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1.
The construction of nests (or beds) for sleeping is a chimpanzee universal, yet little is known about the adaptive function of nest-building. We present an in-depth study of nest-building by unhabituated chimpanzees at the Seringbara study site in the Nimba Mountains, Guinea, West Africa. We recorded 1520 chimpanzee nests over 28 mo during three study periods between 2003 and 2008. We investigated where chimpanzees built their nests, both across the home range and in nest trees, and assessed how altitude and habitat type affected nest site selectivity. We examined whether or not chimpanzees were selective in nest tree choice regarding physical tree characteristics and tree species and assessed plant species preference for both tree- and ground-nesting. We tested three, nonmutually exclusive, hypotheses for the function of arboreal nest-building. We assessed whether selectivity for nest tree characteristics reflected an antipredator strategy, examined whether nesting patterns (both arboreal and terrestrial) and nesting height were influenced by variation in climatic conditions (temperature, humidity, wind), and measured mosquito densities at ground level and in trees at 10 m and related mosquito densities to nesting patterns. Chimpanzees preferred to nest above 1000 m and nested mainly in primary forest. They preferred relatively large trees with a low first branch, dense canopy, and small leaves and showed preference for particular tree species, which was stable across years, whereas plant choice for ground-nesting was largely based on plant availability. We found no support for the antipredation hypothesis, nor did mosquito densities explain arboreal nest-building. The thermoregulation hypothesis was supported, as both nesting patterns and nest-height variation across seasons reflected a humidity-avoidance strategy. Chimpanzees nested higher in trees and at higher altitudes in the wet season. In sum, chimpanzees were selective in their choice of nest sites, locations, and materials, and tree-nesting patterns at Seringbara were best explained by a thermoregulation strategy of humidity avoidance.  相似文献   

2.
The chimpanzees (Pan troglodytes verus) of the Nimba Mountains, Guinea, West Africa, commonly make both elaborate ("night") and simple ("day") nests on the ground. In this study we investigated which factors might influence ground-nesting in this population, and tested two ecological hypotheses: 1) climatic conditions, such as high wind speeds at high altitudes, may deter chimpanzees from nesting in trees; and 2) a lack of appropriate arboreal nesting opportunities may drive the chimpanzees to nest on the ground. In addition to testing these two hypotheses, we explored whether ground-nesting is a sex-linked behavior. Data were collected monthly between August 2003 and May 2004 along transects and ad libitum. To identify the sex of ground-nesting individuals, we used DNA extracted from hair samples. The results showed that the occurrence and distribution of ground nests were not affected by climatic conditions or a lack of appropriate nest trees. Support was found for the notion that ground-nesting is a sex-linked behavior, as males were responsible for building all of the elaborate ground nests and most of the simple ground nests sampled. Elaborate ground nests occurred mostly in nest groups associated with tree nests, whereas simple ground nests usually occurred without tree nests in their vicinity. These results suggest that ground-nesting may be socially, rather than ecologically, determined.  相似文献   

3.
All great apes build nightly a structure ("nest" or "bed") that is assumed to function primarily as a sleeping-platform. However, several other nest function hypotheses have been proposed: antipredation, antipathogen, and thermoregulation. I tested these simple shelter functions of chimpanzee nests in an experiment for which I was the subject in Fongoli, Senegal. I slept 11 nights in chimpanzee nests and on the bare ground to test for differences in sleep quality, potential exposure to disease through bites from possible vectors, and insulation. No difference was found in the total amount of sleep nor in sleep quality; however, sleep was more disturbed on the ground. Differences in sleep disturbance between arboreal and ground conditions seemed primarily due to causes of anxiety and alertness, e.g., vocalizations of terrestrial mammals. Arboreal nest-sleeping seems to reduce risk of bites from possible disease vectors and provide insulation in cold conditions. This preliminary, but direct, test of chimpanzee nest function has implications for the evolutionary transition from limb-roosting to nest-reclining sleep in the hominoids, and from tree-to-ground sleep in the genus Homo.  相似文献   

