Abstract We assessed water-body use by elephants through monitoring elephant signs around them. E... more Abstract We assessed water-body use by elephants through monitoring elephant signs around them. Elephant footprints and dung piles were recorded at 25 water bodies fortnightly for one year. Elephants preferred perennial water bodies and avoided those with temporary human dwellings. Human activities did not significantly affect elephant use of water bodies, suggesting low incidence of activities and behavioral adaptation to them by elephants. Elephant signs at perennial water bodies increased in the dry season.
This paper presents spatiotemporal gait parameters of arboreal locomotion in the colobine< i> Rhi... more This paper presents spatiotemporal gait parameters of arboreal locomotion in the colobine< i> Rhinopithecus bieti</i> in its natural habitat. While adult females used exclusively either extended-elbow vertical climbing or pulse climbing, the much larger adult males preferred the less demanding flexed-elbow vertical climbing on thin trees or on trunks with handholds.
Abstract Flexion/extension moment arms were obtained for the major muscles crossing the hip, knee... more Abstract Flexion/extension moment arms were obtained for the major muscles crossing the hip, knee and ankle joints in the orang-utan, gibbon, gorilla (Eastern and Western lowland) and bonobo. Moment arms varied with joint motion and were generally longer in proximal limb muscles than distal limb muscles. The shape of the moment arm curves (ie the plots of moment arm against joint angle) differed in different hindlimb muscles and in the same muscle in different subjects (both in the same and in different ape species).
The human brain stands out among mammals by being unusually large. The expensive-tissue hypothesi... more The human brain stands out among mammals by being unusually large. The expensive-tissue hypothesis 1 explains its evolution by proposing a trade-off between the size of the brain and that of the digestive tract, which is smaller than expected for a primate of our body size. Although this hypothesis is widely accepted, empirical support so far has been equivocal. Here we test it in a sample of 100 mammalian species, including 23 primates, by analysing brain size and organ mass data.
Abstract We present quantitative data on the hindlimb musculature of Pan paniscus, Gorilla gorill... more Abstract We present quantitative data on the hindlimb musculature of Pan paniscus, Gorilla gorilla gorilla, Gorilla gorilla graueri, Pongo pygmaeus abelii and Hylobates lar and discuss the findings in relation to the locomotor habits of each. Muscle mass and fascicle length data were obtained for all major hindlimb muscles. Physiological cross-sectional area (PCSA) was estimated. Data were normalized assuming geometric similarity to allow for comparison of animals of different size/species.
The processes that underlie the formation of the dominance hierarchy in a group are since long un... more The processes that underlie the formation of the dominance hierarchy in a group are since long under debate. Models of self-organisation suggest that dominance hierarchies develop by the self-reinforcing effects of winning and losing fights (the so-called winner-loser effect), but according to 'the prior attribute hypothesis', dominance hierarchies develop from pre-existing individual differences, such as in body mass.
Abstract Quantitative, accurate data regarding the inertial properties of body segments are of pa... more Abstract Quantitative, accurate data regarding the inertial properties of body segments are of paramount importance when developing musculo-skeletal locomotor models of living animals and, by inference, their ancestors. The limited number of available primate cadavers, and the destructive nature of the post-mortem, result in such data being very rare for primates. This study builds on the work of Crompton et al.(Am. J. Phys. Anthropol.
Summary The choice of an appropriate method for fitting a straight line to data is one of the maj... more Summary The choice of an appropriate method for fitting a straight line to data is one of the major procedural problems in bivariate allometric analysis. Commonly used techniques, such as ordinary least-squares regression and major axis, are derived from the general structural relationship model and thus require some knowledge about the distribution of the data. In this paper, we explore a nonparametric alternative, referred to as “rotation method”, which involves no assumptions about error distributions.
Abstract The maximum capability of a muscle can be estimated from simple measurements of muscle a... more Abstract The maximum capability of a muscle can be estimated from simple measurements of muscle architecture such as muscle belly mass, fascicle length and physiological cross-sectional area. While the hindlimb anatomy of the non-human apes has been studied in some detail, a comparative study of the forelimb architecture across a number of species has never been undertaken.
