Scaling of limb joint surface areas in anthropoid primates and other mammals

Recently it was reported that limb joint surface areas scale positively allometrically with body weight in anthropoid primates. This was attributed to the biomechanics of weight bearing: larger animals must require relatively larger joint surfaces to withstand relatively greater weight-related stresses on the joints. Our data on humeral and femoral geometry and joint surface areas in 73 species belonging to six mammalian orders (including primates) demonstrate that positive allometry of joint surface areas is not a general phenomenon for mammals and cannot have its basis among Anthropoidea in the biomechanics of weight bearing. We argue that, to the extent that positive allometric scaling of joint surface areas occurs in anthropoid primates, it is an artifact of differences in positional behaviour among distinct taxonomic groups that also happen to differ widely in body weight. Furthermore, we argue that, among mammals ranging in body size from sportive lemurs to giant pandas, functionally similar groups tend to exhibit: (1) linear dimensions (especially diameters and shaft circumferences) that scale in direct proportion to each other; and (2) joint surface areas that scale in direct proportion to the squares of linear dimensions and to the 2/3 power of body mass. In other words, limb bones of functionally similar animals fit the theoretical model of geometric similarity (or skeletal isometry). Differences in relative sizes of joint surface areas are related to differences in force transmission and movement potential among functionally distinct groups of animals.     

Allometric scaling and locomotor function in the primate pelvis

Identification of positional behavior adaptation in the pelvis of primates is complicated by possible confounding effects of body size and phylogeny. Previous work on primate pelvic allometry has focused primarily on sexual dimorphism and its relationship to obstetric constraints in species with large fetal size relative to maternal size. This study investigates patterns of pelvic scaling with a specific aim to understand how pelvic scaling relates to locomotor function. Patterns of scaling of nine pelvic dimensions were examined in a broad comparative sample of 40 species of primates, covering both haplorhines and strepsirrhines, while accounting for phylogenetic nonindependence. Phylogenetic reduced major axis regressions on pelvic scaling patterns suggest that primate-wide patterns are reflected in haplorhine- and strepsirrhine-specific analyses. Many measures scale isometrically with pelvis size, but notably, features of the ilium tend to scale allometrically. As predicted, ilium width and lower ilium cross-sectional area scale with positive allometry, while lower iliac height scales with negative allometry. Further regression analyses by locomotor group suggest that these ilium measures, as well as pubic symphysis and ischium lengths, differ in their scaling patterns according to locomotor mode. These results suggest that scaling differences within primates, when present, are related to functional differences in locomotor behavior and mechanics. This study supports recent work that identifies adaptations to locomotor loading in the ilium and highlights the need for a better understanding of the relationship between pelvic structural mechanics and the mechanical requirements of primate locomotion. Am J Phys Anthropol 156:511–530, 2015. © 2015 Wiley Periodicals, Inc.     

Curvature of the forelimb bones of anthropoid primates: Overall allometric patterns and specializations in suspensory species

It has previously been reported that brachiating primates, particularly gibbons, are characterized by distinctively straight forelimb long bones, yet no hypotheses have been proposed to explain why straight limb bones may be adaptive to suspensory locomotion. This study explores quantitatively the curvature of the long bones in 13 species of anthropoid primates and analyzes the functional consequences of curvature in biomechanical terms. These analyses demonstrate that, whereas the humeri of gibbons and spider monkeys are functionally less curved than those of other taxa, the ulnae of brachiators are neither more nor less curved than those of other anthropoids, and the gibbon radius is far more curved than would be predicted from body size alone. The humerus is likely significantly less curved in brachiators because of its torsion-dominated loading regime and the greatly increased stress magnitude developed in torsionally loaded curved beams. The large curvature of the radius is localized in the region of attachment of the supinator muscle. Analysis presented here of muscle mass allometry in catarrhines demonstrates that gibbons are characterized by an extremely massive supinator, and the large radial curvature is therefore most likely due to forearm muscle mechanics. This study also demonstrates that the overall pattern of limb bone curvature for anthropoids is distinct from the pattern reported for mammals as a whole. This distinctive scaling relationship may be related to the increased length of the limb bones of primates in comparison to other mammals.     

Comparative limb bone scaling in turtles: Phylogenetic transitions with changes in functional demands?

