitman, Bernardo R. ...
Ecology,
03/2018, Volume:
99, Issue:
3
Journal Article
Peer reviewed
Open access
Co-occurrence methods are increasingly utilized in ecology to infer networks of species interactions where detailed knowledge based on empirical studies is difficult to obtain. Their use is ...particularly common, but not restricted to, microbial networks constructed from metagenomic analyses. In this study, we test the efficacy of this procedure by comparing an inferred network constructed using spatially intensive co-occurrence data from the rocky intertidal zone in central Chile to a well-resolved, empirically based, species interaction network from the same region. We evaluated the overlap in the information provided by each network and the extent to which there is a bias for co-occurrence data to better detect known trophic or non-trophic, positive or negative interactions. We found a poor correspondence between the co-occurrence network and the known species interactions with overall sensitivity (probability of true link detection) equal to 0.469, and specificity (true non-interaction) equal to 0.527. The ability to detect interactions varied with interaction type. Positive non-trophic interactions such as commensalism and facilitation were detected at the highest rates. These results demonstrate that co-occurrence networks do not represent classical ecological networks in which interactions are defined by direct observations or experimental manipulations. Co-occurrence networks provide information about the joint spatial effects of environmental conditions, recruitment, and, to some extent, biotic interactions, and among the latter, they tend to better detect niche-expanding positive non-trophic interactions. Detection of links (sensitivity or specificity) was not higher for well-known intertidal keystone species than for the rest of consumers in the community. Thus, as observed in previous empirical and theoretical studies, patterns of interactions in co-occurrence networks must be interpreted with caution, especially when extending interaction-based ecological theory to interpret network variability and stability. Co-occurrence networks may be particularly valuable for analysis of community dynamics that blends interactions and environment, rather than pairwise interactions alone.
Fishes are the most diverse group of vertebrates, play key functional roles in aquatic ecosystems, and provide protein for a billion people, especially in the developing world. Those functions are ...compromised by mounting pressures on marine biodiversity and ecosystems. Because of its economic and food value, fish biomass production provides an unusually direct link from biodiversity to critical ecosystem services. We used the Reef Life Survey’s global database of 4,556 standardized fish surveys to test the importance of biodiversity to fish production relative to 25 environmental drivers. Temperature, biodiversity, and human influence together explained 47% of the global variation in reef fish biomass among sites. Fish species richness and functional diversity were among the strongest predictors of fish biomass, particularly for the large-bodied species and carnivores preferred by fishers, and these biodiversity effects were robust to potentially confounding influences of sample abundance, scale, and environmental correlations. Warmer temperatures increased biomass directly, presumably by raising metabolism, and indirectly by increasing diversity, whereas temperature variability reduced biomass. Importantly, diversity and climate interact, with biomass of diverse communities less affected by rising and variable temperatures than species-poor communities. Biodiversity thus buffers global fish biomass from climate change, and conservation of marine biodiversity can stabilize fish production in a changing ocean.
Over the past decade, evidence of abrupt latitudinal changes in the dynamics, structure and genetic variability of intertidal and subtidal benthic communities along central-northern Chile has been ...found consistently at 30-32°S. Changes in the advective and thermal environment in nearshore waters have been inferred from ecological patterns, since analyses of in situ physical data have thus far been missing. Here we analyze a unique set of shoreline temperature data, gathered over 4-10 years at 15 sites between 28-35°S, and combine it with satellite-derived winds and sea surface temperatures to investigate the latitudinal transition in nearshore oceanographic conditions suggested by recent ecological studies. Our results show a marked transition in thermal conditions at 30-31°S, superimposed on a broad latitudinal trend, and small-scale structures associated with cape-and-bay topography. The seasonal cycle dominated temperature variability throughout the region, but its relative importance decreased abruptly south of 30-31°S, as variability at synoptic and intra-seasonal scales became more important. The response of shoreline temperatures to meridional wind stress also changed abruptly at the transition, leading to a sharp drop in the occurrence of low-temperature waters at northern sites, and a concurrent decrease in corticated algal biomass. Together, these results suggest a limitation of nitrate availability in nearshore waters north of the transition. The localized alongshore change results from the interaction of latitudinal trends (e.g., wind stress, surface warming, inertial period) with a major headland-bay system (Punta Lengua de Vaca at 30.25°S), which juxtaposes a southern stretch of coast characterized by upwelling with a northern stretch of coast characterized by warm surface waters and stratification. This transition likely generates a number of latitude-dependent controls on ecological processes in the nearshore that can explain species-specific effects, and add strength to the suggestion of an oceanography-driven, major spatial transition in coastal communities at 30-31°S.
