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• 1. [Article] The Effect of Temperature on Phenotypes of the Invasive European Green Crab: Physiologic Mechanisms that Facilitate Invasion Success

  Invasion physiology is an emerging field that endeavors to understand the influence of physiological traits on the establishment of non-native species in novel environments. The invasive European green ...

  Citation × Citation Citation Abstract Copy Title: The Effect of Temperature on Phenotypes of the Invasive European Green Crab: Physiologic Mechanisms that Facilitate Invasion Success Author: Kelley, Amanda Year: 2013 Invasion physiology is an emerging field that endeavors to understand the influence of physiological traits on the establishment of non-native species in novel environments. The invasive European green crab,Carcinus maenas, is one of the world's most successful aquatic invaders, and is currently distributed across temperate marine ecosystems globally. The work presented here explored the thermal physiology of this species, and has highlighted several physiological traits that have likely influenced establishment success. Intraspecific comparisons of crabs sampled from the northern and southern edges of their recipient, or invaded range on the west coast of North America have identified both organismal and cellular physiological difference with respect to upper and lower thermal tolerances. Crabs sampled from British Columbia, Canada (BC) had a significantly lower mean upper thermal tolerance threshold and heat shock protein synthesis, Hsp70, compared to their warm acclimated conspecifics sampled from California (CA). These differential physiologic responses may be rooted in the disparate natural thermal habitats that each population occupies within their respective environments. The ability of this species to extend its current range limits was also investigated. Range expansion to the south has been limited, and is likely restricted by this species lack of adaptation to warmer temperatures. Because range expansion has been chiefly northward, characterizing this species' response to cold stress can identify whether colder temperatures poleward may limit further range expansion. Cold tolerance capacity was determined in the laboratory, and crabs sampled from Vancouver Island, British Columbia were able to withstand the over-wintering thermal regime that occurs in Sitka, Alaska, a site that is currently beyond the range limits of this species. Furthermore, intraspecific assessments found that the cold acclimated BC population exposed to cold shock significantly down regulated protein levels of cyclin D1, cell cycle modulator. Distinct differences in carapace width (CW) were detected along the thermal gradient present in the green crabs' range. This variation in body size was utilized to the test the temperature size rule hypothesis for ectotherms. Simply stated, the temperature size rule is the tendency for ectotherms to develop slower but mature to a larger body sizes at cooler temperatures. The results supported this hypothesis as crabs sampled from the warm portion of the range were found to be smaller than crabs sampled from the colder portion of the range. This pattern was detected along the native range as well. Differences in body size have the potential to influence the scope of invasion; larger individuals are generally more fecund and longer lived, which can increase both the intensity and frequency of larval dispersal that could further propel range expansion. The physiologic properties that the green crab possesses which may influence invasion success were examined using peer-reviewed literature with the aim of determining if these physiological traits confer invasion success across taxa. This analysis tested four hypotheses: 1) Broad geographic temperature tolerances (thermal width) confer a higher upper thermal tolerance threshold when comparing invasive and native species. 2) The upper thermal extreme experienced in nature is correlated with upper thermal tolerance threshold. 3) Protein chaperone expression, a cellular mechanism underlying thermal tolerance threshold, is greater in invasive organisms than in native ones. 4) Acclimation to higher temperatures can promote a greater range of thermal tolerance for invasives compared to natives. These preliminary results generally support the four stated hypotheses, and provide a solid foundation for further studies to explore and identify physiologic traits that facilitate invasion success. Overall, these studies investigated the thermal physiology ofCarcinus maenasfrom an invasive metapopulation and have brought about significant advances in our understanding of what physiologic traits correlate to invasion success in this species. In addition, the data presented here can aid resource managers in identifying habitats, based on thermal tolerance measurements that fit the criteria for invasion. Understanding how invasive organisms vary with respect to thermal tolerance can aid our understanding the patterns and processes of species invasions. Title: The Effect of Temperature on Phenotypes of the Invasive European Green Crab: Physiologic Mechanisms that Facilitate Invasion Success Author: Kelley, Amanda Year: 2013 Invasion physiology is an emerging field that endeavors to understand the influence of physiological traits on the establishment of non-native species in novel environments. The invasive European green crab,Carcinus maenas, is one of the world's most successful aquatic invaders, and is currently distributed across temperate marine ecosystems globally. The work presented here explored the thermal physiology of this species, and has highlighted several physiological traits that have likely influenced establishment success. Intraspecific comparisons of crabs sampled from the northern and southern edges of their recipient, or invaded range on the west coast of North America have identified both organismal and cellular physiological difference with respect to upper and lower thermal tolerances. Crabs sampled from British Columbia, Canada (BC) had a significantly lower mean upper thermal tolerance threshold and heat shock protein synthesis, Hsp70, compared to their warm acclimated conspecifics sampled from California (CA). These differential physiologic responses may be rooted in the disparate natural thermal habitats that each population occupies within their respective environments. The ability of this species to extend its current range limits was also investigated. Range expansion to the south has been limited, and is likely restricted by this species lack of adaptation to warmer temperatures. Because range expansion has been chiefly northward, characterizing this species' response to cold stress can identify whether colder temperatures poleward may limit further range expansion. Cold tolerance capacity was determined in the laboratory, and crabs sampled from Vancouver Island, British Columbia were able to withstand the over-wintering thermal regime that occurs in Sitka, Alaska, a site that is currently beyond the range limits of this species. Furthermore, intraspecific assessments found that the cold acclimated BC population exposed to cold shock significantly down regulated protein levels of cyclin D1, cell cycle modulator. Distinct differences in carapace width (CW) were detected along the thermal gradient present in the green crabs' range. This variation in body size was utilized to the test the temperature size rule hypothesis for ectotherms. Simply stated, the temperature size rule is the tendency for ectotherms to develop slower but mature to a larger body sizes at cooler temperatures. The results supported this hypothesis as crabs sampled from the warm portion of the range were found to be smaller than crabs sampled from the colder portion of the range. This pattern was detected along the native range as well. Differences in body size have the potential to influence the scope of invasion; larger individuals are generally more fecund and longer lived, which can increase both the intensity and frequency of larval dispersal that could further propel range expansion. The physiologic properties that the green crab possesses which may influence invasion success were examined using peer-reviewed literature with the aim of determining if these physiological traits confer invasion success across taxa. This analysis tested four hypotheses: 1) Broad geographic temperature tolerances (thermal width) confer a higher upper thermal tolerance threshold when comparing invasive and native species. 2) The upper thermal extreme experienced in nature is correlated with upper thermal tolerance threshold. 3) Protein chaperone expression, a cellular mechanism underlying thermal tolerance threshold, is greater in invasive organisms than in native ones. 4) Acclimation to higher temperatures can promote a greater range of thermal tolerance for invasives compared to natives. These preliminary results generally support the four stated hypotheses, and provide a solid foundation for further studies to explore and identify physiologic traits that facilitate invasion success. Overall, these studies investigated the thermal physiology ofCarcinus maenasfrom an invasive metapopulation and have brought about significant advances in our understanding of what physiologic traits correlate to invasion success in this species. In addition, the data presented here can aid resource managers in identifying habitats, based on thermal tolerance measurements that fit the criteria for invasion. Understanding how invasive organisms vary with respect to thermal tolerance can aid our understanding the patterns and processes of species invasions. Close