Behavioral and ecological factors account for variation in the mass-independent energy expenditures of endotherms.

A persistent controversy has concerned the identification of the factors that influence the quantitative variation in the physiological characters of species, an example of which is the basal rate of metabolism of endotherms. The most important factor accounting for its variation is body mass as long as the range in mass is appreciable. But mass never accounts for all of the variation and none if species have the same mass. Most of the residual variation around the mass curve is associated with behavioral characters, ecological factors, and phylogeny, i.e., history. These agents influence energy expenditure by different means and at different stages in the life history of species. Phylogeny describes the historic origin, evolution, and distribution of character states in contemporary species. However, the level of energy expenditure is quantitatively determined by the collective of realized states in combination with conditions in the environment. Therefore, two stages determine energy expenditure: (1) the evolution of character states and (2) their impact in conjunction with conditions on the environment. Behavioral characters and ecological factors, when coupled with log10 mass, usually account for >94 % of the variation in the log10 basal rates of birds and mammals, a capacity not found in phylogenetic analyses. The difficulty of determining a direct impact of phylogeny on physiological characters results from its correlation with behavioral characters. When appropriate, the passerine/non-passerine dichotomy in birds and the sub/infraclass dichotomy in mammals combine with behavioral characters, ecological factors, and log10 mass to increase r (2) to account for 96-99 % of the variation in log10 basal rate. This occurs because dichotomies incorporate factors other than those already in the analyses. The clearest demonstration of the direct impact of character states is the equality of energy expenditure in species convergent with species from other clades without any evidence of the clade to which the species belong. A multifactorial approach depends on the inclusion of life history characteristics of species, but this analysis applies only to continuous, quantitative relationships, which are often modified by discontinuous agents. Distinctive character states distinguish species and therefore must be included in any attempt to account for differences in their level of energy expenditure. No analysis that ignores life history characteristics can account for the quantitative variation in energy expenditure beyond that associated with body mass.

Standard energetics of leaf-nosed bats (Hipposideridae): its relationship to intermittent- and protracted-foraging tactics in bats and birds.

Basal rates of metabolism within the insectivorous genera Hipposideros and Ascelliscus, Old World leaf-nosed bats (Hipposideridae), ranged from 58% to 77% of the mammalian standard. The larger species, Hipposideros diadema and Hipposideros maggietaylori, effectively thermoregulated at ambient temperatures down to 9 degrees C, whereas two smaller species, Hipposideros galeritus and Hipposideros cervinus, occasionally permitted body temperatures to fall below 32 degrees C. The low basal rates of metabolism in hipposiderids correlated with a predatory life-style characterized by intermittent flight from a perch to capture insects, a correlation similar to that found in nonpasserine birds. Intermittent-foraging bats and nonpasserines collectively had basal rates of metabolism that averaged 75% of those that pursue insects during protracted flight. However, no difference in basal rate was found between protracted- and intermittent-foraging passerines, which had basal rates 1.8- and 2.4-times those of protracted-foraging and intermittent-foraging bats and nonpasserines, respectively. Bats, swifts, and caprimulgids that enter torpor have basal rates that are 85% of those of similar species that do not enter torpor. Body mass, order affiliation, foraging mode, and propensity to enter into torpor collectively account for 97% of the variation in basal rate of metabolism in insectivorous bats and birds. Foraging style therefore appears to be a factor contributing to the diversity in endotherm energetics. Minimal thermal conductance in the genus Hipposideros ranged from 75% to 102% of the mammalian standard. Birds have minimal thermal conductances that are 75% of mammals and intermittent foragers have minimal conductances that are 78% of protracted foragers.

The comparative energetics of 'caviomorph' rodents.

The energetics of 11 species of New World hystricognath ('caviomorph') rodents are presented and compared with data from the literature on 19 additional species. Log(10) body mass alone accounts for 94% of the variation in the log(10) basal rate of metabolism in caviomorphs. The residual variation in basal rate is correlated with the stratum on which species live: arboreal species have low basal rates; terrestrial and fossorial species have intermediate basal rates; and aquatic species have high basal rates. When stratum is not included in the analysis, folivores, especially those that are arboreal, have lower basal rates than species with other food habits when combined with log(10) body mass. Small island endemics, all of which are folivores, have basal rates that are 61% of continental species. Log(10) basal rate correlates with family affiliation when combined with log(10) mass, but only if no other factor is included. Therefore, caviomorphs with low basal rates are arboreal, folivorous, live on small islands and belong to the Capromyidae, whereas other character combinations are associated with higher basal rates. These observations demonstrate that the basal rates of caviomorphs reflect many factor interactions. No differences in basal rate were found to reflect climate. Log(10) mass, the only factor to correlate with conductance, accounts for 82% of the variation in log(10) minimal thermal conductance. Mean interspecific body temperature was 36.9 degrees C; it was lowest in aquatic and fossorial species.