Several terrestrial vertebrate clades include lineages that have evolved nearly exclusive use of aquatic habitats. In many cases, such transitions are associated with the evolution of flattened limbs that are used to swim via dorsoventral flapping. Such changes in shape may have been facilitated by changes in limb bone loading in novel aquatic environments. Studies on limb bone loading in turtles found that torsion is high relative to bending loads on land, but reduced compared to bending during aquatic rowing. Release from torsion among rowers could have facilitated the evolution of hydrodynamically advantageous flattened limbs among aquatic species. Because rowing is regarded as an intermediate locomotor stage between walking and flapping, rowing species might show limb bone flattening intermediate between the tubular shapes of walkers and the flattened shapes of flappers. We collected measurements of humeri and femora from specimens representing four functionally divergent turtle clades: sea turtles (marine flappers), softshells (specialized freshwater rowers), emydids (generalist semiaquatic rowers), and tortoises (terrestrial walkers). Patterns of limb bone scaling with size were compared across lineages using phylogenetic comparative methods. Although rowing taxa did not show the intermediate scaling patterns we predicted, our data provide other functional insights. For example, flattening of sea turtle humeri was associated with positive allometry (relative to body mass) for the limb bone diameter perpendicular to the flexion-extension plane of the elbow. Moreover, softshell limb bones exhibit positive allometry of femoral diameters relative to body mass, potentially helping them maintain their typical benthic position in water by providing additional weight to compensate for shell reduction. Tortoise limb bones showed positive allometry of diameters, as well as long humeri, relative to body mass, potentially reflecting specializations for resisting loads associated with digging. Overall, scaling patterns of many turtle lineages appear to correlate with distinctive behaviors or locomotor habits.     

Scaling of postcranial joint size in hominoid primates

The scaling of sixteen articular dimensions in the locomotor skeleton of hominoid primates is examined with special reference to a recently proposed model of geometric similarity. Seven species are included in the analysis (gorillas, common chimpanzees, bonobos, orang-utans, siamang, lar gibbons, and modern humans of European descent); all specimens are adult individuals of known body mass (N=87). No significant sexual dimorphism in the scaling of joint size was observed. Overal results are compatible with the biomechanical model predictions of isometry, and lend additional support to the suggestion that joint stresses are of the same order of magnitude in animals differing vastly in body size and locomotor adaptations. The hindlimb and lumbosacral joints of humans, however, are consistently much larger than expected for their body mass. Full-time bipedality obviously precludes the sharing of weight support and propulsion with the forelimbs, and this fundamental difference is accurately reflected in the relative joint size of humans.     

Evolution of locomotion in Anthropoidea: the semicircular canal evidence

Our understanding of locomotor evolution in anthropoid primates has been limited to those taxa for which good postcranial fossil material and appropriate modern analogues are available. We report the results of an analysis of semicircular canal size variation in 16 fossil anthropoid species dating from the Late Eocene to the Late Miocene, and use these data to reconstruct evolutionary changes in locomotor adaptations in anthropoid primates over the last 35 Ma. Phylogenetically informed regression analyses of semicircular canal size reveal three important aspects of anthropoid locomotor evolution: (i) the earliest anthropoid primates engaged in relatively slow locomotor behaviours, suggesting that this was the basal anthropoid pattern; (ii) platyrrhines from the Miocene of South America were relatively agile compared with earlier anthropoids; and (iii) while the last common ancestor of cercopithecoids and hominoids likely was relatively slow like earlier stem catarrhines, the results suggest that the basal crown catarrhine may have been a relatively agile animal. The latter scenario would indicate that hominoids of the later Miocene secondarily derived their relatively slow locomotor repertoires.     

Craniodental features in male Mandrillus may signal size and fitness: an allometric approach

According to a hypothesis in the broader mammalian literature, secondary sexual characteristics that have evolved to signal fitness and size to other conspecifics should exhibit positive allometry across adult males within a species. Here this hypothesis is tested in the genus Mandrillus. The overbuilding of bony features in larger individuals necessitates a functional explanation as bone is metabolically expensive to produce and maintain. Canine size and size of the maxillary ridge are scaled against a body size surrogate in intraspecific samples of male Mandrillus sphinx (mandrills) and Mandrillus leucophaeus (drills). Areal dimensions are weighted more heavily as they represent the size of a feature as it is viewed by individuals. Measures of the maxillary ridge and canine tooth are significantly correlated with the size surrogate and scale with positive allometry in both samples supporting the hypothesis that these features function to advertise a male's body size and fitness to other males competing for mates and potential discerning females. This is the first study in primates to test for intraspecific positive allometric scaling of bony facial features in adult males based on a theory of fitness signaling and sexual selection.     