4.
The Homo habilis OH 62 partial skeleton has played an important, although controversial role in interpretations of early Homo locomotor behavior. Past interpretive problems stemmed from uncertain bone length estimates and comparisons using external bone breadth proportions, which do not clearly distinguish between modern humans and apes. Here, true cross-sectional bone strength measurements of the OH 62 femur and humerus are compared with those of modern humans and chimpanzees, as well as two early H. erectus specimens-KNM-WT 15000 and KNM-ER 1808. The comparative sections include two locations in the femur and two in the humerus in order to encompass the range of possible section positions in the OH 62 specimens. For each combination of section locations, femoral to humeral strength proportions of OH 62 fall below the 95% confidence interval of modern humans, and for most comparisons, within the 95% confidence interval of chimpanzees. In contrast, the two H. erectus specimens both fall within or even above the modern human distributions. This indicates that load distribution between the limbs, and by implication, locomotor behavior, was significantly different in H. habilis from that of H. erectus and modern humans. When considered with other postcranial evidence, the most likely interpretation is that H. habilis, although bipedal when terrestrial, still engaged in frequent arboreal behavior, while H. erectus was a completely committed terrestrial biped. This adds to the evidence that H. habilis (sensu stricto) and H. erectus represent ecologically distinct, parallel lineages during the early Pleistocene.  相似文献   

5.
The nightly construction of an arboreal sleeping platform (SP) has been observed among every chimpanzee's population studied to date. Here, we report on bioclimatic aspects of SP site choice among dry-habitat chimpanzees (Pan troglodytes schweinfurthii) at the Toro-Semliki Wildlife Reserve, Uganda. We placed a portable weather monitor within 1 m of chimpanzee SPs and compared the microenvironment of this site with terrestrial monitors placed 10 cm above the ground directly underneath the simultaneously studied SP. We calculated physical "comfort levels" of monitored sites using the RayMan thermophysiological model that we modified to take ape body proportions into account. The RayMan tool gauges energy balance using wind speed, temperature, relative humidity, and heat index in conjunction with the study subject's mass and stature to determine whether the individual is in energy balance or homeostasis. We found that (1) terrestrial microclimates have greater homeostatic potential than arboreal microclimates, and (2) there is a significant positive linear relationship between wind speed and height of SP in the forest canopy. Advantages of terrestrial sites are that they require lesser energetic expenditure to stabilize the body when the SP is under construction and perhaps during use as well. We found that terrestrial sites also had better homeostatic potentials. This combination of advantages explains why SPs are so often sited terrestrially in habitats where predation risk is low. Early hominins must have had technological or social measures to avoid or deter predators that were significantly advanced over those found among chimpanzees before they began sleeping on the ground.  相似文献   

6.
In this survey conducted in southern Cameroon, we compared ant-Hemiptera associations on plantations treated with insecticides, on plantations 2 years after insecticide treatments ceased, and on control lots that never received insecticide treatments. By eliminating arboreal-nesting ants, insecticides favored the presence of "ecologically dominant" ground-nesting, arboreal-foraging species that occupied the tree crowns. The reinstallation of arboreal ants was slow as 2 yr after insecticide treatment ceased differences with the control lots were significant. This intermediary period also illustrated that arboreal ants can found and develop colonies on trees occupied by ground-nesting species. Certain arboreal species were more frequent during this intermediary period than on the control lots, showing that the period of installation in the trees was followed by competition between arboreal ants. We confirm that ground-nesting ants tend a wide range of hemipteran families, including well known agricultural pests, whereas arboreal ants, particularly dominant species, were mostly associated with Coccidae and Stictococcidae that do not pose problems to the supporting trees. A tree effect was also noted for both ant and hemipteran distribution. We concluded that because of insecticide use, ground-nesting ants pose problems through their associated Hemiptera. On the contrary, dominant arboreal ants, strong predators, benefit their supporting trees by excluding ground-nesting species and tending mostly nonpest Hemiptera. Nevertheless, certain of them, carpenter species or species likely to tend Pseudococcidae, have to be eliminated through integrated management.  相似文献   

