Long-term supratidal rockpool invertebrate community, Discovery Bay, Jamaica.

The rockpool cluster offers unique characteristics making it a model system for general questions in ecology: (1) all rockpools share biotic history (any species can reach any rockpool); (2) they form a strong gradient of conditions from benign to harsh; (3) 1-day sampling across all rockpools ensures census consistency; (4) rockpools respond to changing conditions within a short (days) time frame; (5) they are easy to manipulate (note: the data are from an unmanipulated rockpool subset), and (6) they may act as a single metacommunity that exhibits consistent species distribution patterns on a broader scale (unpublished). Consequently, the rockpools continue generating insights, with the first publications in 1996. The data represent an intensive rockpool metacommunity monitoring project, making them of considerable value to our understanding of tropical coastal metacommunity dynamics and general ecological processes. The dataset covers surveys of invertebrate fauna in 49, primarily supratidal, rockpools on a fossil coral reef over 25 years. All rockpools occur within a 73 × 47 m array of rocks at a distance of less than 2 m from the nearest neighbor. About 200 other rockpools occur on the same area. They are in a sheltered bay (Discovery Bay, Jamaica) between 0 and 5 m from the ocean. Typically, rockpools are 5-30 cm deep and 40 cm across on average, with elevation from a few centimeters to 300 cm above sea level. Rockpools may drain excess water from precipitation or waves into other rockpools, which allows organisms to disperse passively downstream. Of the 49 rockpools in the survey, 35 are subject to occasional drying up, while the others appear permanent. Most collections (1989-2004) were annual censuses of invertebrate populations, exceeding a total of 475,000 invertebrates counted, with only minor record gaps. In all cases, species level taxonomic information consists of detailed photographs. In some cases, notes are included with the taxonomic data where species identification could not be matched to information available in the literature. Samples from 2005 to 2019 still require organism identification. Abiotic parameters were measured the day before biotic sampling took place as the process of biotic sampling can impact abiotic parameters through stirring, oxygenation and filtering (temperature, pH, turbidity, dissolved oxygen, light intensity, salinity, alkalinity, and nutrients). The cumulative richness in the metacommunity consist of 78 freshwater, marine, and brackish water taxa, with a mean richness per rockpool of 5.5 distinct species. Regarding taxonomic makeup, ostracods dominated in both diversity and number, followed by copepods, and insects. There are no copyright restrictions on the data set; please cite this data paper when using these data in publications.

Interactions between two functionally distinct aquatic invertebrate herbivores complicate ecosystem- and population-level resilience.

Resilience, the capacity for a system to bounce-back after a perturbation, is critical for conservation and restoration efforts. Different functional traits have differential effects on system-level resilience. We test this experimentally in a lab system consisting of algae consumed by zooplankton, snails, or both, using an eutrophication event as a perturbation. We examined seston settlement load, chlorophyll-a and ammonium concentration as gauges of resilience. We find that Daphnia magna increased our measures of resilience. But this effect is not consistent across ecosystem measures; in fact, D. magna increased the difference between disturbed and undisturbed treatments in seston settlement loads. We have some evidence of shifting reproductive strategy in response to perturbation in D. magna and in the presence of Physa sp. These shifts correspond with altered population levels in D. magna, suggesting feedback loops between the herbivore species. While these results suggest only an ambiguous connection between functional traits to ecosystem resilience, they point to the difficulties in establishing such a link: indirect effects of one species on reproduction of another and different scales of response among components of the system, are just two examples that may compromise the power of simple predictions.

Disentangling local, metapopulation, and cross-community sources of stabilization and asynchrony in metacommunities.

