Ontogeny of limb proportions in late through final Jomon period foragers

This study reports on developmental patterning in the intralimb indices of Late/Final Jomon period (4000-2300 BP) people. Jomon foragers represent the descendants of migrants from Northeast Asia, who arrived in the Japanese Islands around 20,000 BP. Among adults, Jomon brachial indices are elevated and similar to warm adapted, low latitude people, while crural indices are intermediate and similar to people from moderate latitudes. Two hypotheses regarding the development of intralimb indices among Jomon period foragers are tested: (1) intralimb indices of Jomon people maintain predicted ecogeographic relationships over ontogeny; (2) greater evolvability will be observed in the brachial index, while greater developmental constraint will be observed in the crural index. Changes in intralimb proportions in a Jomon skeletal growth series are compared to those in two contrasting samples: Inuit from Point Hope (cold adapted) and Nubians from Kulubnarti (warm adapted). A quadratic equation best describes the ontogeny of brachial and crural indices, with high indices in infancy followed by a decline in childhood and an increase in adolescence. Despite these shifts, ecogeographically predicted differences and similarities in the indices are maintained between samples throughout ontogeny. In addition, radial relative to humeral length is significantly less correlated than tibial relative to femoral length. These results suggest genetic conservation of intralimb indices over the course of development. However, radial and humeral lengths are less correlated than tibial and femoral lengths among Jomon subadults and adults, potentially suggesting greater evolvability of the brachial index and more developmental constraint on the crural index.     

Variation in ancient Egyptian stature and body proportions

Stature and the pattern of body proportions were investigated in a series of six time-successive Egyptian populations in order to investigate the biological effects on human growth of the development and intensification of agriculture, and the formation of state-level social organization. Univariate analyses of variance were performed to assess differences between the sexes and among various time periods. Significant differences were found both in stature and in raw long bone length measurements between the early semipastoral population and the later intensive agricultural population. The size differences were greater in males than in females. This disparity is suggested to be due to greater male response to poor nutrition in the earlier populations, and with the increasing development of social hierarchy, males were being provisioned preferentially over females. Little change in body shape was found through time, suggesting that all body segments were varying in size in response to environmental and social conditions. The change found in body plan is suggested to be the result of the later groups having a more tropical (Nilotic) form than the preceding populations.     

Body proportions in ancient Andeans from high and low altitudes

Living human populations from high altitudes in the Andes exhibit relatively short limbs compared with neighboring groups from lower elevations as adaptations to cold climates characteristic of high-altitude environments. This study compares relative limb lengths and proportions in pre-Contact human skeletons from different altitudes to test whether ecogeographic variation also existed in Andean prehistory. Maximum lengths of the humerus, radius, femur, and tibia, and femoral head breadth are measured in sex-specific groups of adult human skeletons (N = 346) from the central (n = 80) and the south-central (n = 123) Andean coasts, the Atacama Desert at 2,500 m (n = 102), and the southern Peruvian highlands at 2,000-3,800 m (n = 41). To test whether limb lengths vary with altitude, comparisons are made of intralimb proportions, limb lengths against body mass estimates derived from published equations, limb lengths against the geometric mean of all measurements, and principal component analysis. Intralimb proportions do not statistically differ between coastal groups and those from the Atacama Desert, whereas intralimb proportions are significantly shorter in the Peruvian highland sample. Overall body size and limb lengths relative to body size vary along an altitudinal gradient, with larger individuals from coastal environments and smaller individuals with relatively longer limbs for their size from higher elevations. Ecogeographic variation in relation to climate explains the variation in intralimb proportions, and dietary variation may explain the altitudinal cline in body size and limb lengths relative to body size. The potential effects of gene flow on variation in body proportions in Andean prehistory are also explored.     

A morphometric study of limb proportions in leaping prosimians

Study of individual limb proportions—most notably, for instance, of the intermembral index—distinguishes from most other primates those various prosimians capable of leaping. The present investigation analyzes 24 measures of limb proportions (taken by F.-K. Jouffroy and J. Lessertisseur) from 161 specimens representing 12 species of prosimians, using the multivariate statistical method of multiple discriminant analysis (carried out by R. German and C.E. Oxnard). The results indicate that there are three major gradients: (1) indriids, (2) cheirogaleines, and (3) galagines plus Tarsius, radiating from a fourth centrally located group of species, Lemur, Lepilemur, and Hapalemur. It is likely that the morphometric relationships displayed here can be associated with behavioral (locomotor) parallels among these various forms. Though leaping is a most important adaptation among prosimians, it seems clear that it has evolved separately a number of times, and it seems at least likely that the biomechanical nature of the adaptation differs in different groups. The results, therefore, present certain new hypotheses about prosimian locomotion that can only be tested with further field and laboratory studies. This study also emphasizes the fact that anatomical adaptation is complex. Though the analysis of each anatomical region separately provides a similar qualitative picture of the variation among genera, it is only in the analysis of all measures taken together that the complete pattern described above appears. Though no new variables have been added in that analysis, new information is obviously contained within the results. Since this information stems from the interactions of the variables, it must be due to associations between these anatomical areas.     

