Research Article |
Corresponding author: Valery M. Gavrilov ( vmgavrilov@mail.ru ) Academic editor: George Sangster
© 2023 Valery M. Gavrilov, Tatiana B. Golubeva, Andrey V. Bushuev.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gavrilov VM, Golubeva TB, Bushuev AV (2023) Metabolic rate, sleep duration, and body temperature in evolution of mammals and birds: the influence of geological time of principal groups divergence. ZooKeys 1148: 1-27. https://doi.org/10.3897/zookeys.1148.93458
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This study contains an analysis of basal metabolic rate (BMR) in 1817 endothermic species. The aim was to establish how metabolic scaling varies between the main groups of endotherms during evolution. The data for all the considered groups were combined and the common exponent in the allometric relationship between the BMR and body weight was established as b = 0.7248. Reduced to the common slope, the relative metabolic rate forms the following series: Neognathae – Passeriformes – 1.00, Neognathae – Non-Passeriformes – 0.75, Palaeognathae – 0.53, Eutheria – 0.57, Marsupialia – 0.44, and Monotremata – 0.26. The main finding is that the metabolic rate in the six main groups of mammals and birds consistently increases as the geological time of the group’s divergence approaches the present. In parallel, the average body temperature in the group rises, the duration of sleep decreases and the duration of activity increases. BMR in a taxon correlates with its evolutionary age: the later a clade diverged, the higher is its metabolic rate and the longer is its activity period; group exponents decrease as group divergence nears present times while with increase metabolic rate during activity, they not only do not decrease but can increase. Sleep duration in mammals was on average 40% longer than in birds while BMR, in contrast, was 40% higher in birds. The evolution of metabolic scaling, body temperature, sleep duration, and activity during the development of endothermic life forms is demonstrated, allowing for a better understanding of the underlying principles of endothermy formation.
Activity, basal metabolic rate, birds, body temperature, mammals, phylogeny, scaling, sleep duration, time of divergence
The aim was to verify the Bennett and Ruben hypothesis that the level of BMR determines the duration of activity of an animal. We wished to consider the increased duration of activity in evolution, and establish how this relates to BMR levels. Duration activity obviously varies between the groups of endotherms, how do we show the change in activity duration in different groups of animals in evolution? We suggest, in general terms, to determine the duration of activity in endothermic animals through the duration of sleep. Overall, the duration of sleep is inversely related to the overall activity, i.e., activity duration equals 24 hours minus sleep duration. Sleep (at least in endotherms) is a natural recurrent state in animals that decreases sensory activity, inhibits most voluntary muscle activity, and causes low levels of interaction with the environment (
We hypothesised that the above groups of endothermic animals which evolved in different geological times should exhibit varying BMRs, body temperature and sleep duration. The timescale is important because it enables the comparison of phylogenesis directly with the evolution of other organisms, and with the events of planetary histories, such as geological or climatic events. Revolutionary advances in molecular biology have permitted the estimation of times of taxon divergence on the basis of molecular clocks. After analysing these data, we found the most realistic the geological time divergence of these groups from the main trunk of vertebrates (see
Basal metabolic rate (BMR) is the minimal metabolic rate of endotherms. Comparative studies in animal energetics are based on the allometric equation between BMR and body mass (m):
BMR = amb or log BMR = log (a) + b * log (m)
where b is a scaling exponent (the slope of the regression line) and a is the allometric coefficient (antilog of the regression intercept). Analysis of studies on shifts in metabolic scaling across different evolutionary transitions in both exponential (
Serious difficulties in discussing the functional meaning of the allometric coefficient a in different groups of animals are created by its strong dependence on the units of measurement and, especially, on the scaling exponent, which varies greatly between taxa, with respect to both different and the same taxonomic rank. In this study, we aimed to develop an effective way to compare BMR across groups, regardless of body size. We believe that the dimensionless ratio of BMR in different clades of endothermic animals will assist comparative studies. It can be obtained if the BMR dependencies on m are held to a common scaling index. We conducted a statistical test of the hypothesis that all six groups can be calculated with one common exponent. The inclusion of such a test is crucial, because only then is it possible to establish a dimensionless BMR ratio for the main selected groups of birds and mammals.