Humans stand out among the apes by having both an extremely large brain and a relatively high rep... more Humans stand out among the apes by having both an extremely large brain and a relatively high reproductive output, which has been proposed to be a consequence of cooperative breeding. Here, we test for general correlates of allomaternal care in a broad sample of 445 mammal species, by examining life history traits, brain size, and different helping behaviors, such as provisioning, carrying, huddling or protecting the offspring and the mother.
For over a century, various neuroanatomical measures have been employed as assays of cognitive ab... more For over a century, various neuroanatomical measures have been employed as assays of cognitive ability in comparative studies. Nevertheless, it is still unclear whether these measures actually correspond to cognitive ability. A recent metaanalysis of cognitive performance of a broad set of primate species has made it possible to provide a quantitative estimate of general cognitive ability across primates. We find that this estimate is not strongly correlated with neuroanatomical measures that statistically control for a possible effect of body size, such as encephalization quotient or brain size residuals. Instead, absolute brain size measures were the best predictors of primate cognitive ability. Moreover, there was no indication that neocortex-based measures were superior to measures based on the whole brain. The results of previous comparative studies on the evolution of intelligence must be reviewed with this conclusion in mind.
In the ongoing discussion about brain evolution in vertebrates, the main interest has shifted fro... more In the ongoing discussion about brain evolution in vertebrates, the main interest has shifted from theories focusing on energy balance to theories proposing social or ecological benefits of enhanced intellect. With the availability of a wealth of new data on basal metabolic rate (BMR) and brain size and with the aid of reliable techniques of comparative analysis, we are able to show that in fact energetics is an issue in the maintenance of a relatively large brain, and that brain size is positively correlated with the BMR in mammals, controlling for body size effects. We conclude that attempts to explain brain size variation in different taxa must consider the ability to sustain the energy costs alongside cognitive benefits.
The correlation between brain size and life history has been investigated in many previous studie... more The correlation between brain size and life history has been investigated in many previous studies, and several viable explanations have been proposed. However, the results of these studies are often at odds, causing uncertainties about whether these two character complexes underwent correlated evolution. These disparities could arise from the mixture of wild and captive values in the datasets, potentially obscuring real relationships, and from differences in the methods of controlling for phylogenetic non independence of species values. This paper seeks to resolve these difficulties by proposing an overarching hypothesis that encompasses many of the previously proposed hypotheses, and (2) testing the predictions of this hypothesis using rigorously compiled data and utilizing multiple methods of analysis. We hypothesize that the adaptive benefit of increased encephalization is an increase in reproductive lifespan or efficiency, which must be sufficient to outweigh the costs due to growing and maturing the larger brain. These costs and benefits are directly reflected in the length of life history stages. We tested this hypothesis on a wide range of primate species. Our results demonstrate that encephalization is significantly correlated with prolongation of all stages of developmental life history except the lactational period, and is significantly correlated with an extension of the reproductive lifespan. These results support the contention that the link between brain size and life history is caused by a balance between the costs of growing a brain and the survival benefits the brain provides. Thus, our results suggest that the evolution of prolonged life history during human evolution is caused by increased encephalization.
