Ecosystem decay exacerbates biodiversity loss with habitat loss.

Although habitat loss is the predominant factor leading to biodiversity loss in the Anthropocene1,2, exactly how this loss manifests-and at which scales-remains a central debate3-6. The 'passive sampling' hypothesis suggests that species are lost in proportion to their abundance and distribution in the natural habitat7,8, whereas the 'ecosystem decay' hypothesis suggests that ecological processes change in smaller and more-isolated habitats such that more species are lost than would have been expected simply through loss of habitat alone9,10. Generalizable tests of these hypotheses have been limited by heterogeneous sampling designs and a narrow focus on estimates of species richness that are strongly dependent on scale. Here we analyse 123 studies of assemblage-level abundances of focal taxa taken from multiple habitat fragments of varying size to evaluate the influence of passive sampling and ecosystem decay on biodiversity loss. We found overall support for the ecosystem decay hypothesis. Across all studies, ecosystems and taxa, biodiversity estimates from smaller habitat fragments-when controlled for sampling effort-contain fewer individuals, fewer species and less-even communities than expected from a sample of larger fragments. However, the diversity loss due to ecosystem decay in some studies (for example, those in which habitat loss took place more than 100 years ago) was less than expected from the overall pattern, as a result of compositional turnover by species that were not originally present in the intact habitats. We conclude that the incorporation of non-passive effects of habitat loss on biodiversity change will improve biodiversity scenarios under future land use, and planning for habitat protection and restoration.

Linking changes in species composition and biomass in a globally distributed grassland experiment.

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.

Long-term changes in temperate marine fish assemblages are driven by a small subset of species.

The species composition of plant and animal assemblages across the globe has changed substantially over the past century. How do the dynamics of individual species cause this change? We classified species into seven unique categories of temporal dynamics based on the ordered sequence of presences and absences that each species contributes to an assemblage time series. We applied this framework to 14,434 species trajectories comprising 280 assemblages of temperate marine fishes surveyed annually for 20 or more years. Although 90% of the assemblages diverged in species composition from the baseline year, this compositional change was largely driven by only 8% of the species' trajectories. Quantifying the reorganization of assemblages based on species shared temporal dynamics should facilitate the task of monitoring and restoring biodiversity. We suggest ways in which our framework could provide informative measures of compositional change, as well as leverage future research on pattern and process in ecological systems.

Effects of site-selection bias on estimates of biodiversity change.

Estimates of biodiversity change are essential for the management and conservation of ecosystems. Accurate estimates rely on selecting representative sites, but monitoring often focuses on sites of special interest. How such site-selection biases influence estimates of biodiversity change is largely unknown. Site-selection bias potentially occurs across four major sources of biodiversity data, decreasing in likelihood from citizen science, museums, national park monitoring, and academic research. We defined site-selection bias as a preference for sites that are either densely populated (i.e., abundance bias) or species rich (i.e., richness bias). We simulated biodiversity change in a virtual landscape and tracked the observed biodiversity at a sampled site. The site was selected either randomly or with a site-selection bias. We used a simple spatially resolved, individual-based model to predict the movement or dispersal of individuals in and out of the chosen sampling site. Site-selection bias exaggerated estimates of biodiversity loss in sites selected with a bias by on average 300-400% compared with randomly selected sites. Based on our simulations, site-selection bias resulted in positive trends being estimated as negative trends: richness increase was estimated as 0.1 in randomly selected sites, whereas sites selected with a bias showed a richness change of -0.1 to -0.2 on average. Thus, site-selection bias may falsely indicate decreases in biodiversity. We varied sampling design and characteristics of the species and found that site-selection biases were strongest in short time series, for small grains, organisms with low dispersal ability, large regional species pools, and strong spatial aggregation. Based on these findings, to minimize site-selection bias, we recommend use of systematic site-selection schemes; maximizing sampling area; calculating biodiversity measures cumulatively across plots; and use of biodiversity measures that are less sensitive to rare species, such as the effective number of species. Awareness of the potential impact of site-selection bias is needed for biodiversity monitoring, the design of new studies on biodiversity change, and the interpretation of existing data.

