Abstract

ABSTRACTThe legacy of the ‘SL > SS principle’, that a single or a few large habitat patches (SL) conserve more species than several small patches (SS), is evident in decisions to protect large patches while down‐weighting small ones. However, empirical support for this principle is lacking, and most studies find either no difference or the opposite pattern (SS > SL). To resolve this dilemma, we propose a research agenda by asking, ‘are there consistent, empirically demonstrated conditions leading to SL > SS?’ We first review and summarize ‘single large or several small’ (SLOSS) theory and predictions. We found that most predictions of SL > SS assume that between‐patch variation in extinction rate dominates the outcome of the extinction–colonization dynamic. This is predicted to occur when populations in separate patches are largely independent of each other due to low between‐patch movements, and when species differ in minimum patch size requirements, leading to strong nestedness in species composition along the patch size gradient. However, even when between‐patch variation in extinction rate dominates the outcome of the extinction–colonization dynamic, theory can predict SS > SL. This occurs if extinctions are caused by antagonistic species interactions or disturbances, leading to spreading‐of‐risk of landscape‐scale extinction across SS. SS > SL is also predicted when variation in colonization dominates the outcome of the extinction–colonization dynamic, due to higher immigration rates for SS than SL, and larger species pools in proximity to SS than SL. Theory that considers change in species composition among patches also predicts SS > SL because of higher beta diversity across SS than SL. This results mainly from greater environmental heterogeneity in SS due to greater variation in micro‐habitats within and across SS habitat patches (‘across‐habitat heterogeneity’), and/or more heterogeneous successional trajectories across SS than SL. Based on our review of the relevant theory, we develop the ‘SLOSS cube hypothesis’, where the combination of three variables – between‐patch movement, the role of spreading‐of‐risk in landscape‐scale population persistence, and across‐habitat heterogeneity – predict the SLOSS outcome. We use the SLOSS cube hypothesis and existing SLOSS empirical evidence, to predict SL > SS only when all of the following are true: low between‐patch movement, low importance of spreading‐of‐risk for landscape‐scale population persistence, and low across‐habitat heterogeneity. Testing this prediction will be challenging, as it will require many studies of species groups and regions where these conditions hold. Each such study would compare gamma diversity across multiple landscapes varying in number and sizes of patches. If the prediction is not generally supported across such tests, then the mechanisms leading to SL > SS are extremely rare in nature and the SL > SS principle should be abandoned.

Highlights

  • The legacy of the ‘SL > SS principle’, that a single or a few large habitat patches (SL) conserve more species than several small patches (SS), is evident in decisions to protect large patches while down-weighting small ones

  • Most predictions of SL > SS derive from the assumption that variation in extinction rate dominates the outcome of the extinction–colonization dynamic

  • At the same time the idea persists that the SL > SS principle is generally valid under some conditions (Table 1), such as for groups of habitat-interior species and in landscapes with low matrix quality (Pfeifer et al, 2017; Fletcher et al, 2018), and that these are the conditions where biodiversity is most threatened

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Summary

Predictions based on extinction–colonization dynamics

(2) Species have minimum patch size requirements.a (3) Negative edge effects accentuate both of the previous mechanisms because patch edge-to-area ratio decreases with patch size This disproportionately reduces patch size and increases demographic stochasticity for small patches compared to large patches.b (4) Higher per-unit-area emigration rate from small than large patches, due to Extinction probability over the SS > SL landscape is lower for SS than SL. Higher edge-to-area ratio, leads to higher dispersal mortality in the matrix over SS than over SL.c (1) Between-patch movements of a competitor/predator/parasitoid are lower than their within-patch movements, and lower than between-patch movements of the affected species. Our aim is to encourage future research in a direction that will resolve the SLOSS dilemma

REVIEW OF SLOSS PREDICTIONS
RESEARCH AGENDA
DISCUSSION
CONCLUSIONS
Findings
VIII. Supporting information
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