American journal of physical anthropology, Jan 1, 2005
Vertical climbing has played an important role in theories about the evolution of habitual bipeda... more Vertical climbing has played an important role in theories about the evolution of habitual bipedalism in early hominids and of locomotor specialization in hominoids. However, empirical data on vertical climbing in nonhuman primates are scarce, especially regarding kinematics. In this paper, the kinematics of flexedelbow vertical climbing of four hominoid species are reported: western lowland gorillas (Gorilla gorilla gorilla), bonobos (Pan paniscus), Sumatran orangutans (Pongo pygmaeus abelii), and yellow-cheeked crested gibbons (Hylobates concolor gabriellae). The climbing sequences were recorded simultaneously by four digital video cameras in European zoos. Analyzed parameters include intersegmental flexion/extension angles of the shoulder, elbow, hip, and knee joints, horizontal abduction/adduction angles of the upper arm and thigh, and distance of the body
Costs and benefits of encephalization are a major topic of debate in the study of primate and hum... more Costs and benefits of encephalization are a major topic of debate in the study of primate and human evolution. Comparative studies provide an opportunity to test the validity of a hypothesis as a general principle, rather than it being a special case in primate or hominid evolution. If a population evolves a larger brain, the metabolic costs of doing so must be paid for by either an increased energy turnover (direct metabolic constraint) or by a trade-off with other energetically expensive costs of body maintenance, locomotion, or reproduction, here referred to as the energy trade-off hypothesis, an extension of the influential Expensive Tissue Hypothesis of Aiello and Wheeler (1995, Curr. Anthropol. 36, 199e 221). In the present paper, we tested these hypotheses on birds using raw species values, family means, and independent contrasts analysis to account for phylogenetic influences. First, we tested whether basal metabolic rates are correlated with brain mass or any other variable of interest. This not being the case, we examined various trade-offs between brain mass and the mass of other expensive tissues such as gut mass, which is approximated by gut length or diet quality. Only weak support was found for this original Expensive Tissue Hypothesis in birds. However, other energy allocations such as locomotor mode and reproductive strategy may also be reduced to shunt energy to an enlarged brain. We found a significantly negative correlation between brain mass and pectoral muscle mass, which averages 18% of body mass in birds and is indicative of their relative costs of flight. Reproductive costs, on the other hand, are positively correlated with brain mass in birds. An increase in brain mass may allow birds to devote more energy to reproduction, although not through an increase in their own energy budget as in mammals, but through direct provisioning of their offspring. The trade-off between locomotor costs and brain mass in birds lets us conclude that an analogous effect could have played a role in the evolution of a larger brain in human evolution.
To explain variation in relative brain size among homoiothermic vertebrates, we propose the Expen... more To explain variation in relative brain size among homoiothermic vertebrates, we propose the Expensive Brain hypothesis as a unifying explanatory framework. It claims that the costs of a relatively large brain must be met by any combination of increased total energy turnover or reduced energy allocation to another expensive function such as digestion, locomotion, or production (growth and reproduction). Focusing on the energetic costs of brain enlargement, a comparative analysis of the largest mammalian sample assembled to date shows that an increase in brain size leads to larger neonates among all mammals and a longer period of immaturity among monotokous precocial species, but not among the polytokous altricial ones, who instead reduce their litter size. Relatively large brained mammals, altricial and precocial, also show reduced annual fertility rates as compared to their smaller brained relatives, but allomaternal energy inputs allow some cooperatively breeding altricial carnivores to produce even more offspring in a shorter time despite having a relatively large brain. Thus, the Expensive Brain framework explains why brain size is linked to life history pace in some, but not all mammalian lineages. This framework encompasses other hypotheses of energetic constraints on brain size variation and is also compatible with the Brain Malnutrition Risk hypothesis, but the absence of a mammal-wide correlation between brain size and immature period argues against the Needing-to-Learn explanation for slower development among large brained mammals.
The expensive brain hypothesis predicts an interspecific link between relative brain size and lif... more The expensive brain hypothesis predicts an interspecific link between relative brain size and life-history pace. Indeed, animals with relatively large brains have reduced rates of growth and reproduction. However, they also have increased total lifespan. Here we show that the reduction in production with increasing brain size is not fully compensated by the increase in lifespan. Consequently, the maximum rate of population increase (r max ) is negatively correlated with brain mass. This result is not due to a confounding effect of body size, indicating that the wellknown correlation between r max and body size is driven by brain size, at least among homeothermic vertebrates. Thus, each lineage faces a 'grey ceiling', i.e. a maximum viable brain size, beyond which r max is so low that the risk of local or species extinction is very high. We found that the steep decline in r max with brain size is absent in taxa with allomaternal offspring provisioning, such as cooperatively breeding mammals and most altricial birds. These taxa thus do not face a lineage-specific grey ceiling, which explains the far greater number of independent origins of large brain size in birds than mammals. We also predict that (absolute and relative) brain size is an important predictor of macroevolutionary extinction patterns.