Molar scaling in strepsirrhine primates

We examined how maxillary molar dimensions change with body and skull size estimates among 54 species of living and subfossil strepsirrhine primates. Strepsirrhine maxillary molar areas tend to scale with negative allometry, or possibly isometry, relative to body mass. This observation supports several previous scaling analyses showing that primate molar areas scale at or slightly below geometric similarity relative to body mass. Strepsirrhine molar areas do not change relative to body mass(0.75), as predicted by the metabolic scaling hypothesis. Relative to basicranial length, maxillary molar areas tend to scale with positive allometry. Previous claims that primate molar areas scale with positive allometry relative to body mass appear to rest on the incorrect assumption that skull dimensions scale isometrically with body mass. We identified specific factors that help us to better understand these observed scaling patterns. Lorisiform and lemuriform maxillary molar scaling patterns did not differ significantly, suggesting that the two infraorders had little independent influence on strepsirrhine scaling patterns. Contrary to many previous studies of primate dental allometry, we found little evidence for significant differences in molar area scaling patterns among frugivorous, folivorous, and insectivorous groups. We were able to distinguish folivorous species from frugivorous and insectivorous taxa by comparing M1 lengths and widths. Folivores tend to have a mesiodistally elongated M1 for a given buccolingual M1 width when compared to the other two dietary groups. It has recently been shown that brain mass has a strong influence on primate dental eruption rates. We extended this comparison to relative maxillary molar sizes, but found that brain mass appears to have little influence on the size of strepsirrhine molars. Alternatively, we observed a strong correlation between the relative size of the facial skull and relative molar areas among strepsirrhines. We hypothesize that this association may be underlain by a partial sharing of the patterning of development between molar and facial skull elements.     

Patterns of joint size dimorphism in the elbow and knee of catarrhine primates

Differences in body size between conspecific sexes may incur differences in the relative size and/or shape of load-bearing joints, potentially confounding our understanding of variation in the fossil record. More specifically, larger males may experience relatively greater limb joint stress levels than females, unless an increase in weight-related forces is compensated for by positive allometry of articular surface areas. This study examines variation in limb joint size dimorphism (JSD) among extant catarrhines to: 1) determine whether taxa exhibit JSD beyond that expected to simply maintain geometric similarity between sexes, and 2) test whether taxa differ in JSD (relative to body size dimorphism) with respect to differences in limb use and/or phylogeny. "Joint size" was quantified for the distal humerus and distal femur of 25 taxa. Analysis of variance was used to test for differences between sexes (in joint size ratios) and among taxa (in patterns of dimorphism). Multiple regression was used to examine differences in JSD among taxa after accounting for variation in body size dimorphism (BSD) and body size. Although degrees of humeral and femoral JSD tend to be the same within species, interspecific variation exists in the extent to which both joints are dimorphic relative to BSD. While most cercopithecoids exhibit relatively high degrees of JSD (i.e., positive allometry), nonhuman hominoids exhibit degrees of JSD closer to isometry. These results may reflect a fundamental distinction between cercopithecoids and hominoids in joint design. Overall, the results make more sense (from a mechanical standpoint) when adjustments to BSD are made to account for the larger effective female body mass associated with bearing offspring. In contrast to other hominoids, modern humans exhibit relatively high JSD in both the knee and elbow (despite lack of forelimb use in weight support). Estimates of BSD based on fossil limb bones will vary according to the extant analogue chosen for comparison.     