7.
Knuckle-walking is a pattern of digitigrade locomotion unique to African apes among Primates. Only chimpanzees and gorillas are specially adapted for supporting weight on the dorsal aspects of middle phalanges of flexed hand digits II–V. When forced to the ground, most orangutans assume one of a variety of flexed hand postures, but they cannot knuckle-walk. Some orangutans place their hands in palmigrade postures which are impossible to African apes. The knuckle-walking hands and plantigrade feet of African apes are both morphologically and adaptively distinct from those of Pongo, their nearest relative among extant apes. These features are associated with a common adaptive shift to terrestrial locomotion and support placing chimpanzees and gorillas in the same genus Pan. It is further suggested than Pan comprises the subgenera (a) Pan, including P. troglodytes and pygmy chimpanzees, and (b) Gorilla, including mountain and lowland populations of P. gorilla. African apes probably diverged from ancestral pongids that were specially adapted for distributing their weight in terminal branches of the forest canopy. Early adjustments to terrestrial locomotion may have involved fist-walking which later evolved into knuckle-walking. Orangutans continued to adapt to feeding and locomotion in the forest canopy and their hands and feet became highly specialized for four-digit prehension. Although chimpanzees retained arboreal feeding and nesting habits, they moved from tree to tree by terrestrial routes and became less restricted in habitat. While adapting to a diet of ground plants gorillas increased in size to the point that arboreal nesting is less frequent among them than among chimpanzees and orangutans. Early hominids probably diverged from pongids that had not developed prospective adaptations to knuckle-walking, and therefore did not evolve through a knuckle-walking stage. Initial adjustments to terrestrial quadrupedal locomotion and resting stance probably included palmigrade hand posturing. Their thumbs may have been already well developed as an adaptation for grasping during arboreal climbing. A combination of selection pressures for efficient terrestrial locomotor support and for object manipulation further advanced early hominid hands toward modern human configuration.  相似文献   

8.
Data on nesting behavior and nests of the Sonso community of chimpanzees (Pan troglodytes schweinfurthii) living in the Budongo Forest, Uganda, were collected between October 1995 and December 1996. Analysis of the selection of nests shows a preference for particular tree species, especially Cynometra alexandri. Male chimpanzees nested lower than females. Day nests were found to be structurally simpler than night nests and were built in tree species used for feeding, at the same height as feeding activity. Day and night nests serve different functions, and should be clearly distinguished in studies of nesting behavior.  相似文献   

9.
A trademark of Homo sapiens is the enormous variation in behavioral patterns across populations. Insight into the development of human cultures can be aided by studies on communities of Pan across Africa that display unique combinations of social behavior and elementary technology. Only cross-population comparisons can reveal whether the diversity reflects differential genetics, environmental constraints, or is a cultural variant. However, the recently recognized and most endangered subspecies, Pan troglodytes vellerosus, remains completely unstudied in this respect. We report first evidence from a new long-term study of Nigerian chimpanzees at Gashaka. Their dietary composition is highly varied and they have to cope with high concentrations of antifeedant defenses of plants against consumption. Gashaka chimpanzees use a varied tool kit for extractive foraging. For example, they harvest insects throughout the year, via digging sticks and probes, to obtain honey from stingless-bee and honeybee nests, dipping wands to prey on army ants, and fishing rods to eat arboreal ants. Tools appeared to be custom-made with a considerable degree of standardization in length, diameter, and preferential use of distal ends. Moreover, compared to the rainy season, tools were longer during the dry season when insects retreat further into their nests. Many of the expressions of subsistence technology seem to be environmentally constrained. Most notably, the absence of termite-eating could reflect a low abundance of mounds. Other traits may represent cultural variation. For example, the chimpanzees did not hammer open 2 types of hard-shelled nuts with tools, unlike what occurs elsewhere in West Africa. The prevalence of elementary technology may indicate that the material culture of Gashaka chimpanzees is most related to core cultural tendencies of Central African populations.  相似文献   

10.
We studied the nesting behavior of the critically endangered West African chimpanzee (Pan troglodytes verus). We assumed that the nesting data stemmed from a single, unhabituated community at the Bagnomba hill site in the savanna-woodlands of southeastern Senegal. The aim of this study was to examine chimpanzees’ nesting habits in terms of the tree species utilized and sleeping nest heights. We recorded a total of 550 chimpanzee nests at Bagnomba between January 2015 and December 2015. The chimpanzees here made nests in particular tree species more often than others. The majority of nests (63%) were in two tree species: Diospyros mespiliformis and Pterocarpus erinaceus. The average height of nesting trees was 10.54 m (SD 3.91, range, 0.0–29.0 m) and average nest height was 7.90 m (SD 3.62, range, 0.0–25.0 m). The result of a linear regression analysis (r = 0.7874; n = 550; p < 0.05) is consistent with a preference for nesting at a particular height. Bagnomba chimpanzees rarely made ground nests (0.36% of nests), but the presence of any ground nesting was unexpected, given that at least one leopard (Panthera pardus) also occupied the hill. This knowledge will enable stakeholders involved in the protection of chimpanzees specifically and of biodiversity in general to better understand chimpanzee ecology and inform a conservation action plan in Senegal where the survival of this species is threatened.  相似文献   

