Allometric scaling of seed retention time in seed dispersers and its application to estimation of seed dispersal potentials of theropod dinosaurs

Seed retention time (SRT), the time interval between seed ingestion and defaecation, is a critical parameter that determines the spatial pattern of seed dispersal created by an animal, and is therefore, an essential component of trait‐based modelling of seed dispersal functions. However, no simple predictive model of SRT for any given animal exists. We explored the linkage between animal traits and SRT. We collected previously published data on mean SRT for 112 species of birds, mammals, reptiles and fishes and investigated the general allometric scaling of mean SRT with body mass for each taxon. Moreover, we analysed the effects of food habit and digestive strategy on mean SRT for birds and mammals. In general, mean SRT increased with body mass in all four taxa, whereas the pattern of allometric scaling varied greatly among the taxa. Birds had a smaller intercept and larger slope than those of mammals, whereas reptiles had a much larger intercept and smaller slope than those of either birds or mammals. For birds, food habit was also detected as an important factor affecting SRT. We applied the allometric scaling that was obtained for birds to estimate mean SRT of extinct Mesozoic dinosaurs (Theropoda) – few of which are assumed to have acted as seed dispersers. SRT for large carnivorous theropods was estimated to be 4–5 days, when considering only body mass. The present study provides allometric scaling parameters of mean SRT for a variety of seed‐dispersing animals, and highlights large variations in scaling among taxa. The allometric scaling obtained could be a critical component of further trait‐based modelling of seed dispersal functions. Further, the potential and limitations of the scaling of animal SRT with body mass and a future pathway to the development of trait‐based modelling are discussed.     

Rapid warming is associated with population decline among terrestrial birds and mammals globally

Animal populations have undergone substantial declines in recent decades. These declines have occurred alongside rapid, human‐driven environmental change, including climate warming. An association between population declines and environmental change is well established, yet there has been relatively little analysis of the importance of the rates of climate warming and its interaction with conversion to anthropogenic land use in causing population declines. Here we present a global assessment of the impact of rapid climate warming and anthropogenic land use conversion on 987 populations of 481 species of terrestrial birds and mammals since 1950. We collated spatially referenced population trends of at least 5 years’ duration from the Living Planet database and used mixed effects models to assess the association of these trends with observed rates of climate warming, rates of conversion to anthropogenic land use, body mass, and protected area coverage. We found that declines in population abundance for both birds and mammals are greater in areas where mean temperature has increased more rapidly, and that this effect is more pronounced for birds. However, we do not find a strong effect of conversion to anthropogenic land use, body mass, or protected area coverage. Our results identify a link between rapid warming and population declines, thus supporting the notion that rapid climate warming is a global threat to biodiversity.     

An exploration of differences in the scaling of life history traits with body mass within reptiles and between amniotes

Allometric relationships linking species characteristics to body size or mass (scaling) are important in biology. However, studies on the scaling of life history traits in the reptiles (the nonavian Reptilia) are rather scarce, especially for the clades Crocodilia, Testudines, and Rhynchocephalia (single extant species, the tuatara). Previous studies on the scaling of reptilian life history traits indicated that they differ from those seen in the other amniotes (mammals and birds), but so far most comparative studies used small species samples and also not phylogenetically informed analyses. Here, we analyzed the scaling of nine life history traits with adult body mass for crocodiles (n = 22), squamates (n = 294), turtles (n = 52), and reptiles (n = 369). We used for the first time a phylogenetically informed approach for crocodiles, turtles, and the whole group of reptiles. We explored differences in scaling relationships between the reptilian clades Crocodilia, Squamata, and Testudines as well as differences between reptiles, mammals, and birds. Finally, we applied our scaling relationships, in order to gain new insights into the degree of the exceptionality of the tuatara's life history within reptiles. We observed for none of the life history traits studied any difference in their scaling with body mass between squamates, crocodiles, and turtles, except for clutch size and egg weight showing small differences between these groups. Compared to birds and mammals, scaling relationships of reptiles were similar for time‐related traits, but they differed for reproductive traits. The tuatara's life history is more similar to that of a similar‐sized turtle or crocodile than to a squamate.     