In this paper, we used a macroevolutionary perspective to look for the tipping points at which the principal clades of endotherms transitioned to new evolutionary allometric scaling relationships. We analysed in detail the parameters of metabolic scaling in six groups of endotherms: Monotremata, Marsupialia and Eutheria from mammals, and Palaeognathae, Neognathae-Non-Passeriformes and Neognathae-Passeriformes from birds. We then aimed to establish the dimensionless BMR ratio for the main groups of birds and mammals. We correlated this dimensionless relationship with the divergence time of the group and other parameters.
We compared the regressions obtained by the phylogenetic method of generalised least squares (PGLS) with the regressions obtained using ordinary least squares (OLS), to determine how phylogeny does affect the dimensionless BMR ratio for the main groups of birds and mammals.
We calculated BMR in the main groups of endotherms from the data published by
To correlate BMR levels and body temperature (TB), we used Clarke’s data (table 10.2,
We expect that the duration of activity varies between the main groups of endotherms. Overall sleep duration is inversely related to the overall activity duration. We calculated sleep duration in the main groups of endotherms from the data in the work of
In spite of some inconsistencies between the palaeontological record and molecular evidence, the sequence of emergence of the extant groups of endotherms may be presented as follows: monotremes 271 mya (
The body mass and BMR data were log10-transformed before analysis to account for allometric scale. All scaling exponents in allometric equations that are used in our study, were based on ordinary least squares (OLS), weighted least squares (WLS) or phylogenetic generalised least squares (PGLS) regressions of log10 (BMR) ~ log10 (m). Because our sample of different endothermic groups varied greatly in the number of species, in some cases we augmented the OLS analysis with WLS by incorporating the sample size information into the regression. Both OLS and WLS regressions were conducted using the basic R function ‘lm’ (
We applied a test of the homogeneity of the slopes, which in our case tests the null hypothesis H0: b1 = b2 = b3 = b4 = b5 = b6. Using the R lm procedure (
The avian phylogeny was extracted from the birdtree.org database (http://www.birdtree.org) using the study by
We calculated the allometric equations for BMR of all major groups of mammals and birds. The regression lines in these groups differ slightly in slope, but differ significantly in intercepts (Fig.
Parameters of allometric equation for basal metabolic rate in principal groups of endothermic animals obtained from OLS analyses.
Group | Number of species | Body mass range, g | OLS: a ± SE | OLS: b ± SE | OLS: R2 |
---|---|---|---|---|---|
Mammalia | 817 | 2.2–4037500 | 3.248±0.107 | 0.735±0.006 | 0.956 |
Monotremata | 3 | 1284–10300 | 5.861± 0.512 | 0.565±0.387 | 0.681 |
Marsupialia | 84 | 5.4–32490 | 2.300±0.152 | 0.753±0.011 | 0.983 |
Eutheria | 730 | 2.2–4037500 | 3.326±0.115 | 0.736±0.006 | 0.956 |
Aves | 1000 | 2.8–92400 | 7.435±0.167 | 0.648±0.005 | 0.940 |
Palaeognathae | 9 | 220.8–92400 | 3.221±1.147 | 0.727±0.041 | 0.978 |
Non-Passeriformes | 404 | 3.2–23370 | 5.507±0.262 | 0.691±0.009 | 0.939 |
Passeriformes | 587 | 5.1–1203 | 7.379±0.256 | 0.668±0.010 | 0.871 |
Parameters of allometric equation for basal metabolic rate in principal groups of endothermic animals obtained from PGLS analysis.
Group | Pagel’s λ | PGLS: a ± SE | PGLS: b ± SE | PGLS: R2 |
---|---|---|---|---|
Mammalia | 0.870 | 2.357±0.632 | 0.735±0.009 | 0.888 |
Monotremata | 0.000 | 5.861±NA | 0.565±0.387 | 0.681 |
Marsupialia | 0.214 | 2.407±0.222 | 0.746±0.013 | 0.976 |
Eutheria | 0.813 | 2.910±0.393 | 0.733±0.011 | 0.874 |
Aves | 0.664 | 5.514±0.605 | 0.679±0.010 | 0.830 |
Palaeognathae | 0.000 | 3.221±0.871 | 0.727±0.041 | 0.978 |
Non-Passeriformes | 0.630 | 4.833±0.589 | 0.708±0.014 | 0.865 |
Passeriformes | 0.443 | 7.818±0.610 | 0.642±0.014 | 0.780 |
It is noteworthy that we are not aiming at revisiting numerous previous research works concerning the effect of phylogeny on metabolic scaling. We are interested in how the results obtained by regressions PGLS and OLS are reflected in the dimensionless ratio BMR, in groups and on scaling options. Here, we present comparisons of slopes and intercepts from both OLS and PGLS regressions of log (BMR) ~ log (m), obtained for several major groups of endotherms.