11.
Arboreality and bipedality in the Hadar hominids   总被引:3,自引:0,他引:3  
Numerous studies of the locomotor skeleton of the Hadar hominids have revealed traits indicative of both arboreal climbing/suspension and terrestrial bipedalism. These earliest known hominids must have devoted part of their activities to feeding, sleeping and/or predator avoidance in trees, while also spending time on the ground where they moved bipedally. In this paper we offer new data on phalangeal length and curvature, morphology of the tarsus and metatarsophalangeal joints, and body proportions that further strengthen the argument for arboreality in the Hadar hominids. We also provide additional evidence on limb and pedal proportions and on the functional anatomy of the hip, knee and foot, indicating that the bipedality practiced at Hadar differed from that of modern humans. Consideration of the ecology at Hadar, in conjunction with modern primate models, supports the notion of arboredality in these earliest australopithecines. We speculate that selection for terrestrial bipedality may have intensified through the Plio-Pleistocene as forests and woodland patches shrunk and the need arose to move increasingly longer distances on the ground. Only with Homo erectus might body size, culture and other factors have combined to 'release' hominids from their dependence on trees.  相似文献   

12.
Small animals must be capable of moving on a wide variety of surfaces; thus, examining the mechanics of locomotion on a wide variety of substrates is necessary to understand how the animal can utilize its habitat. Therefore, locomotor kinetics are examined on arboreal and terrestrial sloped substrates in the marsupial Monodelphis domestica (gray short-tailed opossum). Substrate reaction forces were measured as opossums moved across four trackways: 30 degrees upslope and 30 degrees downslope trackways, which were flat ("terrestrial") or cylindrical ("arboreal"). Regardless of substrate slope, medial limb forces were measured on arboreal trackways and usually lateral limb forces on terrestrial trackways. Otherwise the general patterns of vertical and craniocaudal forces and impulses were similar between same-sloped terrestrial and arboreal trackways. Some significant modifications to these gross patterns occurred: on the arboreal upslope trackway, hindlimbs supported more body weight than on the terrestrial uphill, possibly because hindlimbs were more stably positioned on the upslope arboreal trackway than forelimbs. Furthermore, the difference between fore- and hindlimbs with respect to craniocaudal impulses was less on the arboreal sloped trackways. In conclusion, kinetic patterns can usually be explained by body weight support roles and by the placement of the limbs on the arboreal trackway.  相似文献   

13.
Chimpanzees (Pan troglodytes) make nests for resting and sleeping, which is unusual for anthropoid primates but common to all great apes. Arboreal nesting has been linked to predation pressure, but few studies have tested the adaptive nature of this behavior. We collected data at two chimpanzee study sites in southeastern Senegal that differed in predator presence to test the hypothesis that elevated sleeping platforms are adaptations for predator defense. At Assirik in the Parc National du Niokolo-Koba, chimpanzees face four species of large carnivore, whereas at Fongoli, outside national park boundaries, humans have exterminated almost all natural predators. We quantified the availability of vegetation at the two sites to test the alternative hypothesis that differences in nesting reflect differences in habitat structure. We also examined possible sex differences in nesting behavior, community demographic differences, seasonality and nest age differences as variables also potentially affecting nest characteristics and nesting behavior between the two sites. Chimpanzees at Fongoli nested at lower heights and farther apart than did chimpanzees at Assirik and sometimes made nests on the ground. The absence of predators outside of the national park may account for the differences in nest characteristics at the two sites, given the similarities in habitat structure between Fongoli and Assirik. However, Fongoli chimpanzees regularly build arboreal nests for sleeping, even under minimal predation pressure, and this requires explanation.  相似文献   