Home range, home range overlap, and species energy use among herbivorous mammals

Harestad & Bunnell (1979) showed that, at least for North American species, home ranges of large herbivorous mammals are relatively larger than we would expect on metabolic grounds, and suggested that the productivity of the environment for mammal species decreases with increasing body size. This interpretation assumes that the number of conspecifics that share an individual's home range is independent of body size. Data presented here show that this is not true for the species in their sample; the home range is shared with an increasing number of conspecifics in larger herbivore species. The productivity of the environment for a species is independent of body size and the area available to an individual for its own use scales approximately as do individual metabolic requirements. These results agree with conclusions based upon the scaling of population density with body mass and illustrate the interrelationship between home range and dietary and social organization trends among mammalian herbivores. Individual home range area is a function of the way in which the local population of a species, not merely an individual, exploits the environment.     

Causes and consequences of variation in offspring body mass: meta‐analyses in birds and mammals

Early survival is highly variable and strongly influences observed population growth rates in most vertebrate populations. One of the major potential drivers of survival variation among juveniles is body mass. Heavy juveniles are better fed and have greater body reserves, and are thus assumed to survive better than light individuals. In spite of this, some studies have failed to detect an influence of body mass on offspring survival, questioning whether offspring body mass does indeed consistently influence juvenile survival, or whether this occurs in particular species/environments. Furthermore, the causes for variation in offspring mass are poorly understood, although maternal mass has often been reported to play a crucial role. To understand why offspring differ in body mass, and how this influences juvenile survival, we performed phylogenetically corrected meta‐analyses of both the relationship between offspring body mass and offspring survival in birds and mammals and the relationship between maternal mass and offspring mass in mammals. We found strong support for an overall positive effect of offspring body mass on survival, with a more pronounced influence in mammals than in birds. An increase of one standard deviation of body mass increased the odds of offspring survival by 71% in mammals and by 44% in birds. A cost of being too fat in birds in terms of flight performance might explain why body mass is a less reliable predictor of offspring survival in birds. We then looked for moderators explaining the among‐study differences reported in the intensity of this relationship. Surprisingly, sex did not influence the intensity of the offspring mass–survival relationship and phylogeny only accounted for a small proportion of observed variation in the intensity of that relationship. Among the potential factors that might affect the relationship between mass and survival in juveniles, only environmental conditions was influential in mammals. Offspring survival was most strongly influenced by body mass in captive populations and wild populations in the absence of predation. We also found support for the expected positive effect of maternal mass on offspring mass in mammals (r_pearson = 0.387). As body mass is a strong predictor of early survival, we expected heavier mothers to allocate more to their offspring, leading them to be heavier and so to have a higher survival. However, none of the potential factors we tested for variation in the maternal mass–offspring mass relationship had a detectable influence. Further studies should focus on linking these two relationships to determine whether a strong effect of offspring size on early survival is associated with a high correlation coefficient between maternal mass and offspring mass.     

Determinants of local abundance in a major radiation of Australian passerines (Aves: Meliphagoidea)

Aim  To identify the factors that contribute to variation in abundance (population density), and to investigate whether habitat breadth and diet breadth predict macroecological patterns in a suborder of passerine birds (Meliphagoidea).
Location  Australia (including Tasmania).
Methods  Mean abundance data were collated from site surveys of bird abundance (the Australian Bird Count); range size and latitudinal position data from published distribution maps; and body mass and diet breadth information from published accounts. A diversity index of habitats used (habitat breadth) was calculated from the bird census data. We used bivariate correlation and multiple regression techniques, employing two phylogenetic comparative methods: phylogenetic generalized least squares and independent contrasts.
Results  Body mass and latitude were the only strong predictors of abundance, with larger-bodied and lower-latitude species existing at lower densities. Together, however, body mass and latitude explained only 11.1% of the variation in mean abundance. Range size and habitat breadth were positively correlated, as were diet breadth and body mass. However, neither range size, nor habitat breadth and diet breadth, explained patterns in abundance either directly or indirectly.
Main conclusions  Levels of abundance (population density) in meliphagoid birds are most closely linked to body mass and latitudinal position, but not range size. As with many other macroecological analyses, we find little evidence for aspects of niche breadth having an effect on patterns of abundance. We hypothesize that evolutionary age may also have a determining effect on why species tend to be rarer (less abundant) in the tropics.     

