Warming tolerance across insect ontogeny: influence of joint shifts in microclimates and thermal limits
Résumé
Not all species will experience global warming
equally. First, organisms liv e in microcli mates, the
temperatures of which can deviate substantially from
macroclimate temperatures (Gates 1980, Angilletta
2009). Second, the physiological performance and
tolerance of organisms filters microclimate conditions,
often nonlinearly (Huey and Kingsolver 1993, Huey et
al. 2012). Consequently, the impact of warming on the
persistence and distribution of species cannot be inferred
solely from macroclimatic observations. Instead, what is
required is a mechanistic understanding of both
microclimate and the physiological tolerance of species
(Williams et al. 2008, Helmuth et al. 2010). For instance,
tropical ectotherms were predicted to suffer the most
from global warming because they already live close to
their ther mal li mits, wh ereas temperat e ectother ms
would benefit from a higher warming tolerance: a larger
difference between macroclimate temperature and ther-
mal limits (Deutsch et al. 2008, Tewksbury et al. 2008).
Recently, a study showed that equilibrium body
temperatures of most ectotherms at all latitudes are
higher than both their thermal limits and ambient air
temperature (inferred as the highest monthly mean of
daily maximum air temperature), suggesting the need to
thermo regulat e for the mai ntenanc e of popula tions
(Sunday et al. 2014). However, the importance of
microclimates is often neglected in studies on warming
tolerance of ectotherms, as we will detail (Potter et al.
2013, Scheffers et al. 2013a).
Warming tolerance (WT) is calculated as the devia-
tion between the thermal limit—e.g., the critical
maximal temperature, CTmax, or the uppe r letha l
temperature, ULT—and the habitat temperature, which
usually is taken as a monthly or annual average of
macroscale air temperature (Deutsch et al. 2008). On
that basis, a dire ct r elatio nshi p betwe en warming
tolerance and absolute latitude is expected in ectotherms
because macroclimate temperature increases from the
poles to the equator and there is no clear pattern of
CTmax or ULT over latitudes (Addo-Bediako et al.
2000, Deutsch et al. 2008, Sunday et al. 2011, Diamond
et al. 2012, Arau´ jo et al. 2013). Frequently, this
latitudinal pattern of warming tolerance is used to
forecast potential distribution shifts in ectotherms
caused by global warming (Sunday et al. 2012). Macro-
climate temperature, however, can be uninformative
regarding the actual relationship between physiological
tolerance and latitude, because the pertinent variable
often is the microclimate temperature (Fig. 1; see Huey
et al. 2012, Sunday et al. 2014). Two lines of evidence
suggestthatlackofknowledgeonmicroclimates