Using OLS, coefficient a was significantly different for two of the vertebrate classes (p < 0.001); the slopes of the regression lines also differed (p < 0.001). Using PGLS, for birds and mammals, both a and b were also significantly different (p < 0.001).
For all major groups, the calculation using PGLS had more influence on a, the allometric coefficient, than on the slopes of the regression lines. Paleognathae and Neognathae did not differ in the intercepts of PGLS regressions, but rather they differed at the tendency level (i.e., tending to differ): neognaths tend to have higher BMR. The same can be said about the differences between Passeriformes and non-Passeriformes: the former tend to have higher BMR (more precisely, they tend to differ) and the BMR of Passeriformes is higher.
Pagel’s lambda is a potential measure of “phylogenetic signal”, the extent to which correlations in common traits reflect their shared evolutionary history. The effect of phylogeny was greater for both a and b, if the value of Pagel’s λ for the group was greater. The effect of lambda on scaling options is as follows:
aOLS-aPGLS = – 0.591+ 1.8993 λ, R2 = 0.3113;
bOLS-bPGLS = 0.0504-0.0497 λ, R2 = 0.1397.
With an increase in lambda, the difference in the allometric coefficient a between the OLS and PGLS measurements increases, while the difference in exponent b decreases. The higher the value of Pagel’s λ for the group, the more its effect of phylogeny was greater for this group (Table
We determined that of the three models, according to the BIC criterion, the best model is the one with a common slope and separate intercepts. To obtain the ability to compare the BMR in different groups, we recalculated the equations transformed with a common average b = 0.7248 using the standard OLS procedure.
Reduced to the common slope b = 0.7248, the value of a gradually and understandably increases from Monotremata to Passeriformes, and it is between 1.63 for Monotremata and 6.18 for Passeriformes (Table
Changes in allometric coefficient a when reduced to common slope b = 0.7248 ± 0.0039.
Group | a at original scaling exponent (slope) | R2 | a at common slope b = 0.7248 | R2 | a/aPasseriformes, BMR ratio |
---|---|---|---|---|---|
Monotremata | 5.861 | 0.681 | 1.63 | 0.665 | 0.264 |
Marsupialia | 2.3 | 0.983 | 2.69 | 0.980 | 0.435 |
Eutheria | 3.326 | 0.956 | 3.53 | 0.956 | 0.571 |
Paleognathae | 3.221 | 0.940 | 3.29 | 0.978 | 0.532 |
Non-Passeriformes | 5.507 | 0.9395 | 4.65 | 0.939 | 0.752 |
Passeriformes | 7.379 | 0.79070 | 6.18 | 0.871 | 1.000 |
A the allometric coefficient in six groups of endothermic animals, depending on the geological time of divergence of the clades: by the phylogenetic generalised least squares (PGLS) and ordinary least squares (OLS). a (mlO2/h) is the allometric coefficient (antilog of the regression intercept) B the exponents in six groups of endothermic animals, depending on the geological time of divergence of the clades: by the phylogenetic generalised least squares (PGLS) and ordinary least squares (OLS). b is a scaling exponent (the slope of the regression line).
The analyses of metabolic scaling in the six groups illustrate that the variation in metabolic scaling relationships is systematically related to metabolic level. Metabolic scaling in the main groups of endothermic animals correlates with their evolutionary age: the younger the group is, the higher is its metabolic rate, but it increases more slowly with increasing body weight (Fig.
BMR increases in evolutionarily younger groups (see also Table
A the allometric coefficient a (mlO2/h) in different groups, is formulated by recalculating the equations and transforming with a common average b = 0.7248, depending on the geological time of divergence of the clades B BMR ratio in the six principal groups of endothermic animals are expressed in relation to BMR of Passeriformes (a non-dimensional coefficient of the intercept), dependent on the time of appearance in evolution.
Applying regressions with a common slope b = 0.7248 sharply increase both R2 and the reliability of the regressions, to a higher degree when using the dimensionless BMR ratio than in units of ml O2/h per g (Fig.