Dichotomy of eutherian reproduction and metabolism

How anatomical, physiological and ecological (life history) features scale with body mass is a fundamental question in biology. There is an ongoing debate in the scientific literature whether allometric scaling follows a universal pattern that can be described in a single model, or differs between groups. However, recently some analyses were published demonstrating a change in scaling across the body mass range: brain‐size allometry of mammals indicates that scaling follows a curvilinear pattern in double‐logarithmic space, and a quadratic pattern in double‐logarithmic space was found in one of the largest physiological datasets, on basal metabolic rate (MR) in mammals. Here, we analysed a variety of independent datasets on anatomical, physiological and ecological characteristics in mammals, birds and reptiles to answer the question whether the quadratic scaling is a universal biological law, or a pattern unique to mammals. The pattern was present in mammalian basal and field MR, brain size, and reproduction parameters, but neither in other organ allometries in mammals, nor in the scaling of MR in birds and reptiles. However, the curvature was better explained by separate allometric scaling of three different mammalian reproduction strategies: marsupials, and eutherian mammals with one and with many offspring. The two latter strategies are distributed unequally over the body mass range in eutherian mammals. Our findings show that a quadratic model, as well as a traditional allometric model with a universal scaling exponent (such as 0.67 or 0.75), may be inappropriate in mammals as they are a result of different scalings within these three reproductive groups. We propose that the observed distribution pattern is the result of the eutherian mammal clade's uniquely pronounced dichotomy of reproductive strategies.     

Density dependence in an island population of silvereyes

The population of silvereyes Zosterops lateralis chlorocephalus , on Heron Island, Great Barrier Reef has been monitored accurately since 1965. Between 1979 and 1993, the breeding success of all birds was determined by monitoring nests. The population fluctuated between 225 and 483 individuals. Four cyclones led to substantial mortality. As this data set is long-term, has little observation error, and is from an effectively closed population, it provides an unusual opportunity to examine density dependence in reproduction or mortality. Using a stochastic logistic model, we found clear evidence of density dependence in adult population size. Logistic regression suggested that fledgling survival decreased with the numbers of birds attempting to breed. There was also some suggestion that adult survival might be density dependent. The fitted stochastic logistic model predicts negligible risks of extinction for this population, in contrast to the predictions of a published population viability analysis. Whilst our statistical model including density dependence may provide better predictions of the "usual" behaviour of a population than a population viability analysis, we suggest that caution should be exercised when statistically fitted models are used to predict the behaviour of the population at extremes, such as near extinction.     

Global drivers of population density in terrestrial vertebrates

Aim

Although the effects of life history traits on population density have been investigated widely, how spatial environmental variation influences population density for a large range of organisms and at a broad spatial scale is poorly known. Filling this knowledge gap is crucial for global species management and conservation planning and to understand the potential impact of environmental changes on multiple species.

Location

Global.

Time period

Present.

Major taxa studied

Terrestrial amphibians, reptiles, birds and mammals.

Methods

We collected population density estimates for a range of terrestrial vertebrates, including 364 estimates for amphibians, 850 for reptiles, 5,667 for birds and 7,651 for mammals. We contrasted the importance of life history traits and environmental predictors using mixed models and tested different hypotheses to explain the variation in population density for the four groups. We assessed the predictive accuracy of models through cross‐validation and mapped the partial response of vertebrate population density to environmental variables globally.

Results

Amphibians were more abundant in wet areas with high productivity levels, whereas reptiles showed relatively higher densities in arid areas with low productivity and stable temperatures. The density of birds and mammals was typically high in temperate wet areas with intermediate levels of productivity. The models showed good predictive abilities, with pseudo‐R² ranging between 0.68 (birds) and 0.83 (reptiles).

Main conclusions

Traits determine most of the variation in population density across species, whereas environmental conditions explain the intraspecific variation across populations. Species traits, resource availability and climatic stability have a different influence on the population density of the four groups. These models can be used to predict the average species population density over large areas and be used to explore macroecological patterns and inform conservation analyses.     