Climate change has led to intensification and poleward migration of the Southeastern Pacific Anticyclone, forcing diverging regions of increasing, equatorward and decreasing, poleward coastal ...phytoplankton productivity along the Humboldt Upwelling Ecosystem, and a transition zone around 31° S. Using a 20-year dataset of barnacle larval recruitment and adult abundances, we show that striking increases in larval arrival have occurred since 1999 in the region of higher productivity, while slower but significantly negative trends dominate poleward of 30° S, where years of recruitment failure are now common. Rapid increases in benthic adults result from fast recruitment-stock feedbacks following increased recruitment. Slower population declines in the decreased productivity region may result from aging but still reproducing adults that provide temporary insurance against population collapses. Thus, in this region of the ocean where surface waters have been cooling down, climate change is transforming coastal pelagic and benthic ecosystems through altering primary productivity, which seems to propagate up the food web at rates modulated by stock-recruitment feedbacks and storage effects. Slower effects of downward productivity warn us that poleward stocks may be closer to collapse than current abundances may suggest.
Species richness has dominated our view of global biodiversity patterns for centuries. The dominance of this paradigm is reflected in the focus by ecologists and conservation managers on richness and ...associated occurrence-based measures for understanding drivers of broad-scale diversity patterns and as a biological basis for management. However, this is changing rapidly, as it is now recognized that not only the number of species but the species present, their phenotypes and the number of individuals of each species are critical in determining the nature and strength of the relationships between species diversity and a range of ecological functions (such as biomass production and nutrient cycling). Integrating these measures should provide a more relevant representation of global biodiversity patterns in terms of ecological functions than that provided by simple species counts. Here we provide comparisons of a traditional global biodiversity distribution measure based on richness with metrics that incorporate species abundances and functional traits. We use data from standardized quantitative surveys of 2,473 marine reef fish species at 1,844 sites, spanning 133 degrees of latitude from all ocean basins, to identify new diversity hotspots in some temperate regions and the tropical eastern Pacific Ocean. These relate to high diversity of functional traits amongst individuals in the community (calculated using Rao's Q), and differ from previously reported patterns in functional diversity and richness for terrestrial animals, which emphasize species-rich tropical regions only. There is a global trend for greater evenness in the number of individuals of each species, across the reef fish species observed at sites ('community evenness'), at higher latitudes. This contributes to the distribution of functional diversity hotspots and contrasts with well-known latitudinal gradients in richness. Our findings suggest that the contribution of species diversity to a range of ecosystem functions varies over large scales, and imply that in tropical regions, which have higher numbers of species, each species contributes proportionally less to community-level ecological processes on average than species in temperate regions. Metrics of ecological function usefully complement metrics of species diversity in conservation management, including when identifying planning priorities and when tracking changes to biodiversity values.
Understanding the drivers of geographical variation in species distributions, and the resulting community structure, constitutes one of the grandest challenges in ecology. Geographical patterns of ...species richness and composition have been relatively well studied. Less is known about how the entire set of trophic and non-trophic ecological interactions, and the complex networks that they create by gluing species together in complex communities, change across geographical extents. Here, we compiled data of species composition and three types of ecological interactions occurring between species in rocky intertidal communities across a large spatial extent (~970 km of shoreline) of central Chile, and analyzed the geographical variability in these multiplex networks (i.e., comprising several interaction types) of ecological interactions. We calculated nine network summary statistics common across interaction types, and additional network attributes specific to each of the different types of interactions. We then investigated potential environmental drivers of this multivariate network organization. These included variation in sea surface temperature and coastal upwelling, the main drivers of productivity in nearshore waters. Our results suggest that structural properties of multiplex ecological networks are affected by local species richness and modulated by factors influencing productivity and environmental predictability. Our results show that non-trophic negative interactions are more sensitive to spatially structured temporal environmental variation than feeding relationships, with non-trophic positive interactions being the least labile to it. We also show that environmental effects are partly mediated through changes in species richness and partly through direct influences on species interactions, probably associated to changes in environmental predictability and to bottom-up nutrient availability. Our findings highlight the need for a comprehensive picture of ecological interactions and their geographical variability if we are to predict potential effects of environmental changes on ecological communities.
