Abstract

In order to predict which species can successfully cope with global warming and how other environmental stressors modulate their vulnerability to climate‐related environmental factors, an understanding of the ecophysiology underpinning thermal limits is essential for both conservation biology and invasion biology.Heat tolerance and the extent to which heat tolerance differed with oxygen availability were examined for four native and four alien freshwater peracarid crustacean species, with differences in habitat use across species. Three hypotheses were tested: (1) Heat and lack of oxygen synergistically reduce survival of species; (2) patterns in heat tolerance and the modulation thereof by oxygen differ between alien and native species and between species with different habitat use; (3) small animals can better tolerate heat than large animals, and this difference is more pronounced under hypoxia.To assess heat tolerances under different oxygen levels, animal survival was monitored in experimental chambers in which the water temperature was ramped up (0.25°C min−1). Heat tolerance (CTmax) was scored as the cessation of all pleopod movement, and heating trials were performed under hypoxia (5 kPa oxygen), normoxia (20 kPa) and hyperoxia (60 kPa).Heat tolerance differed across species as did the extent by which heat tolerance was affected by oxygen conditions. Heat‐tolerant species, for example, Asellus aquaticus and Crangonyx pseudogracilis, showed little response to oxygen conditions in their CTmax, whereas the CTmax of heat‐sensitive species, for example, Dikerogammarus villosus and Gammarus fossarum, was more plastic, being increased by hyperoxia and reduced by hypoxia.In contrast to other studies on crustaceans, alien species were not more heat‐tolerant than native species. Instead, differences in heat tolerance were best explained by habitat use, with species from standing waters being heat tolerant and species from running waters being heat sensitive. In addition, larger animals displayed lower critical maximum temperature, but only under hypoxia. An analysis of data available in the literature on metabolic responses of the study species to temperature and oxygen conditions suggests that oxygen conformers and species whose oxygen demand rapidly increases with temperature (low activation energy) may be more heat sensitive.The alien species D. villosus appeared most susceptible to hypoxia and heat stress. This may explain why this species is very successful in colonizing new areas in littoral zones with rocky substrate which are well aerated due to continuous wave action generated by passing ships or prevailing winds. This species is less capable of spreading to other waters which are poorly oxygenated and where C. pseudogracilis is the more likely dominant alien species. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13050/suppinfo is available for this article.

Highlights

  • Freshwater animals face new thermal challenges owing to climate change, local discharge of cooling water and to globalization resulting in redistribution and invasions of species into habitats with a different thermal regime

  • Three hypotheses were tested: (1) Heat and lack of oxygen synergistically reduce survival of species; (2) patterns in heat tolerance and the modulation thereof by oxygen differ between alien and native species and between species with different habitat use; (3) small animals can better tolerate heat than large animals, and this difference is more pronounced under hypoxia

  • Heat tolerance (CTmax) was scored as the cessation of all pleopod movement, and heating trials were performed under hypoxia (5 kPa oxygen), normoxia (20 kPa) and hyperoxia (60 kPa). 4

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Summary

| INTRODUCTION

Freshwater animals face new thermal challenges owing to climate change, local discharge of cooling water and to globalization resulting in redistribution and invasions of species into habitats with a different thermal regime. The problem of insufficient oxygen under warm conditions may be more immediate in water than in air as aquatic gas exchange is challenging due to the lower rate of oxygen diffusion in water and the larger effort required for ventilation as water has a higher density and viscosity (Verberk & Atkinson, 2013) This challenge of breathing underwater is thought to explain the recurrent evolution of air breathing in crustaceans from tropical waters that may be more prone to severe hypoxia (Fusi et al, 2016; Giomi et al, 2014). Alien species may have experienced low or changing oxygen conditions during the invasion process, for example, during ballast water transport or crossing different water bodies, explaining why species that successfully spread outside their native range could better cope with hypoxia and heat (Bates et al, 2013; Jewett, Hines, & Ruiz, 2005; Lenz et al, 2011). We tested whether differences in thermal tolerance between our study species were related to their oxygen demand (e.g. Verberk & Bilton, 2011), by collating published data on oxygen consumption rates in response to temperature and ambient oxygen conditions

| MATERIALS AND METHODS
Findings
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