Species distribution models (SDMs) are a common approach to describing species’ space‐use and spatially‐explicit abundance. With a myriad of model types, methods and parameterization options available, it is challenging to make informed decisions about how to build robust SDMs appropriate for a given purpose. One key component of SDM development is the appropriate parameterization of covariates, such as the inclusion of covariates that reflect underlying processes (e.g. abiotic and biotic covariates) and covariates that act as proxies for unobserved processes (e.g. space and time covariates). It is unclear how different SDMs apportion variance among a suite of covariates, and how parameterization decisions influence model accuracy and performance. To examine trade‐offs in covariation parameterization in SDMs, we explore the attribution of spatiotemporal and environmental variation across a suite of SDMs. We first used simulated species distributions with known environmental preferences to compare three types of SDM: a machine learning model (boosted regression tree), a semi‐parametric model (generalized additive model) and a spatiotemporal mixed‐effects model (vector autoregressive spatiotemporal model, VAST). We then applied the same comparative framework to a case study with three fish species (arrowtooth flounder, pacific cod and walleye pollock) in the eastern Bering Sea, USA. Model type and covariate parameterization both had significant effects on model accuracy and performance. We found that including either spatiotemporal or environmental covariates typically reproduced patterns of species distribution and abundance across the three models tested, but model accuracy and performance was maximized when including both spatiotemporal and environmental covariates in the same model framework. Our results reveal trade‐offs in the current generation of SDM tools between accurately estimating species abundance, accurately estimating spatial patterns, and accurately quantifying underlying species–environment relationships. These comparisons between model types and parameterization options can help SDM users better understand sources of model bias and estimate error.
Background
Studies that attempt to measure shifts in species distributions often consider a single species in isolation. However, understanding changes in spatial overlap between predators and their prey might provide deeper insight into how species redistribution affects food web dynamics.
Predator–prey overlap metrics
Here, we review a suite of 10 metrics [range overlap, area overlap, the local index of collocation (Pianka's O), Hurlbert's index, biomass‐weighted overlap, asymmetrical alpha, Schoener's D, Bhattacharyya's coefficient, the global index of collocation and the AB ratio] that describe how two species overlap in space, using concepts such as binary co‐occurrence, encounter rates, spatial niche similarity, spatial independence, geographical similarity and trophic transfer. We describe the specific ecological insights that can be gained using each overlap metric, in order to determine which is most appropriate for describing spatial predator–prey interactions for different applications.
Simulation and case study
We use simulated predator and prey distributions to demonstrate how the 10 metrics respond to variation in three types of predator–prey interactions: changing spatial overlap between predator and prey, changing predator population size and changing patterns of predator aggregation in response to prey density. We also apply these overlap metrics to a case study of a predatory fish (arrowtooth flounder, Atheresthes stomias) and its prey (juvenile walleye pollock, Gadus chalcogrammus) in the Eastern Bering Sea, AK, USA. We show how the metrics can be applied to understand spatial and temporal variation in the overlap of species distributions in this rapidly changing Arctic ecosystem.
Conclusions
Using both simulated and empirical data, we provide a roadmap for ecologists and other practitioners to select overlap metrics to describe particular aspects of spatial predator–prey interactions. We outline a range of research and management applications for which each metric may be suited.
Research has estimated associations between water temperature and the spatial distribution of marine fishes based upon correlations between temperature and the centroid of fish distribution (centre of gravity, COG). Analysts have then projected future water temperatures to forecast shifts in COG, but often neglected to demonstrate that temperature explains a substantial portion of historical distribution shifts. We argue that estimating the proportion of observed distributional shifts that can be attributed to temperature vs. other factors is a critical first step in forecasting future changes. We illustrate this approach using Gadus chalcogrammus (Walleye pollock) in the Eastern Bering Sea, and use a vector‐autoregressive spatiotemporal model to attribute variation in COG from 1982 to 2015 to three factors: local or regional changes in surface and bottom temperature (“temperature effects”), fluctuations in size‐structure that cause COG to be skewed towards juvenile or adult habitats (“size‐structured effects”) or otherwise unexplained spatiotemporal variation in distribution (“unexplained effects”). We find that the majority of variation in COG (including the north‐west trend since 1982) is largely unexplained by temperature or size‐structured effects. Temperature alone generates a small portion of primarily north–south variation in COG, while size‐structured effects generate a small portion of east–west variation. We therefore conclude that projections of future distribution based on temperature alone are likely to miss a substantial portion of both the interannual variation and interdecadal trends in COG for this species. More generally, we suggest that decomposing variation in COG into multiple causal factors is a vital first step for projecting likely impacts of temperature change.
Indices of abundance are important for estimating population trends in stock assessment and ideally should be based on fishery-independent surveys to avoid problems associated with the hyperstability of the commercial catch per unit effort (cpue) data. However, recent studies indicate that the efficiency of the survey bottom trawl (BT) for some species can be density-dependent, which could affect the reliability of survey-derived indices of abundance. A function qe∼f(u), where qe is the BT efficiency and u the catch rate, was derived using experimentally derived acoustic dead-zone correction and BT efficiency parameters obtained from combining a subset of BT catch data with synchronously collected acoustic data from walleye pollock (Theragra chalcogramma) in the eastern Bering Sea (EBS). We found that qe decreased with increasing BT catches resulting in hyperstability of the index of abundance derived from BT survey. Density-dependent qe resulted in spatially and temporarily variable bias in survey cpue and biased population age structure derived from survey data. We used the relationship qe∼f(u) to correct the EBS trawl survey index of abundance for density-dependence. We also obtained a variance–covariance matrix for a new index that accounted for sampling variability and the uncertainty associated with the qe. We found that incorporating estimates of the new index of abundance changed outputs from the walleye pollock stock assessment model. Although changes were minor, we advocate incorporating estimates of density-dependent qe into the walleye pollock stock assessment as a precautionary measure that should be undertaken to avoid negative consequences of the density-dependent qe.
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