Efectos del Sesgo en la Selección de Sitio sobre las Estimaciones del Cambio en la Biodiversidad Resumen Las estimaciones del cambio en la biodiversidad son esenciales para el manejo y la conservación de los ecosistemas. Las estimaciones precisas dependen de la selección de sitios representativos pero su monitoreo con frecuencia se enfoca en los sitios de interés especial. En su mayoría se desconoce cómo influyen tales sesgos en la selección de sitios sobre las estimaciones del cambio en la biodiversidad. El sesgo en la selección de sitios ocurre potencialmente en cuatro fuentes principales de datos sobre biodiversidad, disminuyendo en probabilidad cuando los datos vienen de la ciencia ciudadana, museos, el monitoreo de los parques nacionales y la investigación académica. Definimos al sesgo en la selección de sitios como la preferencia por sitios que están densamente poblados (es decir, sesgo por abundancia) o que son ricos en especies (es decir, sesgo por riqueza). Simulamos el cambio en la biodiversidad en un paisaje virtual y le dimos seguimiento a la biodiversidad observada en un sitio muestreado. El sitio fue seleccionado al azar o con un sesgo en la selección de sitio. Usamos un modelo simple basado en los individuos y resuelto espacialmente para predecir el movimiento o la dispersión de los individuos dentro y fuera del sitio de muestreo elegido. El sesgo en la selección de sitio exageró las estimaciones de la pérdida de la biodiversidad en los sitios seleccionados con un sesgo en promedio de 300-400% en comparación con sitios seleccionados al azar. Con base en nuestras simulaciones, el sesgo en la selección de sitio derivó en que las tendencias positivas se estimaran como tendencias negativas: se estimó que el incremento en la riqueza fue de 0.1 en sitios seleccionados al azar, mientras que en los sitios seleccionados con un sesgo mostraron un cambio en la riqueza de −0.1 a −0.2 en promedio. Así, el sesgo en la selección de sitio puede indicar erróneamente la existencia de disminuciones en la biodiversidad. Variamos el diseño del muestreo y las características de las especies y encontramos que los sesgos en la selección de sitio estaban más consolidados en las series de tiempo corto, para los granos pequeños, organismos con una baja habilidad de dispersión, grandes patrimonios genéticos de especies regionales y una agregación espacial fuerte. Con base en estos resultados, para lograr minimizar el sesgo en la selección de sitio, recomendamos usar esquemas sistemáticos de selección de sitio; maximizar el área de muestreo; calcular las medidas de biodiversidad acumulativamente en los lotes; y usar las medidas de biodiversidad que son menos sensibles a las especies raras, como el número efectivo de especies. Se necesita tener conciencia sobre el impacto potencial del sesgo en la selección de sitio para el monitoreo de la biodiversidad, el diseño de nuevos estudios sobre el cambio en la biodiversidad y la interpretación de los datos existentes.

Reducing dispersal limitation via seed addition increases species richness but not above-ground biomass.

Seed dispersal limitation, which can be exacerbated by a number of anthropogenic causes, can result in local communities having fewer species than they might potentially support, representing a potential diversity deficit. The link between processes that shape natural variation in diversity, such as dispersal limitation, and the consequent effects on productivity is less well known. Here, we synthesised data from 12 seed addition experiments in grassland communities to examine the influence of reducing seed dispersal limitation (from 1 to 60 species added across experiments) on species richness and productivity. For every 10 species of seed added, we found that species richness increased by about two species. However, the increase in species richness by overcoming seed limitation did not lead to a concomitant increase in above-ground biomass production. This highlights the need to consider the relationship between biodiversity and ecosystem functioning in a pluralistic way that considers both the processes that shape diversity and productivity simultaneously in naturally assembled communities.

Embracing scale-dependence to achieve a deeper understanding of biodiversity and its change across communities.