The metabolism of New Guinean pteropodid bats.

Rate of metabolism and body temperature were measured in eight species of pteropodid bats that live in Papua New Guinea. These data and those from 13 other species available in the literature are examined to determine the factors with which the energetics of pteropodids are correlated. Lowland populations of species that weigh < 35 g generally have low basal rates and often become torpid, whereas their highland populations had standard or high basal rates and were precise thermoregulators, as were all adult pteropodids that weighed more than 60 g. In large pteropodids belonging to the genera Dobsonia and Pteropus, females are smaller and consequently have lower total basal rates than males. Compared to species found on continents and large islands, species limited in distribution to small islands have lower basal rates, both because of a smaller mass and a reduction of metabolism independent of body mass. These trends are most marked in females that belong to small-island species, adjustments that may facilitate reproduction and survival on small oceanic islands with a limited resource base. Minimal thermal conductances are usually low in species that roost exposed in trees, whereas large species that roost in caves have either standard or high conductances.

The influence of body mass, climate, and distribution on the energetics of South Pacific pigeons.

Rate of metabolism and temperature regulation were studied in 16 species of South Pacific pigeons, which constitute 13 fruit-eaters, 1 seed-eater, 1 fruit/nut-eater, and 1 fruit/leaf-eater; 14 tropical and two temperate species; and ten mainland and six intermediate- or small-island species. The data presented here and those from 11 additional columbids indicate in an analysis of covariance that log(10) basal rate of metabolism is correlated with log(10) body mass (P< or =0.0001), distribution (P=0.0023), and climate (P=0.0016). These factors account for 94.3% of the variation in log(10) basal rate of metabolism. In this analysis the lowest basal rates, corrected for body mass, are found in tropical pigeons living on small oceanic islands, whereas the highest basal rates are found in temperate species living on continents. The reduction of basal rate in large columbids facilitates their long-term persistence on small islands characterized by a limited resource base and unstable weather. Some small-island specialists have a smaller mass than their continental relatives, which further reduces resource requirements. The question whether a reduction in basal rate occurs in small columbids on small islands is unresolved. Log(10) minimal thermal conductance is apparently correlated only with log(10) body mass (P< or =0.0001); r(2)=89.4%. The mean nocturnal body temperature of columbids is 39.7 degrees C.

Laboratory and field studies of the energy expenditure of endotherms: A comparison.

Field energy expenditures of endotherms have been estimated by combining laboratory measurements of energy expenditure with field time budgets and by direct measurements in the field through the use of doubly-labelled water. Too much variation occurs in field expenditures to rely on scaling alone to predict energy expenditure without losing the unique characteristics of distinctive species. Field expenditures are most effectively analysed when combined with measurements of the same species under 'standard' conditions in the laboratory. The effect of body size on the expenditures associated with maintenance, locomotion and reproduction are also most clearly established in the laboratory.

Complications inherent in scaling the basal rate of metabolism in mammals.

The scaling of the basal rate of metabolism in mammals is reexamined. Both the power and level of the scaling function are sensitive to various factors that interact with body mass and rate of metabolism, including the precision of temperature regulation, food habits, and activity level. This sensitivity implies that the rate of metabolism is a highly plastic character in the course of evolution. Consequently, the singular effect of mass on the rate of metabolism is most effectively analyzed in ecologically and physiologically uniform sets of species, rather than in taxonomically defined groups, which often are ecologically and physiologically diverse. Otherwise, all fitted curves for mammals integrate a variety of competing factors, thereby reflecting the species used and denying unique analytic significance to the power in scaling relations. Kleiber's eutherian curve may represent a relatively uniform set of data because all the species included were domesticated and because selection for high rates of production (and high rates of metabolism) occurred in the process of domestication. In the analysis of scaling relationships, the standard error of estimate (Sy.x) is a more valuable measure of the residual variation than is (1.0-r2) because r2 is a non-linear measure of the conformation of data to the relation and because Sy.x, unlike r2, is independent of the units used in the scaling relationship. At present the best estimate indicates that total rate of metabolism scales proportionally to approximately m0.60 at small masses (less than 300 g), as long as small species do not enter torpor, and scales proportionally to approximately m0.75 at large masses (greater than or equal to 300 g). Physiological properties other than metabolism are potentially sensitive to secondary factors, so their scaling functions also would be most clearly defined for physiologically uniform groups of species. This view suggests that insight into the significance of scaling relations can be obtained by examining the residual variation around a scaling function as well as by examining conformation to the function.