Variation in limb proportions between Jomon foragers and Yayoi agriculturalists from prehistoric Japan

Variation in limb proportions between prehistoric Jomon and Yayoi people of Japan are explored by this study. Jomon people were the descendents of Pleistocene nomads who migrated to the Japanese Islands around 30,000 yBP. Phenotypic and genotypic evidence indicates that Yayoi people were recent migrants to Japan from continental Northeast Asia who likely interbred with Jomon foragers. Limb proportions of Jomon and Yayoi people were compared using RMA regression and "Quick-Test" calculations to investigate relative variability between these two groups. Cluster and principal components analyses were performed on size-standardized limb lengths and used to compare Jomon and Yayoi people with other groups from various climatic zones. Elongated distal relative to proximal limb lengths were observed among Jomon compared to Yayoi people. Jomon limb proportions were similar to human groups from temperate/tropical climates at lower latitudes, while Yayoi limb proportions more closely resemble groups from colder climates at higher latitudes. Limb proportional similarities with groups from warmer environments among Jomon foragers likely reflect morphological changes following Pleistocene colonization of the Japanese Islands. Cold-derived limb proportions among the Yayoi people likely indicate retention of these traits following comparatively recent migrations to the Japanese Islands. Changes in limb proportions experienced by Jomon foragers and retention of cold-derived limb proportions among Yayoi people conform to previous findings that report changes in these proportions following long-standing evolution in a specific environment.     

Relative variation in human proximal and distal limb segment lengths

The pattern of variation and covariation of proximal and distal limb segment lengths was examined within and between 20 geographically diverse skeletal samples of modern humans. Analyses of variance-covariance matrices (VCMs) of logarithmically transformed (ln) variates of humerus, radius, femur, and tibia length were performed to test the following hypotheses: first, within populations, the distal and proximal segments will have equal relative (i.e., size-independent) variability. However, between populations, the tibia is predicted to be more variable than the other segments. Tests of fit of computed VCMs to theoretical matrices by an iterative procedure (Anderson [1973] Ann. Stat. 1:135-141) reject the equal variance hypotheses, rather suggesting that the relative variances of the distal limb segments are greater than are those of the proximal. Males and females differ somewhat in that within females, the distal segments of both limbs have equal variance, while within males, the tibia has greater relative variance than the radius. The second hypothesis, regarding between-group variability, is somewhat supported in that between human populations, one cannot reject that the tibia has greater relative variance than the other limb segments. However, neither can one reject an alternative hypothesis that both distal limb segments (tibia and radius) are more variable than the proximal segments. Differential growth allometry is explored, and likely plays a major role in differences seen both within and between human populations.     

Limb bone proportions and body mass of the cave bear (Ursus spelaeus)

The European cave bear evolved during the Middle Pleistocene and adapted to mountain environments. Earlier workers have described the cave bear as a robust bear. In this study the cave bears limb bone morphology is compared to the limb bone morphology of extant bears. Body mass estimates for the cave bear are made both based on different limb bone characters and based on dental and cranial characters. The shafts are wider in the cave bear limb bones than in the extant bear limb bones, and consequently the shaft widths give higher weight estimates to the cave bear than the other dimensions. The widened shafts are suggested to be a special adaptation (of presently unknown significance) rather than an indicator of an increased body mass.     

An exploration of adult body shape and limb proportions at Kellis 2, Dakhleh Oasis,Egypt

Several studies have shown that the human body generally conforms to the ecogeographical expectations of Bergmann's and Allen's rules; however, recent evidence suggests that these expectations may not hold completely for some populations. Egypt is located at the crossroads of Sub‐Saharan Africa, Southern Europe, and the Near East, and gene flow among groups in these regions may confound ecogeographical patterning. In this study, we test the fit of the adult physique of a large sample (N = 163) of females and males from the Kellis 2 cemetery (Dakhleh Oasis, Egypt) against ecogeographical predictions. Body shape (i.e., body mass relative to stature) was assessed by the femur head diameter to bicondylar femur length index (FHD/BFL), and brachial and crural indices were calculated to examine intralimb proportions. Body shape in the Kellis 2 sample is not significantly different from high‐latitude groups and a Lower Nubian sample, and intralimb proportions are not significantly different from mid‐latitude and other low‐latitude groups. This study demonstrates the potential uniqueness of body shape and intralimb proportions in an ancient Egyptian sample, and further highlights the complex relationship between ecogeographic patterning and adaptation. Am J Phys Anthropol 153:496–505, 2014. © 2013 Wiley Periodicals, Inc.     