14.
Four chimpanzee (Pan troglodytes)mother—infant dyads and four orangutan (Pongo pygmaeus)motherinfant dyads were studied for the first 11 months of the infants’ lives. For both species, ventroventral contact and nipple contact decreased over time at a similar rate, but total contact decreased earlier in the orangutans and was 50% lower than for the chimpanzees at the end of the study. Social play between the mothers and the infants did not differ in frequency between the species, but orangutans played above the ground and chimpanzees on the ground. Solitary play differed in form between the species and, like social play, reflected their differences in arboreal and terrestrial proclivities. In addition, the orangutans engaged in solitary play considerably more frequently than the chimpanzees during the second half-year of life. The developmental differences in mother-infant contact and solitary play of these apes are consistent with the differences in their speciestypical social organization. The data may reflect, therefore, early development of species differences in the social and relatively solitary natures of chimpanzees and orangutans, respectively. An erratum to this article is available at .  相似文献   

15.
Arboreal, and in particular suspensory, postures may elicit a preference for the strongest limb to be used in postural support in large bodied primates. However, selection may have favored ambilaterality rather than a preference for a particular hand in chimpanzees (Pan troglodytes) fishing arboreally for ants. To investigate the influence of arboreality on hand preference we recorded handedness in seven captive bonobos (Pan paniscus) manipulating a foraging device during terrestrial and arboreal postures in a symmetrical environment, observing 2726 bouts of manipulation. When accessing the foraging device in the arboreal position the bonobos adopted predominantly suspensory postures. There was no population level hand preference for manipulating the foraging device in either the terrestrial or arboreal positions. However, four of seven individuals that interacted with the foraging devices showed a significant preference for one hand (two were left handed, two were right handed) when manipulating the foraging device in the arboreal position whereas only one individual (left handed) showed a preference in the terrestrial position. This suggests that individuals may have a preferred or strongest limb for postural support in a symmetrical arboreal environment, resulting in a bias to use the opposite hand for manipulation. However, the hand that is preferred for postural support differs between individuals. Although our sample is for two captive groups at the same zoo, our findings suggest that the demand of maintaining arboreal postures and environmental complexity influence hand preference.  相似文献   

16.
Homo faber was once proposed as a label for humans specifically to highlight their unique propensity for tool use. However, new observations on complex tool use by the chimpanzees of Loango National Park, Gabon, expand our knowledge about tool-using abilities in Pan troglodytes. Chimpanzees in Loango, when using tools to extract honey from three types of bee nests, were observed to regularly use three- to five-element tool sets. In other words, different types of tools were used sequentially to access a single food source. Such tool sets included multi-function tools that present typical wear for two distinct uses. In addition, chimpanzees exploited underground bee nests and used ground-perforating tools to locate nest chambers that were not visible from the ground surface. These new observations concur with others from Central African chimpanzees to highlight the importance of honey extraction in arguments favoring the emergence of complex tool use in hominoids, including different tool types, expanded tool sets, multifunction tools, and the exploitation of underground resources. This last technique requires sophisticated cognitive abilities concerning unseen objects. A sequential analysis reveals a higher level of complexity in honey extraction than previously proposed for nut cracking or hunting tools, and compares with some technologies attributed to early hominins from the Early and Middle Stone Age. A better understanding of similarities in human and chimpanzee tool use will allow for a greater understanding of tool-using skills that are uniquely human.  相似文献   

17.
We investigated oil palm (Elaeis guineensis) use for feeding in 3 chimpanzee communities: Bossou and Seringbara in Guinea and Yealé in Côte d'Ivoire. Bossou was used as the benchmark for comparison. Bossou chimpanzees (Pan troglodytes verus) exhibit a wide range of oil palm targeted behaviors. We used direct observations of their two tool use, i.e., nut-cracking and pestle pounding, to establish strict and reliable criteria to ascertain the presence of comparable behaviors at the two adjacent Nimba sites. Based on monthly surveys of oil palms across the three sites, significant differences in patterns of use emerged. Bossou chimpanzees demonstrated the greatest frequency of oil palm use, while Seringbara chimpanzees, 6 km away, failed to exhibit any use and Yealé chimpanzees, 12 km away, showed all uses comparable to Bossou chimpanzees except pestle pounding and mature leaf pith-feeding. We examined the density and distribution of oil palms, tool availability for nut-cracking and pestle pounding, fruit, flower and nut availability, competition with sympatric species for fruit and nuts and the diversity of fruit species in the diet across the 3 sites. We found no clear difference in proximate environmental variables underlying observed variations in oil palm use among the 3 sites, yielding the conclusion that the differences are cultural. Assuming individual interchange between communities and the involvement of social learning in the intracommunity transmission and maintenance of oil palm uses, the result raises interesting questions about diffusion of behavior between neighboring chimpanzee communities.  相似文献   