Ontogeny,function, and scaling of the mandibular symphysis in papionin primates

In vivo study of mastication in adult cercopithecine primates demonstrates a link between mandibular symphyseal form and resistance to “wishboning,” or lateral transverse bending. Mechanical consideration of wishboning at the symphysis indicates exponentially higher stresses along the lingual surface with increasing symphyseal curvature. Lengthening the anteroposterior width of the symphysis acts to resist these higher loads. Interspecific adult cercopithecine allometries show that both symphyseal curvature and symphyseal width exhibit positive allometry relative to body mass. The experimental and allometric data support an hypothesis that the cercopithecine mandibular symphysis is designed to maintain functional equivalence—in this case dynamic strain similarity—in wishboning stress and strain magnitudes across adult cercopithecines. We test the hypothesis that functional equivalence during masticatory wishboning is maintained throughout ontogeny by calculating relative stress estimates from morphometric dimensions of the mandibular symphysis in two cercopithecine primates, Macaca fascicularis and M. nemestrina. Results indicate no significant differences in relative stress estimates among the two macaque ontogenies and an interspecific sample of adult papionin primates. Further, relative stress estimates do not change significantly throughout ontogeny in either species. These results offer the first evidence for the maintenance of functional equivalence in stress and strain levels during postnatal growth in a habitually loaded cranial structure. Scaling analyses demonstrate significant slope differences for both symphyseal curvature and width between the ontogenetic and interspecific samples. The distinct interspecific cercopithecine slopes are realized by a series of ontogenetic transpositions in both symphyseal curvature and width. Throughout papionin ontogeny, symphyseal curvature increases with less negative allometry, while symphysis width increases with less positive allometry versus the interspecific pattern. As symphyseal curvature and width are inversely proportional to one another in estimating relative stresses, functionally equivalent stress levels are maintained both ontogenetically and interspecifically, because the relatively slower rate of allometric increase in symphyseal curvature during growth is compensated for by a slower rate of allometric increase in symphyseal width. These results indicate the primacy of maintaining functional equivalence during growth and the need for ontogenetic data in understanding the evolutionary processes that affect form–function relations as well as the interspecific patterning of adult form across a clade. J. Morphol. 235:157–175, 1998. © 1998 Wiley-Liss, Inc.     

Scale Effects between Body Size and Limb Design in Quadrupedal Mammals

Recently the metabolic cost of swinging the limbs has been found to be much greater than previously thought, raising the possibility that limb rotational inertia influences the energetics of locomotion. Larger mammals have a lower mass-specific cost of transport than smaller mammals. The scaling of the mass-specific cost of transport is partly explained by decreasing stride frequency with increasing body size; however, it is unknown if limb rotational inertia also influences the mass-specific cost of transport. Limb length and inertial properties – limb mass, center of mass (COM) position, moment of inertia, radius of gyration, and natural frequency – were measured in 44 species of terrestrial mammals, spanning eight taxonomic orders. Limb length increases disproportionately with body mass via positive allometry (length ∝ body mass^0.40); the positive allometry of limb length may help explain the scaling of the metabolic cost of transport. When scaled against body mass, forelimb inertial properties, apart from mass, scale with positive allometry. Fore- and hindlimb mass scale according to geometric similarity (limb mass ∝ body mass^1.0), as do the remaining hindlimb inertial properties. The positive allometry of limb length is largely the result of absolute differences in limb inertial properties between mammalian subgroups. Though likely detrimental to locomotor costs in large mammals, scale effects in limb inertial properties appear to be concomitant with scale effects in sensorimotor control and locomotor ability in terrestrial mammals. Across mammals, the forelimb''s potential for angular acceleration scales according to geometric similarity, whereas the hindlimb''s potential for angular acceleration scales with positive allometry.     