Ecogeographic variation in human nasal passages

Theoretically, individuals whose ancestors evolved in cold and/or dry climates should have greater nasal mucosal surface area relative to air volume of the nasal passages than individuals whose ancestors evolved in warm, humid climates. A high surface-area-to-volume (SA/V) ratio allows relatively more air to come in contact with the mucosa and facilitates more efficient heat and moisture exchange during inspiration and expiration, which would be adaptive in a cold, dry environment. Conversely, a low SA/V ratio is not as efficient at recapturing heat and moisture during expiration and allows for better heat dissipation, which would be adaptive in a warm, humid environment. To test this hypothesis, cross-sectional measurements of the nasal passages that reflect surface area and volume were collected from a sample of CT scans of patients of European and African ancestry. Results indicate that individuals of European descent do have higher SA/V ratios than individuals of African descent, but only when decongested. Otherwise, the two groups show little difference. This pattern of variation may be due to selection for different SA/V configurations during times of physical exertion, which has been shown to elicit decongestion. Relationships between linear measurements of the skeletal nasal aperture and cavity and cross-sectional dimensions were also examined. Contrary to predictions, the nasal index, the ratio of nasal breadth to nasal height, is not strongly correlated with internal dimensions. However, differences between the nasal indices of the two groups are highly significant. These results may be indicative of different adaptive solutions to the same problem.     

Comparison of gut proportions in four small-bodied Amazonian cebids

Gut proportions in four small-bodied Amazonian cebids (Callicebus caligatus, Callicebus moloch, Pithecia irrorata, and Saimiri madeirae) are reported and compared in the context of known differences in feeding ecology. The gastrointestinal tracts of both Callicebus and Pithecia were found to be relatively undifferentiated, as expected from their predominantly frugivorous diets. In Saimiri, the marked dominance of the small intestine correlates with a highly insectivorous diet. © 1995 Wiley-Liss, Inc.     

Allometry and apparent paradoxes in human limb proportions: Implications for scaling factors

It has been consistently demonstrated that human proximal limb elements exhibit negative allometry, while distal elements scale with positive allometry. Such scaling implies that longer limbs will have higher intralimb indices, a phenomenon not borne out by empirical analyses. This, therefore, creates a paradox within the limb allometry literature. This study shows that these apparently conflicting results are the product of two separate phenomena. First, the use of the geometric mean of limb elements produces allometry coefficients that are not independent, and that when using ordinary least squares regression must yield an average slope of one. This phenomenon argues against using the geometric mean as a size variable when examining limb allometry. While the employment of relevant dimensions independent of those under analysis to calculate the geometric mean--as suggested by Coleman (Am J Phys Anthropol 135 (2008) 404-415)--may be a partial method for resolving the problem, an empirically determined, independent and biologically relevant size variable is advocated. If stature is used instead of the geometric mean as an independent size variable, all major limb elements scale with positive allometry. Second, while limb allometry coefficients do indicate differential allometry in limb elements, and thus should lead to some intralimb index allometry, this pattern appears to be attenuated by other sources of limb element length variation.     

Fore- and hindlimb proportions in Plio-Pleistocene hominids.

Associated fore- and hindlimb parts of five individuals are known from the hominid Plio-Pleistocene fossil collections in Africa. Four of these have been classified as Australopithecus and show definite evidence that in comparison with humans, forelimbs were relatively large and hindlimbs were relatively small. The fourth individual, placed in the genus Homo, has human proportions. These findings do not necessarily imply locomotor differences: the forelimbs may have been relatively long in Australopithecus simply because they were as yet not completely reduced from their generalized hominoid ancestral state.     