On the validity of Bergmann's rule

Aim We reviewed the occurrence of Bergmann's rule in birds (ninety‐four species) and mammals (149 species), using only studies where statistical significance of the results was tested. We also tested whether studies using different characters as surrogates of body size have a different tendency to conform to Bergmann's rule, whether body size and nest type (in birds) have an influence on the tendency to conform to the rule, and whether sedentary birds conform to the rule more than migratory birds. Location Worldwide. Methods We reviewed published data on geographic and temporal variation in body size, using only studies where the statistical significance of the results was tested. We asked how many species conform to the rule out of all species studied in each order and family. Results Over 72% of the birds and 65% of the mammal species follow Bergmann's rule. An overall tendency to follow the rule occurs also within orders and families. Studies using body mass in mammals show the greatest tendency to adhere to Bergmann's rule (linear measurements and dental measurements show a weaker tendency); while in birds, studies using body mass and other surrogates (linear measurements and egg size) show a similar tendency. Birds of different body mass categories exhibit a similar tendency to follow Bergmann's rule, while in mammals the lower body size categories (4–50 and 50–500 g) show a significantly lower tendency to conform to the rule. Sedentary birds tend to conform to Bergmann's rule more than migratory species. Nest type does not affect the tendency to conform to Bergmann's rule. Main conclusions Bergmann's rule is a valid ecological generalization for birds and mammals.     

Dynamics of phenotypic change: wing length declines in a resident farmland passerine despite survival advantage of longer wings

In many taxa, environmental changes that alter resource availability and energetics, such as climate change and land use change, are associated with changes in body size. We use wing length as a proxy for overall structural body size to examine a paradoxical trend of declining wing length within a Yellowhammer Emberiza citrinella population sampled over 21 years, in which it has been previously shown that longer wings are associated with higher survival rates. Higher temperatures during the previous winter (prior to the moult determining current wing length) explained 23% of wing length decrease within our population, but changes may also be correlated with non‐climatic environmental variation such as changes in farming mechanisms linked to food availability. We found no evidence for within‐individual wing length shrinkage with age, but our data suggested a progressive decline in the sizes of immature birds recruiting to the population. This trend was weaker, although not significantly so, among adults, suggesting that the decline in the sizes of recruits was offset by higher subsequent survival of larger birds post‐recruitment. These data suggest that ecological processes can contribute more than selection to observed phenotypic trends and highlight the importance of long‐term studies for providing longitudinal insights into population processes.     

Brief communication: Hair density and body mass in mammals and the evolution of human hairlessness

Humans are unusual among mammals in appearing hairless. Several hypotheses propose explanations for this phenotype, but few data are available to test these hypotheses. To elucidate the evolutionary history of human “hairlessness,” a comparative approach is needed. One previous study on primate hair density concluded that great apes have systematically less dense hair than smaller primates. While there is a negative correlation between body size and hair density, it remains unclear whether great apes have less dense hair than is expected for their body size. To revisit the scaling relationship between hair density and body size in mammals, I compiled data from the literature on 23 primates and 29 nonprimate mammals and conducted Phylogenetic Generalized Least Squares regressions. Among anthropoids, there is a significant negative correlation between hair density and body mass. Chimpanzees display the largest residuals, exhibiting less dense hair than is expected for their body size. There is a negative correlation between hair density and body mass among the broader mammalian sample, although the functional significance of this scaling relationship remains to be tested. Results indicate that all primates, and chimpanzees in particular, are relatively hairless compared to other mammals. This suggests that there may have been selective pressures acting on the ancestor of humans and chimpanzees that led to an initial reduction in hair density. To further understand the evolution of human hairlessness, a systematic study of hair density and physiology in a wide range of species is necessary. Am J Phys Anthropol 152:145–150, 2013. © 2013 Wiley Periodicals, Inc.     

A refined model of body mass and population density in flightless birds reconciles extreme bimodal population estimates for extinct moa

Flightless birds were once the largest and heaviest terrestrial fauna on many archipelagos around the world. Robust approaches for estimating their population parameters are essential for understanding prehistoric insular ecosystems and extinction processes. Body mass and population density are negatively related for extant flightless bird species, providing a method for quantifying densities and population sizes of extinct flightless species. Here we assemble an updated global data set of body mass and population densities for extant flightless birds and estimate the relationship between these variables. We use generalised least squares models that account for phylogenetic relatedness and incorporate the effects of limiting factors (e.g. habitat suitability) on population density. We demonstrate the applicability of this allometric relationship to extinct species by estimating densities for each of the nine species of moa (Dinornithiformes) and generating a combined spatially explicit map of total moa density across New Zealand. To compare our density estimates with those previously published, we summed individual species' abundances to generate a mean national density of 2.02–9.66 birds km⁻² for low- and high-density scenarios, respectively. Our results reconcile the extreme bimodality of previous estimates (< 2 birds km⁻² and > 10 birds km⁻²) and are comparable to contemporary densities of large herbivorous wild mammals introduced into New Zealand about 150 yr ago. The revised moa density has little effect on the harvest rates required to bring about extinction within 150–200 yr, indicating that rapid extinction was an inevitable response to human hunting, irrespective of the initial population of moa.     