Asynchronous fluctuations of populations are essential for maintaining stable levels of bio-mass and ecosystem function in landscapes. Yet, understanding the stabilization of metacommunities by asynchrony is complicated by the existence of multiple forms of asynchrony that are typically studied independently: Community ecologists, for instance, focus on asynchrony within and among local communities, while population ecologists emphasize asynchrony of populations in metapopulations. Still, other forms of asynchrony, such as that which underlies the spatial insurance effect, are not captured by any existing analytical frameworks. We therefore developed a framework that would in one analysis unmask the stabilizing roles of local communities and metapopulations and so unify these perspectives. Our framework shows that metacommunity stabilization arises from one local and two regional forms of asynchrony: (1) asynchrony among species of a local community, (2) asynchrony among populations of a metapopulation, and (3) cross-community asynchrony, which is between different species in different local communities and underlies spatial insurance. For each type of stabilization, we derived links to diversity indices and associated diversity-stability relationships. We deployed this framework in a set of rock pool invertebrate metacommunities in Discovery Bay, Jamaica, to partition sources of stabilization and test their dependence on diversity. Cross-community asynchrony was the dominant form of stabilization, accounting for >60% of total metacommunity stabilization despite being undetectable with existing frameworks. Environmental variation influenced types of stabilization through different mechanisms. pH and dissolved oxygen, for example, increased asynchrony by decorrelating local species, while salinity did so by changing the abundance structure of metapopulations. Lastly, all types of asynchrony depended strongly on different types of diversity (alpha, metapopulation, and beta diversity drove local, metapopulation, and cross-community asynchrony, respectively) to produce multiple diversity-stability relationships within metacommunities. Our new partition of metacommunity dynamics highlights how different elements-from local communities to metapopulations-combine to stabilize metacommunities and depend critically on contrasting environmental regimes and diversities. Understanding and balancing these sources of stability in dynamic landscapes is a looming challenge for the future. We suggest that synthetic frameworks which merge ecological perspectives will be essential for grasping and safeguarding the stability of natural systems.

Biological invasions, ecological resilience and adaptive governance.

In a world of increasing interconnections in global trade as well as rapid change in climate and land cover, the accelerating introduction and spread of invasive species is a critical concern due to associated negative social and ecological impacts, both real and perceived. Much of the societal response to invasive species to date has been associated with negative economic consequences of invasions. This response has shaped a war-like approach to addressing invasions, one with an agenda of eradications and intense ecological restoration efforts towards prior or more desirable ecological regimes. This trajectory often ignores the concept of ecological resilience and associated approaches of resilience-based governance. We argue that the relationship between ecological resilience and invasive species has been understudied to the detriment of attempts to govern invasions, and that most management actions fail, primarily because they do not incorporate adaptive, learning-based approaches. Invasive species can decrease resilience by reducing the biodiversity that underpins ecological functions and processes, making ecosystems more prone to regime shifts. However, invasions do not always result in a shift to an alternative regime; invasions can also increase resilience by introducing novelty, replacing lost ecological functions or adding redundancy that strengthens already existing structures and processes in an ecosystem. This paper examines the potential impacts of species invasions on the resilience of ecosystems and suggests that resilience-based approaches can inform policy by linking the governance of biological invasions to the negotiation of tradeoffs between ecosystem services.

Spatial variation as a tool for inferring temporal variation and diagnosing types of mechanisms in ecosystems.

Ecological processes, like the rise and fall of populations, leave an imprint of their dynamics as a pattern in space. Mining this spatial record for insight into temporal change underlies many applications, including using spatial snapshots to infer trends in communities, rates of species spread across boundaries, likelihood of chaotic dynamics, and proximity to regime shifts. However, these approaches rely on an inherent but undefined link between spatial and temporal variation. We present a quantitative link between a variable's spatial and temporal variation based on established variance-partitioning techniques, and test it for predictive and diagnostic applications. A strong link existed between spatial and regional temporal variation (estimated as Coefficients of Variation or CV's) in 136 variables from three aquatic ecosystems. This association suggests a basis for substituting one for the other, either quantitatively or qualitatively, when long time series are lacking. We further show that weak substitution of temporal for spatial CV results from distortion by specific spatiotemporal patterns (e.g., inter-patch synchrony). Where spatial and temporal CV's do not match, we pinpoint the spatiotemporal causes of deviation in the dynamics of variables and suggest ways that may control for them. In turn, we demonstrate the use of this framework for describing spatiotemporal patterns in multiple ecosystem variables and attributing them to types of mechanisms. Linking spatial and temporal variability makes quantitative the hitherto inexact practice of space-for-time substitution and may thus point to new opportunities for navigating the complex variation of ecosystems.