Because biodiversity is multidimensional and scale-dependent, it is challenging to estimate its change. However, it is unclear (1) how much scale-dependence matters for empirical studies, and (2) if it does matter, how exactly we should quantify biodiversity change. To address the first question, we analysed studies with comparisons among multiple assemblages, and found that rarefaction curves frequently crossed, implying reversals in the ranking of species richness across spatial scales. Moreover, the most frequently measured aspect of diversity - species richness - was poorly correlated with other measures of diversity. Second, we collated studies that included spatial scale in their estimates of biodiversity change in response to ecological drivers and found frequent and strong scale-dependence, including nearly 10% of studies which showed that biodiversity changes switched directions across scales. Having established the complexity of empirical biodiversity comparisons, we describe a synthesis of methods based on rarefaction curves that allow more explicit analyses of spatial and sampling effects on biodiversity comparisons. We use a case study of nutrient additions in experimental ponds to illustrate how this multi-dimensional and multi-scale perspective informs the responses of biodiversity to ecological drivers.

BioTIME: A database of biodiversity time series for the Anthropocene.

MOTIVATION: The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. MAIN TYPES OF VARIABLES INCLUDED: The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. SPATIAL LOCATION AND GRAIN: BioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2). TIME PERIOD AND GRAIN: BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. MAJOR TAXA AND LEVEL OF MEASUREMENT: BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. SOFTWARE FORMAT: .csv and .SQL.

Global reef fish richness gradients emerge from divergent and scale-dependent component changes.

Biodiversity varies from place to place due to environmental and historical factors. To improve our understanding of how history and the environment influence observed patterns, we need to address the limitations of the most commonly used biodiversity metric, species richness. Here, we show that scale-dependent dissections of species richness into components of total abundance, species relative abundances and spatial aggregations of species reveal that two well-known biogeographic reef fish species richness gradients emerge from very different underlying component patterns. Latitudinal richness is underpinned by scale-independent patterns of total and relative abundances, suggesting ecological constraints scale up to determine abundances within communities. In contrast, the longitudinal gradient of species richness typically attributed to historical biogeography only emerges at the largest scale and is accompanied by a similar pattern of relative abundances, suggesting that site-to-site compositional variation leading to species aggregation (i.e. a component of ß-diversity) underlies this gradient. Examining relationships among the components that underpin biodiversity gradients reveals new patterns that can better identify processes influencing patterns of biodiversity.

Remarkable size-spectra stability in a marine system undergoing massive invasion.

The Mediterranean Sea is an invasion hotspot, with non-indigenous species suspected to be a major driver behind community changes. We used size spectra, a reliable index of food web structure, to examine how the influx of Red Sea fishes into the Mediterranean Sea has impacted the indigenous species community. This is the first attempt to use changes in the size spectra to reveal the effect of biological invasions. We used data from trawl catches along Israel's shoreline spanning 20 years to estimate changes in the community size spectra of both indigenous and non-indigenous species. We found that the relative biomass of non-indigenous species increased over the 20 years, especially for small and large species, leading to a convergence with the indigenous species size spectra. Hence, the biomass of indigenous and non-indigenous species has become identical for all size classes, suggesting similar energetic constraints and sensitivities to fishing. However, over this time period the size spectrum of indigenous species has remained remarkably constant. This suggests that the wide-scale invasion of non-indigenous species into the Mediterranean may have had little impact on the community structure of indigenous species.

Aggression, interference, and the functional response of coral-feeding butterflyfishes.

Functional responses describing how foraging rates change with respect to resource density are central to our understanding of interspecific interactions. Competitive interactions are an important determinant of foraging rates; however, the relationship between the exploitation and interference components of competition has received little empirical or theoretical consideration. Moreover, little is known about the relationship between aggressive behavioural interactions and interference competition. Using a natural gradient of consumer and resource densities, we empirically examine how aggressiveness relates to consumer-consumer encounter rates and foraging for four species of Chaetodon reef fish spanning a range of dietary niche breadths. The probability of aggression was most strongly associated with both total consumer and resource densities. In contrast, total encounter rates were best predicted by conspecific consumer density, and were highest for the most specialised consumer (Chaetodon trifascialis), not the most aggressive (Chaetodon baronessa). The most specialised consumer, not the most aggressive, also exhibited the largest reduction in foraging rates with increasing consumer density. Our results support the idea of a positive link between the exploitation and interference components of competition for the most specialised consumer. Moreover, our results caution against inferring the presence of ecological interactions (competition) from observations of behaviour (aggression and agonism) alone.

Experimental evaluation of diversity-productivity relationships in a coral reef fish assemblage.