Of mice and wrens: the relation between abundance and geographic range size in British mammals and birds

We examine the relation between population size and geographic range size for British breeding birds and mammals. As for most other assemblages studied, a strong positive interspecific correlation is found in both taxa. The relation is also recovered once the phylogenetic relatedness of species has been controlled for using an evolutionary comparative method. The slope of the relation is steeper for birds than for mammals, but this is due in large part to two species of mammals that have much higher population sizes than expected from their small geographic ranges. These outlying mammal species are the only ones in Britain to be found only on small offshore islands, and so may be exhibiting density compensation effects. With them excluded, the slope of the abundance–range size relation for mammals is not significantly different to that for birds. However, the elevation of the relation is higher for mammals than for birds, indicating that mammals are approximately 30 times more abundant than birds of equivalent geographic range size. An earlier study of these assemblages showed that, for a given body mass, bats had abundances more similar to birds than to non-volant mammals, suggesting that the difference in abundance between mammals and birds might be due to constraints of flight. Our analyses show that the abundance–range size relation for bats is not different for that from other mammals, and that the anomalously low abundance of bats for their body mass may result because they have smaller than expected geographic extents for their size. Other reasons why birds and mammals might have different elevations for the relation between population size and geographic range size are discussed, together with possible reasons for why the slopes of these relations might be similar.     

Examining the Prey Mass of Terrestrial and Aquatic Carnivorous Mammals: Minimum,Maximum and Range

Predator-prey body mass relationships are a vital part of food webs across ecosystems and provide key information for predicting the susceptibility of carnivore populations to extinction. Despite this, there has been limited research on the minimum and maximum prey size of mammalian carnivores. Without information on large-scale patterns of prey mass, we limit our understanding of predation pressure, trophic cascades and susceptibility of carnivores to decreasing prey populations. The majority of studies that examine predator-prey body mass relationships focus on either a single or a subset of mammalian species, which limits the strength of our models as well as their broader application. We examine the relationship between predator body mass and the minimum, maximum and range of their prey''s body mass across 108 mammalian carnivores, from weasels to baleen whales (Carnivora and Cetacea). We test whether mammals show a positive relationship between prey and predator body mass, as in reptiles and birds, as well as examine how environment (aquatic and terrestrial) and phylogenetic relatedness play a role in this relationship. We found that phylogenetic relatedness is a strong driver of predator-prey mass patterns in carnivorous mammals and accounts for a higher proportion of variance compared with the biological drivers of body mass and environment. We show a positive predator-prey body mass pattern for terrestrial mammals as found in reptiles and birds, but no relationship for aquatic mammals. Our results will benefit our understanding of trophic interactions, the susceptibility of carnivores to population declines and the role of carnivores within ecosystems.     

Energetics, body size, and the limits to endothermy

The scaling rate of metabolism with respect to body mass is analysed. Scaling of heat production implies that scaling also exists between temperature regulation and body mass. Most vertebrates follow a Kleiber relation down to a "critical mass, below which the scaling of metabolism must be changed to ensure the maintenance of endothermy. Such an adjustment is found interspecifically in birds and mammals, and is found intraspecifically in mammals during post-natal growth. If the Kleiber scaling relation is maintained below the critical mass, mammals and birds shiR from endothermic temperature regulation (above critical mass) to endothermy with obligatory torpor (below critical mass). If the Kleiber relation is followed to masses far below the critical mass, ectothermy results. Critical mass varies inversely with the level of energy expenditure, which therefore accounts for the fact that most mammals and birds are endotherms and most reptiles and fish are ectotherms. The same relationship permits the facultative endothermy found in some insects and plants.
The scaling relations existing among rate of metabolism, endothermy, and body mass can be written as a modification of the Kleiber relation. This analysis suggests that any organism, irrespective of phylogenetic position, can be endothermic at any body size, if its rate of metabolism is high enough, or can be endothermic with any rate of metabolism, if it is large enough. Consequently, it is difficult to distinguish minimal endothermy from inertial homoiothermy in animals having a large mass. The boundary conditions for effective endothermy are similar to the relationship described between metabolism and mass in the evolution of endothermy through a decrease in mass in the phylogeny of mammals. Even though endothermy may evolve with an increase in mass, its perfection may always require an evolutionary decrease in mass.     