Local and regional processes in community assembly.

Controversy on whether local (deterministic) or regional (stochastic) factors control the structure of communities persists after decades of research. The main reason for why it has not been resolved may lie in the nature of evidence which largely comes from realized natural communities. In such communities assembly history leaves a mark that may support either set of factors. To avoid the confounding effects of assembly history we controlled for these effects experimentally. We created a null community by mixing 17 rock pool communities. We then divided the null community into replicates and distributed among treatments representing a gradient of factors from local to regional. We hypothesized that if deterministic factors dominate the assembly of communities, community structures should show a corresponding gradient from being very similar and convergent to dissimilar and divergent. In contrast, if local processes are predominantly stochastic in nature, such a gradient of community configurations should emerge even in the homogeneous setting. Our results appear to partially support both hypotheses and thus suggest that both deterministic and stochastic processes contribute to the assembly of communities. Furthermore, we found that to satisfactorily explain patterns observed in natural communities environmental heterogeneity and regional processes must also be considered. In conclusion, although deterministic mechanisms seem to be important in the assembly of communities, in natural systems their signal may be diluted and masked whenever other factors exert meaningful influence. Such factors increase the number of possible paths to the point that the number of paths equals the number of communities in a metacommunity.

Population synchrony decreases with richness and increases with environmental fluctuations in an experimental metacommunity.

Fluctuations of local but connected populations may show correlation or synchrony whenever they experience significant dispersal or correlated environmental biotic and abiotic variability. Synchrony may be an important variable in multispecies systems, but its nature and implications have not been explicitly examined. Because the number of locally coexisting species (richness) affects the population variability of community members, we manipulated richness under different regimes of environmental fluctuation (EF). We predicted that the temporal synchrony of populations in a species should decline with increasing richness of the metacommunity they live in. Additionally, we predicted that specialist species that are sensitive to a specific environmental factor would show higher synchronization when EF increases. We thus created experimental communities with varied richness, EF, and species specialization to examine the synchronizing effects of these factors on three aquatic invertebrate species. We created four levels of richness and three levels of EF by manipulating the salinity of the culture media. Monocultures exhibited higher population synchrony than metacommunities of 2-4 species. Furthermore, we found that species responded differently to EF treatments: high EF enhanced population synchrony for the specialist and intermediate species, but not for the generalist species. Our findings emphasize that the magnitude of EF and species richness both contribute to determine population synchrony, and importantly, our results suggest that biotic diversity may actually stabilize metacommunities by disrupting synchrony.

Contrasts between habitat generalists and specialists: an empirical extension to the basic metacommunity framework.

Emergence of the metacommunity concept has made a substantial contribution to better understanding of the community composition and dynamics in a regional context. However, long-term field data for testing of available metacommunity models are still scarce, and the extent to which these models apply to the real world remains unknown. Tests conducted so far have largely sought to fit data on the entire regional set of species to one of several metacommunity models, implicitly assuming that all species operate similarly over the same set of sites. However, species differ in their habitat use. These differences can, in the most general terms, be expressed as a gradient of habitat specialization (ranging from habitat specialists to habitat generalists). We postulate that such differences in habitat specialization will have implications for metacommunity dynamics. Specifically, we predict that specialists respond more to local processes and generalists respond to regional spatial processes. We tested these predictions using natural microcosm communities for which long-term (nine-year) environmental and population dynamics data were available. We used redundancy analysis to determine the proportion of variation explained by environmental and spatial factors. We repeated this analysis to explain variation in the entire regional set of species, in generalist species only, and in specialists only. We further used ANOVA to test for differences in the proportions of explained variation. We found that habitat specialists responded primarily to environmental factors and habitat generalists responded mainly to spatial factors. Thus, from the metacommunity perspective, the dynamics of habitat specialists are best explained by a combination of species sorting and mass effects, while that of habitat generalists are best explained by patch dynamics and neutral models. Consequently, we infer that a natural metacommunity can exhibit complicated dynamics, with some groups of species (e.g., habitat specialists) governed according to environmental processes and other groups (e.g., habitat generalists) governed mainly by dispersal processes.