Reassessing manual proportions in Australopithecus afarensis

Previous analyses of hand morphology in Australopithecus afarensis have concluded that this taxon had modern human‐like manual proportions, with relatively long thumbs and short fingers. These conclusions are based on the A.L.333 composite fossil assemblage from Hadar, Ethiopia, and are premised on the ability to assign phalanges to a single individual, and to the correct side and digit. Neither assignment is secure, however, given the taphonomy and sample composition at A.L.333. We use a resampling approach that includes the entire assemblage of complete hand elements at Hadar, and takes into account uncertainties in identifying phalanges by individual, side and digit number. This approach provides the most conservative estimates of manual proportions in Au. afarensis. We resampled hand long bone lengths in Au. afarensis and extant hominoids, and obtained confidence limits for distributions of manual proportions in the latter. Results confirm that intrinsic manual proportions in Au. afarensis are dissimilar to Pan and Pongo. However, manual proportions in Au. afarensis often fall at the upper end of the distribution in Gorilla, and very lower end in Homo, corresponding to disproportionately short thumbs and long medial digits in Homo. This suggests that manual proportions in Au. afarensis, particularly metacarpal proportions, were not as derived towards Homo as previously described, but rather are intermediate between gorillas and humans. Functionally, these results suggest Au. afarensis could not produce precision grips with the same efficiency as modern humans, which may in part account for the absence of lithic technology in this fossil taxon. Am J Phys Anthropol 152:393–406, 2013. © 2013 Wiley Periodicals, Inc.     

Predynastic egyptian stature and physical proportions

An attempt has been made to estimate male and female Egyptian stature from long bone length usingTrotter &Gleser negro stature formulae, previous work by the authors having shown that these rather than white formulae give more consistent results with male dynastic material. Evidence is presented that the tibia length should include the spine in the later (1958) formulae and should exclude it in the earlier (1952) formulae. It is also shown that better results are obtained if the constants in the stature formulae are modified so as to conform more exactly with the basic data published byTrotter &Gleser. When consistency has been achieved in this way, predynastic, proportions are founded to be such that distal segments of the limbs are even longer in relation to the proximal segments than they are in modern negroes. Such proportions are termed ?super-negroid?.     

Developmental variation in ecogeographic body proportions

While ecogeographic variation in adult human body proportions has been extensively explored, relatively less attention has been paid to the effect of Bergmann's and Allen's rules on human body shape during growth. The relationship between climate and immature body form is particularly important, as immature mortality is high, mechanisms of thermoregulation differ between young and mature humans, and immature body proportions fluctuate due to basic parameters of growth. This study explores changes in immature ecogeographic body proportions via analyses of anthropometric data from children included in Eveleth and Tanner's (1976) Worldwide Variation in Human Growth, as well as limb proportion measurements in eight different skeletal samples. Moderate to strong correlations exist between climatic data and immature stature, weight, BMI, and bi-iliac breadth; these relationships are as strong, if not stronger, in immature individuals as they are in adults. Correlations between climate and trunk height relative to stature are weak or nonexistent. Altitude also has significant effects on immature body form, with children from higher altitudes displaying smaller statures and lower body weights. Brachial and crural indices remain constant over the course of growth and display consistent, moderate correlations with latitude across ontogeny that are just as high as those detected in adults. The results of this study suggest that while some features of immature body form, such as bi-iliac breadth and intralimb indices, are strongly dictated by ecogeographic principles, other characteristics of immature body proportions are influenced by intrinsic and extrinsic factors such as nutrition and basic constraints of growth.     

Development of patterns of activity in diaphragm of fetal lamb early in gestation

To examine the development of respiratory motor activity early in mammalian development and its relationship to nonrespiratory activity, we recorded spontaneous electromyogram activity from chronically instrumented fetal lambs over the period from 45 to 65 days' gestation (G45 to G65, term = G147). Two distinct forms of motor behavior were observed at G45 in recordings made from the costal diaphragm and longissimus dorsi muscles. The predominant behavior consisted of cycles of sustained, coincident activity in the two muscles alternating with periods of inactivity. The incidence of this type of activity declined between G45 and G65 and the cyclic nature of the discharges disappeared in most animals. The second form of motor behavior at G45 consisted of episodes of repetitive bursting activity lasting up to 20 min that were confined to the diaphragm. These bursts had a duration of 97.5 ± 8.3 ms (mean ± S.E.M.) and frequently occurred as doublets in which two bursts were separated by an intervening period of 100–200 ms. The mean duration of these bursts declined to 69.7 ± 7.7 ms at G65, doublets became rare, and bursts evolved a stereotyped form by G65 that was characterized by an abrupt onset and rapid decline in discharge intensity. Repetitive bursts of this form evolve into the mature respiratory motor pattern over the second half of gestation. At G45, episodes of repetitive bursting were almost always linked with episodes of sustained discharge, while at G65 these two forms of behavior were always segregated. We conclude that the neurons responsible for generating the respiratory rhythm in the lamb are assembled into a functional rhythm generator and make appropriate connections to motor output pathways as early as G45. The generation of the respiratory rhythm at G45 appears to be triggered by episodes of widespread motor activity that occur in both respiratory and nonrespiratory muscles. © 1996 John Wiley & Sons, Inc.