Refuges and ecological traps: Extreme drought threatens persistence of an endangered fish in intermittent streams.

Recent droughts raise global concern over potential biodiversity loss and mitigating impacts to vulnerable species has become a management priority. However, drought impacts on populations are difficult to predict, in part, because habitat refuges can buffer organisms from harsh environmental conditions. In a global change context, more extreme droughts may turn previously suitable habitats into ecological traps, where vulnerable species can no longer persist. Here, we explore the impacts of California's recent record-breaking drought on endangered juvenile Coho salmon. We estimated the variability of cumulative salmon survival using mark-recapture of nearly 20,000 tagged fish in intermittent stream pools during a 7-year period encompassing drought and non-drought conditions. We then determined the relative importance of physical habitat, streamflow, precipitation, landscape, and biological characteristics that may limit survival during drought. Our most striking result was an increase in the number of pools with reduced or zero survival during drought years and a coincident increase in spatial variability in survival among study reaches. In nearly half of the stream pools, salmon survival during drought was similar to mean survival of pools assessed during non-drought years, indicating some pools had remarkable resistance (ability to withstand disturbance) to extreme drought. Lower survival was most attributable to longer duration of disconnection between upstream and downstream habitats, a consequence of increasing drought severity. Our results not only suggest that many pools sustain juvenile salmon in non-drought years transition into ecological traps during drought but also highlight that some pools serve as refuges even under extreme drought conditions. Projected increases in drought severity that lead to longer droughts and greater habitat fragmentation could transform an increasing proportion of suitable habitats into ecological traps. Predicting future impacts of drought on Coho salmon and other sensitive species will require identification and protection of drought refuges and management strategies that prevent further habitat fragmentation.

Context-dependent resistance of freshwater invertebrate communities to drying.

More freshwater ecosystems are drying in response to global change thereby posing serious threat to freshwater biota and functions. The production of desiccation-resistant forms is an important adaptation that helps maintain biodiversity in temporary freshwaters by buffering communities from drying, but its potential to mitigate the negative effects of drying in freshwater ecosystems could vary greatly across regions and ecosystem types. We explored this context dependency by quantifying the potential contribution of desiccation-resistance forms to invertebrate community recovery across levels of regional drying prevalence (defined as the occurrence of drying events in freshwaters in a given region) and ecosystem types (lentic, lotic) in temporary neotropical freshwaters. We first predicted that regional drying prevalence influences the selection of species with desiccation-resistant forms from the regional species pools and thus increases the ability of communities to recover from drying. Second, we predicted lentic freshwaters harbor higher proportions of species with desiccation-resistant forms compared to lotic, in response to contrasted hydrologic connectivity. To test these predictions, we used natural experiments to quantify the contribution of desiccation-resistant forms to benthic invertebrate community recovery in nine intermittent streams and six geographically isolated temporary wetlands from three Bolivian regions differing in drying prevalence. The contribution of desiccation-resistant forms to community recovery was highest where regional drying prevalence was high, suggesting the species pool was adapted to regional disturbance regimes. The contribution of desiccation-resistant forms to community recovery was lower in streams than in wetlands, emphasizing the importance of hydrologic connectivity and associated recolonization processes from in-stream refuges to recovery in lotic systems. In all regions, the majority of functional traits were present in desiccation-resistant taxa indicating this adaptation may help maintain ecosystem functions by buffering communities from the loss of functional traits. Accounting for regional context and hydrologic connectivity in community recovery processes following drying can help refine predictions of freshwater biodiversity response to global change.