18.
As habitat loss and fragmentation place growing pressure on endangered nonhuman primate populations, researchers find increasing evidence for novel responses in behavior. In western Uganda between the Budongo and Bugoma Forests, chimpanzees (Pan troglodytes schweinfurthii) inhabit a mosaic landscape comprising forest fragments, human settlements, and agricultural land. We recorded nests and feeding evidence of unhabituated chimpanzees in this region over a 12-mo period. We found extensive evidence of nesting in introduced tree species, including eucalyptus (Eucalyptus grandis), guava (Psidium guajava), cocoa (Theobroma cacao), and Caribbean pine (Pinus caribaea). In addition, we found instances of ground nesting, nest reuse, and composite nests constructed from branches of multiple trees. This evidence may indicate a lack of suitable nesting trees or attempts by chimpanzees to nest in areas of riparian forest that allow them to avoid human detection. We also found new evidence for eucalyptus bark feeding by chimpanzees. Such evidence suggests chimpanzees respond flexibly to mitigate anthropogenic pressures in human-dominated landscapes. The limits of such flexibility remain unknown. Further research is needed to examine systematically the factors influencing the use of such resources and to understand better the extent to which chimpanzees can persist while relying on them.  相似文献   

19.
Ground-nesting species are vulnerable to a wide range of predators and often experience very high levels of nest predation. Strategies to reduce nest vulnerability can include concealing nests in vegetation and/or nesting in locations in which nests and eggs are camouflaged and less easy for predators to locate. These strategies could have important implications for the distribution of ground-nesting species and the success rates of nests in areas with differing vegetation structure. However, the factors influencing the success of nest concealment and camouflage strategies in ground-nesting species are complex. Here we explore the effects of local vegetation structure and extent of nest concealment on nest predation rates in a range of ground-nesting, sympatric wader species with differing nest concealment strategies (open-nest species: Oystercatcher Haematopus ostralegus, Golden Plover Pluvialis apricaria and Whimbrel Numenius phaeopus; concealed-nest species: Black-tailed Godwit Limosa limosa, Redshank Tringa totanus and Snipe Gallinago gallinago) in south Iceland, in landscapes that comprise substantial variability in vegetation structure at a range of scales. We monitored 469 nests of these six wader species in 2015 and 2016 and ~40% of these nests were predated. Nest predation rates were similar for open-nest and concealed-nest species and did not vary with vegetation structure in the surrounding landscape, but nest-concealing species were ~10% more likely to have nests predated when they were poorly concealed, and the frequency of poorly concealed nests was higher in colder conditions at the start of the breeding season. For concealed-nest species, the reduced capacity to hide nests in colder conditions is likely to reflect low rates of vegetation growth in such conditions. The ongoing trend for warmer springs at subarctic latitudes could result in more rapid vegetation growth, with consequent increases in the success rates of early nests of concealed-nest species. Temperature-related effects on nest concealment from predators could thus be an important mechanism through which climate change affecting vegetation could have population-level impacts on breeding birds at higher latitudes.  相似文献   

20.
We conducted a meta-analysis of 45 studies reporting basal metabolic rate (BMR) data for Homo sapiens and Pan troglodytes to determine the effects of sex, age, and latitude (a proxy for climate, in humans only). BMR was normalized for body size using fat-free mass in humans and body mass in chimpanzees. We found no effect of sex in either species and no age effect in chimpanzees. In humans, juveniles differed significantly from adults (ANCOVA: P < 0.001), and senescent adults differed significantly from adults younger than 50 years (P < 0.001). Europeans differed significantly from tropical populations (P < 0.001). On the basis of these observations, we derived new equations describing the relationship between BMR and body size, and used them to predict total daily energy expenditure (TEE) in four early hominin species. Our predictions concur with previous TEE estimates (i.e. Leonard and Robertson: Am J Phys Anthropol 102 (1997) 265-281), and support the conclusion that TEE increased greatly with H. erectus. Our results show that intraspecific variation in BMR does not affect TEE estimates for interspecific comparisons. Comparisons of more closely related groups such as humans and Neandertals, however, may benefit from consideration of this variation.  相似文献   

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