We performed a meta-analysis of sleep duration in the main taxa of endotherms (Fig.
A sleep duration in the six main groups of endothermic animals depending on the geological time of appearance of the group in evolution. Regression lines and statistics in the figure were calculated using the OLS method. Using the weighted least squares (WLS) method, p = 0.01548 B activity duration (24 minus sleep duration) as a function of geological time since taxon’s divergence, in mya, in different groups of endotherms. Regression lines and statistics in the figure were calculated using the OLS method. Using the weighted least squares (WLS) method, p = 0.01548. The total duration of sleep is an indicator of the reverse value of total activities.
The size-corrected BMR in the six groups of endothermic animals is correlated with the duration of their activity, with this parameter increasing from monotremes to Passeriformes (Fig.
The greater the proportion of the day when animals are active, the higher their BMR is. Placental mammals and palaeognath birds exhibit similar BMR; i.e., terrestrial animals that do not fly have the same BMR level and the same duration of activity.
In this study, we established correlations of various biological traits, with the development of the metabolic level adjusted for body-size effects, by using an analysis of covariance. We determined the effect by comparing the elevations (indicated by Y-intercepts) of multiple size-scaling relationships, which are preliminarily brought to a common slope. This statistical method thus allows comparison of metabolic rate using the allometric coefficient “a” among different groups, which is reasonable since exponent “b” is fixed. We also found the dimensionless ratio of BMP levels in these groups. We found correlations of these levels with the average body temperature at group level; the relationship between the level BMP group and sleep duration; the relationship between levels of BMP in different taxa with the time of their divergence from the main clade of vertebrates; and the relationship between the metabolic levels and duration of sleep-in different groups. Now, we intend to look at how the level of BMR relates to the state of these groups at the present time. We correlate BMR levels and body temperature with known events in the history of the Earth.
Endothermic taxa with higher BMR contain more species. Palaeognaths include 57 extant species. Passeriformes are the largest order of birds and include ca. 6000 species or 60% of the 10,000 extant avian species. The body sizes of passerines vary from the Raven (Corvus corax), whose mass can reach 1.5 kg, to the tiny Short-tailed pygmy tyrant (Myiornis ecaudatus) (4.2 g). However, most birds from this order have body mass between 10 and 80 g, and a mean passerine is smaller than a mean bird of any other order. Neognath non-passerines include ca. 4000 species of 25 orders (
Most extant mammals (99%) belong to the subclass Theria, which consists of 5136 eutherian species and 346 marsupial species. Just five species of monotremes survive today (
We combined data on ectotherms and endotherms using our data from previous works (
How taxon divergence geological time and metabolic rate affect the metabolic scaling in nine groups of vertebrates, the number of extant species in the taxon and their size ranges.
Taxa | Taxon divergence geological time, MYA | a at common slope, mlO2/h for m = 1 g | b | Number of extant species | Body mass range of extant species, g |
---|---|---|---|---|---|
Fishes | 465 | 0.264 | 0.88 | 25000 | 0.004–12000000 |
Amphibia | 365 | 0.366 | 0.88 | 4000 | 0.15–70000 |
Reptilia | 322 | 0.398 | 0.76 | 8000 | 0.14–2000000 |
Monotremata | 217 | 1.63 | 4 | 2000–10000 | |
Marsupialia | 193 | 2.69 | 0.75 | 346 | 20–100000 |
Eutheria | 115 | 3.53 | 0.74 | 5136 | 1.8–150000000 |
Palaeognathae | 110 | 3.29 | 0.73 | 57 | 30–156000 |
Non-Passeriformes | 90 | 4.65 | 0.69 | 4000 | 1.95–40000 |
Passeriformes | 50 | 6.18 | 0.67 | 6000 | 4.2–1500 |
May we emphasise that the patterns of metabolic scaling found by us for endotherms (
As for the number of species in the taxon and their size range, there is no distinct correlation with the time of divergence. The oldest group, fish, have the highest species diversity, the number of species in this paraphyletic group of vertebrate aquatic jawed animals, which differ in gill breathing throughout the postembryonic development of the organism, is comparable to the number of species in other groups of vertebrates together. They are distinguished by the largest size range and the smallest sizes among vertebrates (Table