Theoretical studies have shown that coexistence between competitors can be favored in a spatially heterogeneous environment by a number of mechanisms, which ultimately allow the expression of ...persistent or transitory variation in species competitive abilities, colonization, or reproduction. Four distinctive paradigms to model metacommunities have been identified according to assumptions about the biology of the species and essential aspects of the environment. Missing from these are mechanisms of coexistence that can arise from the dispersal process itself without explicit spatial heterogeneity or biological trade-offs. These mechanisms have only recently received attention, but they may be common in marine communities and other systems in which dispersal is obligatory and modulated by the physical environment. We investigate coexistence in spatially homogeneous metacommunities where there is no partitioning of resources, no competition-colonization trade-off, and no possibility of source-sink dynamics. Coexistence is shown to be possible through three distinct mechanisms related to the dispersal process itself. Firstly, in a neutral scenario, inclusion of temporal variability in the connectivity matrix, emulating an intrinsic attribute of ocean character and other turbulent environments, can promote the invasion of an equally matched competitor and, in a hierarchical competition scenario, the persistence of an otherwise unviable, inferior competitor (the dispersal variability mechanism). Secondly, a sufficiently large difference in the shape of the time-independent dispersal kernels of the two species, which may result from differences in larval-release timing, buoyancy, or behavior, can produce stable coexistence in the center of their shared range (the dispersal-shape mechanism). Thirdly, asymmetry in the dispersal process due to biased advection renders the metapopulation model reactive, such that small variations in the upstream abundances can be sufficient for the subordinate species to stably persist (the dispersal-bias mechanism). These results demonstrate that a subordinate species may persist by occupying a dispersal niche that differs sufficiently from that of the dominant species. Further theoretical research is necessary to develop simple empirical tests for these and other dispersal-based coexistence mechanisms.
How multiple types of non-trophic interactions map onto trophic networks in real communities remains largely unknown. We present the first effort, to our knowledge, describing a comprehensive ...ecological network that includes all known trophic and diverse non-trophic links among >100 coexisting species for the marine rocky intertidal community of the central Chilean coast. Our results suggest that non-trophic interactions exhibit highly nonrandom structures both alone and with respect to food web structure. The occurrence of different types of interactions, relative to all possible links, was well predicted by trophic structure and simple traits of the source and target species. In this community, competition for space and positive interactions related to habitat/refuge provisioning by sessile and/or basal species were by far the most abundant non-trophic interactions. If these patterns are corroborated in other ecosystems, they may suggest potentially important dynamic constraints on the combined architecture of trophic and non-trophic interactions. The nonrandom patterning of non-trophic interactions suggests a path forward for developing a more comprehensive ecological network theory to predict the functioning and resilience of ecological communities.
Understanding the factors that modulate bacterial community assembly in natural soils is a longstanding challenge in microbial community ecology. In this work, we compared two microbial co-occurrence ...networks representing bacterial soil communities from two different sections of a pH, temperature and humidity gradient occurring along a western slope of the Andes in the Atacama Desert. In doing so, a topological graph alignment of co-occurrence networks was used to determine the impact of a shift in environmental variables on OTUs taxonomic composition and their relationships. We observed that a fraction of association patterns identified in the co-occurrence networks are persistent despite large environmental variation. This apparent resilience seems to be due to: (1) a proportion of OTUs that persist across the gradient and maintain similar association patterns within the community and (2) bacterial community ecological rearrangements, where an important fraction of the OTUs come to fill the ecological roles of other OTUs in the other network. Actually, potential functional features suggest a fundamental role of persistent OTUs along the soil gradient involving nitrogen fixation. Our results allow identifying factors that induce changes in microbial assemblage configuration, altering specific bacterial soil functions and interactions within the microbial communities in natural environments.
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