title: Conserved and narrow temperature limits in alpine insects: Thermal tolerance and supercooling points of the ice-crawlers, Grylloblatta (Insecta: Grylloblattodea: Grylloblattidae).
abstract: For many terrestrial species, habitat associations and range size are dependent on physiological limits, which in turn may influence large-scale patterns of species diversity. The temperature range experienced by individuals is considered to shape the breadth of the thermal niche, with species occupying temporally and/or geographically stable climates tolerating a narrow temperature range. High-elevation environments experience large temperature fluctuations, with frequent periods below 0 °C, but Grylloblatta (Grylloblattodea: Grylloblattidae) occupy climatically stable microhabitats within this region. Here we test critical thermal limits and supercooling points for five Grylloblatta populations from across a large geographic area, to examine whether the stable microhabitats of this group are associated with a narrow thermal niche and assess their capacity to tolerate cold conditions. Thermal limits are highly conserved in Grylloblatta, despite substantial genetic divergence among populations spanning 1500 m elevation and being separated by over 500 km. Further, Grylloblatta show exceptionally narrow thermal limits compared to other insect taxa with little capacity to improve cold tolerance via plasticity. In contrast, upper thermal limits were significantly depressed by cold acclimation. Grylloblatta maintain coordinated movement until they freeze, and they die upon freezing. Convergence of the critical thermal minima, supercooling point and lower lethal limits point to adaptation to a cold but, importantly, constant thermal environment. These physiological data provide an explanation for the high endemism and patchy distribution of Grylloblatta, which relies on subterranean retreats to accommodate narrow thermal limits. These retreats are currently buffered from temperature fluctuations by snow cover, and a declining snowpack thus places Grylloblatta at risk of exposure to temperatures beyond its tolerance capacity.
authors.affiliation: University of Wisconsin-Madison, Department of Entomology, 1630 Linden Drive, Madison, WI 53706, USA. Electronic address: schoville2@wisc.edu.
authors.forename: Sean D
authors.initials: SD
authors.lastname: Schoville
authors.affiliation: Department of Zoology, University of Melbourne, Parkville 3010, Australia.
authors.forename: Rachel A
authors.initials: RA
authors.lastname: Slatyer
authors.affiliation: Conservation Biology Center, University of the Wilderness, 919 S. Adams St., Spokane, WA 99204, USA.
authors.forename: James C
authors.initials: JC
authors.lastname: Bergdahl
authors.affiliation: University of Wisconsin-Madison, Department of Entomology, 1630 Linden Drive, Madison, WI 53706, USA.
authors.forename: Glenda A
authors.initials: GA
authors.lastname: Valdez
mesh_terms: D000064:Acclimatization; D000222:Adaptation, Physiological; D000531:Altitude; D000818:Animals; D002980:Climate; D003080:Cold Temperature; D017753:Ecosystem; D007313:Insecta / Q000502:physiology*
keywords: Climate variability hypothesis; Ectotherm; Freeze avoidance; Lethal limits; Physiological tolerance; Plasticity