From our point of view, the acquisition of endothermy increases the minimum size of animals (apparently only in animals of ~ 2 g and above endothermy can be established), while the maximum sizes are controlled by other factors (for example, flying birds obey the rules of aerodynamics). Variation in the scaling exponent (slope) was caused primarily by the fact that mammal and bird datasets included species from the six major groups in various proportions. Different slopes appear due to different BMR values across taxonomic groups, sample sizes, range of body mass across taxa, and different representations of species within each group. For example, the overall slope for all birds, b = 0.674, is obtained due to the highest BMR of passerines, which are concentrated in the lower part of the size range, and the lowest BMR of palaeognaths, which form the upper part of the size range. In addition, the higher the exponent of body weight (m) in the BMR equation, the larger the range of sizes group will be. In this case, the lack of forces for movement (which grows ∝ m0.83) will not be a size limiter. The size range in mammals with an empirical exponent for body weight in the equation for BMR b = 0.735 is two orders of magnitude higher than in birds with an exponent of 0.668. If we consider our results in the broader context of the metabolic optimum of life (
To maintain a constant body temperature, endothermic birds and mammals must maintain a balance between heat production and dissipation. Heat dissipation is the main thermodynamic problem that animals need to solve. To lose the heat produced during activity, the animals need efficient mechanisms based on a well-developed blood circulation system, and the ability to regulate thermal isolation of the skin. Body temperature of extant birds and mammals is similar, which is explained by the thermal conditions that are typical on Earth, protein properties and thermal optimum of biochemical reactions. We suggest that it was not until the mid-Cretaceous that birds and mammals, with a modern level of BMR, formed. Apparently, only a particular level of BMR makes it possible to maintain body temperature that is necessary for homeothermic endothermy. It should be emphasised that
We suggest that Triassic and Jurassic mammals and birds, whose fossil remains are fragmentarily known, did not possess fully developed endothermy, and in this regard were similar to monotremes or those even less advanced. The Mesozoic was the warmest and most stable period in the history of the Earth (
We believe that during the entire Mesozoic era, various clades of mammals and birds evolved, in which homeothermic endothermy developed. In these clades, morpho-physiological traits originated that led to the development of full endothermy. It should be emphasised that all extant orders of birds and mammals evolved after the Cretaceous thermal maximum (
The BMR level in the group correlates with the duration of activity. The BMR level and duration of the group’s activity increase as the geological time of the group’s appearance approaches the present time. Since
Maximum energy consumption has also been studied in birds and mammals. In mammals, maximal oxygen consumption during exercise MMR (ml O2 hour–1) = 17.187m0.8598 R2 = 0.691 (
Other well-documented levels of energy expenditure have been reported in birds. This is averages daily energy expenditure of caged birds for long-term sustained rate of biological activity and the expenditure of energy in flight. In this case, we can compare the ratio of energy expenditure for these activities in passerine and non-passerine birds. Further, the existing energy averages daily energy expenditure of caged birds for long-term sustained rate of biological activity and the expenditure (
We suggest that the advent of angiosperms triggered the rapid development of endothermic animals, i.e., animals are able to maintain at a certain metabolic rate for sustained homeothermy. This process, endothermisation of vertebrates, took place in the branches leading to the mammalian and avian clades (Gavrilov, 2013). Angiosperms evolved in the early Cretaceous (
It was not until the mid-to-late Cretaceous that birds and mammals started to play the leading roles in the communities that they have at present. Birds lost their teeth in the late Cretaceous, and the existing mammals already included both marsupials and placentals. These developments were related to the general evolution of the organic world, which caused the advent of flowering plants in the Mesozoic, along with the related fauna of invertebrates and a high diversity of flying insects. Only the development of angiosperms and the related diversity of invertebrates provided a sufficient food base to supply for the development of endothermic animals. Angiosperms and insects provided food resources that were necessary for endotherms but were not suitable for many reptiles adapted to the utilisation of Mesozoic flora and fauna, which enabled the ecological expansion of endotherms. Birds and mammals replaced reptiles in many mainstream ecological niches, adapted to various habitats and quickly expanded to a broad range of size classes (mammals – eight orders of magnitude by size, birds – six orders of magnitude). This development was further facilitated by the cooling of the planet at that time.