The evolution of body size in birds. II. The role of reproductive power

Given that body mass evolves non-randomly in birds, it is important to ask what factors might be responsible. One suggestion is that the rate at which individuals turn resources into offspring, termed reproductive power, might explain this non-randomness. This is because, in mammals, the body mass with the highest reproductive power is the most common (modal) one. Reproductive power was estimated for birds from data on energetic content of eggs and population productivity. According to the formulation of Brown et al. (1993), reproductive power is composed of two component processes: acquisition (acquiring resources and storing them in reproductive biomass) and conversion (converting reproductive biomass into offspring). As with mammals, estimates of reproductive power indicate that the most common body mass in birds is also the body mass that maximizes reproductive power. The relationship between reproductive power and diversity is different for species smaller than this modal body mass when compared to those that are larger. The relationship of body mass and reproductive power is different between birds and mammals in two ways: (1) the body mass that maximizes reproductive power is smaller in birds (33g) than in mammals (100g), and (2) mammals generate more reproductive power than an equivalent-sized bird. Reproductive power is determined primarily by acquisition in small birds and mammals, while it is determined by conversion in the largest birds and mammals. It is likely that reproductive power is closely tied to the evolution and diversification of body masses because it constrains the ways in which traits affecting fitness can evolve.     

Spatial ecology and conservation of the microendemic ovenbird Cipo Cinclodes (Cinclodes espinhacensis) from the Brazilian highlands

Birds in the genus Cinclodes are habitat specialists, with most restricted to the highlands of South America. The recently described Cipo Cinclodes (C. espinhacensis) is isolated in the southern Espinhaço Range of Brazil and is considered Endangered in Brazil and Near Threatened by the IUCN, but as a subspecies of Long‐tailed Cinclodes (C. pabsti). We examined the population and spatial ecology of Cipo Cinclodes at two geographic scales to improve our understanding of their basic biology and conservation status. We monitored 30 birds at Serra do Breu and found relatively large home ranges (mean = 9.3 ha), a density of paired adults of 0.09/ha, a male‐skewed adult sex ratio (males/total adults = 0.57) due to territories occupied by unpaired males, and long‐term site fidelity. Cipo Cinclodes used all habitat types available in our study area, including rocky outcrops, grasslands, and riparian areas, but habitat selection analyses revealed the importance of riparian areas for foraging and rocky outcrops for nesting. At the species distribution scale, we compiled known and novel recorded occurrence points and used them to calculate the extent of occurrence (EOO) and the area of occupancy (AOO). We used a Maxent species distribution model to generate a binary map to estimate upper limits for EOO (EOO around the model predicted area) and AOO (comprised by the model predicted area within the EOO). We obtained 41 locations, resulting in an EOO of 890.7 km² (up to 1748.7 km²) and an AOO of 100 km² (up to 327.5 km²). The global population is estimated to be between 880 and 2882 birds, which is concerning because small populations are at risk of extinction due to demographic stochasticity, genetic drift, and the interaction of these factors. As such, our results support the designation of Cipo Cinclodes as Endangered on the Brazilian red list.     

Do insect metabolic rates at rest and during flight scale with body mass?

Energetically costly behaviours, such as flight, push physiological systems to their limits requiring metabolic rates (MR) that are highly elevated above the resting MR (RMR). Both RMR and MR during exercise (e.g. flight or running) in birds and mammals scale allometrically, although there is little consensus about the underlying mechanisms or the scaling relationships themselves. Even less is known about the allometric scaling of RMR and MR during exercise in insects. We analysed data on the resting and flight MR (FMR) of over 50 insect species that fly to determine whether RMR and FMR scale allometrically. RMR scaled with body mass to the power of 0.66 (M0.66), whereas FMR scaled with M1.10. Further analysis suggested that FMR scaled with two separate relationships; insects weighing less than 10mg had fourfold lower FMR than predicted from the scaling of FMR in insects weighing more than 10mg, although both groups scaled with M0.86. The scaling exponents of RMR and FMR in insects were not significantly different from those of birds and mammals, suggesting that they might be determined by similar factors. We argue that low FMR in small insects suggests these insects may be making considerable energy savings during flight, which could be extremely important for the physiology and evolution of insect flight.