We present a compilation of endocranial volumes (ECV) for 176 non-human primate species based on ... more We present a compilation of endocranial volumes (ECV) for 176 non-human primate species based on individual data collected from 3813 museum specimens, at least 88% being wild-caught. In combination with body mass data from wild individuals, strong correlations between endocranial volume and body mass within taxonomic groups were found. Errors attributable to different techniques for measuring cranial capacity were negligible and unbiased. The overall slopes for regressions of log ECV on log body mass in primates are 0.773 for least-squares regression and 0.793 for reduced major axis regression. The least-squares slope is reduced to 0.565 when independent contrasts are substituted for species means (branch lengths from molecular studies). A common slope of 0.646 is obtained with logged species means when grade shifts between major groups are taken into account using ANCOVA. In addition to providing a comprehensive and reliable database for comparative analyses of primate brain size, we show that the scaling relationship between brain mass and ECV does not differ significantly from isometry in primates. We also demonstrate that ECV does not differ substantially between captive and wild samples of the same species. ECV may be a more reliable indicator of brain size than brain mass, because considerably larger samples can be collected to better represent the full range of intraspecific variation. We also provide support for the maternal energy hypothesis by showing that basal metabolic rate (BMR) and gestation period are both positively correlated with brain size in primates, after controlling for the influence of body mass and potential effects of phylogenetic relatedness.
Comparative neurobiologists have provided ample evidence that in vertebrates small animals have p... more Comparative neurobiologists have provided ample evidence that in vertebrates small animals have proportionally larger brains: in a double-logarithmic plot of brain weight versus body weight all data points conform quite closely to a straight line with a slope of less than one. Hence vertebrate brains scale allometrically, rather than isometrically, with body size. Here we extend the phylogenetic scope of such studies and the size range of the brains under investigation to the insects, especially ants. We show that the principle of (negative) allometry applies as well, but that ants have considerably smaller brains than any ant-sized vertebrate would have, and that this result holds even if the relatively higher exoskeleton weights of ants (as compared to endoskeleton weights of mammals) are taken into account. Finally, interspecific comparisons within one genus of ants, Cataglyphis, show that species exhibiting small colony sizes (of a few hundred individuals) have significantly smaller brains than species in which colonies are composed of several thousand individuals.
Abstract We assessed water-body use by elephants through monitoring elephant signs around them. E... more Abstract We assessed water-body use by elephants through monitoring elephant signs around them. Elephant footprints and dung piles were recorded at 25 water bodies fortnightly for one year. Elephants preferred perennial water bodies and avoided those with temporary human dwellings. Human activities did not significantly affect elephant use of water bodies, suggesting low incidence of activities and behavioral adaptation to them by elephants. Elephant signs at perennial water bodies increased in the dry season.
This paper presents spatiotemporal gait parameters of arboreal locomotion in the colobine< i> Rhi... more This paper presents spatiotemporal gait parameters of arboreal locomotion in the colobine< i> Rhinopithecus bieti</i> in its natural habitat. While adult females used exclusively either extended-elbow vertical climbing or pulse climbing, the much larger adult males preferred the less demanding flexed-elbow vertical climbing on thin trees or on trunks with handholds.
Abstract Flexion/extension moment arms were obtained for the major muscles crossing the hip, knee... more Abstract Flexion/extension moment arms were obtained for the major muscles crossing the hip, knee and ankle joints in the orang-utan, gibbon, gorilla (Eastern and Western lowland) and bonobo. Moment arms varied with joint motion and were generally longer in proximal limb muscles than distal limb muscles. The shape of the moment arm curves (ie the plots of moment arm against joint angle) differed in different hindlimb muscles and in the same muscle in different subjects (both in the same and in different ape species).
The human brain stands out among mammals by being unusually large. The expensive-tissue hypothesi... more The human brain stands out among mammals by being unusually large. The expensive-tissue hypothesis 1 explains its evolution by proposing a trade-off between the size of the brain and that of the digestive tract, which is smaller than expected for a primate of our body size. Although this hypothesis is widely accepted, empirical support so far has been equivocal. Here we test it in a sample of 100 mammalian species, including 23 primates, by analysing brain size and organ mass data.