The global decline in biodiversity is causing increasing concern about the effects of biodiversity loss on ecosystem services such as productivity. Biodiversity has been hypothesised to be important in maintaining productivity of biological assemblages because niche complementarity and facilitation among the constituent species can result in more efficient use of resources. However, these conclusions are primarily based on studies with plant communities, and the relationship between diversity and productivity at higher trophic levels is largely unknown, especially in the marine environment. Here, we used a manipulative field experiment to test the effects of species richness and species identity on biomass accumulation in coral reef fish assemblages at Lizard Island. Small patch reefs were stocked with a total of 30 juveniles belonging to three planktivorous damselfish (genus Pomacentrus) according to three different levels of fish species richness (one, two and three species) and seven different combinations of fish species. Species richness had no effect on the relative growth in this assemblage after 18 days, but relative growth differed among individual fish species and the different combinations of species. Patterns of increase in biomass were best explained by species-specific differences and variable effects of intra- and interspecific competition on growth. These results suggest that niche complementarity and facilitation are not the most influential drivers of total productivity within this guild of planktivorous fishes. Total productivity may be resilient to declining reef fish biodiversity, but this will depend on which species are lost and on the life-history traits of remaining species.

Synthesis of recovery patterns in microbial communities across environments.

BACKGROUND: Disturbances alter the diversity and composition of microbial communities. Yet a generalized empirical assessment of microbiome responses to disturbance across different environments is needed to understand the factors driving microbiome recovery, and the role of the environment in driving these patterns. RESULTS: To this end, we combined null models with Bayesian generalized linear models to examine 86 time series of disturbed mammalian, aquatic, and soil microbiomes up to 50 days following disturbance. Overall, disturbances had the strongest effect on mammalian microbiomes, which lost taxa and later recovered their richness, but not their composition. In contrast, following disturbance, aquatic microbiomes tended away from their pre-disturbance composition over time. Surprisingly, across all environments, we found no evidence of increased compositional dispersion (i.e., variance) following disturbance, in contrast to the expectations of the Anna Karenina Principle. CONCLUSIONS: This is the first study to systematically compare secondary successional dynamics across disturbed microbiomes, using a consistent temporal scale and modeling approach. Our findings show that the recovery of microbiomes is environment-specific, and helps to reconcile existing, environment-specific research into a unified perspective. Video Abstract.

Synthesis reveals approximately balanced biotic differentiation and homogenization.

It is commonly thought that the biodiversity crisis includes widespread declines in the spatial variation of species composition, called biotic homogenization. Using a typology relating homogenization and differentiation to local and regional diversity changes, we synthesize patterns across 461 metacommunities surveyed for 10 to 91 years, and 64 species checklists (13 to 500+ years). Across all datasets, we found that no change was the most common outcome, but with many instances of homogenization and differentiation. A weak homogenizing trend of a 0.3% increase in species shared among communities/year on average was driven by increased numbers of widespread (high occupancy) species and strongly associated with checklist data that have longer durations and large spatial scales. At smaller spatial and temporal scales, we show that homogenization and differentiation can be driven by changes in the number and spatial distributions of both rare and common species. The multiscale perspective introduced here can help identify scale-dependent drivers underpinning biotic differentiation and homogenization.

Heterospecific aggression and dominance in a guild of coral-feeding fishes: the roles of dietary ecology and phylogeny.

Interspecific competition mediates biodiversity maintenance and is an important selective pressure for evolution. Competition is often conceptualized as being exploitative (indirect) or involving direct interference. However, most empirical studies are phenomenological, focusing on quantifying effects of density manipulations, and most competition theory has characterized exploitation competition systems. The effects on resource use of traits associated with direct, interference competition has received far less attention. Here we examine the relationships of dietary ecology and phylogeny to heterospecific aggression in a guild of corallivorous reef fishes. We find that, among chaetodontids (butterflyfishes), heterospecific aggression depends on a synergistic interaction of dietary overlap and specialization: aggression increases with dietary overlap for interactions between specialists but not for interactions involving generalists. Moreover, behavioral dominance is a monotonically increasing function of dietary specialization. The strong, positive relationship of dominance to specialization suggests that heterospecific aggression may contribute to the maintenance of biodiversity where it promotes resource partitioning. Additionally, we find strong phylogenetic signals in dietary overlap and specialization but not behavioral dominance. Our results support the use of phylogeny as a proxy for ecological similarity among butterflyfishes, but we find that direct measures of dietary overlap and specialization predict heterospecific agression much better than phylogeny.