Body size and scaling of the hands and feet of prosimian primates

The hands and feet of primates fulfill a variety of biological roles linked with food acquisition and positional behavior. Current explanations of shape differences in cheiridial morphology among prosimians are closely tied to body size differences. Although numerous studies have examined the relationships between body mass and limb morphology in prosimians, no scaling analysis has specifically considered hand and foot dimensions and intrinsic proportions. In this study, we present such an analysis for a sample of 270 skeletal specimens distributed over eight prosimian families. The degree of association between size and shape was assessed using nonparametric correlational techniques, while the relationship between each ray element length and body mass (from published data and a body mass surrogate) was tested for allometric scaling. Since tarsiers and strepsirrhines encompass many taxa of varying degrees of phylogenetic relatedness, effective degrees of freedom were calculated, and comparisons between families were performed to partially address the problem of statistical nonindependence and "phylogenetic inertia." Correlational analyses indicate negative allometry between relative phalangeal length (as reflected by phalangeal indices) and body mass, except for the pollex and hallux. Thus, as size increases, there is a significant decrease in the relative length of the digits when considering all prosimian taxa sampled. Regression analyses show that while the digital portion of the rays scales isometrically with body mass, the palmar/plantar portion of the rays often scales with positive allometry. Some but not all of these broadly interspecific allometric patterns remain statistically significant when effective degrees of freedom are taken into account. As is often the case in interspecific scaling, comparisons within families show different scaling trends in the cheiridia than those seen across families (i.e., lorisids, indriids, and lemurids exhibit rather different allometries). The interspecific pattern of positive allometry that appears to best characterize the metapodials of prosimians, especially those of the foot, parallels differences found in the morphology of the volar skin. Indeed, relatively longer metapodials appear to covary with flatter and more coalesced volar pads, which in turn slightly improve frictional force for animals that are at a comparative disadvantage while climbing because of their larger mass. Despite the essentially isometric relationship found between digit length and body mass across prosimians, examination of the residual variation reveals that tarsiers and Daubentonia possess, relative to their body sizes, remarkably long fingers. Such marked departures between body size and finger length observed in these particular primates are closely linked with specialized modes of prey acquisition and manipulation involving the hands.     

Ontoaenetic correlates of diet in anthropoid primates

This study assesses ontogenetic correlates of diet in anthropoid primates. Associations between body weight growth, adult size, and diet are evaluated for a sample of 42 primate species, of which 8 are classifiable as “folivores.” The hypothesis that folivores show a pattern of growth that differs from “nonfolivores” is tested. Ontogenetic variation is summarized through use of parametric and nonparametric regression analysis. Several analytical techniques, including broad interspecific and detailed comparisons among species of similar adult size, are applied. This investigation indicates a clear association between body weight ontogeny and diet: folivorous species grow more rapidly over a shorter duration than comprably sized nonfolivorus species. A positive correlation between adult size and diet is not unambiguously established in this sample. A threshold (at around 1 kg) below which insectivory is very common may adequately characterize the association between adult size and diet in anthropoid primates. Above this threshold, adult size does not appear to covary predictably with diet. Evolutionary correlates of the ontogenetic pattern seen in folivores may include a variety of factors. The distinctive pattern of development in folivores may relate to the profile of ecological and social risks that these species face. Morphophysiological advantages to rapid growth may relate to a need for accelerated alimentary (dental and gut) development. The implications of ontogenetic variation in folivores are discussed. © 1994 Wiley-Liss, Inc.     

Mandibular form and function in North American and European Adapidae and Omomyidae

Previous experimental and comparative studies among a wide variety of primate and nonprimate mammals provide a unique source of information for investigating the functional and phylogenetic significance of variation in the masticatory apparatus of Eocene primates. To provide a quantitative study of mandibular form and function in Eocene primates, the scaling of jaw dimensions and the development of symphyseal fusion was considered in a broad sample of North American and European Adapidae and Omomyidae. Statistical analyses indicate a significant size-related pattern of symphyseal fusion across Eocene primates, with larger taxa often having a greater degree of fusion than smaller species; this trend is also evident at the family level. As adapids are mostly larger than omomyids and these taxa show allometry of symphyseal fusion, this may explain why no omomyids evince complete fusion. Controlling for jaw size, species with greater symphyseal fusion tend to have more robust jaws than those with a lesser amount of fusion. Upon further examination, a primary reason why adapids have more robust mandibles than omomyids is associated with the presence of taxa with fused symphyses, and thus more robust jaws, in the adapid sample, whereas no omomyids have fused symphyses. In addition, there is little indication of a dietary effect, as measured by molar shear-crest development, on symphyseal fusion. Moreover, as there is no correlation between molar shear-crest development and skull size, this also points to the absence of a size-related pattern of dietary preference underlying the allometry of symphyseal fusion. Based on the interspecific and ontogenetic allometry of symphyseal ossification in Eocene primates, jaw-scaling patterns are used to further examine the functional determinants of fusion in this group. This study indicates that greater dorsoventral shear during mastication is a more likely factor than lateral transverse bending (“wishboning”) in the evolution of symphyseal fusion among “late-fusing” mammals like adapids and omomyids. Given that wishboning is an important functional determinant of symphyseal form in recent anthropoids, apparently the evolutionary development of marked wishboning occurs only in taxa that shift the timing of fusion to a growth stage preceding the onset of weaning (before adult masticatory patterns are fully developed) and perhaps first ossified the symphysis to counter elevated dorsoventral shear stress. As early anthropoids probably consisted of members varying interspecifically and ontogenetically in the degree of ossification, it is especially informative to analyze the adaptive setting in which anthropoid symphyseal fusion evolved from a similar primitive “prosimian” perspective. © 1996 Wiley-Liss, Inc.     