Abstract We present quantitative data on the hindlimb musculature of Pan paniscus, Gorilla gorill... more Abstract We present quantitative data on the hindlimb musculature of Pan paniscus, Gorilla gorilla gorilla, Gorilla gorilla graueri, Pongo pygmaeus abelii and Hylobates lar and discuss the findings in relation to the locomotor habits of each. Muscle mass and fascicle length data were obtained for all major hindlimb muscles. Physiological cross-sectional area (PCSA) was estimated. Data were normalized assuming geometric similarity to allow for comparison of animals of different size/species.
The processes that underlie the formation of the dominance hierarchy in a group are since long un... more The processes that underlie the formation of the dominance hierarchy in a group are since long under debate. Models of self-organisation suggest that dominance hierarchies develop by the self-reinforcing effects of winning and losing fights (the so-called winner-loser effect), but according to 'the prior attribute hypothesis', dominance hierarchies develop from pre-existing individual differences, such as in body mass.
Abstract Quantitative, accurate data regarding the inertial properties of body segments are of pa... more Abstract Quantitative, accurate data regarding the inertial properties of body segments are of paramount importance when developing musculo-skeletal locomotor models of living animals and, by inference, their ancestors. The limited number of available primate cadavers, and the destructive nature of the post-mortem, result in such data being very rare for primates. This study builds on the work of Crompton et al.(Am. J. Phys. Anthropol.
Summary The choice of an appropriate method for fitting a straight line to data is one of the maj... more Summary The choice of an appropriate method for fitting a straight line to data is one of the major procedural problems in bivariate allometric analysis. Commonly used techniques, such as ordinary least-squares regression and major axis, are derived from the general structural relationship model and thus require some knowledge about the distribution of the data. In this paper, we explore a nonparametric alternative, referred to as “rotation method”, which involves no assumptions about error distributions.
Abstract The maximum capability of a muscle can be estimated from simple measurements of muscle a... more Abstract The maximum capability of a muscle can be estimated from simple measurements of muscle architecture such as muscle belly mass, fascicle length and physiological cross-sectional area. While the hindlimb anatomy of the non-human apes has been studied in some detail, a comparative study of the forelimb architecture across a number of species has never been undertaken.
Humans stand out among the apes by having both an extremely large brain and a relatively high rep... more Humans stand out among the apes by having both an extremely large brain and a relatively high reproductive output, which has been proposed to be a consequence of cooperative breeding. Here, we test for general correlates of allomaternal care in a broad sample of 445 mammal species, by examining life history traits, brain size, and different helping behaviors, such as provisioning, carrying, huddling or protecting the offspring and the mother.
For over a century, various neuroanatomical measures have been employed as assays of cognitive ab... more For over a century, various neuroanatomical measures have been employed as assays of cognitive ability in comparative studies. Nevertheless, it is still unclear whether these measures actually correspond to cognitive ability. A recent metaanalysis of cognitive performance of a broad set of primate species has made it possible to provide a quantitative estimate of general cognitive ability across primates. We find that this estimate is not strongly correlated with neuroanatomical measures that statistically control for a possible effect of body size, such as encephalization quotient or brain size residuals. Instead, absolute brain size measures were the best predictors of primate cognitive ability. Moreover, there was no indication that neocortex-based measures were superior to measures based on the whole brain. The results of previous comparative studies on the evolution of intelligence must be reviewed with this conclusion in mind.
In the ongoing discussion about brain evolution in vertebrates, the main interest has shifted fro... more In the ongoing discussion about brain evolution in vertebrates, the main interest has shifted from theories focusing on energy balance to theories proposing social or ecological benefits of enhanced intellect. With the availability of a wealth of new data on basal metabolic rate (BMR) and brain size and with the aid of reliable techniques of comparative analysis, we are able to show that in fact energetics is an issue in the maintenance of a relatively large brain, and that brain size is positively correlated with the BMR in mammals, controlling for body size effects. We conclude that attempts to explain brain size variation in different taxa must consider the ability to sustain the energy costs alongside cognitive benefits.