Understanding 'it depends' in ecology: a guide to hypothesising, visualising and interpreting statistical interactions.

Ecologists routinely use statistical models to detect and explain interactions among ecological drivers, with a goal to evaluate whether an effect of interest changes in sign or magnitude in different contexts. Two fundamental properties of interactions are often overlooked during the process of hypothesising, visualising and interpreting interactions between drivers: the measurement scale - whether a response is analysed on an additive or multiplicative scale, such as a ratio or logarithmic scale; and the symmetry - whether dependencies are considered in both directions. Overlooking these properties can lead to one or more of three inferential errors: misinterpretation of (i) the detection and magnitude (Type-D error), and (ii) the sign of effect modification (Type-S error); and (iii) misidentification of the underlying processes (Type-A error). We illustrate each of these errors with a broad range of ecological questions applied to empirical and simulated data sets. We demonstrate how meta-analysis, a widely used approach that seeks explicitly to characterise context dependence, is especially prone to all three errors. Based on these insights, we propose guidelines to improve hypothesis generation, testing, visualisation and interpretation of interactions in ecology.

Regional occupancy increases for widespread species but decreases for narrowly distributed species in metacommunity time series.

While human activities are known to elicit rapid turnover in species composition through time, the properties of the species that increase or decrease their spatial occupancy underlying this turnover are less clear. Here, we used an extensive dataset of 238 metacommunity time series of multiple taxa spread across the globe to evaluate whether species that are more widespread (large-ranged species) differed in how they changed their site occupancy over the 10-90 years the metacommunities were monitored relative to species that are more narrowly distributed (small-ranged species). We found that on average, large-ranged species tended to increase in occupancy through time, whereas small-ranged species tended to decrease. These relationships were stronger in marine than in terrestrial and freshwater realms. However, in terrestrial regions, the directional changes in occupancy were less extreme in protected areas. Our findings provide evidence for systematic decreases in occupancy of small-ranged species, and that habitat protection could mitigate these losses in the face of environmental change.

Looking back on biodiversity change: lessons for the road ahead.

Estimating biodiversity change across the planet in the context of widespread human modification is a critical challenge. Here, we review how biodiversity has changed in recent decades across scales and taxonomic groups, focusing on four diversity metrics: species richness, temporal turnover, spatial beta-diversity and abundance. At local scales, change across all metrics includes many examples of both increases and declines and tends to be centred around zero, but with higher prevalence of declining trends in beta-diversity (increasing similarity in composition across space or biotic homogenization) and abundance. The exception to this pattern is temporal turnover, with changes in species composition through time observed in most local assemblages. Less is known about change at regional scales, although several studies suggest that increases in richness are more prevalent than declines. Change at the global scale is the hardest to estimate accurately, but most studies suggest extinction rates are probably outpacing speciation rates, although both are elevated. Recognizing this variability is essential to accurately portray how biodiversity change is unfolding, and highlights how much remains unknown about the magnitude and direction of multiple biodiversity metrics at different scales. Reducing these blind spots is essential to allow appropriate management actions to be deployed. This article is part of the theme issue 'Detecting and attributing the causes of biodiversity change: needs, gaps and solutions'.

Six years of demography data for 11 reef coral species.

