Tropical birds are relatively long-lived and produce few offspring, which develop slowly and mature relatively late in life, the slow end of the life-history axis, whereas temperate birds lie at the opposite end of this continuum. We tested the hypothesis that tropical birds have evolved a reduced basal metabolic rate (BMR). We measured BMR of 69 species of tropical birds, the largest data set amassed on metabolic rates of tropical birds, and compared these measurements with 59 estimates of BMR for temperate birds. Our analyses included conventional least squares regression, regressions based on phylogenetic independent contrasts, and a comparison of BMR of 13 phylogenetically matched pairs, one species from the tropics and one from northerly temperate areas. Our triptych showed that tropical birds had a reduced BMR, compelling evidence for a connection between the life history of tropical birds and a slow pace of life. Further, tropical migrants breeding in temperate habitats had a lower BMR than did temperate residents, suggesting that these migrants have physiological traits consistent with a slow pace of life. In addition, we determined that tropical birds had a lower cold-induced peak metabolic rate and thermogenic metabolic scope than temperate species, a finding that is consistent with the hypothesis that their environment has not selected for high levels of thermogenesis, or alternatively, that a slow pace of life may be incompatible with high thermogenic capacity. We conclude that physiological function correlates with the suite of life-history traits.basal metabolic rate ͉ life history ͉ peak metabolic rate ͉ tropics ͉ migration L ife-history traits, such as clutch size, age at maturity, rate of growth of offspring, and longevity tend to cluster in ecological space such that species with long lives produce small numbers of offspring that develop slowly and mature relatively late in life (1-3). At the opposite end of this ''slow-fast'' continuum, organisms have low survival rates coupled with large numbers of offspring that develop rapidly and reach sexual maturity at an early age. Among some species, life-history traits also covary with rate of energy expenditure, the pace of life, such that animals with high rates of reproduction have high energy expenditure, and vice versa (4, 5). Support for the connectivity between life history, especially survivorship, and rate of living comes from studies on fruit flies Drosophila (6), houseflies Musca domestica (7), and nematode worms Caenorhabditis elegans (8), as well as meta-analyses on mammals (2, 9, 10). Although less attention has been given to relationships between life history and metabolism in birds, Trevelyan et al. (11) did show an association between resting metabolic rate in birds and their maximum life span.With small clutches (12-14), slower growth as nestlings (15), long periods when fledglings are dependent on parents (16, 17), and higher rates of adult survival than temperate counterparts (15,(18)(19)(20), tropical birds fall along the slow en...

We studied the relationship between one component of immune function and basal metabolic rate (BMR), an indicator of the 'pace-of-life syndrome', among 12 tropical bird species and among individuals of the tropical house wren (Troglodytes aedon), to gain insights into functional connections between life history and physiology. To assess constitutive innate immunity we introduced a new technique in the field of ecological and evolutionary immunology that quantifies the bactericidal activity of whole blood. This in vitro assay utilises a single blood sample to provide a functional, integrated measure of constitutive innate immunity. We found that the bactericidal activity of whole blood varied considerably among species and among individuals within a species. This variation was not correlated with body mass or whole-organism BMR. However, among species, bacteria killing activity was negatively correlated with mass-adjusted BMR, suggesting that species with a slower pace-of-life have evolved a more robust constitutive innate immune capability. Among individuals of a single species, the house wren, bacteria killing activity was positively correlated with mass-adjusted BMR, pointing to physiological differences in individual quality on which natural selection potentially could act.

We tested the hypothesis that birds in arid environments, where primary productivity is low and surface water is scarce, have reduced energy expenditure and water loss compared with their mesic counterparts. Using both conventional least squares regression and regression based on phylogenetically independent contrasts, we showed that birds from desert habitats have reduced basal and field metabolic rates compared with species from mesic areas. Previous work showed that desert birds have reduced rates of total evaporative water loss when exposed to moderate environmental temperatures in the laboratory. We tested whether reduced rates of total evaporative water loss translate into low field water fluxes. Conventional ANCOVA indicated that desert birds have reduced water fluxes, but an analysis based on phylogenetically independent contrasts did not support this finding, despite the wide array of taxonomic affiliations of species in the data set. We conclude that the high ambient temperatures, the low primary productivity, and the water scarcity in desert environments have selected for or resulted in reduced rates of energy expenditure and evaporative water loss in birds that live in these climes.

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Broad-scale comparisons of birds indicate the possibility of adaptive modification of basal metabolic rate (BMR) and total evaporative water loss (TEWL) in species from desert environments, but these might be confounded by phylogeny or phenotypic plasticity. This study relates variation in avian BMR and TEWL to a continuously varying measure of environment, aridity. We test the hypotheses that BMR and TEWL are reduced along an aridity gradient within the lark family (Alaudidae), and investigate the role of phylogenetic inertia. For 12 species of lark, BMR and TEWL decreased along a gradient of increasing aridity, a finding consistent with our proposals. We constructed a phylogeny for 22 species of lark based on sequences of two mitochondrial genes, and investigated whether phylogenetic affinity played a part in the correlation of phenotype and environment. A test for serial independence of the data for masscorrected TEWL and aridity showed no influence of phylogeny on our findings. However, we did discover a significant phylogenetic effect in mass-corrected data for BMR, a result attributable to common phylogenetic history or to common ecological factors. A test of the relationship between BMR and aridity using phylogenetic independent constrasts was consistent with our previous analysis: BMR decreased with increasing aridity.