The correlation between brain size and life history has been investigated in many previous studie... more The correlation between brain size and life history has been investigated in many previous studies, and several viable explanations have been proposed. However, the results of these studies are often at odds, causing uncertainties about whether these two character complexes underwent correlated evolution. These disparities could arise from the mixture of wild and captive values in the datasets, potentially obscuring real relationships, and from differences in the methods of controlling for phylogenetic non independence of species values. This paper seeks to resolve these difficulties by proposing an overarching hypothesis that encompasses many of the previously proposed hypotheses, and (2) testing the predictions of this hypothesis using rigorously compiled data and utilizing multiple methods of analysis. We hypothesize that the adaptive benefit of increased encephalization is an increase in reproductive lifespan or efficiency, which must be sufficient to outweigh the costs due to growing and maturing the larger brain. These costs and benefits are directly reflected in the length of life history stages. We tested this hypothesis on a wide range of primate species. Our results demonstrate that encephalization is significantly correlated with prolongation of all stages of developmental life history except the lactational period, and is significantly correlated with an extension of the reproductive lifespan. These results support the contention that the link between brain size and life history is caused by a balance between the costs of growing a brain and the survival benefits the brain provides. Thus, our results suggest that the evolution of prolonged life history during human evolution is caused by increased encephalization.
American journal of physical anthropology, Jan 1, 2005
Vertical climbing has played an important role in theories about the evolution of habitual bipeda... more Vertical climbing has played an important role in theories about the evolution of habitual bipedalism in early hominids and of locomotor specialization in hominoids. However, empirical data on vertical climbing in nonhuman primates are scarce, especially regarding kinematics. In this paper, the kinematics of flexedelbow vertical climbing of four hominoid species are reported: western lowland gorillas (Gorilla gorilla gorilla), bonobos (Pan paniscus), Sumatran orangutans (Pongo pygmaeus abelii), and yellow-cheeked crested gibbons (Hylobates concolor gabriellae). The climbing sequences were recorded simultaneously by four digital video cameras in European zoos. Analyzed parameters include intersegmental flexion/extension angles of the shoulder, elbow, hip, and knee joints, horizontal abduction/adduction angles of the upper arm and thigh, and distance of the body
Costs and benefits of encephalization are a major topic of debate in the study of primate and hum... more Costs and benefits of encephalization are a major topic of debate in the study of primate and human evolution. Comparative studies provide an opportunity to test the validity of a hypothesis as a general principle, rather than it being a special case in primate or hominid evolution. If a population evolves a larger brain, the metabolic costs of doing so must be paid for by either an increased energy turnover (direct metabolic constraint) or by a trade-off with other energetically expensive costs of body maintenance, locomotion, or reproduction, here referred to as the energy trade-off hypothesis, an extension of the influential Expensive Tissue Hypothesis of Aiello and Wheeler (1995, Curr. Anthropol. 36, 199e 221). In the present paper, we tested these hypotheses on birds using raw species values, family means, and independent contrasts analysis to account for phylogenetic influences. First, we tested whether basal metabolic rates are correlated with brain mass or any other variable of interest. This not being the case, we examined various trade-offs between brain mass and the mass of other expensive tissues such as gut mass, which is approximated by gut length or diet quality. Only weak support was found for this original Expensive Tissue Hypothesis in birds. However, other energy allocations such as locomotor mode and reproductive strategy may also be reduced to shunt energy to an enlarged brain. We found a significantly negative correlation between brain mass and pectoral muscle mass, which averages 18% of body mass in birds and is indicative of their relative costs of flight. Reproductive costs, on the other hand, are positively correlated with brain mass in birds. An increase in brain mass may allow birds to devote more energy to reproduction, although not through an increase in their own energy budget as in mammals, but through direct provisioning of their offspring. The trade-off between locomotor costs and brain mass in birds lets us conclude that an analogous effect could have played a role in the evolution of a larger brain in human evolution.