Non-omnivorous generality promotes population stability.

Using a combination of stable isotope analysis of delta13C and delta15N and long-term census data on population abundances for meiofauna in tropical aquatic rock pools, we provide evidence that species which exhibit greater variation in delta13C, an indication of a greater range of distinct carbon sources in their diet, have more stable populations than species with lower variation in delta13C. This link between increased isotope variability and reduced population variability, however, did not hold for delta15N. This suggests that increases in population stability were due to non-omnivorous feeding on multiple carbon sources within a trophic level rather than omnivorous feeding on multiple carbon sources across trophic levels. Our findings corroborate MacArthur's original hypothesis that populations that can access a greater range of resources are more stable than those which consume a more restricted range of resources.

Are natural microcosms useful model systems for ecology?

Several recent, high-impact ecological studies feature natural microcosms as tools for testing effects of fragmentation, metacommunity theory or links between biodiversity and ecosystem processes. These studies combine the microcosm advantages of small size, short generation times, contained structure and hierarchical spatial arrangement with advantages of field studies: natural environmental variance, 'openness' and realistic species combinations with shared evolutionary histories. This enables tests of theory pertaining to spatial and temporal dynamics, for example, the effects of neighboring communities on local diversity, or the effects of biodiversity on ecosystem function. Using examples, we comment on the position of natural microcosms in the roster of ecological research strategies and tools. We conclude that natural microcosms are as versatile as artificial microcosms, but as complex and biologically realistic as other natural systems. Research to date combined with inherent attributes of natural microcosms make them strong candidate model systems for ecology.

Removing the confounding effect of habitat specialization reveals the stabilizing contribution of diversity to species variability.

Earlier studies have found that diversity, S, stabilizes the relative variability of combined biomass or abundance of species making up a community. However, the effect of S on variability of constituent species has been elusive. We hypothesize that the proportion of specialists increases with S and, because specialists are more variable, this shift in composition will mask the stabilizing effect of S on populations of species making up a community. The test uses data on variability and ecological specialization of species in 49 natural rock pool invertebrate communities. Initial analyses produced inconclusive results similar to earlier studies. However, when variability owing to species' specialization was factored out, S reduced species' abundance variability, although not in all communities. Our study explains why the stabilizing effect of diversity on populations has not been found earlier.

The relative sensitivity of four benthic invertebrates to metals in spiked-sediment exposures and application to contaminated field sediment.

The relative sensitivity of four benthic invertebrates (Hyalella azteca, Chironomus riparius, Hexagenia spp., and Tubifex tubifex) was determined for Cd, Cu, and Ni in water-only and in spiked-sediment exposures. Survival (median lethal concentrations [LC50s] and the concentrations estimated to be lethal to 25% of test organisms [LC25s]), and endpoints for growth and reproduction (mean inhibitory concentrations [IC25s]) were compared. The sensitivities differed depending on the species and metal, although some trends emerged. In water-only exposures, H. azteca is the most sensitive species to cadmium and nickel, with mean LC50s of 0.013 and 3.6 mg/L, respectively; C. riparius is the most sensitive species to copper, with a mean LC50 of 0.043 mg/L. In the spiked-sediment exposures, the order in decreasing sensitivity to copper is Hyalella = Hexagenia < Chironomus < Tubifex for survival and growth/reproduction. For cadmium, the order in decreasing sensitivity is Hyalella = Chironomus < Hexagenia < Tubifex, and for nickel is Hyalella << Hexagenia < Chironomus < Tubifex. Chironomus riparius and Hexagenia spp. survival can be used to distinguish between toxicity caused by different metals. Species test responses in field-collected sediment(Collingwood Harbour, ON, Canada) were examined in an attempt to determine the causative agent of toxicity throughout, using the established species sensitivities. Sediment toxicity was categorized first by comparing species responses to those established for a reference database. Test responses in the field-collected sediment do not support causality by Cu, a suspected toxicant based on comparison of sediment chemistry with sediment quality guidelines.