Phyletic size change and brain/body allometry: A consideration based on the African pongids and other primates

A problematic aspect of brain/body allometry is the frequency of interspecific series which exhibit allometry coefficients of approximately 0.33. This coefficient is significantly lower than the 0.66 value which is usually taken to be the interspecific norm. A number of explanations have been forwarded to account for this finding. These include (1) intraspecificallometry explanations, (2) nonallometric explanations, and (3) Jerison’s “extraneurons” hypothesis, among others. The African apes, which exhibit a lowered interspecific allometry coefficient, are used here to consider previous explanations. These are found to be inadequate in a number of ways, and an alternative explanation is proposed. This explanation is based on patterns of brain and body size change during ontogeny and phytogeny. It is argued that the interspecific allometry coefficient in African apes parallels the intraspecific one because similar ontogenetic modifications of body growth separate large and small forms along each curve. In both cases, body size differences are produced primarily by growth in later postnatal periods, during which little brain growth occurs. Data on body growth, neonatal scaling, and various lifehistory traits support this explanation. This work extends previous warnings that sizecorrected estimates of relative brain size may not correspond very closely to our understanding of the behavioral capacities of certain species in lineages characterized by rapid change in body size.     

Mechanical significance of femoral head trabecular bone structure in Loris and Galago evaluated using micromechanical finite element models

Work on the interspecific and intraspecific variation of trabecular bone in the proximal femur of primates demonstrates important architectural variation between animals with different locomotor behaviors. This variation is thought to be related to the processes of bone adaptation whereby bone structure is optimized to the mechanical environment. Micromechanical finite element models were created for the proximal femur of the leaping Galago senegalensis and the climbing and quadrupedal Loris tardigradus by converting bone voxels from high-resolution X-ray computed tomography scans of the femoral head to eight-noded brick elements. The resulting models had approximately 1.8 million elements each. Loading conditions representing takeoff phase of a leap and more generalized load orientations were applied to the models, and the models were solved using the iterative "row-by-row" matrix-vector multiplication algorithm. The principal strain and Von Mises stress results for the leaping model were similar for both species at each load orientation. Similar hip joint reaction forces in the range of 4.9 x to 12 x body weight were calculated for both species under each loading condition, but the hip reaction values estimated for Loris were higher than predicted based on locomotor behavior. These results suggest that functional adaptation to hip joint loading may not fully explain the differences in femoral head trabecular bone structure in Galago and Loris. The finite element method represents a unique and useful tool for analyzing the functional adaptation of trabecular bone in a diversity of animals and for reconstructing locomotor behavior in extinct taxa.     

Postnatal allometry of the skeleton in Tupaia glis (Scandentia: Tupaiidae) and Galea musteloides (Rodentia: Caviidae)--a test of the three-segment limb hypothesis

During the evolution of therian mammals, the two-segmented, sprawled tetrapod limbs were transformed into three-segmented limbs in parasagittal zig-zag configuration (three-segment limb hypothesis). As a consequence, the functional correspondence of limb segments has changed (now: scapula to thigh, upper arm to shank, fore arm plus hand to foot). Therefore, the scapula was taken into account in the current study of the postnatal growth of the postcranial skeleton in two small mammalian species (Tupaia glis, Galea musteloides). Comparisons were made between the functionally equivalent elements and not in the traditional way between serially homologous segments. This study presents a test of the three-segment limb hypothesis which predicts a greater ontogenetic congruence in the functionally equivalent elements in fore and hind limbs than in the serially homologous elements. A growth sequence, with decreasing regression coefficients from proximal to distal, was observed in both species under study. This proximo-distal growth sequence is assumed to be ancestral in the ontogeny of eutherian mammals. Different reproductive modes have evolved within eutherian mammals. To test the influence of different life histories on ontogenetic scaling during postnatal growth, one species with altricial juveniles (Tupaia glis) assumed to be the ancestral mode of development for eutherians and one species with derived, precocial young (Galea musteloides) were selected. The growth series covered postnatal development from the first successive steps with a lifted belly to the adult locomotory pattern; thus, functionally equivalent developmental stages were compared. The higher number of allometrically positive or isometrically growing segments in the altricial mammalian species was interpreted as a remnant of the fast growth period in the nest without great locomotor demands, and the clearly negative allometry in nearly all segments in the precocial young was interpreted as a response to the demand on early locomotor activity. Different life histories seem to have a strong influence on postnatal ontogenetic scaling; the effects of the developmental differences are still observable when comparing adults of the two species.     