To explain variation in relative brain size among homoiothermic vertebrates, we propose the Expen... more To explain variation in relative brain size among homoiothermic vertebrates, we propose the Expensive Brain hypothesis as a unifying explanatory framework. It claims that the costs of a relatively large brain must be met by any combination of increased total energy turnover or reduced energy allocation to another expensive function such as digestion, locomotion, or production (growth and reproduction). Focusing on the energetic costs of brain enlargement, a comparative analysis of the largest mammalian sample assembled to date shows that an increase in brain size leads to larger neonates among all mammals and a longer period of immaturity among monotokous precocial species, but not among the polytokous altricial ones, who instead reduce their litter size. Relatively large brained mammals, altricial and precocial, also show reduced annual fertility rates as compared to their smaller brained relatives, but allomaternal energy inputs allow some cooperatively breeding altricial carnivores to produce even more offspring in a shorter time despite having a relatively large brain. Thus, the Expensive Brain framework explains why brain size is linked to life history pace in some, but not all mammalian lineages. This framework encompasses other hypotheses of energetic constraints on brain size variation and is also compatible with the Brain Malnutrition Risk hypothesis, but the absence of a mammal-wide correlation between brain size and immature period argues against the Needing-to-Learn explanation for slower development among large brained mammals.
The expensive brain hypothesis predicts an interspecific link between relative brain size and lif... more The expensive brain hypothesis predicts an interspecific link between relative brain size and life-history pace. Indeed, animals with relatively large brains have reduced rates of growth and reproduction. However, they also have increased total lifespan. Here we show that the reduction in production with increasing brain size is not fully compensated by the increase in lifespan. Consequently, the maximum rate of population increase (r max ) is negatively correlated with brain mass. This result is not due to a confounding effect of body size, indicating that the wellknown correlation between r max and body size is driven by brain size, at least among homeothermic vertebrates. Thus, each lineage faces a 'grey ceiling', i.e. a maximum viable brain size, beyond which r max is so low that the risk of local or species extinction is very high. We found that the steep decline in r max with brain size is absent in taxa with allomaternal offspring provisioning, such as cooperatively breeding mammals and most altricial birds. These taxa thus do not face a lineage-specific grey ceiling, which explains the far greater number of independent origins of large brain size in birds than mammals. We also predict that (absolute and relative) brain size is an important predictor of macroevolutionary extinction patterns.
We present a compilation of endocranial volumes (ECV) for 176 non-human primate species based on ... more We present a compilation of endocranial volumes (ECV) for 176 non-human primate species based on individual data collected from 3813 museum specimens, at least 88% being wild-caught. In combination with body mass data from wild individuals, strong correlations between endocranial volume and body mass within taxonomic groups were found. Errors attributable to different techniques for measuring cranial capacity were negligible and unbiased. The overall slopes for regressions of log ECV on log body mass in primates are 0.773 for least-squares regression and 0.793 for reduced major axis regression. The least-squares slope is reduced to 0.565 when independent contrasts are substituted for species means (branch lengths from molecular studies). A common slope of 0.646 is obtained with logged species means when grade shifts between major groups are taken into account using ANCOVA. In addition to providing a comprehensive and reliable database for comparative analyses of primate brain size, we show that the scaling relationship between brain mass and ECV does not differ significantly from isometry in primates. We also demonstrate that ECV does not differ substantially between captive and wild samples of the same species. ECV may be a more reliable indicator of brain size than brain mass, because considerably larger samples can be collected to better represent the full range of intraspecific variation. We also provide support for the maternal energy hypothesis by showing that basal metabolic rate (BMR) and gestation period are both positively correlated with brain size in primates, after controlling for the influence of body mass and potential effects of phylogenetic relatedness.
Comparative neurobiologists have provided ample evidence that in vertebrates small animals have p... more Comparative neurobiologists have provided ample evidence that in vertebrates small animals have proportionally larger brains: in a double-logarithmic plot of brain weight versus body weight all data points conform quite closely to a straight line with a slope of less than one. Hence vertebrate brains scale allometrically, rather than isometrically, with body size. Here we extend the phylogenetic scope of such studies and the size range of the brains under investigation to the insects, especially ants. We show that the principle of (negative) allometry applies as well, but that ants have considerably smaller brains than any ant-sized vertebrate would have, and that this result holds even if the relatively higher exoskeleton weights of ants (as compared to endoskeleton weights of mammals) are taken into account. Finally, interspecific comparisons within one genus of ants, Cataglyphis, show that species exhibiting small colony sizes (of a few hundred individuals) have significantly smaller brains than species in which colonies are composed of several thousand individuals.
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Papers by Karin Isler