Scleractinian corals are colonial animals with a range of life-history strategies, making up diverse species assemblages that define coral reefs. We tagged and tracked ~30 colonies from each of 11 species during seven trips spanning 6 years (2009-2015) to measure their vital rates and competitive interactions on the reef crest at Trimodal Reef, Lizard Island, Australia. Pairs of species were chosen from five growth forms in which one species of the pair was locally rare (R) and the other common (C). The sampled growth forms were massive (Goniastrea pectinata [R] and G. retiformis [C]), digitate (Acropora humilis [R] and A. cf. digitifera [C]), corymbose (A. millepora [R] and A. nasuta [C]), tabular (A. cytherea [R] and A. hyacinthus [C]) and arborescent (A. robusta [R] and A. intermedia [C]). An extra corymbose species with intermediate abundance, A. spathulata was included when it became apparent that A. millepora was too rare on the reef crest, making the 11 species in total. The tagged colonies were visited each year in the weeks prior to spawning. During visits, two or more observers each took two or three photographs of each tagged colony from directly above and on the horizontal plane with a scale plate to track planar area. Dead or missing colonies were recorded and new colonies tagged to maintain ~30 colonies per species throughout the 6 years of the study. In addition to tracking tagged corals, 30 fragments were collected from neighboring untagged colonies of each species for counting numbers of eggs per polyp (fecundity); and fragments of untagged colonies were brought into the laboratory where spawned eggs were collected for biomass and energy measurements. We also conducted surveys at the study site to generate size structure data for each species in several of the years. Each tagged colony photograph was digitized by at least two people. Therefore, we could examine sources of error in planar area for both photographers and outliners. Competitive interactions were recorded for a subset of species by measuring the margins of tagged colony outlines interacting with neighboring corals. The study was abruptly ended by Tropical Cyclone Nathan (Category 4) that killed all but nine of the more than 300 tagged colonies in early 2015. Nonetheless, these data will be of use to other researchers interested in coral demography and coexistence, functional ecology, and parametrizing population, community, and ecosystem models. The data set is not copyright restricted, and users should cite this paper when using the data.

Widespread shifts in body size within populations and assemblages.

Biotic responses to global change include directional shifts in organismal traits. Body size, an integrative trait that determines demographic rates and ecosystem functions, is thought to be shrinking in the Anthropocene. Here, we assessed the prevalence of body size change in six taxon groups across 5025 assemblage time series spanning 1960 to 2020. Using the Price equation to partition this change into within-species body size versus compositional changes, we detected prevailing decreases in body size through time driven primarily by fish, with more variable patterns in other taxa. We found that change in assemblage composition contributes more to body size changes than within-species trends, but both components show substantial variation in magnitude and direction. The biomass of assemblages remains quite stable as decreases in body size trade off with increases in abundance.

Risk spreading, connectivity, and optimal reserve spacing.

Two important processes determining the dynamics of spatially structured populations are dispersal and the spatial covariance of demographic fluctuations. Spatially explicit approaches to conservation, such as reserve networks, must consider the tension between these two processes and reach a balance between distances near enough to maintain connectivity, but far enough to benefit from risk spreading. Here, we model this trade-off. We show how two measures of metapopulation persistence depend on the shape of the dispersal kernel and the shape of the distance decay in demographic covariance, and we consider the implications of this trade-off for reserve spacing. The relative rates of distance decay in dispersal and demographic covariance determine whether the long-run metapopulation growth rate, and quasi-extinction risk, peak for adjacent patches or intermediately spaced patches; two local maxima in metapopulation persistence are also possible. When dispersal itself fluctuates over time, the trade-off changes. Temporal variation in mean distance that propagules are dispersed (i.e., propagule advection) decreases metapopulation persistence and decreases the likelihood that persistence will peak for adjacent patches. Conversely, variation in diffusion (the extent of random spread around mean dispersal) increases metapopulation persistence overall and causes it to peak at shorter inter-patch distances. Thus, failure to consider temporal variation in dispersal processes increases the risk that reserve spacings will fail to meet the objective of ensuring metapopulation persistence. This study identifies two phenomena that receive relatively little attention in empirical work on reserve spacing, but that can qualitatively change the effectiveness of reserve spacing strategies: (1) the functional form of the distance decay in covariance among patch-specific demographic rates and (2) temporal variation in the shape of the dispersal kernel. The sensitivity of metapopulation recovery and persistence to how covariance of vital rates decreases with distance suggests that estimating the shape of this function is likely to be as important for effective reserve design as estimating connectivity. Similarly, because temporal variation in dispersal dynamics influences the effect of reserve spacing, approaches to reserve design that ignore such variation, and rely instead on long-term average dispersal patterns, are likely to lead to lower metapopulation viability than is actually achievable.