Comparative analysis of encephalization in mammals reveals relaxed constraints on anthropoid primate and cetacean brain scaling

There is a well-established allometric relationship between brain and body mass in mammals. Deviation of relatively increased brain size from this pattern appears to coincide with enhanced cognitive abilities. To examine whether there is a phylogenetic structure to such episodes of changes in encephalization across mammals, we used phylogenetic techniques to analyse brain mass, body mass and encephalization quotient (EQ) among 630 extant mammalian species. Among all mammals, anthropoid primates and odontocete cetaceans have significantly greater variance in EQ, suggesting that evolutionary constraints that result in a strict correlation between brain and body mass have independently become relaxed. Moreover, ancestral state reconstructions of absolute brain mass, body mass and EQ revealed patterns of increase and decrease in EQ within anthropoid primates and cetaceans. We propose both neutral drift and selective factors may have played a role in the evolution of brain-body allometry.     

Ontogenetic and interspecific organ weight allometry in Old World monkeys

The importance of allometry as an analytic tool is well recognized in the literature of primate morphology. However, a number of recent studies have illustrated how interpretive difficulties can arise when researchers confound different types of allometric data. Such confusion is due less to carelessness than to uncertainty about how different types of allometry are related. The present study examines the relationship between two types—ontogenetic and interspecific allometry–in the case of organ weight scaling in six species of Old World monkeys. Accepting the interpretation of interspecific allometry as a reflection of functional scaling constraints, the results of this analysis indicate how ontogenetic patterns have been modified in different-sized species to maintain compliance with these constraints. Specifically, for the heart and lungs it appears that vertical transpositions of individual species' ontogenies are dictated by isometric interspecific allometry, while in the case of the kidneys and liver, the relation of negative allometry across species entails alteration of the relative growth coefficients of the individual species. While these conclusions can at present only be applied to organ weight scaling, the approach of examining interspecific patterns in light of developmental differences between species should prove very helpful in our efforts to understand the phenomena of size and scaling.     

Curvature, length, and cross-sectional geometry of the femur and humerus in anthropoid primates

The aims of this study were to describe the curvature of anthropoid limb bones quantitatively, to determine how limb bone curvature scales with body mass, and to discuss how bone curvature influences static measures of bone strength. Femora and humeri in six anthropoid genera of Old World monkeys, New World monkeys, and gibbons were used. Bone length, curvature, and cross-sectional properties were incorporated into the analysis. These variables were obtained by a new method using three-dimensional morphological data reconstructed from consecutive CT images. This method revealed the patterns of curvature of anthropoid limb bones. Log-transformed scaling analyses of the characters revealed that bone length and especially bone curvature strongly reflected taxonomic/locomotor differences. As compared with Old World monkeys, New World monkeys and gibbons in particular have a proportionally long and less curved femur and humerus relative to body mass. It is also revealed that the section modulus relative to body mass varies less between taxonomic/locomotor groups in anthropoids. Calculation of theoretical bending strengths implied that Old World monkeys achieve near-constant bending strength in accordance with the tendency observed in general terrestrial mammals. Relatively shorter bone length and larger A-P curvature of Old World monkeys largely contribute to this uniformity. Bending strengths in New World monkeys and gibbons were, however, a little lower under lateral loading and extremely stronger and more variable under axial loading as compared with Old World monkeys, due to their relative elongated and weakly curved femora and humeri. These results suggest that arboreal locomotion, including quadrupedalism and suspension, requires functional demands quite dissimilar to those required in terrestrial quadrupedalism.