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Metabolic rate is a key component of energy budgets that scales with body size and varies with large-scale environmental geographical patterns. Here we conduct an analysis of standard metabolic rates (SMR) of marine ectotherms across a 70° latitudinal gradient in both hemispheres that spanned collection temperatures of 0-30 °C. To account for latitudinal differences in the size and skeletal composition between species, SMR was mass normalized to that of a standard-sized (223 mg) ash-free dry mass individual. SMR was measured for 17 species of calcified invertebrates (bivalves, gastropods, urchins and brachiopods), using a single consistent methodology, including 11 species whose SMR was described for the first time. SMR of 15 out of 17 species had a mass-scaling exponent between 2/3 and 1, with no greater support for a 3/4 rather than a 2/3 scaling exponent. After accounting for taxonomy and variability in parameter estimates among species using variance-weighted linear mixed effects modelling, temperature sensitivity of SMR had an activation energy (Ea) of 0.16 for both Northern and Southern Hemisphere species which was lower than predicted under the metabolic theory of ecology (Ea 0.2-1.2 eV). Northern Hemisphere species, however, had a higher SMR at each habitat temperature, but a lower mass-scaling exponent relative to SMR. Evolutionary trade-offs that may be driving differences in metabolic rate (such as metabolic cold adaptation of Northern Hemisphere species) will have important impacts on species abilities to respond to changing environments.

Animal responses to changing environments are most commonly studied in relation to temperature change. The current paradigm for marine ectotherms is that temperature limits are set through oxygen limitation. Oxygen limitation leads to progressive reductions in capacity to perform work or activity, and these are more important and proximate measures of a population's ability to survive. Here we measured the ability of a large Antarctic clam to rebury when removed from sediment at temperatures between -1.5 and 7.5 degrees C and at three oxygen concentrations, 10.2, 20.5 and 27.7%. The proportion of the population capable of reburying declined rapidly and linearly with temperature from around 65% at 0 degrees C to 0% at 6 degrees C in normoxia (20.5% O2). Decreasing oxygen to 10.2% reduced temperature limits for successful burial by around 2 degrees C, and increasing oxygen to 27.7% raised the limits by 1-1.5 degrees C. There was an interactive effect of body size and temperature on burying: the temperature limits of larger individuals were lower than smaller animals. Similarly, these size limits were increased by increasing oxygen availability. Considering data for all temperatures and oxygen levels, the fastest burying rates occurred at 3 degrees C, which is 2 degrees C above the maximum summer temperature at this site.

Seasonal dormancy is a widespread mechanism for reducing energy expenditure during periods of low energy availability. Seasonal variation in activity and the cost of pumping water through the siphons were investigated to estimate the importance of activity regulation to the seasonal energy budget of the Antarctic clam, Laternula elliptica. In the laboratory, a metabolic rate of 26.35 micromol O(2) h(-1) was estimated for a 50-mm shell length L. elliptica pumping water at -0.4 degrees C. In the field, the proportion of time siphons were visible at the sediment surface varied seasonally (32% visible in June/July compared to 86% in December/January). L. elliptica were actively pumping for a minimum of 19% of each 24-h period during winter (August) compared to a summer maximum when animals were actively pumping for 73% of the time (February). This resulted in a 3.7-fold seasonal difference in the calculated energy consumption of a 50-mm L. elliptica (19.2 micromol O(2) h(-1) in February versus 5.0 micromol O(2) h(-1) in August), which closely matches the 3.0-fold seasonal variation in metabolic rate found previously. Seasonal variation in activity could therefore be responsible for much of the seasonal difference in energy consumption of L. elliptica. Inter-annual variation in timing of the seasonal activity maxima (January 2004 and March 1999) was correlated with variation in the timing of the summer plankton bloom in Ryder Bay. In the laboratory, periods of extended siphon closure (133 +/- 114 min, mean +/- SD) were accompanied by long periods of heart arrhythmia (167 +/- 135 min), during which time blood oxygen levels dropped to values close to zero. Heart arrhythmia is most likely part of a hypo-metabolic adaptation to reduce energy costs during extended periods of siphon closure. Physiological and behavioural dormancy, with associated hypoxia tolerance, appear to be key mechanisms controlling the seasonal energy budget of L. elliptica.