Endothermy has formed in birds and mammals independently and in different morpho-physiological bases. However, in both groups, endothermy originated as an effect of selection for aerobic metabolism improvement, that provided a higher level of activity. As a result, these groups developed a basal metabolic rate that increased over time. The advantages of having a high and stable body temperature, which is inevitably related to metabolism intensification, led to the development of thermoregulatory adaptations, such as fur and feathers. Thus, metabolic heat production may be retained and heat absorption may be reduced in hot environments. The emergence of endothermy with an aerobic supply of motion activity, which regulates the level of metabolism and heat loss, has created many opportunities for endothermic animals. Achieving such a level of energy utilisation has allowed these animals to maintain activity for a long period, whereas its sensory support led to the complication and diversification of the behavioural repertoire of birds and mammals. All of these adaptations lead to the penetration of animals into those places on the planet that were previously unsuitable for them.
Our analysis of the most complete data on standardised energetic costs of endothermic animals, available from the literature based on 1817 measurements of species (817 data points from mammals and 1000 data points from birds). We analysed factors that are responsible for energy expenditure by birds and mammals, and look for important life history aspects that are common for a large number of species. Ecological and behavioural factors that govern BMR in birds and mammals are nearly identical, even though each of the two clades possessed some specific features that influenced their energetics, including evolution of endothermy and flight in birds, and endothermy and viviparity in mammals. We used these data to estimate scaling coefficients and intercepts for the main groups. In all groups, BMR varies with body size but with significantly different intercepts. We provide two ways to compare BMR levels, regardless of body size. We determined the common scaling slope for all groups, recalculated the original data with this slope, and obtained the new intercept values, which were then correlated with the intercept to passerines that have the highest BMR and obtained the dimensionless BMR ratio for the selected groups. BMR in groups increase and group exponents decrease as group divergence nears present times while with increase metabolic rate during activity, group scaling exponents not only do not decrease but can increase. We considered BMR variation and the duration of activity in three mammalian subclasses: monotremes, marsupials and eutherians, and in three groups of birds: Palaeognathae, non-Passeriformes and Passeriformes, depending on the evolutionary age of these groups. Activity duration varies between the main groups of endotherms. A high level of activity is related to high BMR. Eutheria and Palaeognathes have similar BMR, i.e., terrestrial lifestyle without flight is based on nearly equal BMR and these groups evolved at practically almost the same time. We calculated sleep duration in the main groups of endotherms on the basis of data from the literature. BMR in a taxon correlates with its evolutionary age: the later a clade diverged, the higher is its metabolic rate and the longer is its activity period. We suggest that each group formed its taxon-specific BMR, depending on the ability to maintain thermal homeostasis under the environmental conditions that prevailed during its emergence. Monotremes were the first to branch off from the basal mammals and have the lowest BMR and the lowest TB. The next level allows marsupials to maintain thermal homeostasis under a broader range of conditions, and have a more protracted period of activity. Finally, the metabolic rate and TB typical of placentals and Palaeognathae formed in the mid Cretaceous, and allowed these groups to occupy a broader range of terrestrial niches. Immediately when the development of blood circulation and respiratory systems made it possible to reach the BMR that allowed maintaining TB of 37 °C, the explosive radiation of mammals and birds started. In the mid and late Cretaceous, birds and mammals started to occupy the leading positions in the ecosystems. Further, at last, some 50 mya passerines that have the highest BMR (nearly 50% higher than eutherians and palaeognath birds), adapted to the forest habitats and gained TB of ca. 40 °C, which is at the upper physiological limit. The duration of activity grew in parallel to the BMR. Ecological expansion of birds and mammals resulted in their worldwide geographic distribution.
We are grateful to the researchers of the Department of Vertebrate Zoology and to its head, L.P. Korzun, for the opportunity to perform this research and for assistance and helpful discussions. We are extremely grateful to N.V. Zelenkov and A.P. Rasnitsyn for valuable critical comments and consultations on issues of paleontology. We thank E.N. Solovyeva for the construction of the avian phylogenetic tree. Our special thanks also go to N.S. Chernetsov for giving time and energy to the translation into English, and the reviewer A. Makarieva for providing extensive and constructive comments. We are extremely grateful to Dr George Sangster, Subject Editor, for his amiability and patience with our manuscript. We would like to thank A.I. Shilov for help with the Figures. This study was supported by RSF-FWO grant #20-44-01005.
Mammalian Basal metabolic rate (BMR) dafrom Genoud et al. (2017)
Data type: table (excel document)
Aves BMR
Data type: table (PDF file)
Sleep duration for endothermic species (by review Cambell & Tobler, 1984) and others
Data type: table (PDF file)